Female control of sperm ejection and retention in the cornsilk fly Euxesta eluta (Diptera: Ulidiidae).

Promiscuous mating systems provide the opportunity for females to bias fertilization toward particular males. However, distinguishing between male sperm competition and active female sperm choice is difficult for species with internal fertilization. Nevertheless, species that store and use sperm of different males in different storing structures and species where females are able to expel all or part of the ejaculates after copulation may be able to bias fertilization. We report a series of experiments aimed at providing evidence of female sperm choice in Euxesta eluta (Hendel), a species of ulidiid fly that expels and consumes ejaculates after copulation. We found no evidence of greater reproductive success for females mated singly, multiply with the same male, or mated multiply with different males. Female E. eluta possesses two spherical spermathecae and a bursa copulatrix for sperm storage, with a ventral receptacle. There was no significant difference in storing more sperm in spermathecae 24 h after copulation than immediately after copulation. Females mated with protein-fed males had greater reproductive success than similar females mated to protein-deprived males. Protein-fed females prevented to consume the ejaculate, retained more sperm when mated to protein-fed males than when mated to protein-deprived males. Our results suggest that female E. eluta can exert control of sperm retention of higher quality males through ejaculate ejection.

Maximum Entropy (MaxEnt) as extreme distribution indicator of two Neotropical fruit fly parasitoids in irrigated drylands of Argentina.

The figitid Ganaspis pelleranoi and the braconid Doryctobracon areolatus (Hym: Braconidae, Opiinae) are wide-ranging (from Florida, USA to Argentina) fruit fly parasitoids with tropical and subtropical distribution with a wet and temperate climate. In Argentina, both parasitoid species are thought to be restricted to the subtropical rainforests of the northwest and northeast, locally known as 'Yungas' and 'Paranaense' forests, respectively. However, these species recently have been recorded at the Monte and Thistle of the Prepuna eco-region, an arid region of central-western Argentina. Despite the extreme environmental conditions, anthropic artificial irrigation seems to be playing a fundamental role in fostering the presence and persistence of these species. Maximum Entropy (MaxEnt) models were developed to assess the suitability of these areas to harbor both species. The present work is a first approach to identify suitable areas for the distribution of these two fruit fly biological control agents in the American continent; based on 19 bioclimatic variables. Furthermore, the models resulting from including the new records in the 'Monte' eco-region suggest that local populations may become adapted to particular micro-environmental conditions generated by artificial irrigation. Models revealed that these artificial oases are suitable for G. pelleranoi but seem to be unsuitable for D. areolatus. This first and new approach to the area suitability of these species invites to produce models that reflect actual distribution including more records of presence in oases with similar conditions, thus decreasing the bias of the model generated by over reliance on areas with higher humidity (forest), which correspond to the distribution known before the inclusion of the new records.

Testing the potential contribution of Wolbachia to speciation when cytoplasmic incompatibility becomes associated with host-related reproductive isolation.

Endosymbiont-induced cytoplasmic incompatibility (CI) may play an important role in arthropod speciation. However, whether CI consistently becomes associated or coupled with other host-related forms of reproductive isolation (RI) to impede the transfer of endosymbionts between hybridizing populations and further the divergence process remains an open question. Here, we show that varying degrees of pre- and postmating RI exist among allopatric populations of two interbreeding cherry-infesting tephritid fruit flies (Rhagoletis cingulata and R. indifferens) across North America. These flies display allochronic and sexual isolation among populations, as well as unidirectional reductions in egg hatch in hybrid crosses involving southwestern USA males. All populations are infected by a Wolbachia strain, wCin2, whereas a second strain, wCin3, only co-infects flies from the southwest USA and Mexico. Strain wCin3 is associated with a unique mitochondrial DNA haplotype and unidirectional postmating RI, implicating the strain as the cause of CI. When coupled with nonendosymbiont RI barriers, we estimate the strength of CI associated with wCin3 would not prevent the strain from introgressing from infected southwestern to uninfected populations elsewhere in the USA if populations were to come into secondary contact and hybridize. In contrast, cytoplasmic-nuclear coupling may impede the transfer of wCin3 if Mexican and USA populations were to come into contact. We discuss our results in the context of the general paucity of examples demonstrating stable Wolbachia hybrid zones and whether the spread of Wolbachia among taxa can be constrained in natural hybrid zones long enough for the endosymbiont to participate in speciation.

Hybridization, Behavioral Patterns, and Pre- and Postzygotic Isolation Between Two Recently Derived Species of Walnut-Infesting Rhagoletis Fruit Flies in the Highlands of Mexico.

Episodes of isolation and secondary contact among populations of insects of Nearctic origin during Pleistocene glacial/postglacial climatic cycles had a strong evolutionary influence on the diversity of flies in the genus Rhagoletis in mountainous areas of Mexico. As a series of experiments undertaken to gather support for phylogenetic hypotheses on the origin of three walnut-infesting species in the suavis group, we examined pre- and postzygotic isolation between Rhagoletis completa Cresson, 1929 and R. ramosae Hernández-Ortiz, 1985. Despite morphological, biological, and behavioral differences, these two species were found to be capable of hybridization. Mating experiments in large enclosures revealed asymmetric sexual isolation. There were notable differences in male sexual behavior. While R. ramosae males mated exclusively on host fruit, R. completa males used fruit and alternative mating locations. During fruit-guarding and male-male contests, R. completa and R. ramosae males adopted markedly different wing postures. R. completa females were more reluctant to copulate with heterospecific males than R. ramosae females. During no choice crosses in small enclosures, there was a reduction of egg hatch for the hybrid cross of R. completa males × R. ramosae females. Our results and previous studies on reproductive isolation between other species pairs in the suavis group support a clade in which R. ramosae, R. zoqui Bush, 1966, and R. completa are close relatives all still capable of hybridizing.

Offseason Medfly Trapping Using Makeshift Fruit-Based and Wine Vinegar Baits.

Development of cost-effective traps and attractants is important for sustainable pest management. In the case of the Medfly, Ceratitis capitata Wiedemann, a worldwide pest of fruit production, recent interest in development of mass trapping strategies and low-cost easy-to-get artisanal traps for resource poor grower use has prompted renewed interest in research on attractants and traps. Additionally, such interest is also rooted on the need for effective female attractants to monitor wild population dynamics under male-only sterile fly releases. Response of wild flies and sterile males to makeshift traps (modified polyethylene terephthalate (pet) bottles) baited with fruit juice and wine vinegar dilutions with and without the addition of sugar revealed that a 50% dilution of wine vinegar or sweetened orange juice were more attractive than the standard Torula baits during the offseason (after harvest) in a peach orchard in arid irrigated valleys of San Juan Argentina. Such results suggest that volatiles from orange peel and juice are good candidates for development of Medfly attractants, and that pet bottles baited with sweetened orange juice and wine vinegar may be used by small growers and homeowners as low-cost-effective traps for Medfly control. Our results also suggest that Medfly response to food-based and fruit-based volatiles may be seasonally dynamic, a finding with important pest management implications.

Evidence for spatial clines and mixed geographic modes of speciation for North American cherry-infesting Rhagoletis (Diptera: Tephritidae) flies.

An important criterion for understanding speciation is the geographic context of population divergence. Three major modes of allopatric, parapatric, and sympatric speciation define the extent of spatial overlap and gene flow between diverging populations. However, mixed modes of speciation are also possible, whereby populations experience periods of allopatry, parapatry, and/or sympatry at different times as they diverge. Here, we report clinal patterns of variation for 21 nuclear-encoded microsatellites and a wing spot phenotype for cherry-infesting Rhagoletis (Diptera: Tephritidae) across North America consistent with these flies having initially diverged in parapatry followed by a period of allopatric differentiation in the early Holocene. However, mitochondrial DNA (mtDNA) displays a different pattern; cherry flies at the ends of the clines in the eastern USA and Pacific Northwest share identical haplotypes, while centrally located populations in the southwestern USA and Mexico possess a different haplotype. We hypothesize that the mitochondrial difference could be due to lineage sorting but more likely reflects a selective sweep of a favorable mtDNA variant or the spread of an endosymbiont. The estimated divergence time for mtDNA suggests possible past allopatry, secondary contact, and subsequent isolation between USA and Mexican fly populations initiated before the Wisconsin glaciation. Thus, the current genetics of cherry flies may involve different mixed modes of divergence occurring in different portions of the fly's range. We discuss the need for additional DNA sequencing and quantification of prezygotic and postzygotic reproductive isolation to verify the multiple mixed-mode hypothesis for cherry flies and draw parallels from other systems to assess the generality that speciation may commonly involve complex biogeographies of varying combinations of allopatric, parapatric, and sympatric divergence.

Promiscous female Euxesta eluta derive nutrients for ovarian development by expelling and consuming ejaculates.

Seminal gifts range from important material donations to items that provide little direct benefit to females. Promiscuous, female silk corn flies Euxesta eluta expel and consume male ejaculates immediately after mating. The evolution and function of this peculiar behavior are currently unknown. We performed a series of experiments aimed to: determine if females under different dietary regimes derive nutrients or water for survival and/or reproduction from ejaculate consumption, if males suffer a fitness cost from supplying females with ejaculates, and if females prefer to mate and/or are more likely to store sperm from well fed than nutritionally stressed presumably inferior males. Experiments revealed that protein deprived E. eluta females derive nutrients for ovarian development through consumption of ejaculates of protein fed males. No seminal products affecting survival appear to be transferred in the consumed ejaculate. However, ovarian development, in contrast to testes growth, occurs in detriment of longevity. Females preferred to mate with protein fed males, yet sperm retention in spermathecae was extremely rare after a single mating. This finding suggests that females could be exerting post copulatory control. A key question that remained to be addressed for the understanding of this puzzling and promiscuous mating system is what ecological factors or male traits drive females to retain sperm from one or several males in order to achieve and/or maximize fertilization potential.

Geographic and Ecological Dimensions of Host Plant-Associated Genetic Differentiation and Speciation in the Rhagoletis cingulata (Diptera: Tephritidae) Sibling Species Group.

Ascertaining the causes of adaptive radiation is central to understanding how new species arise and come to vary with their resources. The ecological theory posits adaptive radiation via divergent natural selection associated with novel resource use; an alternative suggests character displacement following speciation in allopatry and then secondary contact of reproductively isolated but ecologically similar species. Discriminating between hypotheses, therefore, requires the establishment of a key role for ecological diversification in initiating speciation versus a secondary role in facilitating co-existence. Here, we characterize patterns of genetic variation and postzygotic reproductive isolation for tephritid fruit flies in the Rhagoletis cingulata sibling species group to assess the significance of ecology, geography, and non-adaptive processes for their divergence. Our results support the ecological theory: no evidence for intrinsic postzygotic reproductive isolation was found between two populations of allopatric species, while nuclear-encoded microsatellites implied strong ecologically based reproductive isolation among sympatric species infesting different host plants. Analysis of mitochondrial DNA suggested, however, that cytoplasmic-related reproductive isolation may also exist between two geographically isolated populations within R cingulata. Thus, ecology associated with sympatric host shifts and cytoplasmic effects possibly associated with an endosymbiont may be the key initial drivers of the radiation of the R. cingulata group.

The Effect of Seasonal Humidity on Survival and Duration of Dormancy on Diverging Mexican Rhagoletis pomonella (Diptera: Tephritidae) Populations Inhabiting Different Environments.

Phytophagous insects synchronize emergence with plant phenology by engaging in dormancy during periods of host scarcity and environmental stress. Regulation of dormancy is achieved through response to seasonal cues. While temperature and photoperiod are important cues in temperate latitudes, seasonal humidity, such as the onset of rains, can be a reliable cue to for synchronization of emergence and affects survival of overwintering insects. We compared response of Mexican Rhagoletis pomonella (Walsh) (Diptera: Tephritidae) populations inhabiting subtropical environments differing in humidity patterns, to seasonal humidity regimes. Both populations emerged as adults in high proportions and suffered lower mortality under humidity regimes ending with a humid summer, but the effect was more pronounced for the Eje Volcanico Trans Mexicano (EVTM) population, which inhabits a dryer environment and undergoes longer dormancy. While there were no differences among pupae from the Sierra Madre Oriental (SMO) in percent of non-emerged pupae surviving and engaging in prolonged dormancy after a year, EVTM pupae exposed to an initial humid period engaged in prolonged dormancy in higher proportions than those exposed to other regimes. Seasonal humidity had little effect on the duration of dormancy, but EVTM pupae exposed to consecutive dry periods took longer to emerge than those exposed to other regimes. Our results suggest that rather than being used as a token stimulus, humidity affected survival of overwintering R. pomonella, especially at the end of dormancy when energy reserves are depleted and there is an increase in metabolic rate that renders EVTM pupae more susceptible to desiccation.

The Effect of Winter Length on Duration of Dormancy and Survival of Rhagoletis completa (Diptera: Tephritidae) and Associated Parasitoids From Northeastern Mexico.

The walnut husk fly Rhagoletis completa (Cresson), native to the Midwestern United States and Mexico, is invasive in California and Europe. It is one of the most important pests of walnuts in areas gathering 30% of the world production. Knowledge of life-history regulation is important for the design of management strategies. Research on dormancy has been performed on invasive populations, and not on populations at the southern extreme of its native range. Here, we examined the effect of winter length on fly and parasitoid emergence, survival, and duration of dormancy. Percent emergence was higher for chill periods at 5°C ranging from 8 to 20 wk. No or insufficient chill resulted in low emergence and a significant proportion of individuals in prolonged dormancy (>1 yr). Duration of dormancy was longer for pupae at constant temperatures and a 4-wk chill period than longer winter durations. Dormancy was longer for Mexican than that reported for U.S. populations, suggesting the existence of a latitudinal cline where populations at southern latitudes have evolved slower metabolic rates. Three parasitoid species were found associated with R. completa (Aganaspis alujai (Wharton and Ovruski) (Hymenoptera: Figitidae), Diachasmimorpha juglandis Muesebeck, and Diachasmimorpha mellea Gahan) (Hymenoptera: Braconidae). Results suggest that rearing of R. completa is possible by subjecting pupae to chill periods between 8 and 20 wk. Overwintering mortality of flies and A. alujai could be further reduced above 5°C. Our findings can contribute for the accurate development of predictive models on invasion potential, development, fly and parasitoid rearing, and biological control.

Experimental hybridization and reproductive isolation between two sympatric species of tephritid fruit flies in the Anastrepha fraterculus species group.

Among tephritid fruit flies, hybridization has been found to produce local adaptation and speciation, and in the case of pest species, induce behavioral and ecological alterations that can adversely impact efficient pest management. The fraterculus species group within Anastrepha (Diptera: Tephritidae), is a rapidly radiating aggregate, which includes cryptic species complexes, numerous sister species, and several pest species. Molecular studies have highlighted the possibility of introgression between A. fraterculus and A. obliqua. Reproductive isolation has been studied among morphotypes of the A. fraterculus species complex as a tool for species delimitation. Here we examined the existence and strength of prezygotic and postzygotic isolation between sympatric populations of two closely related species within the highly derived fraterculus group (A. fraterculus and A. obliqua), coexisting in nature. Although adults of both species showed a strong tendency for assortative mating, a small proportion of hybrid pairings in both directions were observed. We also observed asymmetric postzygotic isolation, with one hybrid cross displaying a strong reduction in fecundity and F1 egg fertility. Survival was greater for the progeny of homotypic and hybrid crosses in the maternal host. There was a marked female biased sex ratio distortion for both F1 hybrid adults. Hybridization between A. fraterculus and A. obliqua in nature may be difficult but possible; these two species display stronger reproductive isolation than all pairs of species previously examined in the A. fraterculus species complex. Asymmetric postzygotic isolation is suggestive of Wolbachia mediated cytoplasmic incompatibilities that may be exploited in area-wide pest management.

Temporal Diversity and Abundance Patterns of Parasitoids of Fruit-Infesting Tephritidae (Diptera) in the Argentinean Yungas: Implications for Biological Control.

A 4-yr study was done to analyze seasonal patterns underlying host plant-fruit fly-parasitoid interactions in a secondary forest in the Argentinean Yunga and its importance for the implementation of conservation and augmentative biological control. Larval-pupal hymenopteran parasitoids associated with all host plants and fruit fly species were identified and the seasonal occurrence of fruit, infestation levels, parasitism percentage, and relative parasitoid abundance were determined. Three fruit fly species in two genera were found in association with surveyed plants, two of which (Ceratitis capitata (Wiedemann) and Anastrepha fraterculus (Wiedemann)) are of major economic importance. Infestation levels were strongly influenced by environmental factors and peak fruit availability. Five fruit fly parasitoid species were recovered from fly pupae, four braconid species, and one figitid. Time windows for fruit fly population growth were pinpointed. Based on results, the present analysis proposes an effective fruit fly biological control strategy tailored for the northwestern Argentinean citrus-producing area.

Sexual Competitiveness of Anastrepha ludens (Diptera: Tephritidae) Males Exposed to Citrus aurantium and Citrus paradisi Essential Oils.

Males of the Mediterranean fruit fly (Ceratitis capitata (Wiedemann)) display increased mating competitiveness following exposure to the odor of certain host and nonhost plants, and this phenomenon has been used in the sterile insect technique to boost the mating success of released, sterile males. Here, we aimed to establish whether males of the Mexican fruit fly (Anastrepha ludens (Loew)) gain a mating advantage when exposed to the aroma of two preferred hosts, grapefruit (Citrus paradisi Macfadyen) and bitter orange (Citrus aurantium L.). Under seminatural conditions, we observed that, in trials using wildish males (from a young laboratory colony started with wild flies) exclusively, exposure to the aroma of bitter orange had no effect on male mating success but exposure to the odor grapefruit oil increased male mating success significantly. In a separate test involving both exposed and nonexposed wildish and mass-reared, sterile males, although wildish males were clearly more competitive than sterile males, exposure to grapefruit oil had no detectable effect on either male type. Exposure to oils had no effect on copulation duration in any of the experiments. We discuss the possibility that the positive effect of grapefruit essential oils on wildish male competitiveness may have been linked to exposure of females to grapefruit as a larval food, which may have imprinted them with grapefruit odors during pupal eclosion and biased their response as adults to odors of their maternal host.

A new species of Anastrepha (Diptera: Tephritidae) from Euphorbia tehuacana (Euphorbiaceae) in Mexico.

Anastrepha tehuacana, a new species of Tephritidae (Diptera) from Tehuacán, Puebla, Mexico reared from seeds of Euphorbia tehuacana (Brandegee) V.W. Steinm. (Euphorbiaceae), is described and illustrated. Its probable relationship to A. relicta Hernández-Ortiz is discussed.

Status of Rhagoletis (Diptera: Tephritidae) pests in the NAPPO countries.

The North American Plant Protection Organization (NAPPO) is an organization comprising plant protection regulatory officials of the three signatory countries: the United States, Canada, and México. NAPPO develops Regional Standards for Phytosanitary Measures (RSPMs) as well as discussion papers on important issues related to plant protection. The Fruit Panel of NAPPO organized a Technical Assistance Group (TAG) to develop a discussion paper on the status of pest species of Rhagoletis (Diptera: Tephritidae) within the NAPPO countries. As regulations are developed to prevent the spread of these pests both within and outside of the NAPPO countries, it was important to understand the pest status of these species. This article reviews the basic biology, host range, distribution, potential to spread, management, and regulatory status of pest species of Rhagoletis within the NAPPO countries.

Effect of host plant chemistry on genetic differentiation and reduction of gene flow among Anastrepha fraterculus (Diptera: Tephritidae) populations exploiting sympatric, synchronic hosts.

Herbivore host specialization includes changes in behavior, driven by locally induced adaptations to specific plants. These adaptations often result in sexual isolation that can be gauged through detection of reduced gene flow between host associated populations. Hypothetically, reduced gene flow can be mediated both by differential response to specific plant kairomones and by the influence of larval diet on some adult traits such as pheromone composition. These hypotheses could serve as a model to explain rapid radiation of phytophagous tephritid fruit flies, a group that includes several complexes of cryptic species. The South American Fruit Fly Anastrepha fraterculus (Wiedemann) is a complex of at least seven cryptic species among which pheromone mediated sexual isolation resulted in rapid differentiation. Cryptic species also exhibit differences in host affiliation. In search of a model explaining rapid radiation in this group, we studied host plant chemical composition and genetic structure of three host associated sympatric populations of A. fraterculus. Chemical composition among host plant fruit varied widely both for nutrient and potentially toxic secondary metabolite content. Adaptation to plant chemistry appears to have produced population differentiation. We found host mediated differentiation to be stronger between populations exploiting sympatric synchronic hosts differing in chemical composition, than between populations that exploit hosts that fruit in succession. Gene flow among such host associated populations was extremely low. We propose as a working hypothesis for future research, that for those differences to persist over time, isolating mechanisms such as male produced sex pheromones and female preferences resulting from adaptation to different larval diets should evolve.

Developing diagnostic SNP panels for the identification of true fruit flies (Diptera: Tephritidae) within the limits of COI-based species delimitation.

BACKGROUND: Rapid and reliable identification of quarantine pests is essential for plant inspection services to prevent introduction of invasive species. For insects, this may be a serious problem when dealing with morphologically similar cryptic species complexes and early developmental stages that lack distinctive characters useful for taxonomic identification. DNA based barcoding could solve many of these problems. The standard barcode fragment, an approx. 650 base pairs long sequence of the 5'end of the mitochondrial cytochrome oxidase I (COI), enables differentiation of a very wide range of arthropods. However, problems remain in some taxa, such as Tephritidae, where recent genetic differentiation among some of the described species hinders accurate molecular discrimination. RESULTS: In order to explore the full species discrimination potential of COI, we sequenced the barcoding region of the COI gene of a range of economically important Tephritid species and complemented these data with all GenBank and BOLD entries for the systematic group available as of January 2012. We explored the limits of species delimitation of this barcode fragment among 193 putative Tephritid species and established operational taxonomic units (OTUs), between which discrimination is reliably possible. Furthermore, to enable future development of rapid diagnostic assays based on this sequence information, we characterized all single nucleotide polymorphisms (SNPs) and established "near-minimal" sets of SNPs that differentiate among all included OTUs with at least three and four SNPs, respectively. CONCLUSIONS: We found that although several species cannot be differentiated based on the genetic diversity observed in COI and hence form composite OTUs, 85% of all OTUs correspond to described species. Because our SNP panels are developed based on all currently available sequence information and rely on a minimal pairwise difference of three SNPs, they are highly reliable and hence represent an important resource for developing taxon-specific diagnostic assays. For selected cases, possible explanations that may cause composite OTUs are discussed.

Mass rearing history and irradiation affect mating performance of the male fruit fly, Anastrepha obliqua.

As an initial step to improve the efficiency of the sterile insect technique applied to eradicate, suppress, and control wild Anastrepha obliqua (Macquart) (Diptera: Tephritidae) in mango producing areas of Mexico, the effect of radiation dose and mass rearing history on male mating performance was examined. Field cage tests in which both male and female laboratory flies were irradiated at different doses (0, 40, and 80 Gy) were released with cohorts of wild flies of both sexes, revealing that both mass rearing history and irradiation affected male mating performance. Laboratory males were accepted for copulation by wild females less frequently than wild males. Copulations involving laboratory males were shorter than those involving wild males. Irradiated males mated less frequently with wild females than wild males, and irradiated females appeared to be less able to reject courting males of both origins. High levels of fertility for untreated laboratory females crossed with males irradiated at different doses may reflect problems in mass rearing affecting homogeneity of pupal age before irradiation, and possibly masked a dose effect. Proposed remedial measures to improve male mating performance are discussed.

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.

Effect of larval host food substrate on egg load dynamics, egg size and adult female size in four species of braconid fruit fly (Diptera: Tephritidae) parasitoids.

Life history theory predicts that individuals will allocate resources to different traits so as to maximize overall fitness. Because conditions experienced during early development can have strong downstream effects on adult phenotype and fitness, we investigated how four species of synovigenic, larval-pupal parasitoids that vary sharply in their degree of specialization (niche breadth) and life history (Diachasmimorpha longicaudata, Doryctobracon crawfordi, Opius hirtus and Utetes anastrephae), allocate resources acquired during the larval stage towards adult reproduction. Parasitoid larvae developed in a single host species reared on four different substrates that differed in quality. We measured parasitoid egg load at the moment of emergence and at 24 h, egg numbers over time, egg size, and also adult size. We predicted that across species the most specialized would have a lower capacity to respond to changes in host substrate quality than wasps with a broad host range, and that within species, females that emerged from hosts that developed in better quality substrates would have the most resources to invest in reproduction. Consistent with our predictions, the more specialized parasitoids were less plastic in some responses to host diet than the more generalist. However, patterns of egg load and size were variable across species. In general, there was a remarkable degree of reproductive effort-allocation constancy within parasitoid species. This may reflect more "time-limited" rather than "egg-limited" foraging strategies where the most expensive component of reproductive success is to locate and handle patchily-distributed and fruit-sequestered hosts. If so, egg costs, independent of degree of specialization, are relatively trivial and sufficient resources are available in fly larvae stemming from all of the substrates tested.