Phylogeny of the parasitic wasp subfamily Euphorinae (Braconidae) and evolution of its host preferences

The braconid subfamily Euphorinae is a large, cosmopolitan group of endoparasitoid wasps. The majority of species attack adult hosts, a strategy that is rare among parasitic wasps, but there are also many species that attack nymphs and larval stages. Euphorine hosts may belong to a variety of insect orders (Coleoptera, Hemiptera, Hymenoptera, Neuroptera, Psocoptera, Orthoptera and Lepidoptera) although most euphorine tribes are confined to Coleoptera. Here we investigate the phylogenetic relationships of the Euphorinae based on molecular data (3 kb of nucleotide data from four markers: 18S, 28S, CAD and COI) and propose a higher‐level classification based upon the resulting phylogeny. We also infer the evolution of host associations and discuss the diversification of the Euphorinae. Results from both Bayesian inference and maximum‐likelihood analysis show that the subfamily, as previously circumscribed, is paraphyletic. We propose that the subfamily be expanded to include the tribes Meteorini and Planitorini (Mannokeraia + Planitorus), so that it corresponds to a clade that is strongly supported as monophyletic in our analyses. Based on our results, a revised higher classification of the Euphorinae is proposed, in which 52 extant genera and 14 tribes are recognized. We reinstate the genus Microctonus belonging to the tribe Perilitini, and synonymize Ussuraridelus with Holdawayella, Sinuatophorus with Eucosmophorus. Furthermore, we propose the following tribal rearrangements: Spathicopis and Stenothremma are transferred to Perilitini; Tuberidelus, Eucosmophorus and Plynops to Cosmophorini; Ecclitura to Dinocampini; Chrysopophthorus, Holdawayella and Wesmaelia to Helorimorphini; Proclithroporus and Heia to Townesilitini. The monotypic tribe Cryptoxilonini is synonymized with Cosmophorini. The genera Pygostolus and Litostolus are placed in a separate tribe, Pygostolini, previously recognized as a subtribe among the Centistini. Parsimony‐based ancestral state reconstructions suggest that the ancestor of Euphorinae was a parasitoid of lepidopteran larvae, and that a host shift to larval Coleoptera occurred only in one clade of the Meteorini, some members of which secondarily shifted back to larval lepidopteran hosts. In the remainder of the subfamily, there was an initial shift from larval to adult coleopterans, followed by subsequent shifts to adults or larvae of Hemiptera, Hymenoptera, Neuroptera, Orthoptera and Psocoptera.     

Recruitment of native parasitoids by an exotic leaf miner, Cameraria ohridella: host-parasitoid synchronization and influence of the environment

Abstract

• 1 The horse‐chestnut leaf miner, Cameraria ohridella, is a moth of unknown origin that has recently invaded Europe and severely defoliates the European horse‐chestnut, an important ornamental tree.
• 2 Several indigenous parasitoids have colonized this new host, but parasitism remains low. One of the hypotheses suggested to explain the low parasitism is that candidate parasitoids emerge too early in spring to attack the first host generation and, thus, need early‐occurring leaf miners as alternate hosts. This hypothesis was tested by observing the synchronization between the phenology of the moth and that of its main parasitoids, and by comparing parasitism rates and parasitoid richness in different environments with various levels of biological diversity.
• 3 In spring, the bulk of the parasitoids emerge at least 5 weeks before the occurrence of the first suitable larvae of C. ohridella whereas most parasitoid adults reared outdoors die within 5 weeks after emergence.
• 4 Parasitism rates and parasitoid richness do not increase with biological diversity, suggesting that most parasitoids attacking the first generation of C. ohridella do not come from alternate hosts. Parasitism does not increase later in the year in the subsequent generations, when host‐parasitoid synchronization becomes less critical.
• 5 We conclude that, although the spring emergence of parasitoids is not synchronized with the phenology of C. ohridella, the parasitoids attacking the first generation are probably old or late‐emerging adults of the overwintering generation. The lack of synchronization is probably not the only reason for the poor recruitment of native parasitoids by C. ohridella.

     

The effect of host developmental stage at parasitism on sex‐related size differentiation in a larval endoparasitoid

• 1 For their larval development, parasitoids depend on the quality and quantity of resources provided by a single host. Therefore, a close relationship is predicted between the size of the host at parasitism and the size of the emerging adult wasp. This relationship is less clear for koinobiont than for idiobiont parasitoids.
• 2 As size differentiation in host species exhibiting sexual size dimorphism (SSD) is likely to occur already during larval development, in koinobiont larval endoparasitoids the size of the emerging adult may also be constrained based on the sex of the host caterpillar.
• 3 Sex‐specific growth trajectories were compared in unparasitised Plutella xylostella caterpillars and in second and fourth instar hosts that were parasitised by the solitary larval koinobiont endoparasitoid Diadegma semiclausum. Both species exhibit SSD, where females are significantly larger than males.
• 4 Healthy female P. xylostella caterpillars developed significantly faster than their male conspecifics. Host regulation induced by D. semiclausum parasitism depended on the instar attacked. Parasitism in second‐instar caterpillars reduced growth compared to healthy unparasitised caterpillars, whereas parasitism in fourth‐instar caterpillars arrested development. The reduction in growth was most pronounced in hosts producing male D. semiclausum.
• 5 Parasitism itself had the largest impact on host growth. SSD in the parasitoid is mainly the result of differences in growth rate of the parasitoid–host complex producing male and female wasps and differences in exploitation of the host resources. Female wasps converted host biomass more efficiently into adult biomass than males.

     

Seasonal incidence of two co-occurring adult parasitoids of Acalymma vittatum in New York State: Centistes (Syrrhizus) diabroticae and Celatoria setosa

We report the incidence of striped cucumber beetle (Acalymma vittatum [Fabricius]) (Coleoptera: Chrysomelidae) parasitism in bi-weekly samples from diverse farm locations in central New York State over the course of two growing seasons. Two parasitoid species not previously verified in this geographic region were found: Celatoria setosa (Coquillett) (Diptera: Tachinidae) and Centistes (Syrrhizus) diabroticae (Gahan) (Hymenoptera: Braconidae). Parasitism by C. setosa, up to 43%, [mean 8.67%] was comparable to previously published rates. Contrary to the expectations of earlier authors, however, C. diabroticae was found to be ubiquitous and maximum rates of parasitism (~54%) were >threefold greater than the only other published report for this species [C. diabroticae mean 14.36%]. In laboratory assays, we investigated C. diabroticae larval development. We also tested the effects of wasp clone and host beetle sex on C. diabroticae larval development. This is the first documentation of life cycle information for this braconid.     

外寄生性花绒寄甲的寄生选择及其发育表现

"选择-表现"假说认为,成虫应该选择有利于子代发育的高品质寄主,但在寄主选择中,除了寄主品质外,其他因素也可能影响寄主选择决策。寄主选择研究通常以成虫为对象,而对那些初龄幼虫选择寄主的寄生性昆虫很少关注。以1龄幼虫积极搜寻寄主的寄生性花绒寄甲为模式生物,采用双选试验设计,观察了花绒寄甲初孵幼虫在不同体重青杨天牛幼虫之间、在已被寄生与健康的黄粉虫蛹之间的寄生选择性;然后采用回归设计,观察了花绒寄甲寄生若干不同体重的青杨天牛幼虫后的发育表现。研究结果表明,花绒寄甲1龄幼虫对体型较大的青杨天牛幼虫的选择偏好显著大于对体型较小的寄主幼虫的选择,选择大体型幼虫的比值比是选择小体型幼虫的4.55倍;对已被寄生的寄主黄粉虫蛹的选择偏好显著大于对健康寄主蛹的选择,选择已被寄生寄主的比值比是选择健康寄主的12.57倍。寄生青杨天牛幼虫的花绒寄甲幼虫发育历期平均为11.49 d、蛹历期为26.67 d、幼虫发育至成虫的羽化率50%,这些发育表现与寄生时青杨天牛幼虫的体重没有显著关系。但刚羽化寄甲成虫体重与寄生时寄主的体重存在显著的正直线关系:寄生时的寄主体重每增大0.01 g,羽化出的寄甲成虫体重增大近0.08%;方差分析寄甲成虫体重在不同寄主体重水平之间的差异表明,从体型较大寄主中羽化的寄甲成虫体重显著大于从体型较小寄主中羽化的成虫。研究结果说明,花绒寄甲初孵幼虫在寄主选择决策时,在寄主体型大小与被寄生状态之间可能采取折衷对策,而且对体型大小不同的寄主选择与子代发育适合度表现存在一致性,从而支持"选择-表现"假说。     

Cotesia plutellae parasitizing Plutella xylostella: Host-age dependent parasitism and its effect on host development and food consumption

The braconid Cotesia plutellae(Kurdjumov) (Hymenoptera: Braconidae) is amajor solitary, larval endoparasitoid of thediamondback moth, Plutella xylostella(L.) (Lepidoptera: Plutellidae). Parasitism oflarvae of different host instars and fourdevelopmental ages of the 4th instar ofthe pest was examined. The effects of hostinstar at initial parasitization on thedevelopment, survival, size and fecundity ofthe parasitoid were determined in thelaboratory at 25 °C. The effects ofparasitism on host development and foodconsumption were investigated at 28 °C.Cotesia plutellae could parasitize larvaeof all four instars of P. xylostella, butpreferred 2nd and 3rd instars. In achoice test, the relative parasitism indicesfor 2nd, 3rd and 4th instarswere 0.37, 0.39 and 0.24, respectively.Parasitism decreased sharply with increasinghost age in the 4th instar and approachedzero in host larvae that had gone beyond 37%of 4th stadium. The development time andthe final adult size of the parasitoid variedwith the host instar at initial parasitization.Parasitoids with initial parasitism in the4th instar hosts had the shortestdevelopment time, followed by those in the3rd instar, and then by those in the2nd instar. Parasitoids startingparasitism in 2nd instar hosts weresmaller in body size than those starting in the3rd or 4th instar. However, resultantfemales starting parasitism in 3rd instarhosts had the highest fecundity. Parasitizedlarvae exhibited longer development time andincreased food consumption compared withunparasitized ones. This study presents thefirst record that a solitary parasitoidregulates host behavior leading to an increasein food consumption by the host.     

Parasitism and olfactory responses of <Emphasis Type="Italic">Dastarcus helophoroides</Emphasis> (Coleoptera: Bothrideridae) to different Cerambycid hosts

Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae) is an important natural enemy of longhorned beetles (Coleoptera: Cerambycidae). It is distributed throughout most Provinces in China. We investigated whether there were differences among D. helophoroides populations collected from different hosts in different geographic locations. Results showed that different D. helophoroides populations displayed different olfactory responses to larval frass from different longhorned beetle species. All populations were significantly attracted to the frass of their original hosts. Parasitism rates of different populations also varied when supplied with host larvae of the same longhorned beetle species. These results indicate that the three D. helophoroides populations tested differed in host-related behaviors. Therefore, the population of D. helophoroides must be taken into consideration when implementing biological control programs for different species of longhorned beetle.     

Targeted sugar provision promotes parasitism of the cereal leaf beetle Oulema melanopus

• 1 Parasitoids may often lack access to sugar (e.g. floral nectar) in agricultural settings. Strategically timed spraying of host plants with sugar solution may provide one means of enhancing parasitism at the same time as minimizing nontarget effects (e.g. benefiting the pest itself).
• 2 Sucrose was sprayed in wheat fields of northern Utah (U.S.A.) to assess the effects on parasitism of the cereal leaf beetle Oulema melanopus by the larval parasitoid Tetrastichus julis.
• 3 Early‐season sugar provisioning, when larvae of the pest were first hatching and parasitoid adults were newly emerged, did not affect the numbers of cereal leaf beetle larvae that matured in treated plots but increased parasitism rates of beetle larvae by four‐fold in 2006 and by seven‐fold in 2007.
• 4 No net influx of adult parasitoids into plots was detected after the application of sugar. Locally‐emerging parasitoids may have spent less time searching for their own food needs versus hosts. A laboratory experiment also confirmed that access to sucrose significantly increased parasitoid longevity.
• 5 The field experimental results obtained demonstrate that applications of sugar, implemented to target a key time of the growing season when benefits are maximized for parasitoids and minimized for their hosts, can strongly promote parasitism of the cereal leaf beetle in wheat fields.

     

The biology of the phonotactic parasitoid, Homotrixa sp. (Diptera: Tachinidae), and its impact on the survival of male Sciarasaga quadrata (Orthoptera: Tettigoniidae) in the field

Abstract.

• 1 Unlike most parasitoids, tachinid flies of the tribe Ormiini use sound to locate their hosts. Although thought to exert selection pressure on their host's calling behaviour, little is known about the biology of ormiines. Accordingly, this study reports the biology and impact of the ormiine Hornotrixa sp. upon calling males of the univoltine bushcricket Sciarasaga quadrata Rentz in south-western Australia.
• 2 Populations of adult S.quadrata were monitored in the field over two successive calling seasons. Females, which do not call, were not parasitized by Hornotrixa sp., but the risk of parasitism for males increased as the 3-month calling season progressed. Parasitism did not commence until c. 2 weeks into the calling season, but by the end of the season up to 87% of surviving males were parasitized.
• 3 Parasitized males lived for 14 days and were found singing until their penultimate evening before death in the field. Unparasitized males lived on average 69 days and a maximum of 119 days.
• 4 Multiparasitism of hosts was common, with up to sixteen fly larvae found within parasitized males. The number of fly larvae within hosts significantly increased at the end of the season. However, successful emergence of fly larvae from hosts, as well as pupal size, significantly decreased as more than one fly larva developed within the host.
• 5 Hornotrixa sp. has a long pupal duration of 30–31 days at 20°C. As a consequence, only one complete fly generation, which overwinters in the pupal stage, is likely within each host generation.
• 6 No evidence for differential (size-bias) mortality by Hornotrixa sp. on male S.quadrata was found. The size of parasitized and unparasitized males collected in the field was not significantly different.
• 7 It is concluded that Hornotrixa sp. is a significant mortality factor acting on the survival of adult male S.quadrata.

     

Heterogeneity and density dependence in a field study of a tephritid-parasitoid interaction

Abstract.

• 1 The spatial distributions of two tephritid flies (Urophora stylata (Fabricius) and Terellia serratulae L.) attacking thistle flower heads and the levels of parasitism from their associated parasitoid guilds were studied over a 7-year period.
• 2 Using these data it is possible to seek both temporal, density dependent relationships between average levels of parasitism and host density per generation, and also any spatial patterns of parasitism contributing to stability that may be operating within the same field system.
• 3 Parasitism by the two most important generalist parasitoids of T.serratulae is a direct function of average T.serratulae density per year. There is little evidence of any stabilizing heterogeneity arising from the spatial distribution of parasitism within generations.
• 4 Temporal density dependence of Urophora stylata cannot be confirmed from the 7 years of study but there is evidence of spatial heterogeneity which may have an important effect on the dynamics of the host population.

     

Dynamics of parasitism in the organ-pipe wasp, Trypoxylon politurn: effects of spatial scale on parasitoid functional response

Abstract.

• 1 Life tables and rates of parasitism were tabulated from mud nests built by Trypoxylon politum (Hymenoptera: Sphecidae) at nine different nesting sites from Missouri and Mississippi.
• 2 Most developmental mortality occurred either during the first two instars of development, or during the inactive prepupal phase. The majority (76%) of deaths were caused by insect parasitoids and cleptoparasites. Levels of parasitism and survivorship varied among nesting sites, and among locations within the two sites surveyed at a fine spatial scale.
• 3 Total developmental mortality, K, was positively associated with the number of hosts (immature T.politum) per site. Within one of two sites sampled at a fine spatial scale, K was negatively associated with the local density of hosts. Levels of total parasitism were positively associated with host population size, and negatively associated with local host density within one of the two sites sampled at a fine spatial scale.
• 4 Levels of parasitism by Melittobia (Hymenoptera: Eulophidae) were positively associated with the number of hosts per site, but negatively associated with the local density of hosts within sites. Melittobia parasitism was also negatively associated with the local density of old nesting material within sites.
• 5 Parasitism by Melittobia was a function of both the numbers of nests per quadrat and the mean nest size per quadrat at one of the two sites surveyed at a fine scale. At the other site, parasitism by Melittobia was a function of mean nest size per quadrat.
• 6 The life cycle and nesting behaviour of T.politum, in relation to the regulation of its numbers, is discussed.

     

Spatial density‐dependent parasitism and specificity in the robber fly Mallophora ruficauda (Diptera: Asilidae)

The density‐dependence in parasitism by the robber fly Mallophora ruficauda (Diptera: Asilidae) on scarab beetle larvae (Coleoptera: Scarabaeidae) populations was studied in the present research. Mallophora ruficauda is a pestiferous species common in the open grasslands of the Pampas region of South America. Adults are predators of insects and larvae are solitary parasitoids of third instar larvae of several species of scarab beetle (Coleoptera: Scarabaeidae). In contrast with most studied host‐parasitoid interactions, host searching by M. ruficauda is carried out by both larvae and adults. Typically, robber fly females lay eggs on tall grasses from where larvae drop to the ground, and attack hosts which are buried in the soil. We carried out our study at two spatial scales close to 14 apiaries located in the provinces of Buenos Aires and Entre Ríos (Argentina). We found that parasitism is density‐independent at the larger spatial scale and inversely density‐dependent at the smaller one. We also found that M. ruficauda selects Cyclocephala signaticollis among several scarab beetle species. Specificity is observed both at large and small spatial scales. We discuss the implications of both host specificity and host searching behaviour on the observed parasitism patterns.     

A new biological method to control Anobium punctatum,by using the parasitoid wasp Spathius exarator

Biological pest control by means of beneficial organisms is for long part of agriculture and Integrated Pest Management (IPM). A new and efficient strategy to control the most common timber pest species in churches and museums, the furniture beetle Anobium punctatum (De Geer) (Coleoptera: Anobiidae), is based on the parasitoid wasp species Spathius exarator (Linnaeus) (Hymenoptera: Braconidae). Once this braconid wasp detects its host species beneath the surface, it pierces the wood with its ovipositor to lay one single egg onto the beetle larva. After hatching, the wasp larva feeds on the beetle larva thereby killing it. Afterwards, it pupates and emerges through a self-gnawed hole as an adult wasp. The tiny, 0.5-mm-wide exit hole can easily be distinguished from the 1- to 2-mm-wide exit hole of A. punctatum. Laboratory tests revealed that female wasps have an average life span of 85 days and produce a total of 24 offspring, when nutrition is provided. Between 2012 and 2019, braconid wasps were introduced into 54 different A. punctatum infested buildings. Treatment success was monitored by examining exit holes of new beetles and wasps thereby calculating corresponding parasitism rates. After the first year of treatment, parasitism rates were significantly higher with a mean value of 0.15 when compared to untreated objects with a natural parasitism and a mean value of 0.08 (n = 54). Following treatment of three objects over a period of eight years, parasitism rates continuously increased from 0.02 up to 0.31. In a church organ, which was treated during this period, the monitoring revealed a complete suppression of pest activity. These data prove that this biological method of pest control is an efficient, sustainable and non-toxic option to manage the common furniture beetle.     

Searching strategies and attack rates of parasitoids of the ash bark beetle (Leperisinus varius) and its relevance to biological control

Abstract.

• 1 Point estimates of parasitoid behaviour indicated that braconid parasitoids of the bark beetle, Leperisinus varius, spend 70% of their host searching time stationary on the bark surface, in sharp contrast to the chalcidoid parasitoids which spend 60–70% of their host searching time moving across the bark surface.
• 2 Aggressive encounters between searching parasitoids are common and the data suggest that they do not occur at random.
• 3 The chalcidoid parasitoids, Cheiropachus quadrum and Eurytoma morio, are more frequently involved in aggressive encounters than expected and can displace ovipositing individuals of other species.
• 4 Attack rates for the parasitoids were estimated to vary from two hosts per day for the braconid Coeloides filiformis to four hosts per day for E.morio, and net reproductive rates vaned from 6.3 for C.filiformis to 0.8 for Coeloides melanotus.
• 5 These data suggest that while all parasitoids of the scolytid can act as primary parasitoids, at least some of the chalcidoid parasitoids are facultative cleptoparasitoids and should be avoided in classical biological control introductions.

     

The spatial patterns of parasitism of eumenid wasps,Anterhynchium flavomarginatum andOrancistrocerus drewseni by the miltogrammine flyAmobia distorta

1. Spatial patterns of parasitism of eumenid wasps Anterhynchium flavomarginatum and Orancistrocerus drewseni by the miltogrammine fly Amobia distorta were studied in Kyoto, Japan during 1980–1984.
2. In generations of low (<5%) and medium (5–20%) parasitism, percent parasitism per shed (the habitat of the hosts) increased as a function of host density. Conversely, in generations of high (>20%) parasitism, percent parasitism was rather constant over different host densities.
3. The spatial distributions of adult miltogrammine flies among sheds were censused in generations of low and medium parasitism. The frequency of observations of adult miltogrammine flies was higher at sheds of higher host density (aggregative behavioral response), but on the other hand, the adult miltogrammine flies distributed in an underdispersed (or regular) manner in relation to other conspecifics.
4. The spatially density independent relationship between host density and percent parasitism in generations of high parasitism was explained in relation to parasitoid dispersal from patches of high parasitoid density.

     

Parasitization by Edovum puttleri (Hymenoptera: Eulophidae) in relation to host density in the field

Abstract.

• 1 The relationship between parasitization by Edovum puttleri Grissell and density of eggs of the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae), was studied on two spatial scales (eggs mass and 6 m² cage).
• 2 For both scales, rates of parasitism were generally inversely related to host density for periods ranging from 2 to 8 days after parasitoid release. Thereafter, parasitism became independent of host density.
• 3 The initial inverse-density relationship and subsequent shift to density independence may result from several factors: (1) ambient temperatures, (2) the parasitoid's limited egg production, (3) differential times of exposure of egg masses to parasitoids, and/or (4) the parasitoid's patterns of host feeding and oviposition.
• 4 Although overall levels of parasitism were relatively low, total mortality of L.decemlineata eggs (including nonviable and cannibalized eggs, and those killed by parasitoid feeding) in parasitized egg masses was consistently high (?70–90%).

     

Plant and host effects on the leafroller parasitoid Dolichogenidia tasmanica

Parasitism by the braconid wasp Dolichogenidia tasmanica of first instar larvae of the lightbrown apple moth Epiphyas postvittana, established on four different species of potted host plants, was assessed after 2 weeks of field exposure in an apple orchard. Parasitism varied significantly between larval host plants (apple 58%, broom 59%, clover 30%, poplar 19% ). Parasitism on potted apple seedlings of the co-evolutionary host, E. postvittana, was compared in a field trial with that of two native New Zealand leafroller species (to which D. tasmanica has had exposure for 5 decades only). Parasitism varied significantly with larval host (E. postvittana, 83%; Ctenopseustis herana, 58%;Planotortrix octo, 26% ). Larval collections were made from mature apple trees and identification of larvae was achieved by DNA analysis for the leafroller species using PCR-RFLP of ITS1 + ITS2, and for the parasitoid by specific PCR of partial 18S. Parasitism under natural field conditions on mature apple trees was not different between larval hosts (mean 32.5% ). In laboratory studies, more P. octo larvae departed in response to parasitoid probing behaviour than E. postvittana, which is likely to contribute to the difference in parasitism rates. This study conclusively shows that D. tasmanica parasitises native New Zealand leafrollers, despite their different evolutionary origins.     

Short Distance Cues Used by the Adult Parasitoid <Emphasis Type="Italic">Microctonus hyperodae</Emphasis> Loan (Hymenoptera: Braconidae,Euphorinae) for Host Selection of a Novel Host <Emphasis Type="Italic">Listronotus oregonensis</Emphasis> (Coleoptera: Curculionidae)

Insect parasitoids use host kairomone to detect their hosts. However, in parasitoid species that attack adult hosts, the mobility of adult insect may mean that the host can move away for kairomone sources. The effect of Listronotus oregonensis LeConte (Coleoptera: Curculionidae) adult sex, feces and movement on host selection behavior by Microctonus’’ hyperodae Loan (Hymenoptera: Braconidae; Euphorinae) females was evaluated in the laboratory. We hypothesized that, in addition of using host kairomones, parasitoids of adult stage should use host movement for host selection. The sex of L. oregonensis did not affect the host selection behavior of M. hyperodae. However, host feces decreased the number of weevil antennations done by M. hyperodae. Microctonus hyperodae stopped less frequently near immobile L. oregonensis than near walking ones and these latter were frequently pursued by M. hyperodae. Host movement was the stimulus that elicited oviposition by M. hyperodae. The adaptive implications of these results are discussed.     

Responses of a native beetle to novel exotic plant species with varying invasion history

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Inter‐specific competition and competition‐free space in the tephritid parasitoids Utetes anastrephae and Doryctobracon areolatus (Hymenoptera: Braconidae: Opiinae)

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