The effect of spatial scale on interactions between two weevils and their parasitoid

1. The effect of spatial scale on the interactions between three hymenopteran parasitoids and their weevil hosts was investigated. The parasitoid Mesopolobus incultus (Walker) parasitised Gymnetron pascuorum Gyll.; the parasitoids Entodon sparetus (Walker) and Bracon sp. parasitised Mecinus pyraster Herbst. Both of these weevils develop inside the seedhead of Plantago lanceolata L. but occupy different niches. Seedheads were sampled annually from 162 plants at each of two experimental sites consisting of a series of habitat patches of two distinct sizes. Data were analysed from three site‐years. 2. Parasitoid densities at each site‐year were closely related to the abundance of their respective weevil hosts. The overall proportion of hosts parasitised was more variable for M. incultus than for E. sparetus and Bracon sp. 3. Changes in spatial scale affected the variability of parasitoid densities. For M. incultus, there was generally a greater degree of additional heterogeneity for all increases of scale; for E. sparetus, this was true only at the largest scales; for Bracon sp., all components of variance were negative. 4. The rate of parasitism was related to host density in different ways at different spatial scales. Mesopolobus incultus exhibited inverse density dependence at the finest (seedhead) scale, direct density dependence at the intermediate (plant) scale, and density independence at the large (habitat area 729 m²) scale. Entodon sparetus showed no response to variation in host density at any spatial scale. Bracon sp. showed direct density dependence only at the intermediate and largest scales. 5. Parasitoids E. sparetus and Bracon sp. seemed able to detect more than one M. pyraster individual in seedheads with multiple host occupancy; a greater incidence of conspecific parasitoids than expected emerged from such seedheads.     

Biology of the solitary wasp Trypoxylon (Trypargilum) agamemnon Richards 1934 (Hymenoptera: Crabronidae) in trap‐nests

Buschini, M.L.T. and Fajardo, S. 2009. Biology of the solitary wasp Trypoxylon (Trypargilum) agamemnon Richards 1934 (Hymenoptera: Crabronidae) in trap‐nests. —Acta Zoologica (Stockholm) 91 : 426–432. Some Trypoxylon species build their nests in preexisting tubular cavities like hollow stems and beetle borings in branches. Study of the biology of these insects is relatively easy because the females of these wasps nest with enormous success in trap‐nests. The aim of this study was to investigate the abundance, seasonality and life‐history of Trypoxylon agamemnon. For capture of these insects, trap‐nests were installed in the Parque Municipal das Araucárias in araucaria forest, grassland and swamp, from December, 2001 to December, 2005. Two hundred and ninety seven nests were obtained. They were constructed more often during the summer (from December to April). The nests were built only in araucaria forest and consisted of a linear series of cells, divided by mud partitions, whose number varied from 1 to 7. Normally they have only one vestibular cell. The inner cells had been provisioned, usually with spiders of Anyphaenidae family. Sex‐ratio was strongly female biased. Its main natural enemies included Chrysididae, Ichneumonidae and Tachinidae.     

Fitness effects of sex ratio response to host quality and size in the parasitoid wasp Spalangia cameroni

The parasitoid wasp Spalangia cameroni oviposited a greaterproportion of daughters in stable fly pupae than in house flypupae, even when I controlled for stable flies being smallerthan house flies. Sex ratio manipulation in response to hostquality has been modeled as being adaptive through an effectof host quality on the size and hence offspring production ofdaughters. 5. cameronis response to host species may insteadbe adaptive through an effect on larval survivorship, the developmenttime of daughters, and the size of sons. There was greater survivalof daughters than sons on stable flies. Controlling for hostsize, I found that development time of daughters was about 2%less on stable flies than on house flies. The decrease in developmenttime corresponds to a 2% increase in fitness as estimated byr, the intrinsic rate of increase, and is equivalent to abouta 9% increase in offspring production. Sons were about 2% largerfrom house flies than stable flies, which may increase offspringproduction by up to 3%. Host species had no consistent effecton size of daughters or development time of sons. In additionto the response to host species, mothers oviposited a greaterproportion of daughters in larger stable fly hosts. Whetherthis behavior is adaptive is unclear. Although offspring werelarger when they developed on larger stable flies, the rateof increase was less for daughters dian for sons. Effects ofstable fly size on offspring development time were negligible.     

Strongly female‐biased sex allocation in a trivoltine population of Trypoxylon (Trypargilum) opacum Brèthes (Hymenoptera,Crabronidae)

Buschini, M.L.T. and Bergamaschi, A.C.B. 2009. Strongly female‐biased sex allocation in a trivoltine population of Trypoxylon (Trypargilum) opacum Brèthes (Hymenoptera, Crabronidae). —Acta Zoologica (Stockholm) 91 : 433–439. This study was carried in southern Brazil from December 2001 to December 2004. The aim of this paper is to provide additional information on the life‐history and sex allocation of this little known species. This wasp’s species has two alternative life histories: either they pupated immediately and emerged as adults later in the same season (direct development) or they entered diapause, overwintering and pupating the following spring (delayed development). The numerical sex ratio of overwintering and of direct developing wasps were strongly female biased in 2002, 2003 and 2004.     

Effect of host availability on reproduction and survival of the parasitoid wasp Trichogramma minutum

Abstract.

• 1 We tested the hypothesis that females of the egg parasitoid, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), could adjust their fecundity schedule according to host availability and that there was a negative correlation between reproduction and survival in these wasps.
• 2 Newly-emerged females were provided with an unlimited or limited number of hosts in the first trial and with either unlimited, limited or zero hosts in the second trial.
• 3 When hosts were unlimited, wasps had the highest rate of reproduction in the first day, which decreased dramatically thereafter. When hosts were limited, wasps from the two trials differed in their response. In Trial I, females with limited hosts had lower first-day fecundity than, and the same subsequent-day fecundity as, those with unlimited hosts. However, in Trial II, females with limited host had a lower first-day but a higher subsequent-day fecundity than those with unlimited hosts. This indicates variation in Trichogramma's ability to shift its fecundity schedule in response to host availability.
• 4 There was a positive (rather than a negative) correlation between reproduction and survival. Wasps that oviposited (in host-unlimited treatment) had greater longevity than those that could not (in host-unavailable treatment).
• 5 The sex ratio of the progeny produced by wasps in both host-unlimited and limited treatments shifted gradually from a female to a male bias as the wasps aged.
• 6 We consider the ability of parasitoids to adjust their fecundity schedule as an adaptation to changing host resources and discuss our findings with regard to theories of life history evolution.

     

The dynamics of egg production, oviposition and resorption in a parasitoid wasp

1. The extent to which parasitoid wasps are limited by their egg supply is very important in understanding their reproductive strategies. Egg reserves are dynamic, with most wasps maturing new eggs throughout their life (synovigeny) and many species resorbing eggs that are not used in oviposition. We investigated the extent to which a parasitoid modulates its egg reserves in the light of its experience in finding hosts.
2. The egg dynamics of the Encyrtid Wasp Leptomastix dactylopii , a solitary parasitoid of mealybugs, were studied in the laboratory. This species is synovigenic and practises egg resorption.
3. We allowed newly emerged wasps to experience one of four environments of increasing value in terms of reproductive opportunities. We proposed that wasps that experienced good quality environments would maintain more mature eggs ready for oviposition. Dissection of wasps subject to different periods of host deprivation after the experimental treatment failed to confirm the hypothesis: egg load was independent of experience.
4. We also proposed that any adjustment of egg supply to make up for eggs oviposited would be effected through a reduction in egg resorption. Instead, we found that the wasp quickly made up for eggs oviposited by increased egg production.     

Egg limitation and the functional response of the parasitoid Campoletis grioti (Hym: Ichneumonidae)

The hypothesis that mature egg availability during a parasitoid's lifetime affects the functional response, as well as a parasitoid's handling time and searching efficiency, was tested. Campoletis grioti Blanchard (Hym: Ichneumonidae) female parasitoids and Spodoptera frugiperda Smith (Lep: Noctuidae) host larvae were used as an example. Each female of C. grioti was confined with different host densities (6, 10, 30, 50, 80, 120 and 160 larvae/day) of first and second S. frugiperda instars until the female died. We found that C. grioti showed a type II functional response, with a reduction in the number of parasitized hosts in relation to female ageing. Handling time and searching efficiency increased over time. When we standardized the data simulating unlimited egg supply, the functional response, handling time and searching efficiency became constant throughout the parasitoid's life. We discuss the relationship between egg availability and functional response and its associated parameters, and its consequence on biological control programs.     

Interpopulation variation of hibernal-aestival-diapause in the egg parasitoid wasp Anagrus takeyanus: adaptation to seasonal host-plant alternation of the tingid host, Stephanitis takeyai

The egg parasitoid wasp, Anagrus takeyanus Gordh & Dunbar (Hymenoptera: Mymaridae), is one of the few natural enemies of the andromeda lace bug, Stephanitis takeyai Drake & Maa (Heteroptera: Tingidae). Studying this wasp in two habitats, I found that at Kyoto, where the bug alternates its host plants seasonally, most wasp individuals in the overwintering eggs of the bug entered summer diapause immediately after winter diapause. However, at Nara, where the bug does not alternate its host plant, the wasp proved to have no summer diapause. Changes in the parasitization rate between generations differed at the two study sites. In Kyoto, the overwintered generation suffered the highest parasitism among the three generations of the year. In contrast, the first and second generations suffered higher parasitization rates in Nara. This difference in level of parasitization between the two study sites corresponded to the different diapause regime of the wasp and the seasonal population trends of the lace bug. Laboratory experiments showed that short photoperiod in combination with low temperature could terminate the long diapause at the time the non-aestivating individuals terminate winter diapause. As a life cycle without host plant alternation is likely to be an ancestral character of the bug, the aestivation of the wasp is thought to have been appended to overwintering as an adaptation to the evolution of seasonal host plant alternation in the lace bug.     

The titres of lectins in the haemolymph of Lacanobia oleracea and the effects of parasitism by the ectoparasitoid wasp Eulophus pennicornis

Serum from larvae of Lacanobia oleracea L. (Lepidoptera; Noctuidae) parasitized by Eulophus pennicornis (Hymenoptera; Eulophidae) and from normal non‐parasitized larvae is capable of agglutinating rabbit, sheep, calf, goat, chicken, horse and human erythrocytes, but not yeast. Studies with a range of inhibitory carbohydrates showed that serum lectins(s) had specificity for sugars containing galactose and for rhamnose, and for the glycosubstances fetuin and asialofetuin. Lectin activity is heat‐labile and is not dependent on calcium. Parasitism by E. pennicornis caused an increase in the agglutination titre of the serum from larvae of L. oleracea but not an increase in specific activity (titre per mg protein per ml). However, when venom from the venom gland of female wasps was injected into L. oleracea larvae, both the agglutinating activity and the specific activity of the larval serum increased. The possible causes of this increase are discussed. It is suggested that venom contains antigenic components which, when injected into the haemocoel of the L. oleracea larva, may be increasing lectin synthesis and/or release into the serum.     

The effect of superparasitism on development of the solitary parasitoid wasp, Venturia canescens (Hymenoptera: Ichneumonidae)

Abstract.

• 1 The effects of superparasitism on the rate of development, adult size and mortality of Venturia canescens (Hymenoptera: Ichneumonidae) were investigated. Parasitoids were reared from third (L3) and fifth (L5) instars of one of its hosts, Plodia interpunctella (Lepidoptera: Pyralidae) containing one, two or four parasitoid eggs.
• 2 Superparasitism increased the development time of Venturia reared from both instars, but the developmental delay was more evident in wasps from L5 hosts.
• 3 The size of parasitoids from L3 hosts was unaffected by egg number, but wasps from both superparasitized L5 treatments were significantly smaller than those from singly parasitized hosts.
• 4 Parasitoid mortality was significantly higher in L5 than L3, but within instars did not differ significantly with egg number.
• 5 The results confirm that superparasitism may affect the fitness of both the adult female wasp and her progeny, and should therefore be incorporated into models of superparasitism as an adaptive foraging strategy.

     

Variation in selected life-history parameters of the parasitoid wasp,Aphidius ervi: Influence of host developmental stage

We determined the age-specific fecundity and survival of the solitary parasstoid wasp,Aphidius ervi Haliday (Hymenoptera: Aphidiidae), under constant laboratory conditions. Wasps were reared in each of the four nymphal instars of apterous pea aphids,Acyrthosiphon pisum (Harris) (Homoptera: Aphididae): L₁ (age 24h), L₂ (48 h), L₃ (72 h), and L₄ (120 h). Age-specific survival (l_x) and fecundity (m_x) differed between parasitoids developing in different aphid instars. The wasps’ life-time reproductive success, as indexed by the intrinsic rate of population increase (r_m), varied non-linearly with adult biomass and host size at parasitization. A close agreement between larval growth rates in different host instars and adult reproductive performance suggests that, inA. ervi, fitness correlates may be significantly influenced by larval ontogeny and trade-offs in resource allocation.     

The effects of parasitism by an encyrtid parasitoid, Anagyrus pseudococci on the survival, reproduction and physiological changes of the mealybug, Planococcus citri

Survival, reproduction and some biochemical changes in the citrus mealybug, Planococcus citri (Risso) parasitized by the endoparasitoid, Anagyrus pseudococci (Girault) are reported. The parasitized younger mealybugs were more likely to die than be successfully parasitized. The mean duration (±SE) required for mortality other than mummification were 1.85 ± 0.06, 2.48 ± 0.14 and 2.89 ± 0.17 days for first, second and third instar P. citri respectively. Hosts parasitized in the first instar did not produce any mummies while the second and third instar nymphs and adult stages of the mealybugs were mummified on average 8 to 11 days after attack. Parasitization caused cessation of normal fecundity as well as induction of early egg maturation of the mealybugs. The mealybugs parasitized in the adult stage produced a few eggs but the fecundity of unparasitized mealybugs was about 40 fold higher than that of parasitized mature adults. Parasitized pre-ovipositing and mature adults laid eggs within 24 h of attack. SDS-PAGE analysis of the body content of the parasitized mealybugs after 24 h and 48 h of parasitization and before egg hatch revealed an increase in the concentration of some proteins. Response to wound reactions was not visible either as formation of extra protein bands or increased density of the existing bands. The possible effect of parasitism on the changes of the reproductive physiology of the host is discussed.     

Isolation and characterisation of 26 polymorphic microsatellite markers for the parasitoid wasp Trichogramma dendrolimi (Hymenoptera: Trichogrammatidae)

Polymorphic microsatellite markers were developed for the parasitoid wasp Trichogramma dendrolimi. Forty-four microsatellites were isolated from a (TCG)₈-enriched genomic library and characterised. Twenty-six of these loci were polymorphic and could be used for evaluation of the genetic variation of T. dendrolimi.     

Functional endogenous viral elements in the genome of the parasitoid wasp Cotesia congregata: insights into the evolutionary dynamics of bracoviruses

Bracoviruses represent the most complex endogenous viral elements (EVEs) described to date. Nudiviral genes have been hosted within parasitoid wasp genomes since approximately 100 Ma. They play a crucial role in the wasp life cycle as they produce bracovirus particles, which are injected into parasitized lepidopteran hosts during wasp oviposition. Bracovirus particles encapsidate multiple dsDNA circles encoding virulence genes. Their expression in parasitized caterpillars is essential for wasp parasitism success. Here, we report on the genomic organization of the proviral segments (i.e. master sequences used to produce the encapsidated dsDNA circles) present in the Cotesia congregata parasitoid wasp genome. The provirus is composed of a macrolocus, comprising two-thirds of the proviral segments and of seven dispersed loci, each containing one to three segments. Comparative genomic analyses with closely related species gave insights into the evolutionary dynamics of bracovirus genomes. Conserved synteny in the different wasp genomes showed the orthology of the proviral macrolocus across different species. The nudiviral gene odv-e66-like1 is conserved within the macrolocus, suggesting an ancient co-localization of the nudiviral genome and bracovirus proviral segments. By contrast, the evolution of proviral segments within the macrolocus has involved a series of lineage-specific duplications.     

Aggregation of parasitoids and density-independence of parasitism in field populations of the wasp Aphytis melinus and its host, the red scale Aonidiella aurantii

ABSTRACT.

• 1 Repeated counts were made of the number of adult Aphytis melinus (DeBach) wasps per fruit on Valencia oranges in an orchard over two successive periods. Resulting rates of parasitism per fruit were measured at the end of each period.
• 2 For both periods, corresponding to high and low mean numbers of adult parasites, there was a significant positive regresssion of adult wasps per fruit on the number of available hosts per fruit. However, there was a high level of variability about the regression, and the overall aggregative response appears to be weak.
• 3 For both periods, rates of parasitism per fruit were independent of host density per fruit and they showed a high level of variability at all densities. Similar patterns were found in another, commercial, orchard over a wide range of mean host densities.
• 4 There was no evidence for aggregation of parasites or density dependence of parasitism at a patch size corresponding to the whole tree.
• 5 Suggestions based on some host-parasitoid models, that aggregations of parasite attack in areas of high host density are necessary for effective biological control, are rejected. However, the model of Hassell (1982), incorporating aggregation of parasites and limitations on the effectiveness of the parasite, seems to fit the data quite well.

     

Bacterial infections associated with the son-killer trait in the parasitoid wasp Nasonia (= Mormoniella) vitripennis (Hymenoptera: Pteromalidae)

The son-killer (sk) trait in the parasitoid wasp, Nasonia vitripennis, causes the production of very female-biased sex ratios through the mortality of males. Some 80% of all male eggs fail to hatch. The trait is both maternally and contagiously transmitted. Here evidence is presented that the trait is associated with systemic and chronic bacterial infections in adult wasps. Infections develop trans-stadially, originating in the midgut of larvae and subsequently spreading to other tissues, including the brain, fat body, muscles, eyes, and hemocytes. Female reproductive tracts often show infections but infections are absent in male reproductive organs; other sex differences occur as well. The bacteria involved are pleomorphic, with straight rods being the most common form.     

Agistemus exsertus Gonzalez (Acari: Stigmaeidae) as a predator of two scale insects of the family Diaspididae (Homoptera: Diaspididae)

Abstract

The predacious mite Agistemus exsertus Gonzalez completed its life-span when fed on eggs and crawlers of the Florida red scale Chrysomphalus ficus Ashmead and the white date scale Parlatoria blanchardi (Targioni). The development was faster when individuals were maintained on eggs and crawlers of C. ficus, compared with both stages of P. blanchardi. The average number of eggs/female/day was 2.5 and 1.1 on eggs and crawlers of C. ficus, respectively. Eggs and crawlers of P. blanchardi were an unsuitable food for egg laying for A. exsertus. Life table parameters showed that A. exsertus preferred eggs of C. ficus to the crawlers as prey. The population of the predator feeding on eggs and crawlers of Florida red scale multiplied 45 and 7 times in a generation time of 23 and 20.5 days, respectively. Under these conditions, the intrinsic rate of increase (r _m) was (0.17 and 0.098) individuals/female/day on eggs and crawlers of C. ficus, while the finite rate of increase (λ) was (1.18 and 1.11) on both stages of C. ficus.     

The influence of spatial scale on the genetic structure of a widespread tropical wetland tree, Pterocarpus officinalis (Fabaceae)

Identifying factors that cause genetic differentiation in plant populations and the spatial scale at which genetic structuring can be detected will help to understand plant population dynamics and identify conservation units. In this study, we determined the genetic structure and diversity of Pterocarpus officinalis, a widespread tropical wetland tree, at three spatial scales: (1) drainage basin “watershed” (<10 km), (2) within Puerto Rico (<100 km), and (3) Caribbean-wide (>1000 km) using AFLP. At all three spatial scales, most of the genetic variation occurred within populations, but as the spatial scale increased from the watershed to the Caribbean region, there was an increase in the among population variation (Φ_ST=0.19 to Φ_ST=0.53). At the watershed scale, there was no significant differentiation (P=0.77) among populations in the different watersheds, although there was some evidence that montane and coastal populations differed (P<0.01). At the island scale, there was significant differentiation (P<0.001) among four populations in Puerto Rico. At the regional scale (>1000 km), we found significant differentiation (P<0.001) between island and continental populations in the Caribbean region, which we attributed to factors associated with the colonization history of P. officinalis in the Neotropics. Given that genetic structure can occur from local to regional spatial scales, it is critical that conservation recommendations be based on genetic information collected at the appropriate spatial scale.