Mineral nutrition of the hyperparasitic mistletoe Viscum articulatum Burm. f. (Viscaceae) in tropical Brunei Darussalam

A plant parasite parasitizing another plant parasite is known as a hyperparasite. Information is scarce regarding the ecophysiology of hyperparasites and their hosts despite their potential to illuminate processes of host–parasite solute flux. Here we present mineral profiles and stable isotopic data for two associations of the hyperparasite Viscum articulatum and its primary mistletoe and tree hosts. Acting as the terminal sink, the hyperparasite had consistently higher contents of all major and minor elements evaluated compared to the primary parasite and the proximal portion of the tree host branch. The primary parasite had lower contents of Cu, Mg, Mn, N, and Z relative to the proximal portion of the tree host branch, suggesting nutritional stress applied by the hyperparasite. Interestingly Fe and Cu showed no consistent pattern between host and primary parasite, while the osmotically active elements P and K increased from tree host, to primary mistletoe, and finally the hyperparasitic mistletoe. The δ¹³C partitioning patterns for hyperparasites, primary parasites, and hosts were non‐linear in contrast to linear patterns reported from the literature for autoparasitic mistletoe associations, demonstrating fundamental differences between nutrition in hyperparasites and autoparasites.     

A comparative study of the searching efficiencies of a parasite and a hyperparasite

The searching efficiencies of a primary parasite (Diaeretiella rapae (McIntosh )) and a hyperparasite (Alloxysta brassicae (Ash. )) were investigated and compared. In both species, at all parasite densities, there was a curvilinear relationship (P<0.001) between the number of hosts parasitised and the host density. A linear regression (log a=log Q−m log P) was fitted for log area of discovery against log parasite density (P<0.001). The area of discovery for its immediate (i.e. primary) host (viz. Diaeretiella for the hyperparasite and aphid for Diaeretiella) is lower in the hyperparasite than in the primary parasite. In Diaeretialla both the searching efficiency and the mutual interference constant increased (but not significantly, P>0.05) in the presence of its males.     

The parasite complex ofSaissetia nigra in Malaysia

The percentage parasitism ofSaissetia nigra (Nietner) by chalcidoid parasites and the relative abundance of individual parasites were studied in the field usingHibiscus rosa-sinensis L. as the host plant. In order of abundance (on the basis of percentage recorded) the primary parasites wereAnysis saissetiae Ash.,Aneristus ceroplastae How., andMicroterys newcombi (Gir)., while the hyperparasites wereMarietta exitiosa Comp.,Cheiloneurus saissetiae Noyes & Chua andEupelmus catoxanthae Ferr. The efficiency ofA. saissetiae (the larvae of which feed on the scale eggs) to control the scale population is doubtful because it is often parasitised by all 3 hyperparasites mentioned and each larva during development causes only 58% egg mortality of the parasitised host, leaving many scale eggs unconsumed.A. ceroplastae appears to be more useful parasite because it attacks the 2nd instar scales which cause significant damage to the host plant and it is free from attack by hyperparasites.     

Flexible patch time allocation by the leafminer parasitoid, Opius dimidiatus

Abstract.

• 1 The ability to use flexible decision rules can be an advantage to parasitoid females searching for patchily-distributed hosts. In a series of laboratory experiments the hypothesis that Opius dimidiatus, a solitary parasitoid of the chrysanthemum leafminer (Liriomyza trifolii), adjusts the time she allocates to searching for her larval hosts in response to both patch qualities and experiences with hosts was tested by varying such patch parameters as area, presence of host mines and density of host mines, and by allowing ovipositions and encounters with parasitized hosts.
• 2 Though leaf area was not a factor, the presence of host mines in a leaf did increase the time a female O.dimidiatus spent searching, over time spent on unmined leaves.
• 3 When host mine density was increased, females responded by increasing their search period in a density-dependent manner, suggesting a perception of patch quality.
• 4 Ovipositions in hosts caused females to reset their‘giving-up time’(GUT), or increase search intensity, by adding an amount of search time that increased with each successive oviposition. Conversely, encounters with parasitized (unsuitable) hosts incremented the GUT, but by an amount that decreased with each successive encounter.

     

Brood sex ratios of the solitary parasitoid wasp, Coccophagus atratus

Abstract.

• 1 Female eggs of Coccophagus atratus are deposited within the haemolymph of coccoid scale insects. Male eggs are deposited on to late larval and prepupal stages of parasitoids of scale insects, including conspecifics.
• 2 When presented with either one host type or a combination of both host types, female C.atratus deposit all their available eggs, assigning the appropriate sex egg to each host encountered. Brood sizes are not adjusted for different combinations of hosts.
• 3 Behavioural observations show that females do not move away from patches of hosts until all their eggs are laid, regardless of the host type.
• 4 Brood sex ratios varied with changes in the relative availability of hosts for males and hosts for females. When both host-types were present in equal numbers, male biased sex ratios resulted (mean ±SEM =0.71 ± 0.009) and when 70% of hosts provided were suitable for female eggs, mostly female-biased sex ratios resulted (mean ± SEM = 0.37±0.01).
• 5 Our results do not fit predictions based on the assumption that a sex ratio of 0.5 should be expected in C.atratus. Observed sex ratios indicate that the unusual life histories of these parasitoids need to be taken into account in explanations of their sex ratios.

     

Spread of an introduced parasite across the Hawaiian archipelago independent of its introduced host

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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.

     

Ecological studies on the parasite complex associated with typhlocybine leafhoppers (Homoptera, Cicadellidae)

Abstract.

• 1 The parasite complex associated with nymphal and adult typhlocybine leafhoppers consists of the following: Dryinidae (Aphelopus species), Pipunculidae (Chalarus species) and Diapriidae (Ismarus dorsiger Curtis), the last being parasitic on species of Aphelopus.
• 2 The life cycles and temporal distribution patterns of twenty-nine species of Typhlocybinae are summarized together with the life cycles of British Aphelopus and Chalarus.
• 3 Closely related primary parasite species differ in their adult emergence times. Species differences in adult emergence times of diapausing Chalarus are correlated with species differences in host relations.
• 4 Each primary parasite genus contains both monophagous and polyphagous species. Each polyphagous species shows a distinct ‘preference’ for a particular range of hosts.
• 5 The parasite complexes of different leafhopper communities are compared, and it is concluded that it is the taxonomic composition of the leafhopper communities which is chiefly responsible for the structure and taxonomic composition of their associated parasite complexes.
• 6 Individual species of both Aphelopus and Chalarus show a degree of sensitivity to the physiology of their hosts, so achieving a high degree of synchrony.

     

Sex ratios in field populations of Epidinocarsis lopezi, an exotic parasitoid of the cassava mealybug, in Africa

Abstract.

• 1 In cassava fields in Africa, population sex ratios of Epidinocarsis fopezi vaned from 0.44 (males to total parasitoids) at low host densities to highly male-biased ratios of 0.70 at high host densities.
• 2 This variability is caused by the difference in allocation of sons and daughters to hosts of different sizes, through the following mechanisms: (a) small, i.e. second instar, hosts are mainly used for the production of male offspring, whereas in large, i.e. third instar, hosts a variable, female-biased sex ratio is produced; (b) E.fopezi does not selectively oviposit into large hosts but always accepts both small and large hosts for oviposition upon encountering; (c) in the field, this parasitoid is time-limited, and not egg-limited. On the basis of an optimal diet model, such general host acceptance is shown to be the best strategy.
• 3 Thus, sex ratio increases with host density for three reasons: the proportion of small hosts encountered in the field increases with increasing host density, small hosts are used for male production, and hosts are always accepted when encountered.

     

Superparasitism and mutual interference in the egg parasitoid Anagrus delicatus (Hymenoptera: Mymaridae)

Abstract.

• 1 In nature, interference among Anagrus delicatus (Hymenoptera: Mymaridae) parasitoids reduced the per-capita number of hosts parasitized. Interference increased with parasitoid density.
• 2 Anagrus delicatus did not avoid parasitizing hosts that had recently been parasitized by conspecific wasps. Evidence indicated that this superparasitism was largely a random process, increasing with the ratio of parasitized to unparasitized hosts.
• 3 Individual parasitoid efficiency, the number of hosts killed per wasp per unit time, decreased with increasing wasp density. This occurred whether wasps searched the patch together (simultaneously) or one by one (sequentially), and was the result of an increase in time spent superparasitizing hosts at higher wasp density. This is known as indirect mutual interference.
• 4 Increasing numbers of parasitoids together on the same patch caused a significant decline in the rate and per-capita number of hosts parasitized. However, there was not a correspondent decline in searching efficiency with increasing wasp density (i.e. no direct mutual interference).
• 5 These forms of parasitoid density dependence should contribute to the stability of the host—parasitoid interaction.

     

Evolutionary history predicts high‐impact invasions by herbivorous insects

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Temperature and host diet jointly influence the outcome of infection in a Daphnia-fungal parasite system

1. Climate change has the potential to shape the future of infectious diseases, both directly and indirectly. In aquatic systems, for example, elevated temperatures can modulate the infectivity of waterborne parasites and affect the immune response of zooplanktonic hosts. Moreover, lake warming causes shifts in the communities of primary producers towards cyanobacterial dominance, thus lowering the quality of zooplankton diet. This may further affect host fitness, resulting in suboptimal resources available for parasite growth.
2. Previous experimental studies have demonstrated the respective effects of temperature and host diet on infection outcomes, using the zooplankter Daphnia and its microparasites as model systems. Although cyanobacteria blooms and heat waves are concurrent events in nature, few attempts have been made to combine both stressors in experimental settings.
3. Here, we raised the zooplankter Daphnia (two genotypes) under a full factorial design with varying levels of temperature (the standard 19°C and elevated 23°C), food quality (Scenedesmus obliquus as high-quality green algae, Microcystis aeruginosa and Planktothrix agardhii as low-quality cyanobacteria) and exposed them to the parasitic yeast Metschnikowia bicuspidata. We recorded life history parameters of the host as well as parasite traits related to transmission.
4. The combination of low-quality cyanobacterial diets and elevated temperature resulted in additive detrimental effects on host fecundity. Low-quality diets reduced parasite output, while temperature effects were context dependent. Overall, we argue that the combined effects of elevated water temperature and poor-quality diets may decrease epidemics of a common fungal parasite under a climate change scenario.

     

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.

     

A link between host dispersal and parasite diversity in two sympatric cichlids of Lake Tanganyika

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

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Overdispersion of Allothrombium pulvinum larvae (Acari: Trombidiidae) parasitic on Aphis gossypii (Homoptera: Aphididae) in cotton fields

Abstract.

• 1 Dispersion patterns of the protelean parasite, Allothrombium pulvinum Ewing, among individuals of an aphid host, Aphis gossypii Glover, were examined during spring 1991 in several cotton fields in Jiangsu Province, China.
• 2 The variance-to-mean ratios (i.e. dispersion index) of larval mites per host were greater than 1, indicating that the mite parasites were overdispersed among aphid hosts. The variance increased with the mean according to the power law, variance = 1.51 mean¹⁰⁶, which explained 99.7% of the variation in the data.
• 3 The negative binomial distribution adequately describes the patterns of larval mite dispersion among aphid hosts in eight out of ten populations. The degree of clumping (1/k) decreased curvilinearly with parasite density (mites per host).
• 4 Mites were more clumped among adult aphids than among immature ones.
• 5 Ecological and evolutionary consequences of mite overdispersion within host populations are discussed. The role of Allothrombium in pest control is also discussed.

     

Functional response to host density in a parasitic wasp,with reference to population regulation

1. The functional response to, and preference for, the host density in a parasite were examined experimentally using an icheumon wasp, Exidechthis canescens, and its host Cadra cautella under controlled conditions.
2. Wasps were more active in host-searching at higher than lower host densities. Percent parasitism increased rapidly with initial increments in host density and then tended to increase more slowly at higher host densities. A sigmoid functional response curve is indicated, which implies that the parasite is able to control its host even at low densities.
3. Wasps actively selected areas of high host density in which to concentrate host-searching behavior.
4. Host-searching by E. canescens is stimulated by the odor of the host when present, and by food in which hosts have developed but have been removed.
5. Both the functional response and the host-density preference of the parasite are mediated by its host-searching behavior. This relationship is discussed in the context of population regulation.

     

Host location by the fruit fly parasitoid Diachasmimorpha krausii: role of fruit fly species,life stage and host plant

• 1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts.
• 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively.
• 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato.
• 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae.
• 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present.
• 6 We conclude that herbivore‐induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly‐identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

     

Context dependency of infectious disease: the cyanobacterium Microcystis aeruginosa decreases white bacterial disease in Daphnia magna

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Parasite infection alters host stable‐isotope composition under controlled feeding

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