Tactics of parasitic American coots: host choice and the pattern of egg dispersion among host nests

Summary I examined the tactics adopted by a conspecific brood parasite, the American coot (Fulica americana), and the degree to which these tactics reflect sources of mortality for parasitic eggs. Only 8% of parasitic eggs produced independent offspring, compared to a 35% success rate for non-parasitic eggs, and most mortality was due to egg-rejection by hosts or the consequences of laying eggs too late in the host's nesting cycle. Parasites usually laid parasitically before initiating their own nests and usually parasitized immediate neighbours. Parasites did not remove host eggs before laying their own egg, and egg disappearance in general was not more common at parasitized nests. I found no evidence for non-random host choice, either on the basis of stage of the host's nesting cycle or the host's brood size. The absence of adaptive host choice is likely a consequence of the fact that, due to host limitation, only a small proportion of parasites had meaningful variation among potential hosts to choose from. The pattern of egg dispersion among host nests by individual parasites appears to be a compromise between constraints imposed by host limitation and the increased success obtained from spreading eggs among nests. Most females laying fewer than five parasitic eggs laid them in a single host nest while females laying five or more eggs normally parasitized two or more hosts. An examination of egg rejection and survival rates showed that parasites would maximize success by laying a single egg per host nest, and the pattern of laying several eggs per host nest is likely a consequence of host limitation. However, no egg that was the fifth laid, or later, parasitic egg in a host nest was ever successful and this probably explains why most females laying five or more eggs parasitized more than one host.     

Complex host marking in the cabbage root fly

Summary. The cabbage root fly marks an oviposition site with two different chemical messages. Plants that were exposed to ovipositing flies are less acceptable than control plants, while sand particles that were closely associated with an actual oviposition site stimulate oviposition. By combining the information from these opposing messages, the cabbage root fly may be able to optimise the size of its egg clusters in relation to the food available for the larvae. The findings might account for the aggregated oviposition observed in this species.     

Relationships among host microbiota,parasite resistance or tolerance,and host fitness

Article impact statement: Parasite-induced shifts in host microbiota that lead to parasite resistance or tolerance may have unintended consequences for host fitness.     

植物内生菌与宿主关系研究进展

植物内生菌（endophyte）是指存活于健康植物组织内部,而又不引发宿主植物表现出明显感染症状的微生物类群,主要包括真菌、细菌和放线菌。其在植物体中的分布具有普遍性、多样性的特点。在目前研究过的所有植物中均发现有内生菌,它们可存在于植物的根、茎、叶、花、果实等各个部位。研究发现内生菌除了随植物遗传进行传播外,还可通过多种途径对宿主植物进行侵染,比如可以以内生菌体或孢子形式通过变形、吸器、或渗透等途径侵染植物,也可以通过分解植物表皮细胞壁或通过各种自然开口（包括侧根发生处,气孔,水孔等）或伤口（包括土壤对根的磨损,病虫对植物的损害及收割多年生植物造成的伤口等）等传播途经进入植物。内生菌由于与宿主植物长期共处,进而形成了一种复杂、特殊的关系。它们有的是互利共生关系,而有的是无害或微害寄生关系。两种关系可随多种因素变化而相互转化。本文主要从内生菌在宿主植物中的分布特点、侵染特性及与宿主植物共处方式等方面,对近十年来植物内生菌与宿主关系的研究进展进行综述和展望,以期为植物内生菌资源开发研究提供参考。     

The patchiness of epifoliar fungi in tropical forests: host range, host abundance, and environment

Fungal symbioses affect the diversity, dynamics, and spatial patterns of trees in tropical forests. Their ecological importance is partly driven by their inherent patchiness. We used epifoliar fungi, a guild of common, benign, obligate, fungal symbionts of plants, as a model system to evaluate the relative importance of host phylogeny, host relative abundance, and microclimate on the three-dimensional distribution of plant-fungus symbioses. In parallel studies in rainforests in Panama and Australia, most epifoliar fungi were able to colonize several plant lineages but showed significant host preferences within the local plant community. More closely related plant species were not more likely to share fungal symbionts. Instead, fungal species were more likely to be shared by more abundant hosts, which supported a greater number and diversity of fungi. Environmental conditions strongly affected spatial distributions, with sites in the dark understory 2.5- to fourfold more likely to have epifoliar fungi than in the exposed forest canopy. In the understory, fungal incidence increased with canopy openness. Canopy trees supported only a subset of the fungal symbionts found in the understory, suggesting that adult trees are not reservoirs of these fungal symbionts for understory juveniles.     

Does supernormal stimulus influence parental behaviour of the cuckoo's host?

The supernormal stimulus hypothesis (SSH) states that a cuckoo chick should obtain more parental care than host young by means of exaggerated sensory signals. We tested the SSH by comparing parental care by reed warblers at parasitized and non-parasitized nests. A comparison of feeding rates to parasite and host chicks of the same size showed that parasitized nests received more food than non-parasitized ones with one host chick. There was an interesting relationship between average prey length and the mass of a cuckoo chick: prey length first increased with chick mass, but decreased after the cuckoo chick outgrew the average-sized host brood (three to four young at fledging). This might be expected if fosterers reduced the selectivity of their foraging behaviour when trying to satisfy the supernormal food demands of the parasitic chick. This suggestion is supported by the finding that the relationship between nestling mass and proportion of less economical small prey is inverse to the relationship between nestling mass and prey size. These results suggest that the parental behaviour of reed warblers is adjusted by selection to the needs of an average-sized brood. The overall proportion of insect orders was significantly different between the parasitic and host chicks. This result probably reflects the opportunistic foraging habits of the host. The qualitative difference (proportion of insect orders) between host and cuckoo nestling diets is partly a by-product of unequal length distribution of members of different taxonomic groups. The results of this study are consistent with the SSH.     

Quantifying the burden of managing wildlife diseases in multiple host species

Mitigation of infectious wildlife diseases is especially challenging where pathogens affect communities of multiple host species. Although most ecological studies recognize the challenge posed by multiple-species pathogens, the implications for management are typically assessed only qualitatively. Translating the intuitive understanding that multiple host species are important into practice requires a quantitative assessment of whether and how secondary host species should also be targeted by management and the effort this will require. Using a multiple-species compartmental model, we determined analytically whether and how intensively secondary host species should be managed to prevent outbreaks in focal hosts based on the reproduction number of individual host species and between-species transmission rates. We applied the model to the invasive pathogenic fungus Batrachochytrium salamandrivorans in a 2-host system in northern Europe. Avoiding a disease outbreak in the focal host (fire salamanders [Salamandra salamandra]) was impossible unless management also heavily targeted the secondary host (alpine newts [Ichthyosaura alpestris]). Preventing an outbreak in the community required targeted removal of at least 80% of each species. This proportion increased to 90% in the presence of an environmental reservoir of B. salamandrivorans and when the proportion of individuals removed could not be adjusted for different host species (e.g., when using traps that are not species specific). We recommend the focus of disease-mitigation plans should shift from focal species to the community level and calculate explicitly the management efforts required on secondary host species to move beyond the simple intuitive understanding that multiple host species may all influence the system. Failure to do so may lead to underestimating the magnitude of the effort required and ultimately to suboptimal or futile management attempts.     

Endemic hantavirus infection impairs the winter survival of its rodent host

The influence of pathogens on host fitness is one of the key questions in infection ecology. Hantaviruses have coevolved with their hosts and are generally thought to have little or no effect on host survival or reproduction. We examined the effect of Puumala virus (PUUV) infection on the winter survival of bank voles (Myodes glareolus), the host of this virus. The data were collected by monitoring 22 islands over three consecutive winters (a total of 55 island populations) in an endemic area of central Finland. We show that PUUV infected bank voles had a significantly lower overwinter survival probability than antibody negative bank voles. Antibody negative female bank voles from low-density populations living on large islands had the highest survival. The results were similar at the population level as the spring population size and density were negatively correlated with PUUV prevalence in the autumn. Our results provide the first evidence for a significant effect of PUUV on host survival suggesting that hantaviruses, and endemic pathogens in general, deserve even more attention in studies of host population dynamics.     

Induction of host specificity in larvae of Manduca sexta: chemical dependence controlling host recognition and developmental rate

Summary. The tobacco hornworm, Manduca sexta L. (Lepidoptera: Sphingidae) is a specialist on Solanaceae. This host specificity is induced as the larva feeds on solanaceous foliage, so that solanaceous-reared larvae will refuse to feed on other plants. Experiments were designed to determine the role of dietary constituents on the induction of host specificity and the effects of these on development of M. sexta. Choice assays using leaf discs of cowpea, Vigna sinensis (Fabaceae), were used to monitor the isolation of relevant chemical cues from foliage of potato. An aqueous extract of potato foliage strongly stimulated feeding. This extract was partitioned with n-butanol under alkaline conditions to obtain a highly active butanol extract. Reversed phase flash chromatography with a water-methanol gradient gave an active fraction that was used as a supplement for wheat germ-based artificial diet. Larvae reared on this s-diet became dependent on potato allelochemicals for initiation and continuation of feeding activity. These larvae also developed faster than larvae reared on the control p-diet, but no effect on adult mass was detected. Further flash chromatography of the active fraction under alkaline conditions provided a highly active sub-fraction, and semi-preparative HPLC using gradients of water and acetonitrile resulted in the isolation of a single compound. Bioassays indicated that this compound alone can account for host recognition by solanaceous-reared larvae. The results suggest that the mechanism of induced host specificity in M. sexta involves development of dependence on this compound. Received 21 December 1999; accepted 14 March 2000     

Learning and natal host influence host preference,handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, na?ve female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness. Electronic Publication     

Transmission of symbiotic dinoflagellates through the sexual cycle of the host scyphozoan Linuche unguiculata

Intracellular symbiotic dinoflagellates are associated with the tropical scyphozoan Linuche unguiculata (Swartz, 1788) throughout all stages of the host's life cycle. During sexual reproduction, eggs are released in mucus strands that contain symbiotic dinoflagellates. Fertilization and development take place externally in the water column. Epifluorescence and transmission electron microscopy showed that unfertilized eggs did not contain intracellular algae, but that infection of the developing embryo was generally successful by the 128-cell stage (10 h after fertilization at 23° C). However, experiments with artificially provided Cellufluor-labeled algae demonstrated that older embryos and planulae could be infected by algae through at least 24 h post-fertilization, indicating that the L. unguiculata symbiosis represents a semi-closed system. This novel mode of symbiont acquisition results in most sexually-produced offspring becoming infected with maternally-transmitted algae during early development, but allows for acquisition of non-maternally-provided algae later in development. Most of the algal symbionts during the early stages of embryonic and larval development are located within ectodermal cells. This is in contrast to the other life-cycle stages of L. unguiculata (i.e., scyphistoma, medusa, ephyra), where symbionts are found within the gastrodermis of the host.     

Daphne gnidium, a possible native host plant of the European grapevine moth Lobesia botrana, stimulates its oviposition. Is a host shift relevant?

Summary. We tested the hypothesis that Daphne gnidium is an ancestral host plant of Lobesia botrana. To this end, we studied the oviposition response of this moth to various aerial organs of this plant. Our results show that females prefer to lay eggs on daphne berries rather than on grapes (cv. chasselas) and that polar surface extracts of daphne berries and leaves strongly stimulate oviposition in a dosedependant way, whereas flower extracts are weakly stimulant and branch extracts are inactive or deterrent for oviposition. Furthermore, a total extract of daphne berries stimulates oviposition in semi-natural conditions when applied onto fresh grapes. Oviposition stimulants from berries were isolated by HPLC coupled with a dual-choice oviposition bioassay. The ecological value of D. gnidium for L. botrana is discussed and the possible use of oviposition stimulants from daphne, contrasting with the oviposition signal of the cultivated host plant, is proposed in the perspective of developing behavioural manipulation methods such as stimulo-deterrent diversionary strategies compatible with IPM programs.     

Rational host country use of a severance tax

This paper addresses the basic question: How should a host country use a constant severance tax to control a foreign-owned mining firm? The basic rule is derived, and comparative static analysis is performed to show the effects of price, cost, and interest changes on the optimal tax policy.     

Bromeliad-living spiders improve host plant nutrition and growth

Although bromeliads are believed to obtain nutrients from debris deposited by animals in their rosettes, there is little evidence to support this assumption. Using stable isotope methods, we found that the Neotropical jumping spider Psecas chapoda (Salticidae), which lives strictly associated with the terrestrial bromeliad Bromelia balansae, contributed 18% of the total nitrogen of its host plant in a greenhouse experiment. In a one-year field experiment, plants with spiders produced leaves 15% longer than plants from which the spiders were excluded. This is the first study to show nutrient provisioning in a spider-plant system. Because several animal species live strictly associated with bromeliad rosettes, this type of facultative mutualism involving the Bromeliaceae may be more common than previously thought.     

Parasite transmission in a migratory multiple host system

A transmission model was devised for trichostrongyloid nematodes of saiga antelopes and domestic sheep in Kazakhstan. The framework extends previous models by including seasonal migration of saigas, contact with separate populations of sheep, and climate-driven stochasticity in herbage biomass and in the development, survival and migration onto herbage of free-living larvae. The model was parameterised for the contrasting life histories of Marshallagia, Haemonchus and Nematodirus, three important parasites of saigas and sheep in the region, and was successful at predicting broad qualitative patterns of infection dynamics in sheep and saigas. Parasite transmission between saigas and sheep was predicted to be most important for Marshallagia (from sheep to saigas in the south in winter, and onward transmission to sheep in the north in summer) and Haemonchus (from sheep in the north in summer via saigas to sheep further south in autumn). Model predictions for winter transmission of Marshallagia infection in saigas were consistent with field data, which showed that saigas culled before they have grazed the winter range carry lower burdens of this parasite than older animals. The model provides a mechanistic explanation for its predictions, which will assist hypothesis formation, and further the epidemiological basis of efforts to control parasite transmission between wildlife and livestock in both directions. A similar modelling approach could prove useful in other situations where detailed mechanistic models of parasite transmission are inappropriate in the face of parameter uncertainty and spatio-temporal variation in climate and host density. This is likely to include the majority of wildlife-parasite systems.     

Distribution of larval host plants of the fruit piercing moth,Othreis fullonia

Summary The adult fruit piercing moth,Othreis fullonia, a native of the indo-Malaysian region, causes severe damage to fruits grown throughout the tropical and subtropical belt from Africa through Asia and Australia to the Pacific Islands. Plants of the family Menispermaceae and the genusErythrina (Fabaceae) serve as larval hosts but the adult moths prefer Menispermaceae plants for oviposition. In Africa, Asia and Australia, the moth does not lay eggs onErythrina since members of the Menispermaceae are abundant. However in the insular Pacific region, where most islands have few or no species of Menispermaceae, the introduced fruit piercing moth utilizesErythrina as an alternate larval host, and either depletes, endangers or causes the possible extinction of Menispermaceae.     

Cowbird removals unexpectedly increase productivity of a brood parasite and the songbird host

Generalist brood parasites reduce productivity and population growth of avian hosts and have been implicated in population declines of several songbirds of conservation concern. To estimate the demographic effects of brood parasitism on Bell's Vireos (Vireo bellii), we removed Brown-headed Cowbirds (Molothrus ater) in a replicated switchback experimental design. Cowbird removals decreased parasitism frequency from 77% and 85% at unmanipulated plots to 58% and 47% at removal plots in 2004 and 2005, respectively. Vireo productivity per pair was higher at cowbird removal plots when years were pooled (mean = 2.6 +/- 0.2 [SE] young per pair) compared to unmanipulated plots (1.2 +/- 0.1). Nest desertion frequency was lower at cowbird removal plots (35% of parasitized nests) compared to unmanipulated plots (69%) because removal of host eggs was the proximate cue for nest desertion, and vireos experienced lower rates of egg loss at cowbird removal plots. Nest success was higher among unparasitized than parasitized nests, and parasitized nests at cowbird removal plots had a higher probability of success than parasitized nests at unmanipulated plots. Unexpectedly, cowbird productivity from vireo pairs was higher at cowbird removal plots (mean = 0.3 +/- 0.06 young per pair) than at unmanipulated plots (0.1 +/- 0.03) because fewer parasitized nests were deserted and the probability of nest success was higher. Our study provides the first evidence that increases in cowbird productivity may be an unintended consequence of cowbird control programs, especially during the initial years of trapping when parasitism may only be moderately reduced. Thus, understanding the demographic impacts of cowbird removals requires an informed understanding of the behavioral ecology of host-parasite interactions.     

Influence of host plants on sexual communication in the herbivorous bug Lygocoris pabulinus

Summary. Host plant volatiles may be involved in the sexual communication of insects in several ways. In the pheromone-producing sex, these volatiles may affect pheromone production or release and, in the receptive sex, plant volatiles may have a synergistic effect on the attraction to sex pheromone. We conducted three types of experiments to determine if and how plant volatiles are involved in the sexual communication of Lygocoris pabulinus (L.) (Heteroptera: Miridae), the females of which attract males. In a one-choice cylinder bioassay, females were offered to males on two different plants, belonging to different plant families, i.e. potato leaves and goosefoot leaves, to determine if specific plant volatiles were involved in the attraction of males towards females. Females on potato leaves were as attractive as females on goosefoot leaves, but significantly more attractive than females without plant material. The latter result suggests an interaction between females and potato leaves. However, in two-choice flying and walking bioassays, using delta traps in a wind tunnel and a vertical Y-track olfactometer, males were attracted to females irrespective of the presence of potato leaves. This difference in result is probably due to the fact that in the latter assays females were confined with pollen as an alternative food source, while females in the one-choice assay had access to water only, so that they may have suffered from malnutrition. Males in the one-choice assay were also attracted to potato leaves from which females had been removed, indicating that attractive components from females are deposited and adsorbed to the substrate. Plants are probably only indirectly involved in sexual communication, their surface merely functioning as a substrate from which pheromone is released. Males may subsequently be attracted to such plants or substrates. Clean plant material was not attractive to L. pabulinus males, hence plant volatiles alone do not seem to be used by these males as possible mate location cues.     

Offspring sex ratio response to host size in the parasitoid wasp Spalangia endius

The host size model, an adaptive model for maternal manipulation of offspring sex ratio, was examined for the parasitoid wasp Spalangia endius. In a Florida strain, as the model predicts, daughters emerged from larger hosts than sons, but only when mothers received both small and large hosts simultaneously. The pattern appeared to result from the mother's ovipositional choice and not from differential mortality of the sexes during development. If sex ratio manipulation is adaptive in the Florida strain, it appears to be through a benefit to daughters of developing on large hosts rather than through a benefit to sons of developing on small hosts. Both female and male parasitoids were larger when they developed on larger hosts. For females, developing on a larger host (1) increased offspring production, except for the largest hosts, (2) increased longevity, (3) lengthened development, and (4) had no effect on wing loading. For males, development on a larger host had no effect on any measure of male fitness – mating success, longevity, development duration, or wing loading. In contrast, a strain from India showed no difference in the size of hosts from which daughters versus sons emerged, although both female and male parasitoids were larger when they developed on larger hosts. These results together with previous studies of Spalangia reveal no consistent connection between host-size-dependent sex ratio and host-size-dependent parasitoid size among strains of S. endius or among species of Spalangia. Received: 28 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999