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The distribution of parasites within host populations and communities, and the mechanisms responsible for these patterns, are poorly understood aspects of wildlife parasitology. Here, we evaluate the influence of the average abundance of endoparasite variance, using endoparasites of lizards from the Caatinga domain (semiarid region), north-eastern Brazil. We hypothesized that, due to the high number of generalist endoparasite species, they may occur randomly throughout host populations in an aggregate pattern. In addition, we evaluated the degree to which sample variance is influenced by the average abundance of endoparasite species, patterns of co-occurrence and dominance among endoparasite species and similarities between abundance and the richness of endoparasite infracommunities in several host species. Between September 2015 and February 2016, 2141 lizards (1233 infected) from 16 species were collected from six Caatinga areas. In total, 25,687 endoparasites were collected, which belonged to 13 species including nematodes, pentastomids, cestodes, trematodes and acanthocephalans. Parasite–host associations documented here included 39 newly identified interactions. Endoparasites occurred in a typical aggregate pattern of distribution within their hosts; there was no measurable preference related to the acquisition of hosts by endoparasites. Despite the new records, endoparasites found were commonly associated with lizards in Caatinga environments, which may reflect fauna composed of generalist endoparasite species.
Comparing parasitic infection among individuals of wildlife populations can provide insight into factors that influence wildlife disease ecology. Strongylids are parasitic worms that infect the intestinal tract of vertebrates, and infection with strongylids can be approximated by counting strongylid eggs in dung samples. Here we tested for correlations between strongylid egg counts and 18 different individual characteristics, environmental and social factors in individually known wild African elephants. We counted more eggs in the dung samples of younger elephants and females relative to mature elephants and males. We also found that elephants spending more time outside reserves shed more strongylid eggs than elephants that were more often within reserves. Elephants that were less socially integrated, as measured by how much aggression they received from other elephants, shed fewer strongylid eggs; relatedly, socially isolated orphan elephants that had left their family shed fewer strongylid eggs than elephants that remained with their family. Our results suggest that landscapes altered by livestock grazing and social disruption caused by humans may impact parasitic infection in wildlife.
Several factors influence the dynamics and structure of parasite communities. Our goal was to investigate how the community composition, prevalence and abundance of parasites change across seven populations of the exotic lizard Hemidactylus mabouia in Northeast Brazil, and to describe ontogenetic and sex variations. We found differences in the composition of component communities and patterns of infection according to the host body size across the lizard populations. We did not find any variation between the sexes regarding epidemiological patterns, which can probably be explained by the similar diet and habitat use of male and female H. mabouia. An unusually high abundance and prevalence of trematodes infecting this host lizard was apparent when we compared other native lizard hosts, and we suggest that local environmental conditions might be advantageous to the development and life cycle of these parasites due to the abundance of all the intermediate and definitive hosts.
Certain species of parasites have the apparent ability to alter the behaviour of their host in order to facilitate the completion of their own life cycle. While documented in hairworms (phylum Nematomorpha), the ability for mermithid parasites (from the sister phylum Nematoda) to force hosts to enter water remains more enigmatic. Here, we present the first experimental evidence in a laboratory setting that an insect which normally never enters open water (the European earwig Forficula auricularia) will readily enter the water when infected with a mermithid nematode (Mermis nigrescens). Only adult mermithids appear capable of inducing this polarising shift in behaviour, with mermithid length being a very strong predictor of whether their host enters water. However, mermithid length was only weakly associated with how long it took an earwig to enter water following the beginning of a trial. Considering the evidence presented here and its alignment with a proteomic investigation on the same host–parasite system, this study provides strong evidence for adaptive behavioural manipulation and a foundational system for further behavioural and mechanistic exploration.
Coccidial infections reduce fat-soluble vitamin status and bone mineralisation in broiler chickens. We hypothesised that broilers infected with Eimeria maxima would benefit from increased dietary supplementation with vitamin D (vitD) or with 25-hydroxycholecalciferol (25(OH)D3 or 25D3). Broilers were assigned to diets with low (L) or commercial (M) vitD levels (25 v. 100 μg/kg) supplemented as cholecalciferol (D3) or 25D3. At day 11 of age, birds were inoculated with water or 7000 E. maxima oocysts. Pen performance was calculated over the early (days 1–6), acute (days 7–10) and recovery periods (days 11–14) post-infection (pi). At the end of each period, six birds per treatment were dissected to assess long bone mineralisation, plasma levels of 25D3, Ca and P, and intestinal histomorphometry. Parasite replication and transcription of cytokines IL-10 and interferon-γ (IFN-γ) were assessed at day 6 pi using quantitative PCR. Performance, bone mineralisation and plasma 25D3 levels were significantly reduced during infection (P < 0·05). M diets or diets with 25D3 raised plasma 25D3, improved performance and mineralisation (P < 0·05). Offering L diets compromised feed efficiency pi, reduced femur breaking strength and plasma P levels at day 10 pi in infected birds (P < 0·05). Contrastingly, offering M diets or diets with 25D3 resulted in higher parasite loads (P < 0·001) and reduced jejunal villi length at day 10 pi (P < 0·01), with no effect on IL-10 or IFN-γ transcription. Diets with M levels or 25D3 improved performance and mineralisation, irrespective of infection, while M levels further improved feed efficiency and mineralisation in the presence of coccidiosis.
Immunity to microbial infections is well understood; however, information regarding the immunity to parasitic multicellular organisms remains lacking. To understand innate host cellular immunity to nematodes, we compared the cellular response of the greater wax moth (Galleria mellonella) larvae against the non-parasitic, bacterial-feeding nematode Caenorhabditis elegans and pathogenic nematode Heterorhabditis bacteriophora. When intact first-instar or dauer larvae of C. elegans were injected into a G. mellonella larva, most of the nematodes were alive and not confined by the surrounding reaction by insect haemocytes (encapsulation), similarly as the pathogenic nematode, whereas most of the heat-killed nematodes of both species were severely encapsulated by 24 h after inoculation. Other non-parasitic nematodes were also not encapsulated. Surprisingly, C. elegans injected into the insect haemocoel grew and propagated in the live insect, resulting in death of the host insect. Our results suggest that C. elegans has some basic mechanisms to evade immunity of G. mellonenlla and grow in the haemocoel.
Variation in parasitism risk among hosts can arise from between-patch and within-patch factors, but considerably less information is known about the latter. This study investigated how distributions of the oriental fruit fly Bactrocera dorsalis influenced its parasitism by the pupal parasitoid Dirhinus giffardii in the laboratory. Because B. dorsalis larvae pupate underground, pupation depth was considered as an important factor that affects the risk of parasitism. When the density of B. dorsalis larvae was varied (1, 10, and 100 larvae per arena), average pupation depth increased with the density. When the depth of pupae was manipulated, the rate of parasitism differed by depths. Parasitism at 0 cm differed from the random parasitoid model expectation, but parasitism at 1 cm was not different from the model expectation. Few pupae at 2 cm were parasitized. In another experiment, when pupae were simultaneously presented at 0 cm and 1 cm depths, parasitism at 1 cm was weakened by the presence of puape at 0 cm. These results imply that the density of the host influences pupation depth as well as the distribution of parasitism and plays an important role in host-parasitoid dynamics.
Horizontal gene transfer (HGT) has played an important role in the evolution of nematodes. Among candidate genes, cyanase, which is typically found only in plants, bacteria and fungi, is present in more than 35 members of the Phylum Nematoda, but absent from free-living and clade V organisms. Phylogenetic analyses showed that the cyanases of clade I organisms Trichinella spp., Trichuris spp. and Soboliphyme baturini (Subclass: Dorylaimia) represent a well-supported monophyletic clade with plant cyanases. In contrast, all cyanases found within the Subclass Chromadoria which encompasses filarioids, ascaridoids and strongyloids are homologous to those of bacteria. Western blots exhibited typical multimeric forms of the native molecule in protein extracts of Trichinella spiralis muscle larvae, where immunohistochemical staining localized the protein to the worm hypodermis and underlying muscle. Recombinant Trichinella cyanase was bioactive where gene transcription profiles support functional activity in vivo. Results suggest that: (1) independent HGT in parasitic nematodes originated from different Kingdoms; (2) cyanase acquired an active role in the biology of extant Trichinella; (3) acquisition occurred more than 400 million years ago (MYA), prior to the divergence of the Trichinellida and Dioctophymatida, and (4) early, free-living ancestors of the genus Trichinella had an association with terrestrial plants.
This work provides evidence that Subject Island violation effects vanish if subject-embedded gaps are made as frequent and pragmatically felicitous as non-island counterpart controls. We argue that Subject Island effects are caused by the fact that subject-embedded gaps are pragmatically unusual – as the informational focus does not usually correspond to a dependant of the subject phrase – and therefore are highly contrary to comprehenders’ expectations about the distribution of filler–gap dependencies (Chaves 2013, Hofmeister, Casasanto & Sag 2013). This not only explains why sentences with subject-embedded gaps often become more acceptable ‘parasitically’, in the presence of a second gap outside the island, but also explains why some Subject Island violations fail to exhibit any amelioration with repetition (Sprouse 2009, Crawford 2011, Goodall 2011); some ameliorate marginally (Snyder 2000, 2017) or moderately (Hiramatsu 2000, Clausen 2011, Chaves & Dery 2014), and others become fully acceptable, as in our case. This conclusion extends to self-paced reading Subject Island studies (Stowe 1986, Kurtzman & Crawford 1991, Pickering, Barton & Shillcock 1994, Phillips 2006), which sometimes find evidence of gap filling and sometimes do not.
We performed a long-term natural experiment investigating the impact of the diphyllobotriidean cestode Schistocephalus solidus on the body condition and clutch size (CS) of threespine stickleback Gasterosteus aculeatus, its second intermediate host, and the growth of larval parasites in host fish. We tested the hypothesis that single S. solidus infections were more virulent than multiple infections. We also asked whether the metrics of mean and total parasite mass (proxies for individual and total volume, respectively) were consistent with predictions of the resource constraints or the life history strategy (LHS) hypothesis for the growth of, hence exploitation by, larval helminths in intermediate hosts. The samples were drawn from Walby Lake, Alaska in eight of 11 years. Host body condition and CS (egg number per spawning bout) decreased significantly with intensity after adjustments for host size and parasite index. Thus, infections have an increasingly negative impact on measures of host fitness with greater intensity, in contrast to the hypothesis that single infections are more harmful than multiple infections. We also found that mean parasite mass decreased with intensity while total parasite mass increased with intensity as predicted by the LHS hypothesis.
Mermithid nematodes (Nematoda: Mermithida: Mermithidae) parasitize a wide range of both terrestrial and aquatic invertebrate hosts, yet are recorded in bumble bees (Insecta: Hymenoptera: Apidae: Bombus) only six times historically. Little is known about the specific identity of these parasites. In a single-season nationwide survey of internal parasites of 3646 bumble bees, we encountered six additional instances of mermithid parasitism in four bumble bee species and genetically characterized them using two regions of 18S to identify the specific host–parasite relationships. Three samples from the northeastern USA are morphologically and genetically identified as Mermis nigrescens, whereas three specimens collected from a single agricultural locality in the southeast USA fell into a clade with currently undescribed species. Nucleotide sequences of the V2–V6 region of 18S from the southeastern specimens were 2.6–3.0% divergent from one another, and 2.2–4.0% dissimilar to the nearest matches to available data. The dearth of available data prohibits positive identification of this parasite and its affinity for specific bumble bee hosts. By doubling the records of mermithid parasitism of bumble bee hosts and providing genetic data, this work will inform future investigations of this rare phenomenon.
We examined the life history consequences of cornicle secretion by Aphis fabae Scopoli in second and fourth instars, and its effects on host suitability for its parasitoid, Lysiphlebus fabarum (Marshall). Cornicle secretion did not affect aphid fecundity, but secretion in the second instar enhanced life table parameters, whereas secretion in the fourth instar affected them negatively, suggesting a higher cost of secretion in later instars. Secretion in either instar improved host suitability for L. fabarum. Although control and treated aphids were parasitized at similar rates, and with similar success, wasps developed faster and emerged as larger adults in aphids that had secreted, regardless of instar. Transgenerational effects were also evident. Progeny emergence was higher when parental wasps developed in fourth instars than in seconds, whether aphids secreted or not, and progeny were larger when parental hosts secreted in the second instar, but not in the fourth. Secreting fourth instars were preferred to controls by L. fabarum females in choice tests, but not secreting second instars, and fourth-instar secretion improved wasp emergence. When control aphids were attacked, second instars were more likely to secrete than fourth instars, whereas the latter were more likely to kick the parasitoid. Cornicle secretion reduced the probability of subsequent secretion events and the frequency of other aphid defensive behaviors, indicating energetic tradeoffs among defensive tactics. Overall, our results revealed that cornicle secretion by immature A. fabae exacts both physiological and behavioral costs and results in improved host suitability for its parasitoid.
Temperature is expected to modulate the responses of organisms to stress. Here, we aimed to assess the influence of temperature on the interaction between parasitism and fungicide contamination. Specifically, using the cladoceran Daphnia as a model system, we explored the isolated and interactive effects of parasite challenge (yeast Metschnikowia bicuspidata) and exposure to fungicides (copper sulphate and tebuconazole) at two temperatures (17 and 20 °C), in a fully factorial design. Confirming a previous study, M. bicuspidata infection and copper exposure caused independent effects on Daphnia life history, whereas infection was permanently suppressed with tebuconazole exposure. Here, we show that higher temperature generally increased the virulence of the parasite, with the hosts developing signs of infection earlier, reproducing less and dying at an earlier age. These effects were consistent across copper concentrations, whereas the joint effects of temperature (which enhanced the difference between non-infected and infected hosts) and the anti-parasitic action of tebuconazole resulted in a more pronounced parasite × tebuconazole interaction at the higher temperature. Thus, besides independently influencing parasite and contaminant effects, the temperature can act as a modulator of interactions between pollution and disease.
Hosts face mortality from parasitic and environmental stressors, but interactions of parasitism with other stressors are not well understood, particularly for long-lived hosts. We monitored survival of flour beetles (Tribolium confusum) in a longitudinal design incorporating cestode (Hymenolepis diminuta) infection, starvation and exposure to the pesticide diatomaceous earth (DE). We found that cestode cysticercoids exhibit increasing morphological damage and decreasing ability to excyst over time, but were never eliminated from the host. In the presence of even mild environmental stressors, host lifespan was reduced sufficiently that extensive degradation of cysticercoids was never realized. Median host lifespan was 200 days in the absence of stressors, and 3–197 days with parasitism, starvation and/or DE. Early survival of parasitized hosts was higher relative to controls in the presence of intermediate concentrations of DE, but reduced under all other conditions tested. Parasitism increased host mortality in the presence of other stressors at times when parasitism alone did not cause mortality, consistent with an interpretation of synergy. Environmental stressors modified the parasite numbers needed to reveal intensity-dependent host mortality, but only rarely masked intensity dependence. The longitudinal approach produced observations that would have been overlooked or misinterpreted if survival had only been monitored at a single time point.
The population density of the diamondback moth (DBM), Plutella xylostella (L.), the species diversity of its predators, parasitism on eggs, larvae and pupae, and associated yield of canola were evaluated in experimental plots where spring canola was grown either as a monoculture or strip-intercropped with annual alfalfa in the ratio of 3C:3A, 6C:3A and 9C:3A over a 2-year period. The egg density was significantly higher in monoculture than in intercrops. The highest and lowest densities of larvae and pupae of DBM were recorded in monoculture (2.9–3.2 larvae per plant and 1.14–1.20 pupae per plant) and intercrop 3C:3A (0.7–0.6 larvae per plant and 0.34–0.29 pupae per plant), respectively. Shannon diversity index (H') for species composition of the predators of DBM immature stages was lower in monoculture than in intercrops, and was similar amongst the three intercrops. Moreover, the percent parasitism for eggs, larvae and pupae was higher in intercrops than in monoculture. The dry seed weight loss was higher in monocrop (37.6–40.1%) compared to 3C:3A (7.9–8.6%), 6C:3A (19.5–21.4%) and 9C:3A (21.6–25.4%). Our results indicate that intercropping canola with annual alfalfa, especially in the ratio 3C:3A, can increase the species diversity, parasitism rates of DBM immature stages, and enhance the yield of canola. The implications of these findings, in relation to integrated pest management (IPM) in canola cropping systems, are discussed.
There are several possible causes of aggregated nesting in solitary Aculeata, one being joint defense against parasites. We tested whether females prefer nesting in aggregations, even if they consist of heterospecifics. We compared the colonization and nesting parasitism of trap-nests with and without a red mason bee aggregation. The results did not support our hypothesis that females prefer nesting in aggregations. The numbers of wild Aculeata nests did not differ between trap-nests with and without an aggregation. Unexpectedly, parasitism rates were higher in trap-nests with aggregations. When analyzing only nests of wild insects (mostly wasps), the differences in parasitism disappeared. Natural nesting sites may be such a limited resource that females nested in the first trap-nest they encountered and did not discriminate between our treatments, or wasps might share too few parasites species with bees to benefit from joint nest defense.
In laboratories, the parasitism rate of Ostrinia furnacalis (Güenée) (Lepidoptera: Crambidae) eggs by Trichogramma dendrolimi Matsumura (Hymenoptera: Trichogrammatidae) is low; however, efforts to control O. furnacalis with T. dendrolimi in the field have been successful. In this study, the effects of the number of attacks by T. dendrolimi against O. furnacalis eggs and diet of O. furnacalis larva on wasp development were investigated. The results indicated that more attacks increased significantly not only the successful parasitism rate of O. furnacalis eggs by T. dendrolimi, but also the percentage of host eggs that failed to develop into either O. furnacalis larvae or T. dendrolimi. Both the size and female proportion of T. dendrolimi offspring decreased as the number of attacks increased. The number of T. dendrolimi eggs laid in per host egg increased significantly as the ratio of wasps to host eggs increased from 1:5 to 3:5. Host diet also significantly affected the developmental time of immaturity and the emergence rate of adults of T. dendrolimi. These results illustrate how inundative releases of T. dendrolimi can successfully control O. furnacalis despite the fact that pest parasitism by the subsequent wasp generation decreases sharply in the field. The suitability of O. furnacalis eggs to T. dendrolimi and the superparasitism effects on offspring of T. dendrolimi are discussed.
Cosmopolitan pests such as Brevicoryne brassicae, Lipaphis pseudobrassicae, and Myzus persicae (Aphididae) cause significant damage to Brassicaceae crops. Assessment of the important biotic and abiotic factors that regulate these pests is an essential step in the development of effective Integrated Pest Management programs for these aphids. This study evaluated the influence of leaf position, precipitation, temperature, and parasitism on populations of L. pseudobrassicae, M. persicae, and B. brassicae in collard greens fields in the Triângulo Mineiro region (Minas Gerais state), Brazil. Similar numbers of B. brassicae were found on all parts of the collard green plants, whereas M. persicae and L. pseudobrassicae were found in greatest numbers on the middle and lower parts of the plant. While temperature and precipitation were positively related to aphid population size, their effects were not accumulative, as indicated by a negative interaction term. Although Diaeretiella rapae was the main parasitoid of these aphids, hyperparasitism was dominant; the main hyperparasitoid species recovered from plant samples was Alloxysta fuscicornis. Parasitoids seem to have similar distributions on plants as their hosts. These results may help predict aphid outbreaks and gives clues for specific intra-plant locations when searching for and monitoring aphid populations.
Many plant tissues contain plant secondary compounds (PSC), which have long been recognised as defensive chemicals that deter herbivory via their toxic effects. However, herbivores may also benefit from including PSC into their diets. Plant-derived phenolics, terpenes and alkaloids have antiparasitic properties and sesquiterpene lactones have antibacterial, antifungal and antiparasitic properties. These actions are in part a consequence of the negative actions that PSC exert across several trophic levels, including the bacteria, parasites and fungi that inhabit herbivores’ bodies. Given the dual action, toxin and medicine, it is possible to hypothesise that self-selection of PSC by herbivores should occur when the benefits outweigh the costs of PSC ingestion. Recent research suggests that sheep and goats self-medicate against parasitic infections. They increase preference for condensed tannin-containing foods when experiencing a parasitic burden. This behaviour improves health; it is triggered by parasitism and weakens when parasitism subsides. However, the causes underlying these responses are not straightforward when viewed under a unidimensional cost–benefit analysis. This is because the intensity of antinutritional/toxic and medicinal effects of PSC is not static or just dependent upon the isolated post-ingestive effects of single PSC. Nutrient–PSC and PSC–PSC interactions, social models, as well as feeding patterns, all influence the perceived net benefit of incorporating medicines into a diet. A better understanding of the net benefit of self-medication in complex feeding environments will allow for the development of innovative managing strategies aimed at providing the food alternatives and conditions for improving the nutrition, health and welfare of grazing animals.
In reproducing ewes, a periparturient breakdown of immunity is often observed to result in increased fecal egg excretion, making them the main source of infection for their immunologically naive lambs. In this study, we expanded a simulation model previously developed for growing lambs to explore the impact of the genotype (performance and resistance traits) and host nutrition on the performance and parasitism of both growing lambs and reproducing ewes naturally infected with Teladorsagia circumcincta. Our model accounted for nutrient-demanding phases, such as gestation and lactation, and included a supplementary module to manage the age structure of the ewe flock. The model was validated by comparison with published data. Because model parameters were unknown or poorly estimated, detailed sensitivity analysis of the model was performed for the sheep mortality and the level of infection, following a preliminary screening step. The parameters with the greatest effect on parasite-related outputs were those driving animal growth and milk yield. Our model enables different parasite-control strategies (host nutrition, breeding for resistance and anthelmintic treatments) to be assessed on the long term in a sheep flock. To optimize in silico exploration, the parameters highlighted by the sensitivity analysis should be refined with real data.