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Aggregation of macroparasites among hosts is a near-universal pattern, and has important consequences for the stability of host-parasite associations and the impacts of disease. Identifying which potential drivers are contributing to levels of aggregation observed in parasite-host associations is challenging, particularly for observational studies. We apply beta regressions in a Bayesian framework to determine predictors of aggregation, quantified using Poulin's index of discrepancy ( D ), for 13 species of parasites infecting Icelandic Rock Ptarmigan ( Lagopus muta ) collected over 12 years. 1,140 ptarmigan were collected using sampling protocols maximizing consistency of sample sizes and of composition of host ages and sexes represented across years from 2006-2017. Parasite species, taxonomic group (insect, mite, coccidian, or nematode), and whether the parasite was an ecto- or endoparasite were tested as predictors of aggregation, either alone or by modulating an effect of parasite mean abundance on D . Parasite species was an important predictor of aggregation in models. Despite variation in D across samples and years, relatively consistent aggregation was demonstrated for each specific host-parasite association, but not for broader taxonomic groups, after taking sample mean abundance into account. Furthermore, sample mean abundance was consistently and inversely related to aggregation among the nine ectoparasites, however no relationship between mean abundance and aggregation was observed among the four endoparasites. We discuss sources of variation in observed aggregation, sources both statistical and biological in nature, and show that aggregation is predictable, and distinguishable, among infecting species. We propose explanations for observed patterns and call for the review and re-analysis of parasite and other symbiont distributions using beta regression to identify important drivers of aggregation-both broad and association-specific., Competing Interests: The authors declare there are no competing interests., (©2022 Morrill et al.)

Parasite species often show differential fitness on different host species. We developed an equation-based model to explore conditions favouring host species exploitation and discrimination. In our model, diploid infective stages randomly encountered hosts of two species; the parasite's relative fitness in exploiting each host species, and its ability to discriminate between them, was determined by the parasite's genotype at two independent diallelic loci. Relative host species frequency determined allele frequencies at the exploitation locus, whereas differential fitness and combined host density determined frequency of discrimination alleles. The model predicts instances where populations contain mixes of discriminatory and non-discriminatory infective stages. Also, non-discriminatory parasites should evolve when differential fitness is low to moderate and when combined host densities are low, but not so low as to cause parasite extinction. A corollary is that parasite discrimination (and host-specificity) increases with higher combined host densities. Instances in nature where parasites fail to discriminate when differential fitness is extreme could be explained by one host species evolving resistance, following from earlier selection for parasite non-discrimination. Similar results overall were obtained for haploid extensions of the model. Our model emulates multi-host associations and has implications for understanding broadening of host species ranges by parasites.This article is part of the themed issue 'Opening the black box: re-examining the ecology and evolution of parasite transmission'., (© 2017 The Authors.)

Background: To understand variation in resistance to parasites within host populations, researchers have examined conditions under which immunity is induced and/or is costly. Both host sex and age have been found to influence immune expression and subsequently are likely factors influencing the costs of resistance. The purpose of this study was to examine immune expression and associated survival costs for two age groups (newly emerged and sexually mature individuals) of the damselfly, Enallagma boreale Selys. Survival was assessed for experimentally challenged and control damselflies, housed initially at 22°C and then subjected to low temperatures (15°C) associated with reduced foraging activity and food deprivation. Experimental conditions emulated natural local variation in bouts of good weather followed by inclement weather (successions of days with hourly mean temperatures around 15°C and/or rainy weather). Results: At least one of three immune traits was induced to higher levels for both newly emerged and mature E. boreale challenged by Lippopolysaccharide (LPS) relative to saline-injected controls, when housed at 22°C. The immune traits assayed included haemocyte concentration, Phenoloxidase activity and antibacterial activity and their induction varied among ages and between males and females. For matures, those injected with LPS had lowered survivorship compared to saline-injected controls that were housed initially at 22°C and subsequently at 15°C. Newly emerged LPS-injected damselflies did not show reduced survivorship relative to newly-emerged controls, despite showing immune induction. Conclusion: Reduced longevity following induction of immunity was observed for reproductively mature damselflies, but not for newly emerged damselflies. Costs of resistance depend only partly on the immune trait induced and more on the age (but not sex) of the host. In four years, we often observed bouts of inclement weather following good days and these bouts occurred primarily during the emergence periods, but also during the flight periods, of E. boreale. The duration of these bouts appear sufficient to compromise survival of mature damselflies that responded immunologically to LPS challenge. We further suggest the environmental conditions likely experienced by different ages of damselflies, following resistance expression, has influenced optimal immune investment by individuals in different age classes and the likelihood of detecting costs of resistance. [ABSTRACT FROM AUTHOR]