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The dilution effect was originally proposed to describe the negative effect of increased host diversity on parasite or pathogen abundance; with greater host diversity, parasite or pathogen levels per host are predicted to be lower due to a higher probability of dispersing parasites or pathogens encountering noncompetent hosts. Dilution effects could occur in many mutualisms if dispersing symbionts encounter hosts that vary in their competence. Introduced, non‐native hosts can change the community competence of a local group of host species. Crayfish introductions are occurring world‐wide and these introductions are likely disrupting native crayfish‐symbiont systems. Branchiobdellidan symbionts declined on native Cambarus crayfish co‐occurring with non‐native Faxonius crayfish in the New River watershed, USA. We performed an experiment investigating the effect of host density (1 vs. 2 native hosts) and host diversity (1 native host and 1 introduced host) on branchiobdellidan abundance. The introduced Faxonius cristavarius is a noncompetent host for these worms. Six C. ingens were stocked on a Cambarus chasmodactylus in each treatment and worm numbers were followed over 34 days. Worm numbers decreased over time on C. chasmodactylus alone and in the treatment in which a C. chasmodactylus was paired with an F. cristavarius. Worm numbers remained highest in the 2 C. chasmodactylus treatment. There was no effect of host diversity on worm reproduction. Crayfish invasions may have negative effects on mutualistic symbionts depending on the competence of introduced hosts. Loss of native symbionts is one of the potential hidden, negative effects of invasions on native freshwater diversity. [ABSTRACT FROM AUTHOR]

Numerous mutualisms have been described from terrestrial and marine communities and many of these mutualisms have significant effects on community structure and function. In contrast, there are far fewer examples of mutualisms from freshwater habitats and there is no evidence that any mutualism has community‐wide or ecosystem‐level consequences. Northern hemisphere crayfish are host to a variety of ectosymbiotic worms called branchiobdellidans. The association between some of these "crayfish worms" and their hosts is a mutualism. The outcome of the association is context dependent and can be influenced by host size, symbiont number, and the environment. Here we document in two experiments that the mutualism between crayfish and these worms alters the effect of crayfish on stream community structure and sediment deposition, an important ecosystem variable. We enclosed crayfish stocked with 0 worms and intermediate (3–6) and high worm densities (12) in cages in streams in Boone, North Carolina and Clemson, South Carolina, United States. At both locations, there was a negative relationship between initial worm density and final macroinvertebrate abundance. There was a significant effect of worm treatment on macroinvertebrate community structure in both the Boone and Clemson experiments. In Boone, there were effects on both overall macroinvertebrate abundance and community composition, whereas in Clemson, changes to community structure were primarily driven by changes in total abundance. There was a negative relationship between benthic sediment volume and initial worm density in both experiments, primarily later in the experiments, though these effects were influenced by sediment deposition rates. Our results are the first to demonstrate strong effects of a mutualism on freshwater communities. Both members of this mutualism are found throughout the northern hemisphere, so similar impacts may occur in many other waterways. Given that various species in addition to crayfish function as keystone species and ecosystem engineers in freshwater systems throughout the world, mutualisms involving these strongly interacting species may be as important to the structure and functioning of freshwater systems as comparable mutualisms in marine and terrestrial systems. [ABSTRACT FROM AUTHOR]

Animal fitness is influenced by diverse assemblages of internal and external symbionts. These assemblages often change throughout host ontogeny, but the mechanisms that underlie these changes and their consequences for host fitness are seldom revealed. Here we examine a cleaning symbiosis between crayfish and an assemblage of ectosymbiotic branchiobdellidan worms to uncover what mechanisms drive changes in symbiont composition during host ontogeny and the consequences of these changes for both the host and symbionts. In surveys of a North Carolina river, the dominant worm species shifted from Cambarincola philadelphicus to Cambarincola ingens as crayfish ( Cambarus bartonii) increased in size. We demonstrate that this shift is a function of host regulation by small crayfish and exclusion by a dominant symbiont on large crayfish. In a controlled lab experiment, small crayfish often removed their symbionts but C. ingens was removed at a higher rate than C. philadelphicus. In contrast, C. ingens had higher survivorship and reproduction than C. philadelphicus on large crayfish. We also measured the effect of each worm species on crayfish growth through ontogeny; neither worm species had an effect on small crayfish but both species had similar positive effects on the growth of large crayfish relative to controls. Evidence from another experiment suggested that intraguild predation by C. ingens caused a decline in C. philadelphicus on large crayfish. We have shown that shifts in partner fitness are a function of host size and that these shifts can involve the succession of symbionts. Further, our results suggest that changes in the outcome of symbioses can remain robust throughout host ontogeny despite interactive mechanisms that lead to shifts in symbiont community structure. [ABSTRACT FROM AUTHOR]

Symbiont community assembly is driven by host-symbiont and symbiont-symbiont interactions. The effects that symbionts exert on their hosts are often context-dependent, and existing theoretical frameworks of symbiont community assembly do not consider the implications of variable outcomes to assembly processes., We hypothesized that symbiont-symbiont interactions become increasingly important along a parasitism/mutualism continuum because; (i) negative outcomes favour host resistance which in turn reduces symbiont colonization and subsequently reduce symbiont-symbiont interactions, whereas (ii) positive host outcomes favour tolerance and consequently higher symbiont colonization rates, leading to stronger interactions among symbionts. We found support for this hypothesis in the cleaning symbiosis between crayfish and ectosymbiotic branchiobdellidan worms., The symbiosis between crayfish and their worms can shift from parasitism/commensalism to mutualism as crayfish age. Here, field surveys identified changes in worm density, diversity and composition that were concomitant to changing symbiosis outcomes. We conducted several laboratory experiments and behavioural assays to relate patterns from the field to their likely causal processes., Young crayfish typically hosted only two relatively small worm species. Older crayfish hosted two additional larger species. In laboratory experiments, young crayfish exhibited a directed grooming response to all worm species, but were unable to remove small species. Conversely, adult crayfish did not exhibit grooming responses to any worm species. Relaxed grooming allowed the colonization of large worm species and initiated symbiont-symbiont intraguild predation that reduced the abundance and altered the behaviour of small worm species. Thus, the dominant processes of symbiont community assembly shifted from host resistance to symbiont-symbiont interactions through host ontogeny and a concomitant transition towards mutualism., This work shows that host resistance can have a prevailing influence over symbiont community assembly when symbiosis is disadvantageous to the host. However, when symbiosis is advantageous and resistance is relaxed, symbiont colonization rate and consequently abundance and diversity increases and interactions among symbionts become increasingly important to symbiont community assembly. [ABSTRACT FROM AUTHOR]

Direct horizontal transmission of pathogenic and mutualistic symbionts has profound consequences for host and symbiont fitness alike. While the importance of contact rates for transmission is widely recognized, the processes that underlie variation in transmission during contact are rarely considered. Here, we took a symbiont's perspective of transmission as a form of dispersal and adopted the concept of condition-dependent dispersal strategies from the study of free-living organisms to understand and predict variation in transmission in the cleaning symbiosis between crayfish and ectosymbiotic branchiobdellidan worms. Field study showed that symbiont reproductive success was correlated with host size and competition among worms for microhabitats. Laboratory experiments demonstrated high variability in transmission among host contacts. Moreover, symbionts were more likely to disperse when host size and competition for microhabitat created a fitness environment below a discrete minimum threshold. A predictive model based on a condition-dependent symbiont dispersal strategy correctly predicted transmission in 95% of experimental host encounters and the exact magnitude of transmission in 67%, both significantly better than predictions that assumed a fixed transmission rate. Our work provides a dispersal-based understanding of symbiont transmission and suggests adaptive symbiont dispersal strategies can explain variation in transmission dynamics and complex patterns of host infection. [ABSTRACT FROM AUTHOR]

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When the interests of mutualists are not perfectly aligned, control mechanisms that modulate interactions can maintain mutually beneficial outcomes and stabilize mutualisms over evolutionary time. However, the costs and benefits of symbiosis often change with ontogeny and whether control mechanisms are adjusted to reflect ontogenetic changes is largely unknown. We examined the recently described cleaning symbiosis between crayfish Cambarus chaugaensis and ectosymbiotic annelids ( Xironodrilus appalachius) for evidence of ontogenetic changes in symbiont control. Xironodrilus appalachius provide a beneficial cleaning service to C. chaugaensis by removing epibiotic accumulations from the gills, but crayfish also incur costs via density-dependent facultative parasitism of gill tissue. A series of laboratory experiments using crayfish from three size (age) - classes demonstrated that crayfish use grooming to limit cleaner density and grooming effects on cleaners varied with crayfish age. Small crayfish quickly removed essentially all of their cleaners. Intermediate crayfish removed most of their cleaners, but some cleaners persisted at a location apparently inaccessible to grooming and far from the gill chamber. Large crayfish removed a smaller proportion of cleaners and cleaners were allowed access to the gill chamber, thus initiating the cleaning symbiosis. Cleaner removal was not dependent on cleaner density, suggesting that crayfish do not regulate cleaners to a specific density. Experimental results were corroborated by patterns observed during a field survey. We argue decreased cleaner removal and relaxed control of cleaner attachment sites corresponds to ontogenetic changes in the costs and benefits of symbiosis. This study integrates two major theoretical perspectives from ecological literature; control mechanisms and ontogenetic shifts, and illustrates how changes in control mechanisms with ontogeny may favor life-long positive outcomes of symbiosis. Ontogenetic shifts in the costs and benefits of symbiosis may be common; therefore future theoretical and empirical studies of symbioses should incorporate both perspectives. [ABSTRACT FROM AUTHOR]

Cleaning symbioses provide net benefits by improving each partner's fitness. Ectosymbiotic Cambarincola spp. (branchiobdellidans) can increase growth and survival of Cambarus chasmodactylus (New River Crayfish), but the nature of the symbiosis might change with female reproductive state because brooding offspring (eggs, young) and worms inhabit the same surfaces. Here, we present the results of field surveys that examined whether the number and location of branchiobdellidans on New River Crayfish varies as a function of female crayfish reproductive state. Reproducing female New River Crayfish had fewer total worms, an absence of cocoons, and a relatively greater proportion of worms on lateral body surfaces than non-reproducing crayfish. The altered distribution and reduced abundance of worms suggest that the symbiosis changes with female reproductive status, but additional experiments will be needed to identify the mechanism responsible. [ABSTRACT FROM AUTHOR]