Adaptive host-abandonment of ectoparasites before fledging? Within-brood distribution of nest mites in house sparrow broods

We studied the within-brood distribution of a haematophagous mite Pellonyssus reedi living on nestling house sparrows (Passer domesticus) near the time of fledging. We measured the natural level of infestation of individual nestlings, and determined the feeding efficiency of mites, by scoring their feeding status. Within-brood distribution of mite loads was unrelated to nestling body mass, tarsus length, or immunocompetence. These results did not support parasite preference for large or susceptible hosts. Mite feeding-efficiency was also unrelated to these nestling characteristics, confirming that large nestlings or nestlings with less-developed immunocompetence did not provide superior feeding conditions for mites. Therefore, our results do not support the hypothesis that within-brood distribution of avian ectoparasites is explained by the parasites' preferences for characteristics, such as large body size or low immunocompetence, that make nestlings suitable hosts. On the other hand, we found that mite loads were negatively correlated with nestling age and feather length, suggesting that nestlings closer to fledging harbored fewer mites then their less-developed nestmates. Furthermore, feather length had a stronger relationship with parasite distribution than did nestling age. We presume, therefore, that feather characteristics, i.e., length, may serve as a signal for mites to perceive the ready-to-fledge state of nestlings, inducing abandonment behavior. These results support another, largely neglected hypothesis, i.e., that the avoidance or abandonment of those nestlings that are close to fledging may also explain the parasites' distribution in a brood. This hypothesis is based on the argument that many nest-dwelling ectoparasites breed in the nest material and emerge only periodically to feed on nestlings. In such parasites, the ability to recognize and avoid mature fledglings can be adaptive because this may help the parasites to avoid their removal from the nest so they can continue to reproduce by feeding on unfledged chicks of the current or later broods. Our results suggest that adaptive host-abandonment by nest-dwelling ectoparasites can influence within-brood parasite distributions around the time of fledging.     

Nestling sex predicts susceptibility to parasitism and influences parasite population size within avian broods

Vertebrate hosts differ in their level of parasite susceptibility and infestation. In avian broods, variation in susceptibility of nestlings to ectoparasites may be associated with non‐uniform distributions of parasites among brood mates, with parasites concentrating feeding on the most vulnerable hosts. The presence of a highly susceptible nestling in a brood can benefit the remaining young by reducing the parasite pressure they experience; however, from a parasite’s perspective, broods with fewer susceptible hosts may provide effectively fewer resources than broods of the same size containing a greater abundance of susceptible hosts, and this could limit the number of parasites that a host brood can sustain. To test whether variation in number of susceptible hosts affects the number of parasites in bird nests, we first examined the role of host sex and induced immunity (via methionine supplementation) on susceptibility of mountain bluebirds Sialia currucoides to parasitism by blow flies Protocalliphora spp. We then assessed the effect of variation in number of susceptible hosts on the number of parasites inhabiting the nest. Only females showed a benefit of methionine supplementation, gaining mass more rapidly following supplementation compared to males. This suggests that females are more susceptible to parasites in this system; this was further supported by parasite feeding trials, in which parasites extracted larger blood meals from female than male hosts. Finally, the abundance of parasites in nests was predicted by brood sex ratio: broods containing more female young harboured more parasites. Hence, within‐brood variation in host susceptibility to parasites can not only influence the costs of parasitism for individual nestlings, but may also have consequences for the size of parasite populations within nests. If patterns of maternal investment affect the abundance of nest‐dwelling parasites, these interactions may be important for understanding fitness consequences of maternal resource allocation in many vertebrate hosts.     

Parasite-mediated growth patterns and nutritional constraints in a cavity-nesting bird

1. Trade-offs between growth and immunity of nestling birds can be influenced by parasites, but the magnitude of these effects may depend on availability of critical dietary nutrients. Owing to their importance for both immune system function and growth, dietary carotenoids have the potential to mediate parasite-induced developmental strategies of avian hosts. 2. The effects of ectoparasitic blow flies Protocalliphora spp. and dietary carotenoids (lutein and zeaxanthin) on immune function and patterns of growth in nestling mountain bluebirds Sialia currucoides were investigated by combining parasite removal and carotenoid supplementation treatments in a 2 x 2 design. 3. Supplemental carotenoids enhanced nestlings' T-cell-mediated immune response following intradermal injection of phytohaemagglutinin. 4. The effect of carotenoid supplementation on rate of mass gain depended on whether broods were exposed to parasites: among parasitized broods, those receiving supplemental carotenoids gained mass more rapidly than nonsupplemented broods, whereas there was no effect of supplemental carotenoids on growth of mass in broods that had parasites removed. This suggests that additional dietary carotenoids allowed nestlings to compensate for the otherwise detrimental effects of parasites on mass gain. For length of the eighth primary feather at fledging, early and late broods differed in their response to parasitism: early broods showed an increase in feather length when parasites were removed, while nestlings in late broods had shorter feathers in the absence of parasites. We suggest that this may reflect within-season variation in parasite-mediated growth strategies of nestlings. 5. Maternal condition was positively associated with mass, condition and rate of feather growth of offspring under all conditions, and also influenced nestling immunocompetence, but only in the absence of parasites. 6. We conclude that dietary carotenoids alleviate some of the detrimental effects of parasites on nestling birds; however, parasites also appear to specifically influence other growth and resource allocation strategies, and possibly constrain maternal or genetic effects on offspring phenotype, irrespective of dietary carotenoid availability.     

Intraseasonal variation in immune defence,body mass and hematocrit in adult house martins Delichon urbica

The intensity of parasite infections often increases during the reproductive season of the host as a result of parasite reproduction, increased parasite transmission and increased host susceptibility. We report within‐individual variation in immune parameters, hematocrit and body mass in adult house martins Delichon urbica rearing nestlings in nests experimentally infested with house martin bugs Oeciacus hirundinis and birds rearing nestlings in initially parasite‐free nests. From first to second broods body mass and hematocrit of breeding adult house martins decreased. In contrast leucocytes and immunoglobulins became more abundant. When their nests were infested with ectoparasites adults lost more weight compared with birds raising nestlings in nests treated with pyrethrin, whereas the decrease in hematocrit was more pronounced during infection with blood parasites. Neither experimental infestation with house martin bugs nor blood parasites had a significant effect on the amount of immune defences.     

Influence of hatching asynchrony and within-brood parental investment on size, condition, and immunocompetence in nestling red-billed choughs

Studies of hatching asynchrony have rarely assessed the effect of parental investment strategies on the development of intra-brood hierarchies using indicators of nestling quality. The influence of hatching asynchrony and other variables related to parental investment on immunocompetence was evaluated measuring T-cell mediated immune response in broods of red-billed choughs ( Pyrrhocorax pyrrhocorax ), a large long-living passerine with a pronounced sexual size dimorphism. The results showed that T-cell mediated immune response depends on parental effects as shown by the differences between broods on hatching asynchrony. Male nestlings were both heavier and larger than female nestlings, but there was no effect of hatching order on these traits, nor on sex differences in immunocompetence. Differential investment strategy in relation to laying order did not favour older offspring or either of the sexes. Successful reproduction in this species might be so unpredictable and infrequent that strategies of parental investment in the brood could have evolved to attempt to maximize the survival of all nestlings by avoiding within-brood hierachies of size, body condition, and immunocompetence.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 675–684.     

Experimental evidence for host preference in a tick parasitizing songbird nestlings

Mechanisms of host preference in ectoparasites are important to the understanding of host‐parasite interactions. Since ectoparasites negatively affect the condition of their hosts, while the hosts’ condition itself may affect the parasites’ choice, separating the factors that drive host preference from parasite impact asks for experiments. We combined the data of two choice experiments to investigate the preference of the nidicolous tick Ixodes arboricola when exposed to the nestlings of a passerine bird (Parus major). In the first experiment, in which complete broods at hatching were exposed to an ecologically relevant number of ticks, the relationship between tick loads and nestlings’ developmental status was characterized by a distribution with the highest tick loads on the more developed nestlings. Host preference became more apparent at a smaller brood size, suggesting a role for host density. In a second experiment we evaluated host choice in a pairwise choice experiment, exposing pairs of siblings with contrasting developmental status to eight ticks. In the first and the second pair, a median developed nestling was linked with the most developed and the least developed nestling, respectively. Seventy‐two h after tick exposure we measured the innate constitutive humoral immunity and haematocrit. No differences were found in innate immunity, but the least developed nestlings had on average a lower haematocrit than the median and most developed nestlings. Significantly fewer ticks attached on the least developed nestling compared to the median nestling, and this difference was more pronounced when the innate immunity of the median developed nestling was higher. No difference in tick load was found among the median and best developed nestlings. The linkage between host preference and host physiological condition provide further insight in the mechanisms driving ectoparasite aggregation, which is important for the population dynamics of host, ticks and tick‐transmitted pathogens.     

Immune function and survival of great tit nestlings in relation to growth conditions

Life history theory predicts a trade-off between number and quality of offspring. Reduced quality with increasing brood size may arise from a decrease in body condition or in immunocompetence that would be important in fighting off virulent parasites by immunologically naive offspring. We tested the effect of rearing conditions on immune function of nestling great tits (Parus major) by reducing or increasing broods by two hatchlings. In the middle of the nestling period (on day 8), nestlings from enlarged broods developed lower T cell responses [as measured from the cutaneous swelling reaction to injection with phytohaemagglutinin (PHA)] and tended to have lower total leukocyte and lymphocyte concentrations in their peripheral blood than nestlings from reduced broods. Brood size manipulation affected the PHA response of nestlings most strongly in small clutches, suggesting that nestling immune function was dependent on their parents’ condition, as estimated by original clutch size. Intra-brood differences in nestling mortality were unrelated to immune parameters, but nestlings in broods without mortality had a stronger PHA response, higher concentration of lymphocytes and higher body mass on day 15 than nestlings in broods with mortality. These results support the prediction that the immune function of altricial birds is affected by rearing conditions, and that growth and immune parameters are related to inter-brood differences in nestling survival. Received: 1 February 1999 / Accepted: 19. July 1999     

Physiological and haematological consequences of a novel parasite on the red-rumped swallow Hirundo daurica

Parasite virulence has been hypothesised to increase with the degree of host sociality because highly social hosts have a greater probability of encountering horizontal transmission of parasites and experiencing infections with multiple strains of the same parasites than do solitary hosts. As compared with the defences of closely related social host species, we predicted that solitary hosts should have relatively weak defences against parasites, thus being relatively more affected when parasitised by a novel parasite. We tested this prediction by either experimentally infesting 12 nests of the solitarily nesting red-rumped swallow Hirundo daurica with 50 individuals of the generalist martin bug Oeciacus hirundinis or by fumigation of nine nests. Nestlings 13 days old from the parasite addition group experienced increased mortality, attained lower body mass and tended to have shorter tarsi compared to nestlings from fumigated nests. Surprisingly, nestlings from the parasite addition group had higher packed cell volume (cellular fraction of blood) and lower levels of heat shock proteins (HSP60) than nestlings from the fumigation group. A measure of immunocompetence was not significantly affected by treatment, but its magnitude was positively related to packed cell volume and negatively related to level of HSP60. Solitary hosts like the red-rumped swallow have weak immune responses and low levels of heat shock proteins when infested with ectoparasites while highly social hosts have strong immune responses and high levels of heat shock proteins when infested. These findings partially support the hypothesis that potential host species with weak defences are more susceptible to infection and the deleterious effects of evolving parasites than potential hosts with strong defences.     

Sex-related effects of maternal egg investment on offspring in relation to carotenoid availability in the great tit

1. Maternal carotenoids in the egg yolk have been hypothesized to promote maturation of the immune system and protect against free radical damages. Depending on availability, mothers may thus influence offspring quality by depositing variable amounts of carotenoids into the eggs. Sex allocation theory predicts that in good quality environments, females should invest into offspring of the sex that will provide larger fitness return, generally males. 2. In a field experiment we tested whether female great tits bias their investment towards males when carotenoid availability is increased, and whether male offspring of carotenoid-supplemented mothers show higher body condition. We partially cross-fostered hatchlings to disentangle maternal effects from post-hatching effects, and manipulated hen flea Ceratophyllus gallinae infestation to investigate the relationship between carotenoid availability and resistance to ectoparasites. 3. As predicted, we found that carotenoid-supplemented mothers produced males that were heavier than their sisters at hatching, while the reverse was true for control mothers. This suggests that carotenoid availability during egg production affects male and female hatchlings differentially, possibly via a differential allocation to male and female eggs. 4. A main effect of maternal supplementation became visible 14 days after hatching when nestlings hatched from eggs laid by carotenoid-supplemented mothers had gained significantly more mass than control nestlings. Independently of the carotenoid treatment, fleas impaired mass gain of nestlings during the first 9 days in large broods only and reduced tarsus length of male nestlings at an age of 14 days, suggesting a cost to mount a defence against parasites. 5. Overall, our results suggest that pre-laying availability of carotenoids affects nestling condition in a sex-specific way with potentially longer-lasting effects on offspring fitness.     

Hatching asynchrony and brood reduction influence immune response in Common Kestrel Falco tinnunculus nestlings

The onset of incubation before the end of laying imposes asynchrony at hatching and, therefore, a size hierarchy in the brood. It has been argued that hatching asynchrony might be a strategy to improve reproductive output in terms of quality or quantity of offspring. However, little is known about the mediating effect of hatching asynchrony on offspring quality when brood reduction occurs. Here, we investigate the relationship between phenotypic quality and hatching asynchrony in Common Kestrel Falco tinnunculus nestlings in Spain. Hatching asynchrony did not increase breeding success or nestling quality. Furthermore, hatching asynchrony and brood reduction had different effects on nestlings’ phytohaematogglutinin (PHA)‐mediated immune response and nestling growth. In asynchronous and reduced broods (in which at least one nestling died), nestlings showed a stronger PHA‐mediated immune response and tended to have a smaller body size compared with nestlings raised in synchronous and reduced broods. When brood reduction occurred in broods hatched synchronously, there was no effect on nestling size, but nestlings had a relatively poor PHA‐mediated immune response compared with nestlings raised in asynchronous and reduced broods. We suggest that resources for growth can be directed to immune function only in asynchronously hatched broods, resulting in improved nestling quality, as suggested by their immune response. We also found that males produced a greater PHA‐mediated immune response than females only in brood‐reduced nests without any effect on nestling size or condition, suggesting that females may trade off immune activities and body condition, size or weight. Overall, our results suggest that hatching pattern and brood reduction may mediate resource allocation to different fitness traits. They also highlight that the resolution of immune‐related trade‐offs when brood reduction occurs may differ between male and female nestlings.     

What makes a host profitable? Parasites balance host nutritive resources against immunity

Numerous host qualities can modulate parasite fitness, and among these, host nutritive resources and immunity are of prime importance. Indeed, parasite fitness increases with the amount of nutritive resources extracted from the host body and decreases with host immune response. To maximize fitness, parasites have therefore to balance these two host components. Yet, because host nutritive resources and immunity both increase with host body condition, it is unclear whether parasites perform better on hosts in prime, intermediate, or poor condition. We investigated blood meal size and survival of the ectoparasitic louse fly Crataerina melbae in relation to body condition and cutaneous immune response of their Alpine swift (Apus melba) nestling hosts. Louse flies took a smaller blood meal and lived a shorter period of time when feeding on nestlings that were experimentally food deprived or had their cutaneous immune response boosted with methionine. Consistent with these results, louse fly survival was the highest when feeding on nonexperimental nestlings in intermediate body condition. Our findings emphasize that although hosts in poor condition had a reduced immunocompetence, parasites may have avoided them because individuals in poor condition did not provide adequate resources. These findings highlight the fact that giving host immunocompetence primary consideration can result in a biased appraisal of host-parasite interactions.     

Impacts of parasites in early life: contrasting effects on juvenile growth for different family members

Parasitism experienced early in ontogeny can have a major impact on host growth, development and future fitness, but whether siblings are affected equally by parasitism is poorly understood. In birds, hatching asynchrony induced by hormonal or behavioural mechanisms largely under parental control might predispose young to respond to infection in different ways. Here we show that parasites can have different consequences for offspring depending on their position in the family hierarchy. We experimentally treated European Shag (Phalacrocorax aristoteli) nestlings with the broad-spectrum anti-parasite drug ivermectin and compared their growth rates with nestlings from control broods. Average growth rates measured over the period of linear growth (10 days to 30 days of age) and survival did not differ for nestlings from treated and control broods. However, when considering individuals within broods, parasite treatment reversed the patterns of growth for individual family members: last-hatched nestlings grew significantly slower than their siblings in control nests but grew faster in treated nests. This was at the expense of their earlier-hatched brood-mates, who showed an overall growth rate reduction relative to last-hatched nestlings in treated nests. These results highlight the importance of exploring individual variation in the costs of infection and suggest that parasites could be a key factor modulating within-family dynamics, sibling competition and developmental trajectories from an early age.     

Ectoparasite load is linked to ontogeny and cell-mediated immunity in an avian host system with pronounced hatching asynchrony

Several contrasting hypotheses have been proposed to account for host age-biased parasite distribution, with some of them suggesting a key role of ectoparasites in the evolution and maintenance of weight hierarchies within broods. We examined parasite distribution among individual hosts across the whole period of host exposure to the parasite in a host system that shows distinct within-brood differences in age and age-related mortality. By contrast to previous hypotheses, we found that the abundance of a haematophagous, mobile ectoparasite Carnus haemapterus on nestling European rollers ( Coracias garrulus ) was highest approximately during the mid-nestling stage of their host, coinciding with the inflection point of the host growth phase. Parasite load increased neither with absolute resource availability (i.e. body size), nor body condition index. By contrast to previous evidence, higher parasite load under natural conditions was associated with a stronger cell-mediated immune response. However, this association was moderated by low parasite densities, as well as a better brood body condition index. Overall, although we revealed remarkable host ontogenetic effects on parasite distribution, the present study suggests that a highly mobile ectoparasite generally prefers healthier hosts. We propose that, in host systems with a marked asynchrony of hatching and background mortality within the brood, parasites favour persistence rather than nutritional attractiveness of the host.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 463–473.     

Relative importance of factors affecting nestling immune response differs between junior and senior nestlings within broods of hoopoes Upupa epops

Animals should invest in the immune system to protect themselves from parasites, but the cost of immune responses may limit investment depending on resource availability. In birds' broods, senior and junior chicks in size hierarchies face different rearing conditions, and thus we predicted that factors affecting immune response should differ between them. In asynchronously hatched hoopoe Upupa epops broods, we found that the immune response of senior nestlings was not related to their body condition, but positively related to risk of parasitism (which was indirectly estimated by laying date). This suggests that their immunocompetence is not limited by access to resources, and they can differentially invest in immune response with increasing risk of parasitism. On the other hand, immune response of junior nestlings was related to their body condition, but secondarily also to risk of parasitism. Our results agree with previous studies that have found significant influence of nutritional status and risk of parasitism on nestlings immune defence, but show that the effects of these environmental factors on nestling immunocompetence differ between nestlings occupying high and low rank positions in size hierarchies. The possible influence of maternal effects on the results found is also discussed.     

Behavioural responses to ectoparasites in pied flycatchers Ficedula hypoleuca: an experimental study

Nests of cavity‐nesting birds usually harbor some species of haematophagous ectoparasites that feed on the incubating adults and nestlings. Given the negative impact of ectoparasites on nestlings there will be selection on hosts to reduce parasite infestations through behavioural means. We have experimentally reduced the abundance of all ectoparasites in nests of pied flycatchers Ficedula hypoleuca to explore both whether there are changes in the frequency and duration of putative anti‐parasite behaviours by tending adults, as well as whether such anti‐parasite behaviours are able to compensate for the deleterious effects that parasites may have on nestlings. Heat treatment of nests substantially decreased the density of ectoparasites, and thereby positively affected nestling growth. The frequency and intensity of female grooming and nest sanitation behaviours during the incubation and nestling periods decreased as a consequence of the experimental reduction of ectoparasite infestation. Although nestlings begged more intensely in infested nests, the experiment had no significant effect on parental provisioning effort. Reduction of parasites resulted in larger nestlings shortly before fledging and increased fledging success. This study shows a clear effect of a complete natural nest ectoparasite fauna on parental behaviour at the nest and nestling growth in a cavity‐nesting bird. Although ectoparasites induce anti‐parasite behaviours in females, these behaviours are not able to fully remove parasite's deleterious effects on nestling growth and survival.     

Experimentally Constrained Virulence is Costly for Common Cuckoo Chicks

Chicks of some avian brood parasites show high virulence by eliminating all host progeny in the nest whereas others develop in the presence of host nestmates. Common cuckoo ( Cuculus canorus ) chicks are typically highly virulent parasites as they attempt to evict all host eggs and chicks soon after hatching. However, several features of nest design, including steep walls and/or cavity nests, may effectively prevent cuckoo hatchlings from evicting nestmates. A previous observational study showed low success of cuckoo chicks in evicting progeny of a cavity nester host, the redstart ( Phoenicurus phoenicurus ) but cuckoo chicks showed low survival both when reared alone or in mixed broods with host nestmates. Whether poor cuckoo performance was caused by eviction costs and/or by the effect of presence of host chicks per se remains unclear. We experimentally cancelled any potential eviction costs by removing host eggs immediately after the cuckoo hatched and creating mixed broods 5 days later when the eviction instinct of the cuckoo already ceased. Cuckoos that were forced to compete with host nestlings experienced lower provisioning rates, poorer growth, and lower fledging success than control lone cuckoos. Cuckoos in mixed broods that survived until fledging fledged later, and at lower masses, than those in the sole cuckoo group. Thus, the cuckoo gens specializing on redstarts is similar to other cuckoo gentes, whose chicks are more successful in evicting host nestmates, and it does not benefit from the presence of host brood. Cohabitation with host nestlings then should be viewed as a maladaptive by-product of host cavity nest design.     

Habitat-specific effects of a food supplementation experiment on immunocompetence in Eurasian Magpie Pica pica nestlings

Parasite pressure and nutrition are two of the most important factors affecting the trade-off between nestling growth and immune development. During development, energy and nutrients are often limited, and nestlings should only dedicate differentially more valuable resources to their immune system when the associated benefits are high (i.e. in situations of an increased risk of parasitism). In this study, we manipulated nutritional condition of Eurasian Magpie Pica pica nestlings by providing a food supplement. Additionally, the study area was subdivided into two categories of habitat (irrigated and arid) based on the presence of irrigation canals. Nestling diet composition was more varied in the irrigated compared with in the arid habitat. In addition, nestlings of the irrigated habitat showed a significantly higher infestation of both ectoparasites and blood parasites and a significantly higher cell-mediated immunity, but lower tarsus length compared with nestlings of the arid habitat. Food supplementation to nestlings did not affect tarsus length, but increased nestling cell-mediated immunity in the arid habitat only. Based on the recent demonstrated trade-off between growth and immunocompetence in nestlings, we suggest that differences between habitats in nestling diet and parasite prevalence may have caused different priority rules in the allocation of resources between both fitness traits.     

Low survival of parasite chicks may result from their imperfect adaptation to hosts rather than expression of defenses against parasitism

Host parents exhibit a variety of behaviors toward avian brood parasites, but not all of their actions have necessarily evolved in response to costs imposed by parasites. To investigate whether common waxbills (Estrilda astrild) have evolved defenses specifically against parasitic pin-tailed whydahs (Vidua macroura), I studied the specificity and flexibility of host behaviors toward nestlings at two sites that differed significantly in parasitism rates and intensities. I focused on documenting nestling survival because V. macroura young match the elaborate gape morphology of E. astrild nestlings, a pattern that suggests hosts may possess unique defenses against parasite chicks. Parasite young survived significantly worse than host young in mixed broods. However, this apparent discrimination was not associated with parasitism risk as would be expected if defenses had evolved specifically to counter parasitism. Parasite young may have survived poorly compared to host young because individual chicks were less able to stimulate sufficient care from foster parents or because they were more susceptible to nestling competition, disease, or reduced provisioning by hosts. Mortality may have also been exacerbated by poor timing of parasite egg laying. In nonparasitized and parasitized nests, rates of nestling survival were similar, further suggesting that parenting behaviors that result in chick mortality did not evolve solely in response to parasite young. In addition, orange-breasted waxbills (Amandava subflava) and zebra finches (Taeniopygia guttata), rarely parasitized and nonparasitized relatives of E. astrild, experience similar levels of nestling mortality presumably as a result of phylogenetically widespread parenting strategies. Despite the similarity of parasitic V. macroura nestlings and E. astrild nestlings, I found no evidence that E. astrild parents possess defenses that allow for specific discrimination against parasite chicks during the nestling period. Rather than being subject to host defenses evolved in an arms race, Vidua chicks may simply be imperfectly adapted to life in the nests of their hosts.     

Exposure to ectoparasites increases within-brood variability in size and body mass in the sand martin

Parasites often have detrimental effects on their hosts, and only host individuals able to cope with parasitism are likely to display induced or genetic resistance. Hosts may respond to parasitism by differential investment in offspring depending on their ability to cope with parasitism, because offspring that perform better than their siblings are themselves likely to have superior induced or genetic resistance. We tested whether nestlings of the highly colonial sand martin Riparia riparia were affected by the haematophagous tick Ixodes lividus by experimentally manipulating parasite loads of nests [nests sprayed with pyrethrum to remove parasites (sprayed), or nests sprayed with water (control)] at three stages of the breeding season. Prevalence and intensity of ticks were significantly affected by treatments. Breeding success was not significantly affected by treatment, although post-fledging survival was twice as high among nestlings from sprayed nests than from controls. Mean phenotypic traits of nestlings generally did not differ significantly among treatments, while within-brood variance in keel length (a skeletal character) and body mass were higher in control treatment broods than sprayed ones. Sedimentation rate, which reflects blood protein and immunoglobulin content, was significantly higher and less variable in sprayed than control broods. These findings are consistent with the suggestion that parasitism effects on host reproductive success act through an increase in the variance of offspring quality.     

Perceived risk of ectoparasitism reduces primary reproductive investment in tree swallows Tachycineta bicolor

Many birds have been shown to reduce their reproductive investment in response to infestation of nest sites by ectoparasites. Nest-dwelling parasite populations increase throughout the breeding season, and can reduce the condition and future survival of both breeding adults and their offspring. Thus, avian hosts should be capable of assessing early cues that predict future ectoparasitism risk, and should be able to facultatively adjust their primary reproductive investment in response to anticipated future costs of parasites. We tested this hypothesis in the tree swallow Tachycineta bicolor , a cavity nesting passerine, by presenting a visual cue of avian fleas on the outer surface of nest boxes. This treatment manipulated perceived ectoparasitism risk without exposing birds to parasites, thereby allowing us to examine facultative responses in the absence of early physiological effects of parasites on female reproductive investment. During one of the study years, birds preferentially occupied control boxes, however, across both years, birds nesting in treatment boxes produced significantly smaller clutches, resulting in smaller broods at hatching, relative to those in control boxes. This difference in clutch size could not be explained by differences in phenotypic quality of breeding birds, indicating that for cavity nesting birds such as tree swallows, the perception of future ectoparasitism risk may be sufficient to induce a facultative reduction in reproductive investment early in the breeding season, before nest-dwelling parasite populations have grown very large.