The role of host sex in parasite dynamics: field experiments on the yellow-necked mouse Apodemus flavicollis

We investigated the role of host sex in parasite transmission and questioned: ‘Is host sex important in influencing the dynamics of infection in free living animal populations?’ We experimentally reduced the helminth community of either males or females in a yellow‐necked mice (Apodemus flavicollis) population using an anthelmintic, in replicated trapping areas, and subsequently monitored the prevalence and intensity of macroparasites in the untreated sex. We focussed on the dominant parasite Heligmosomoides polygyrus and found that reducing parasites in males caused a consistent reduction of parasitic intensity in females, estimated through faecal egg counts, but the removal of parasites in females had no significant influence on the parasites in males. This finding suggests that males are responsible for driving the parasite infection in the host population and females may play a relatively trivial role. The possible mechanisms promoting such patterns are discussed.     

Sharing roosts but not ectoparasites: high host-specificity in bat flies and wing mites of Miniopterus schreibersii and Rhinolophus ferrumequinum (Mammalia: Chiroptera)

Schreiber’s bent-winged bat Miniopterus schreibersii and the greater horseshoe bat Rhinolophus ferrumequinum are widespread and common cavernicolous species across southern Europe that host numerous specialized ectoparasite species. The objective of this study was to characterize the species assemblage, genetic diversity, and host specificity of bat flies (Nycteribiidae, Diptera) and wing mites (Spinturnicidae, Acari) found on these bat hosts in Serbia and Bosnia and Herzegovina. Notably, while bat flies lay puparia on the cave walls and can thus be transmitted indirectly, wing mites require direct body contact for transmission. Morphological identification and sequencing of a 710-bp fragment of cytochrome oxidase I gene of 207 bat flies yielded 4 species, 3 on M. schreibersii and 1 on R. ferrumequinum. Sequencing of a 460-bp small subunit ribosomal RNA fragment, in all 190 collected wing mites revealed 2 species, 1 per host. In no case was a parasite associated with 1 host found on the other host. Species and genetic diversity of flies were higher in M. schreibersii, likely reflecting their host’s larger colony sizes and migratory potential. Mite species of both hosts showed similarly low diversity, likely due to their faster life history and lower winter survival. Our findings highlight a remarkably high host-specificity and segregation of ectoparasite species despite direct contact among their hosts in the roost, suggesting a defined host preference in the investigated ectoparasite species. Furthermore, the differences in ectoparasite genetic diversity exemplify the interplay between host and parasite life histories in shaping parasite population genetic structure.     

Sexual selection and infection by ectoparasites in Wellington tree weta,Hemideina crassidens (Orthoptera: Anostostomatidae)

Abstract Sexual selection can affect the prevalence and intensity of infection of individuals by ectoparasitic mites. According to this theory, males should exhibit greater infection by parasites than females and juveniles should be less infected than adults. In the wild, I investigated whether prevalence and intensity of the chyzeriid mite, Nothotrombicula deinacridae (Dumbleton) differed between the sexes and between developmental stages in Wellington tree weta, Hemideina crassidens. Despite being under strong sexual selection, male tree weta did not exhibit greater parasitism and there was some evidence that adults and juveniles differed in prevalence. The sexual selection hypothesis was not supported in this study.     

Host nest preference and nest choice in the cuckoo paper wasppolistes sulcifer (Hymenoptera: Vespidae)

Polistes sulcifer is a cuckoo paper wasps, an obligate social parasite which must usurp a colony of another species in order to reproduce. Field data show thatP. sulcifer females usurp exclusively nests belonging to one species only (P. dominulus). Moreover, they are more frequently found on large and puparich nests. A series of laboratory trials, in which parasite females were offered a binary choice of nests with different characteristics, confirm both the species specificity betweenP. sulcifer andP. dominulus and the parasite's preference to usurp larger nests containing pupae. The data support the hypothesis thatP. sulcifer females choose between available nests. The biological meaning of these findings is discussed.     

Co-evolution between ectoparasites and their insect hosts: a simulation study of a damselfly–water mite interaction

1. A simulation model investigating the co‐evolution of water mites infesting their aquatic insect hosts during emergence is presented. The model is based on field and experimental studies of the ectoparasitic water mite Arrenurus cuspidator and the damselfly Coenagrion puella. 2. Three scenarios were studied: (1) Only the host was allowed to evolve timing of emergence, while the timing of the parasites' infestation opportunity was held constant. (2) Both host and parasite were allowed to evolve. (3) Only the parasite's timing was allowed to evolve, while the host was constrained completely. 3. In the first two scenarios, parasite abundances decreased in the course of evolution and reached values well below those found in the field, whereas in the third scenario, parasite abundances were maintained at a level close to that found in the field. In the second scenario (co‐evolution), the host seemed to be the leader in the evolutionary race. 4. It is concluded that water mite parasitism is capable of shaping emergence patterns in aquatic insects and, despite the same life‐cycle length for host and parasite, the parasite evolves fast enough to shape its hatching pattern to match the emergence pattern of its host.     

Facultative pupal mating in Heliconius erato: Implications for mate choice,female preference,and speciation

Mating systems have broad impacts on how sexual selection and mate choice operate within a species, but studies of mating behavior in the laboratory may not reflect how these processes occur in the wild. Here, we examined the mating behavior of the neotropical butterfly Heliconius erato in the field by releasing larvae and virgin females and observing how they mated. H. erato is considered a pupal‐mating species (i.e., males mate with females as they emerge from the pupal case). However, we observed only two teneral mating events, and experimentally released virgins were almost all mated upon recapture. Our study confirms the presence of some pupal‐mating behavior in H. erato, but suggests that adult mating is likely the prevalent mating strategy in this species. These findings have important implications for the role of color pattern and female mate choice in the generation of reproductive isolation in this diverse genus.     

Host preference by Allothrombium pulvinum (Acari: Trombidiidae) larvae on aphids: Macrosiphum rosae,Aphis gossypii and Hyalopterus amygdali (Homoptera: Aphididae)

In this study aphid-plant association and its effect on host preference of parasitic Allothrombium pulvinum larvae was examined with multiple-choice tests. Host species selection, host size selection and superparasitism with mite larvae were studied with two-choice tests. Three aphid species were used: Macrosiphum rosae, Aphis gossypii and Hyalopterus amygdali. In multiple-choice tests, larvae of A. pulvinum showed no significant preference for any aphid-plant association when given M. rosae on rose, A. gossypii on cucumber and H. amygdali on apricot simultaneously. Two-choice tests showed that larval mites preferred H. amygdali to A. gossypii, but had no preference when offered a choice between A. gossypii and M. rosae or between H. amygdali and M. rosae. In host size selection and superparasitism tests, significantly more mites selected the larger host (M. rosae). Furthermore, parasitised H. amygdali were preferred to unparasitised ones. This revised version was published online in July 2006 with corrections to the Cover Date.     

Origin-related, environmental, sex, and age determinants of immunocompetence, susceptibility to ectoparasites, and disease symptoms in the barn owl

Knowledge of the role of origin‐related, environmental, sex, and age factors on host defence mechanisms is important to understand variation in parasite intensity. Because alternative components of parasite defence may be differently sensitive to various factors, they may not necessarily covary. Many components should therefore be considered to tackle the evolution of host–parasite interactions. In a population of barn owls (Tyto alba), we investigated the role of origin‐related, environmental (i.e. year, season, nest of rearing, and body condition), sex, and age factors on 12 traits linked to immune responses [humoral immune responses towards sheep red blood cells (SRBC), human serum albumin (HSA) and toxoid toxin TT, T‐cell mediated immune response towards the mitogen phytohemagglutinin (PHA)], susceptibility to ectoparasites (number and fecundity of Carnus haemapterus, number of Ixodes ricinus), and disease symptoms (size of the bursa of Fabricius and spleen, proportion of proteins that are immunoglobulins, haematocrit and blood concentration in leucocytes). Cross‐fostering experiments allowed us to detect a heritable component of variation in only four out of nine immune and parasitic parameters (i.e. SRBC‐ and HSA‐responses, haematocrit, and number of C. haemapterus). However, because nestlings were not always cross‐fostered just after hatching, the finding that 44% of the immune and parasitic parameters were heritable is probably an overestimation. These experiments also showed that five out of these nine parameters were sensitive to the nest environment (i.e. SRBC‐ and PHA‐responses, number of C. haemapterus, haematocrit and blood concentration in leucocytes). Female nestlings were more infested by the blood‐sucking fly C. haemapterus than their male nestmates, and their blood was less concentrated in leucocytes. The effect of year, season, age (i.e. reflecting the degree of maturation of the immune system), brood size, position in the within‐brood age hierarchy, and body mass strongly differed between the 12 parameters. Different components of host defence mechanisms are therefore not equally heritable and sensitive to environmental, sex, and age factors, potentially explaining why most of these components did not covary. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 703–718.     

The effect of Jacobin Cuckoo Clamator jacobinus parasitism on the body mass and survival of young in a new host species

The southern African subspecies of Jacobin Cuckoo Clamator jacobinus serratus is a brood parasite of a range of host species. While Jacobin Cuckoos do not evict host young, previous research has found that host young rarely survive the nestling period. Here we provide the first records of Jacobin Cuckoo parasitism of a new host species, the Southern Pied Babbler Turdoides bicolor. We investigate rates of brood parasitism and the survival of host young. The Southern Pied Babbler is one of the largest recorded hosts for Jacobin Cuckoos and, unusually, we find that host young tend to survive the nestling period and maintain similar body mass to host young in unparasitized broods. However, host young were less likely to survive to independence than young raised in unparasitized nests, suggesting a post‐fledging reproductive cost to hosts.     

Host use of a hemiparasitic plant: no trade-offs in performance on different hosts

To examine putative specialization of a hemiparasitic plant to the most beneficial host species, we studied genetic variation in performance and trade-offs between performance on different host species in the generalist hemiparasite, Rhinanthus serotinus. We grew 25 maternal half-sib families of the parasite on Agrostis capillaris and Trifolium pratense and without a host in a greenhouse. Biomass and number of flowers of the parasite were the highest when grown on T. pratense. There were significant interactions between host species and R. serotinus seed-family indicating that the differences in performance on the two hosts and without a host varied among the families. However, we found no significant negative correlations between performance of R. serotinus on the host species or between performance on the two hosts and autotrophic performance. Thus, the genetic factors studied here are not likely to affect the evolution of specialization of R. serotinus to the most beneficial host.     

Infestation and seasonal activity of Ixodes vespertilionis Koch, 1844 (Acari: Ixodidae) on the Maghreb mouse‐eared bat,Myotis punicus Felten, 1977, in northeastern Algeria

Infestation of Ixodes vespertilionis Koch, 1844 on Myotis punicus Felten, 1977 from two sites (Trios Tunnel and Sidi Trad cave) in northeastern Algeria was studied. An overall infestation of 41.4% for all stages was found among bats collected from both sites. By stage, a total of eight females, 70 nymphs, and 107 larvae were recovered from both populations. The number of females recovered per bat at Sidi Trad ranged from 0‐1, for nymphs 0‐2, and for larvae 0‐2. While no female ticks were collected at Trios Tunnel, the number of nymphs ranged from 0‐2 and for larvae 0‐2. At Trios Tunnel, the number of nymphs was significantly higher during April and June but not for July and September. On the other hand, the number of larvae increased from July to November, while at Sidi Trad cave, female ticks were recovered during April and May and then disappeared until the end of the study period. Significant differences were noted during all the months when compared with all stages. Nymphs infested bats significantly during April and May, declined in June and July, and then became steady until October. Larvae peaked in July, with low frequency in April, and then fluctuated from August to November.     

Cuckoos, cowbirds and hosts: adaptations, trade-offs and constraints

The interactions between brood parasitic birds and their host species provide one of the best model systems for coevolution. Despite being intensively studied, the parasite-host system provides ample opportunities to test new predictions from both coevolutionary theory as well as life-history theory in general. I identify four main areas that might be especially fruitful: cuckoo female gentes as alternative reproductive strategies, non-random and nonlinear risks of brood parasitism for host individuals, host parental quality and targeted brood parasitism, and differences and similarities between predation risk and parasitism risk. Rather than being a rare and intriguing system to study coevolutionary processes, I believe that avian brood parasites and their hosts are much more important as extreme cases in the evolution of life-history strategies. They provide unique examples of trade-offs and situations where constraints are either completely removed or particularly severe.     

Microhabitat distributions and species interactions of ectoparasites on the gills of cichlid fish in Lake Victoria,Tanzania

Heterogeneous exposure to parasites may contribute to host species differentiation. Hosts often harbour multiple parasite species which may interact and thus modify each other’s effects on host fitness. Antagonistic or synergistic interactions between parasites may be detectable as niche segregation within hosts. Consequently, the within-host distribution of different parasite taxa may constitute an important axis of infection variation among host populations and species. We investigated the microhabitat distributions and species interactions of gill parasites (four genera) infecting 14 sympatric cichlid species in Lake Victoria, Tanzania. We found that the two most abundant ectoparasite genera (the monogenean Cichlidogyrus spp. and the copepod Lamproglena monodi) were non-randomly distributed across the host gills and their spatial distribution differed between host species. This may indicate microhabitat selection by the parasites and cryptic differences in the host–parasite interaction among host species. Relationships among ectoparasite genera were synergistic: the abundances of Cichlidogyrus spp. and the copepods L. monodi and Ergasilus lamellifer tended to be positively correlated. In contrast, relationships among morphospecies of Cichlidogyrus were antagonistic: the abundances of morphospecies were negatively correlated. Together with niche overlap, this suggests competition among morphospecies of Cichlidogyrus. We also assessed the reproductive activity of the copepod species (the proportion of individuals carrying egg clutches), as it may be affected by the presence of other parasites and provide another indicator of the species specificity of the host–parasite relationship. Copepod reproductive activity did not differ between host species and was not associated with the presence or abundance of other parasites, suggesting that these are generalist parasites, thriving in all cichlid species examined from Lake Victoria.     

Social competition, corticosterone and survival in female lizard morphs

We examined the selective consequences of variation in behaviour and endocrine physiology in two female throat-colour morphs of the lizard, Uta stansburiana in the wild. Female morphs differed in home-range distribution patterns and corticosterone levels in relation to the density and frequency of their female neighbours. Levels of plasma corticosterone of yellow-throated females increased with increased density of both morphs. In contrast, orange-throated females had reduced levels of corticosterone in response to increased density of orange females. Additionally, females with lower corticosterone survived poorly, suggesting that social interactions and high local densities of orange females may be potentially costly for orange females. These results are consistent with decreased fitness effects and suppression of immune function previously reported for orange female morphs surrounded by more orange neighbours. These correlations, in conjunction with previous work in this system, indicate that corticosterone is likely to be an important physiological mechanism regulating female fitness in nature.     

Host preference and survival in selected lines of a Drosophila parasitoid,Asobara tabida

Host selection behaviour of parasitoids has important fitness consequences, if hosts of different quality are available. Here the host selection behaviour, the ability to distinguish between hosts differing in their suitability, of the Drosophila parasitizing wasp Asobara tabida was studied. Females from five lines selected for higher survival in the encapsulating host species D. melanogaster ( 16 ) were compared with females from control lines. Females from all five selected lines more readily accepted the encapsulating host species for oviposition when offered together with a nonencapsulating host species than females from the control lines. We found no evidence for pleiotropic effects and suggest that host selection behaviour evolved parallel to the ability to escape encapsulation in the hosts. Our results also suggest that given the appropriate selection pressures, host selection behaviour can quickly evolve in parasitoids, enabling them to adapt fast to changing circumstances.     

Seasonal abundance of Allothrombium monochaetum and Allothrombium pulvinum in Navarra-Nafarroa (northern Spain), with notes on larval host preference and rate of parasitism

In this paper we report on the seasonal abundance of velvet mite larvae of Allothrombium pulvinum Ewing and Allothrombium monochaetum Goldarazena and Zhang in a meadow located in the mediterranean Navarra-Nafarroa region (northern Spain). Larvae of A. monochaetum peaked in numbers on Aphis fabae in mid-June while larvae of A. pulvinum peaked in the third week of July. Both species preferred the aphid thorax as attachment site. We also provide a list of aphid hosts of both mites and observed density of parasites on the hosts.     

Reproduction in captive lion tamarins (Leontopithecus): Seasonality,infant survival,and sex ratios

Diversity in reproductive and social systems characterizes the primate family Callitrichidae. This paper contributes to our appreciation of this diversity by presenting the first detailed comparative analysis of captive breeding in three species of lion tamarins (Leontopithecus chrysomelas, L. chrysopygus, and L. rosalia) housed at the Centro de Primatologia do Rio de Janeiro. The annual pattern of reproduction in all three species of Leontopithecus was markedly seasonal, with births occurring during the spring, summer, and fall months from August through March. While modal number of litters produced per female per year was 1, approximately 20% of breeding females produced two litters per year. The onset of breeding activity in years when two litters are produced was significantly earlier than in years when only one litter was produced. The cumulative number of offspring surviving to 3 months of age did not differ between years with one vs. two breeding attempts. Like other callitrichids, postnatal mortality was highest during the first week of life, and there were pronounced species differences in offspring survival through 1 year, with significantly lower survivorship in L. chrysomelas. Infant survivorship was affected by a number of experiential factors. Survivorship up to 30 days of life was higher in groups in which the breeding female had previous experience with infants as a nonbreeding helper than in groups in which the female lacked previous helping experience. Likewise, survivorship to 30 days of life was higher for infants born to multiparous females than for infants born to primiparous females. When parity and previous helping experience were analyzed concurrently, the lowest survivorship was associated with offspring produced by inexperienced primiparous females. Genus-wide, there was no significant departure from a 50:50 sex ratio at any point during the first year of life, nor was there evidence for differential mortality for male and female infants. However, L. chrysopygus produced significantly more male infants at birth (65:44) and had male-biased litters (approximately 60% males) throughout the first year of life, while L. chrysomelas showed a nonsignificant tendency toward female-biased litters. © 1996 Wiley-Liss, Inc.