Hemosporidian blood parasites in seabirds—a comparative genetic study of species from Antarctic to tropical habitats

Whereas some bird species are heavily affected by blood parasites in the wild, others reportedly are not. Seabirds, in particular, are often free from blood parasites, even in the presence of potential vectors. By means of polymerase chain reaction, we amplified a DNA fragment from the cytochrome b gene to detect parasites of the genera Plasmodium, Leucocytozoon, and Haemoproteus in 14 seabird species, ranging from Antarctica to the tropical Indian Ocean. We did not detect parasites in 11 of these species, including one Antarctic, four subantarctic, two temperate, and four tropical species. On the other hand, two subantarctic species, thin-billed prions Pachyptila belcheri and dolphin gulls Larus scoresbii, were found infected. One of 28 thin-billed prions had a Plasmodium infection whose DNA sequence was identical to lineage P22 of Plasmodium relictum, and one of 20 dolphin gulls was infected with a Haemoproteus lineage which appears phylogenetically clustered with parasites species isolated from passeriform birds such as Haemoproteus lanii, Haemoproteus magnus, Haemoproteus fringillae, Haemoproteus sylvae, Haemoproteus payevskyi, and Haemoproteus belopolskyi. In addition, we found a high parasite prevalence in a single tropical species, the Christmas Island frigatebird Fregata andrewsi, where 56% of sampled adults were infected with Haemoproteus. The latter formed a monophyletic group that includes a Haemoproteus line from Eastern Asian black-tailed gulls Larus crassirostris. Our results are in agreement with those showing that (a) seabirds are poor in hemosporidians and (b) latitude could be a determining factor to predict the presence of hemosporidians in birds. However, further studies should explore the relative importance of extrinsic and intrinsic factors on parasite prevalence, in particular using phylogenetically controlled comparative analyses, systematic sampling and screening of vectors, and within-species comparisons.     

Deviance partitioning of host factors affecting parasitization in the European brown hare (Lepus europaeus)

Deviance partitioning can provide new insights into the ecology of host-parasite interactions. We studied the host-related factors influencing parasite prevalence, abundance, and species richness in European brown hares (Lepus europaeus) from northern Spain. We defined three groups of explanatory variables: host environment, host population, and individual factors. We hypothesised that parasite infection rates and species richness were determined by different host-related factors depending on the nature of the parasite (endo- or ectoparasite, direct or indirect life cycle). To assess the relative importance of these components, we used deviance partitioning, an innovative approach. The explained deviance (ED) was higher for parasite abundance models, followed by those of prevalence and then by species richness, suggesting that parasite abundance models may best describe the host factors influencing parasitization. Models for parasites with a direct life cycle yielded higher ED values than those for indirect life cycle ones. As a general trend, host individual factors explained the largest proportion of the ED, followed by host environmental factors and, finally, the interaction between host environmental and individual factors. Similar hierarchies were found for parasite prevalence, abundance, and species richness. Individual factors comprised the most relevant group of explanatory variables for both types of parasites. However, host environmental factors were also relevant in models for indirect life-cycle parasites. These findings are consistent with the idea of the host as the main habitat of the parasite; whereas, for indirect life-cycle parasites, transmission would be also modulated by environmental conditions. We suggest that parasitization can be used not only as an indicator of individual fitness but also as an indicator of environmental quality for the host. This research underlines the importance of monitoring parasite rates together with environmental, population, and host factors.     

Sex,age, spleen size,and kidney fat of red deer relative to infection intensities of the lungworm <Emphasis Type="Italic">Elaphostrongylus cervi</Emphasis>

We analyzed the relationships among spleen size, body condition (measured as kidney fat), and larval counts of the nematode Elaphostrongylus cervi in red deer (Cervus elaphus). The aim was to investigate the interaction between host body condition and intensity of infection with parasites. As red deer are highly polygynous, we also tested whether these relationships varied with sex and age of the hosts. Kidney fat and spleen size were positively correlated in subadults (2–3 years old) and adults (>3 years old), but not in calves (<1 year old) or yearlings (1–2 years old). Spleen size was negatively associated with nematode load in subadult females and in adult males. These two age classes are potentially the most nutritionally stressed, as subadult hinds are still growing and often engaging in rearing their first calf, and adult stags were sampled just after the rut, which is recognized as a substantial energy drain in this age–sex class, as they compete to hold females during the mating season. Body condition related negatively to parasite count only in adult males. In the context of red deer life history, these findings suggest that spleen size is dependent on body condition and that it could be affected by variation in resource partitioning among immune defense, growth, and reproductive effort in red deer. For the first time in a wild mammal, the spleen mass is shown to be positively related to body condition and negatively related to parasite infection. We conclude that elucidating whether spleen mass reflects immune defense investment or a measure of general body condition should contribute to understanding topical issues in mammal ecology.     

Social Hackers: Integration in the Host Chemical Recognition System by a Paper Wasp Social Parasite

 Obligate social parasites in the social insects have lost the worker caste and the ability to establish nests. As a result, parasites must usurp a host nest, overcome the host recognition system, and depend on the host workers to rear their offspring. We analysed cuticular hydrocarbon profiles of live parasite females of the paper wasp social parasite Polistes sulcifer before and after usurpation of host nests, using the non-destructive technique of solid-phase micro-extraction. Our results reveal that hydrocarbon profiles of parasites change after usurpation of host nests to match the cuticular profile of the host species. Chemical evidence further shows that the parasite queen changes the odour of the nest by the addition of a parasite-specific hydrocarbon. We discuss the possible role of this in the recognition and acceptance of the parasite and its offspring in the host colony. Received: 18 November 1999 / Accepted in revised form: 22 December 1999     

Colour change in a structural ornament is related to individual quality,parasites and mating patterns in the blue tit

Carry-over effects refer to processes that occur in one season and influence fitness in the following. In birds, two costly activities, namely reproduction and moult, are restricted to a small time window, and sometimes overlap. Thus, colour in newly moulted feathers is likely to be affected by the costs of reproduction. Using models of bird vision we investigated male colour change in a free-living population of blue tits (Cyanistes caeruleus) in three sampling occasions: spring 1, winter and spring 2. We related crown, tail, breast and cheek feather colouration after the moult (winter) to the intensity of infections by blood parasites during reproduction (spring 1). In the following spring (spring 2), we explored mating patterns with respect to changes in feather colour (springs 1 vs. 2). Males that were less intensely infected by the malaria parasite Plasmodium while breeding showed purer white cheek feathers in winter, which may indicate higher feather quality. Increased brightness in the white cheek was associated with better body condition during reproduction. In the following season, males with brighter cheeks paired with females that had noticeably brighter cheek patches compared to the male’s previous mate. These results suggest that the conditions experienced during reproduction are likely to affect moult and thus feather colouration, at least in the white patch. High quality individuals may allocate resources efficiently during reproduction increasing future reproductive success through variation in mating patterns. Carry-over effects from reproduction might extend not only to the non-breeding phase, but also to the following breeding season.     

Integrated pest management of Liriomyza trifolii: Influence of avermectin,cyromazine, and methomyl on leafminer ecology in celery

The impact of weekly applications of avermectin, cyromazine and methomyl on Lyriomyza trifolii (Burgess) and an associated complex of six parasite species was evaluated in celery, Apium graveolens L. Avermectin suppressed leafminer populations, but did not adversely affect (1) seasonal per cent parasitism, (2) adult parasite mortality, or (3) survival and emergence of immature parasites from treated foliage. Although cyromazine use also reduced L. trifolii density, significant reductionsin survival and emergence of immature parasites greatly diminished the potential for biological control by lowering the seasonal per cent parasitism and increasing the L. trifolii: parasite ratio. Methomyl application for control of lepidopterous larvae increased leafminer density and reduced the seasonal percent parasitism by reducing survival of adult parasites.Species composition of the parasite complex was least affected by avermectin and methomyl. Cyromazine significantly reduced the relative abundance of some key parasite species. Major parasite species included species in the family Eulophidae: Diglyphus intermedius (Girault), D. begini (Ashmead), Chrysonotomyia (Achrysocharella) punctiventris (Crawford), Chrysocharis parksi Crawford and C. ainsliei Crawford, and the family Pteromalidae: Halticoptera circulus (Walker). The potential for incorporation of these compounds into an integrated pest management program for celery and tomatoes, Lycopersicon esculentum Mill., is discussed.     

Carotenoids,immune response and the expression of sexual ornaments in male greenfinches (<Emphasis Type="Italic">Carduelis chloris</Emphasis>)

Allocation trade-offs of carotenoids between their use in the immune system and production of sexual ornaments have been suggested as a proximate mechanism maintaining honesty of sexual signals. To test this idea, we experimentally examined whether carotenoid availability in the diet was related to variation in antibody response to novel antigens in male greenfinches (Carduelis chloris aurantiiventris), a species with extensive carotenoid-dependent plumage colouration. We also measured the cost of mounting a humoral response in terms of circulating carotenoids. Finally, we examined the relationship between plumage colour, immune response and circulating carotenoids. We found that males with carotenoid-supplemented diets showed stronger antibody response than non-supplemented birds. We also found that activation of the immune system significantly reduced circulating carotenoids (24.9% lower in immune-challenged birds than in control birds). Finally, intensity (chroma) of ventral plumage colouration of males, a character directly related to concentration of total carotenoids in feathers, was negatively correlated with the immune response and circulating carotenoids in winter. These results support the idea that carotenoids are a limiting resource and that males trade ornamental colouration against immune response.     

Explaining postnatal growth plasticity in a generalist brood parasite

Selection of a particular host has clear consequences for the performance of avian brood parasites. Experimental studies showed that growth rate and fledging mass of brood parasites varied between host species independently of the original host species. Finding correlates of this phenotypic plasticity in growth is important for assessing adaptiveness and potential fitness consequences of host choice. Here, I analyzed the effects of several host characteristics on growth rate and fledging mass of the young of brown-headed cowbird (Molothrus ater), a generalist, non-evicting brood parasite. Cowbird chicks grew better in fast-developing host species and reached higher fledging mass in large hosts with fast postnatal development. A potential proximate mechanism linking fast growth and high fledging mass of cowbird with fast host development is superior food supply in fast-developing foster species. So far, we know very little about the consequences of the great plasticity in cowbird growth for later performance of the adult parasite. Thus, cowbird species could become interesting model systems for investigating the role of plasticity and optimization in the evolution of growth rate in birds.     

An acanthocephalan parasite increases the salinity tolerance of the freshwater amphipod <Emphasis Type="Italic">Gammarus roeseli</Emphasis> (Crustacea: Gammaridae)

Studies of the influence of parasites on host fitness generally conclude that parasites have a strong negative effect on their hosts. In this study, we have investigated experimentally the role of Polymorphus minutus, an acanthocephalan parasite, on the salinity tolerance of the freshwater amphipod Gammarus roeseli, one of its intermediate hosts. Unexpectedly, P. minutus-infected gammarids were more tolerant to salinity stress than uninfected ones. The mean lethal salt concentrations for 50% mortality of hosts tested were 17.3 (infected) and 9.7 g/L (uninfected). The parasitic load (one or two parasites per host) did not affect the result. The size of hosts had no significant influence on the salinity tolerance of either infected or uninfected gammarids. The mobility of all types of gammarid decreased when the salinity exceeded 9.0 g/L, but there was no significant difference between infected and uninfected gammarids. We discuss the higher salinity tolerance of infected amphipods in relation to O₂ consumption and osmoregulation. Finally, we demonstrate that the salinity tolerance is enhanced in the parasitized amphipod but without a significant change in behavior or an osmoregulatory adjustment.     

Effects of life-history traits on parasitism in a monogamous mammal, the eastern rock sengi (Elephantulus myurus)

The distribution of parasites is often characterised by substantial aggregation with a small proportion of hosts harbouring the majority of parasites. This pattern can be generated by abiotic and biotic factors that affect hosts and determine host exposure and susceptibility to parasites. Climate factors can change a host’s investment in life-history traits (e.g. growth, reproduction) generating temporal patterns of parasite aggregation. Similarly, host age may affect such investment. Furthermore, sex-biased parasitism is common among vertebrates and has been linked to sexual dimorphism in morphology, behaviour and physiology. Studies exploring sex-biased parasitism have been almost exclusively conducted on polygynous species where dimorphic traits are often correlated. We investigated the effects of season and life-history traits on tick loads of the monogamous eastern rock sengi (Elephantulus myurus). We found larger tick burdens during the non-breeding season possibly as a result of energetic constraints and/or climate effects on the tick. Reproductive investment resulted in increased larval abundance for females but not males and may be linked to sex-specific life-history strategies. The costs of reproduction could also explain the observed age effect with yearling individuals harbouring lower larval burdens than adults. Although adult males had the greatest larval tick loads, host sex appears to play a minor role in generating the observed parasite heterogeneities. Our study suggests that reproductive investment plays a major role for parasite patterns in the study species.     

Comparative morphology and secretion chemistry of the scoli in caterpillars of<Emphasis Type="Italic"> Hyalophora cecropia</Emphasis>

The morphology of the variously coloured scoli (bristle-bearing structures on the integument, producing an exocrine secretion) on caterpillars of Hyalophora cecropia and the secondary chemistry of the discharged secretions have been investigated for the first time and compared. According to our scanning electron microscopic study, the red/orange, yellow and blue coloured groups of these glands differ morphologically. Gas chromatographic-mass spectrometric analyses showed that the patterns of secondary compounds in the respective glandular secretions are also different. Furthermore, the secretion of the penultimate larval instar is chemically distinct from that of the last instar, as are both secretions from the respective haemolymph. The results favour the idea that the differences in scoli colour, morphology and chemistry could affect various predator species differently.Electronic Supplementary Material  Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

On the ecology of vectors

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Ornamental comb colour predicts T-cell-mediated immunity in male red grouse <Emphasis Type="Italic">Lagopus lagopus scoticus</Emphasis>

Sexual ornaments might reliably indicate the ability to cope with parasites and diseases, and a better ability to mount a primary inflammatory response to a novel challenge. Carotenoid-based ornaments are amongst the commonest sexual signals of birds and often influence mate choice. Because carotenoids are immuno-stimulants, signallers may trade-off allocating these to ornamental colouration or using them for immune responses, so carotenoid-based ornaments might be particularly useful as honest indicators of immuno-compentence. Tetraonid birds, such as the red grouse Lagopus lagopus scoticus, exhibit supra-orbital yellow-red combs, a conspicuous ornament which functions in intra- and inter-sexual selection. The colour of combs is due to epidermal pigmentation by carotenoids, while their size is testosterone-dependent. In this study, I investigated whether comb characteristics, and in particular, comb colour, indicated immuno-competence in free-living male red grouse. I assessed T-cell-mediated immunity using a standardised challenge with phytohaemagglutinin. Red grouse combs reflect in the red and in the ultraviolet spectrum of light, which is not visible to humans but that grouse most likely see, so I measured comb colour across the whole bird visible spectrum (300-700 nm) using a reflectance spectrometer. I found that males with bigger and redder combs, but with less ultraviolet reflectance, had greater T-cell-mediated immune response. Comb colour predicted T-cell-mediated immune response better than comb size, indicating that the carotenoid-based colouration of this ornament might reliably signal this aspect of male quality.     

Blue-green eggshell coloration is not a sexually selected signal of female quality in an open-nesting polygynous passerine

It has been proposed that blue-green egg colours have evolved as a post-mating signal of female quality, selected by males allocating their parental effort in response to the strength of this signal. We tested two main assumptions of the sexually selected egg coloration hypothesis: (1) whether the intensity of eggshell blue-green chroma (BGC) reflects female quality; and (2) whether males make their decisions on the level of parental care that they provide according to the intensity of eggshell BGC. As a model species, we chose the facultatively polygynous great reed warbler (Acrocephalus arundinaceus). In this species, females simultaneously paired with the same male, compete for his nest attendance and could benefit from signalling their quality through egg coloration. However, we found no association between the variation in eggshell BGC and the measures of female quality (physical condition, mean egg volume and age). Moreover, great reed warbler males did not adjust their investment (as measured in terms of nest defence against a brood parasite) in relation to the eggshell BGC. We conclude that blue-green egg coloration in this open-nesting passerine is unlikely to have a signalling function. Rather, the large colour variation among clutches of individual females may depend on yearly fluctuations in environmental conditions.     

Eggshell colour does not predict measures of maternal investment in eggs of <Emphasis Type="Italic">Turdus</Emphasis> thrushes

The striking diversity of avian eggshell colour has long fascinated biologists. Recently, it has been proposed that the blue-green colour of some eggs may function as a post-mating sexually selected signal of female phenotypic quality to their mates to induce higher allocation of paternal care. It has been suggested that maternally deposited yolk carotenoids may be the specific aspect of reproductive quality that the female is signalling via eggshell colour. We use the known properties of the thrush visual system (Turdus sp.) to calculate photon capture for the four single cone photoreceptors, and the principal member of the double cone class for eggs in clutches of two introduced European thrush species (Turdus merula and Turdus philomelos) in New Zealand. We show that differences in the avian-perceived colours of individual eggs are not consistently correlated with different measures of maternal investment in the egg. Given the growing extent of the knowledge between maternal quality, parental investment and eggshell pigmentation across avian taxa, we encourage the use of avian perceptual modelling for testing alternative non-signalling explanations for the structural and physiological basis of these relationships.     

Growth conditions affect carotenoid-based plumage coloration of great tit nestlings

 Carotenoid-based integument colour in animals has been hypothesised to signal individual phenotypic quality because it reliably reflects either foraging efficiency or health status. We investigated whether carotenoid-derived yellow plumage coloration of fledgling great tits (Parus major) reflects their nestling history. Great tit fledglings reared in a poor year (1998) or in the urban habitat were less yellow than these reared in a good year (1999) or in the forest. The origin of nestlings also affected their coloration since nestlings from a city population did not improve their coloration when transferred to the forest. Brood size manipulation affected fledgling colour, but only in the rural population, where nestlings from reduced broods developed more yellow coloration than nestlings from increased and control broods. Effect of brood size manipulation on fledgling plumage colour was independent of the body mass, indicating that growth environment affects fledgling body mass and plumage colour by different pathways. Received: 20 March 2000 / Accepted in revised form: 4 September 2000     

First evidence of regular common cuckoo, <Emphasis Type="Italic">Cuculus canorus</Emphasis>, parasitism on eastern olivaceous warblers, <Emphasis Type="Italic">Hippolais pallida elaeica</Emphasis>

Coevolution is defined as specialized relationships between species that lead to a reciprocal evolutionary change. A particularly suitable model system for studying coevolution is the interactions between obligate avian brood parasites and their hosts. The common cuckoo (Cuculus canorus, hereafter cuckoo) is a well-known brood parasite, which utilizes a range of smaller passerines as hosts. However, warblers of the genus Hippolais have rarely been reported as being victims of cuckoos, and furthermore, few data exist on the occurrence of antiparasite defenses in these hosts. In this study, we examined possible host–parasite coevolution between cuckoos and eastern olivaceous warblers (Hippolais pallida elaeica, hereafter olivaceous warblers) in three closely situated areas in northwestern Bulgaria. The olivaceous warbler has never been reported to be a regular cuckoo host. However, the present study, carried out in 2001–2003 shows that the olivaceous warbler is regularly and heavily parasitized by the cuckoo in this area. Parasitism rate was high (26.6%, 34/128) and consistent among years, with some variation between areas. The cuckoo egg mimicry was moderately good, and olivaceous warbler rejection rate of such eggs was 50%. Cuckoo eggs laid in olivaceous warbler nests had a whitish to whitish-green ground color, and the majority appeared to be distinctly different from cuckoo eggs found in other host species in the area. The olivaceous warbler proved to be a rather good host for cuckoos as 20.6% (7/34) of cuckoo eggs laid produced fledglings, a breeding success comparable to other suitable hosts in Europe. This is the first in-depth study of brood parasitism in a warbler of the genus Hippolais, and cuckoos parasitizing olivaceous warblers probably represent a previously unknown gens.     

营养盐对长江口2种主要赤潮原因藻光合色素和光合作用影响的比较研究

比较研究了不同营养条件对中肋骨条藻和东海原甲藻的光合色素含量和光合作用的影响.结果发现：①低浓度营养盐处理组培养的中肋骨条藻和东海原甲藻的叶绿素a （chlorophyll a,Chl-a）、叶绿素c （chlorophyll c,Chl-c）和总类胡萝卜素（total coloured carotenoids,TCC）含量均低于高浓度营养盐处理组;②初始N/P比值一定（16/1）的条件下, 2种藻的光合速率变化不同,对于中肋骨条藻,低浓度处理组以单位细胞表示的光合速率从第5 d开始显著低于高浓度处理组,至试验结束时, 2组的光合速率(以O₂计,下同)分别为0.031×10^-4 μmol·(cell·h)^-1和0.13×10^-4 μmol·(cell·h)^-1,以单位叶绿素a（μg）表示的光合速率与高浓度处理组之间差异不显著,分别为12.92 μmol·(μg·h)^-1和13.03 μmol·(μg·h)^-1,而对于东海原甲藻,低浓度处理组的以单位细胞和单位叶绿素a表示的光合速率均显著高于高浓度处理组;P源充足低N处理组的2种藻的单位细胞表示和以单位叶绿素a表示的光合速率均高于高N处理组;N源充足低P处理组的2种藻以单位细胞表示和以单位叶绿素a表示的光合速率低于高P处理组;2种藻的光合速率与胞内磷酸盐浓度之间存在显著正相关关系;③相同营养条件下东海原甲藻比中肋骨条藻含有更多的单位细胞光合色素含量和更高的光合速率.试验从2种藻的光合特性角度,进一步论证了东海原甲藻比中肋骨条藻更能适应营养盐浓度较低的环境.     

The parasite connection in ecosystems and macroevolution

In addition to their obvious negative effects (“pathogens”), endoparasites of various kinds play an important role in shaping and maintaining modern animal communities. In the long-term, parasites including pathogens are indispensable entities of any ecosystem. To understand this, it is essential that one changes the viewpoint from the host’s interests to that of the parasite. Together with geographic isolation, trophic arms race, symbiosis, and niche partitioning, all parasites (including balance strategists, i.e. seemingly non-pathogenic ones) modulate their hosts’ population densities. In addition, heteroxenic parasites control the balance between predator and prey species, particularly if final and intermediate hosts are vertebrates. Thereby, such parasites enhance the bonds in ecosystems and help maintain the status quo. As the links between eukaryotic parasites and their hosts are less flexible than trophic connections, parasite networks probably contributed to the observed stasis and incumbency of ecosystems over geologic time, in spite of continuous Darwinian innovation. Because heteroxenic parasites target taxonomic levels above that of the species (e.g. families), these taxa may have also become units of selection in global catastrophies. Macroevolutionary extrapolations, however, are difficult to verify because endoparasites cannot fossilize.     

Does contrast between eggshell ground and spot coloration affect egg rejection?

Obligate avian brood parasitic species impose the costs of incubating foreign eggs and raising young upon their unrelated hosts. The most common host defence is the rejection of parasitic eggs from the nest. Both egg colours and spot patterns influence egg rejection decisions in many host species, yet no studies have explicitly examined the role of variation in spot coloration. We studied the American robin Turdus migratorius, a blue-green unspotted egg-laying host of the brown-headed cowbird Molothrus ater, a brood parasite that lays non-mimetic spotted eggs. We examined host responses to model eggs with variable spot coloration against a constant robin-mimetic ground colour to identify patterns of rejection associated with perceived contrast between spot and ground colours. By using avian visual modelling, we found that robins were more likely to reject eggs whose spots had greater chromatic (hue) but not achromatic (brightness) contrast. Therefore, egg rejection decision rules in the American robin may depend on the colour contrast between parasite eggshell spot and host ground coloration. Our study also suggests that egg recognition in relation to spot coloration, like ground colour recognition, is tuned to the natural variation of avian eggshell spot colours but not to unnatural spot colours.