Relationship between host diversity and parasite diversity: flea assemblages on small mammals

Aim We examined the relationship between host species richness and parasite species richness using simultaneously collected data on small mammals (Insectivora, Rodentia and Lagomorpha) and their flea parasites. Location The study used previously published data on small mammals and their fleas from 37 different regions. All the world's main geographical regions other than Australasia and Wallacea were represented in the study, i.e. neotropical, nearctic, palaearctic, oriental and afrotropical realms. Methods We controlled the data for the area sampled and sampling effort and then tested this relationship using both cross‐region conventional analysis and the independent contrasts method (to control for the effects of biogeographic historical relationships among different regions). Brooks parsimony analysis was used to construct a region cladogram based on the presence/absence of a host species and host phylogeny. Results Both cross‐region and independent contrasts analyses showed a positive correlation between host species richness and flea species richness. Conventional cross‐region regression under‐ or overestimated fleas species richness in the majority of regions. Main conclusions When the regression derived by the independent contrasts method was mapped onto the original tip data space, points that deviated significantly from the regression originated from Kenya, Mississippi and southern California (lower than expected flea richness) and Chile, Idaho, south‐western California and Kyrgyzstan (higher than expected flea richness). These deviations can be explained by the environmental mediation of host–flea relationships and by a degree of environmental variety in sampled areas.     

Ectoparasite loads decrease the fitness of alpine marmots (Marmota marmota) but are not a cost of sociality

Parasites arc often considered an unavoidable cost of livingin groups. We examined this hypothesis by investigating theectoparasite fauna of a free-living alpine marmot population,a highly social sciurid species. Only a single important ectoparasitewas found, the mite Echinonyssus blanchardi. Heavy infestationwith this mite correlated with reproductive success of marmotsin two ways. Infant winter mortality increased with ectoparasiteload, probably because ectoparasites reduced the ability ofparents and alloparents to warm infants during hibernation.Litters produced by females exposed to a high number of mitesduring the preceding winter were weaned later. Late weanershad a lower chance of surviving hibernation. However, we foundno relationship between ectoparasite load and group size ormarmot density. Instead, the number of E. blanchardi per animalwas lower in areas where marmot groups were clumped, i.e., infavorable habitats. Although ectoparasites most likely decreasethe fitess of alpine marmots to some extent, they should notbe considered a cost of social life in this species.     

Contrasting responses of beta diversity components to environmental and host-associated factors in insect ectoparasites

1. The present study investigated whether different components (species replacement and species gains/losses) of compositional and phylogenetic beta diversity of insect ectoparasites responded similarly to environmental and host-associated gradients using a large dataset on distribution of fleas and their rodent hosts in Mongolia. 2. Generalised dissimilarity modelling was applied to investigate whether environmental variables or host dissimilarity was the best predictor of species/lineage replacement and species/lineage gains/losses (= richness difference) components of compositional and phylogenetic flea beta diversity. 3. The total compositional beta diversity of fleas was influenced mainly by the gradient in air temperature and, to a lesser degree, by total host beta diversity, with the former effect being associated with the richness difference component and the latter effect being associated with species replacement component. The total phylogenetic beta diversity of fleas was best explained by the total phylogenetic beta diversity of hosts, with this effect being mainly associated with the lineage replacement component, whereas the lineage richness difference component responded mainly to the temperature gradient. 4. The results of the present indicate that not only multiple beta diversity facets are driven by different factors, but also different components of the same beta diversity facet respond to different environmental (for parasites, including host-associated) gradients. These patterns were masked when only total beta diversity was analysed. 5. This emphasizes the importance of considering the components of insect beta diversity separately. Ignoring the separate components of beta diversity can lead to potentially erroneous inferences about the relative contribution of abiotic and biotic effects on beta diversity.     

Geographical variation in the 'bottom-up' control of diversity: fleas and their small mammalian hosts

Aim We searched for signs of the ‘bottom‐up’ diversity effect in the association between fleas (Siphonaptera) and their small mammalian hosts (Rodentia, Insectivora and Lagomorpha). We asked (1) whether a strong dependence of flea species richness on host species richness is characteristic for both Palaeoarctic and Nearctic realms; (2) if yes, whether the ratio of host species per flea species along the host diversity gradient is similar between the Palaeoarctic and Nearctic; and (3) whether factors other than host species richness (i.e. geographical position, climate and landscape) might better explain variation in flea species richness than host species richness. Location The study used previously published data on species richness of fleas and their small mammalian hosts from 26 Palaeoarctic and 19 Nearctic regions. Methods We regressed the number of flea species on the number of small mammal species across regions, separately for Palaeoarctic and Nearctic realms, using both non‐transformed data as well as data corrected for the confounding effects of host sampling effort and sampling area. To test whether flea species richness is determined by external factors unrelated to the host, we used stepwise multiple regressions of flea species richness against host species richness and parameters describing the geographical position, climate and relief of a region. Results When non‐transformed data were analysed, flea species richness was positively correlated with host species richness in both the Palaeoarctic and Nearctic, although the slopes of the two regressions differed significantly. After removal of the confounding effects of host sampling effort and sampling area, Palaeoarctic flea species richness remained strongly positively correlated with host species richness, whereas in the Nearctic, flea species richness appeared to be completely independent of host species richness. Results of the multiple regressions using corrected data demonstrated that in the Palaeoarctic, flea species richness was correlated with both the number of host species and the mean altitude of the region, whereas in the Nearctic, flea species richness only tended to be weakly correlated with latitude (however, this correlation turned out to be non‐significant after Bonferroni correction). Main conclusions We found evidence of bottom‐up control of flea diversity in the Palaeoarctic regions only, and not in the Nearctic. We explore several potential explanations for the different patterns observed in the two biogeographical realms, including differences in (1) levels of host specialization, (2) history of host–parasite associations and (3) landscape effects on flea diversification. We conclude that these factors combine to create different macroecological patterns in different biogeographical realms, and that diversity is not governed by the same forces everywhere.     

Latitudinal gradients of parasite species richness in primates

Infectious disease risk is thought to increase in the tropics, but little is known about latitudinal gradients of parasite diversity. We used a comparative data set encompassing 330 parasite species reported from 119 primate hosts to examine latitudinal gradients in the diversity of micro and macroparasites per primate host species. Analyses conducted with and without controlling for host phylogeny showed that parasite species richness increased closer to the equator for protozoan parasites, but not for viruses or helminths. Relative to other major parasite groups, protozoa reported from wild primates were transmitted disproportionately by arthropod vectors. Within the protozoa, our results revealed that vector‐borne parasites showed a highly significant latitudinal gradient in species richness. This higher diversity of vector‐borne protozoa near the tropics could be influenced by a greater abundance or diversity of biting arthropods in the tropics, or by climatic effects on vector behaviour and parasite development. Many vector‐borne diseases, such as leishmaniasis, trypanosomiasis, and malaria pose risks to both humans and wildlife, and nearly one‐third of the protozoan parasites from free‐living primates in our data set have been reported to infect humans. Because the geographical distribution and prevalence of many vector‐borne parasites are expected to increase because of global warming, these results are important for predicting future parasite‐mediated threats to biodiversity and human health.     

外来种入侵与物种多样性

在入侵生态学研究方面 ,物种多样性与生物入侵之间的关系已成为当前研究和争论的焦点。自Elton的经典假说提出以来 ,物种丰富度高的群落比物种贫乏的群落更能抵抗外来种入侵的观点得到广泛接受。一些理论模型和多样性处理实验支持了该假说。但现在越来越多的野外观测和实验研究开始对这一经典假说提出异议 ,甚至反对。同时 ,在入侵生态学广泛受到关注的今天 ,大量的实验研究也提出了一些新的观点。本文对Elton经典假说提出以来全球有关物种多样性与生物入侵关系的主要研究及其观点进行了评述 ,以期为我国有关研究工作的开展提供参考。     

Helminth parasite richness among vertebrates

The numbers of intestinal helminth species (parasite richnesS) recorded from each of 488 vertebrate host species are compared using data compiled from the published literature. Associations between parasite richness, sampling effort, host size and host habitat (aquatic versus terrestrial) are assessed using a method designed to control for phylogenetic association. Parasite richness increases with the number of surveys on which each estimate of parasite richness is based (sampling effort). When the effects of sampling effort are controlled for, there remains a strong positive relationship between parasite richness and host body size. There is no tendency for aquatic hosts to harbour more parasite species than terrestrial hosts independently of differences in sampling effort and body size. The results are interpreted in the context of hosts representing habitats for parasite colonization, resource allocation between parasite species, and the age of the major mammalian radiations.     

Elevation gradients of diversity for rodents and bats in Oaxaca, Mexico

1  This study documents patterns of rodent and bat diversity related to abiotic and biotic factors along elevational gradients in the Sierra Mazateca (640–2600 m a.s.l.) and Sierra Mixteca (700–3000 m a.s.l.) in Oaxaca, Mexico.
2  The two transects share similar faunas: 17 and 23 rodent species were captured in the sierras Mazateca and Mixteca, respectively, 14 of which occurred on both transects. Rodent species richness was similar in the wet season and the dry season along both transects. Rodent species richness peaked at 1025–1050 m in tropical semi-deciduous forest on both transects. Endemic species were restricted to high-elevation habitats.
3  Sixteen and 17 bat species were captured in the sierras Mazateca and Mixteca, respectively; 11 occurred on both transects. Bat species richness was higher in the wet season than in the dry season in the Sierra Mazateca. Bat species richness peaked at 1850 m in pine–oak forest in the Sierra Mazateca, and at 750 m and 1050 m in tropical semi-deciduous forest in the Sierra Mixteca, decreasing abruptly at higher elevations on both transects.
4  Patterns of trophic diversity of rodents and bats coincided with those of species richness on each transect. Species richness increased with increasing habitat diversity; increased with increasing rainfall and productivity; increased with increasing resource diversity; and increased in areas with high rates of speciation (rodents only).
5  The need for conservation action in Oaxaca is urgent and proponents should promote establishment of protected areas linking lowland habitats with high species richness to high-elevation habitats harbouring large numbers of endemic forms.     

Impact of municipal effluents and hydrological regime on myxozoan parasite communities of fish

Increased productivity from sewage effluents can enhance species richness locally. Results from a study of spottail shiners (Notropis hudsonius) in 1999 showed that prevalence and the mean number of myxozoan parasite species per host were higher downstream of the wastewater outflow from the Island of Montreal than upstream in the St. Lawrence River, Quebec, Canada. This was attributed to organic enrichment of the sediments which presumably lead to increased densities of oligochaetes, the alternate hosts, downstream of Montreal. Spottail shiners subsequently were collected every August/early September in 2001–2004 to examine the stability and repeatability of these patterns. Prevalence and mean number of myxozoan species per fish typically were again higher downstream of the sewage source each year compared to upstream, although there was no significant difference in these measurements pooled across years between localities immediately upstream and downstream of the effluent outflow. Density of the oligochaete Limnodrilus hoffmeistereri, a common alternate host of myxozoans, was much higher at two downstream localities than at an upstream one. At a larger spatial scale, mean myxozoan infracommunity richness across sites in the St. Lawrence River was negatively correlated with mean water levels measured in the 3 months prior to fish sampling. Results suggest that on a local scale, variations in prevalence and diversity among localities are influenced by municipal effluents, but that at a landscape scale annual variations across sites are affected by the hydrological regime and climate. In effect, water level fluctuation had a landscape-wide impact that was superimposed over pollution-induced local variations.     

Host ecology and life-history traits associated with blood parasite species richness in birds

Identifying host traits associated with the number of different parasite species or strains harboured by a particular host species can have important implications for understanding the impact of parasitism on hosts. We investigated associations between host ecology and life history, and parasite richness and prevalence of the four major avian blood parasite genera. We used an extensive data on blood parasite infections and host ecology in 263 bird species from the Western Palearctic, combining species-specific data with a comparative approach to control for similarity in phenotype among host species due to the effects of common phylogenetic descent. Adult survival rate negatively correlated with the number of parasite species infecting a host species when controlling for similarity due to common descent and body mass. In addition, the prevalence of Haemoproteus, Plasmodium and Leucocytozoon was higher in species harbouring a richer parasite assemblage. These results suggest that the impact on host fitness caused by avian haematozoa may be underestimated in natural populations if the exacerbated virulence associated with exposure to multiple parasites is not taken into account.     

Latitudinal and altitudinal diversity patterns and Rapoport effects in north-west European land snails and their causes

The latitudinal and altitudinal range sizes of north-west European land-snail species increase with increasing latitude/altitude. These Rapoport effects are not caused by northern/high-altitude species with wider latitudinal/altitudinal ranges and southern/low-altitude species with narrower latitudinal/altitudinal ranges, as predicted by the climatic variability hypothesis. They are instead caused mainly by different northern/upper borders of species occurring in the south part of the study area or at low and intermediate altitudes, respectively. This pattern indicates that the observed Rapoport effects are the result mainly of differential northward/upward expansion of species that were restricted to southern/low or intermediate altitude refugia during the glacials. Although all species occurring in a refugium experienced the same climatic conditions, there is stochastic variation in their climatic tolerance. Species with broader climatic tolerance were able to expand farer northwards/upwards postglacial. The altitudinal distribution of species richness in the analysed alpine faunas cannot be explained by the Rapoport-rescue hypothesis, because species richness peaks at intermediate altitudes and because there is no negative correlation between the number of range borders and altitude. The Rapoport-rescue hypothesis alone is probably also insufficient to explain the decrease in species richness with increasing latitude.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 309–323.     

Density, body mass and parasite species richness of terrestrial mammals

We investigated the relationships between helminth species richness and body mass and density of terrestrial mammals. Cross-species analysis and the phylogenetically independent contrast method produced different results. A non-phylogenetic approach (cross-species comparisons) led to the conclusion that parasite richness is linked to host body size. However, an analysis using phylogenetically independent contrasts showed no relationship between host body size and parasite richness. Conversely, a non-phylogenetic approach generated a negative relationship between parasite richness and host density, whereas the independent contrast method showed the opposite trend – that is, parasite richness is positively correlated with host density. From an evolutionary perspective, our results suggest that opportunities for parasite colonization depend more closely on how many hosts are available in a given area than on how large the hosts are. From an epidemiological point of view, our results confirm theoretical models which assume that host density is linked to the opportunity of a parasite to invade a population of hosts. Our findings also suggest that parasitism may be a cost associated with host density. Finally, we provide some support for the non-linear allometry between density and mammal body mass (Silva and Downing, 1995), and explain why host density and host body mass do not relate equally to parasite species richness.     

物种多样性和功能群多样性与生态系统生产力的关系——以内蒙古短花针茅草原为例

生物多样性与生态系统生产力之间的关系是当前生态学领域的热点问题。短花针茅(Stipa breviflora)草原是内蒙古荒漠草原的主要类型, 生态系统脆弱, 气候波动剧烈, 研究内蒙古短花针茅草原生物多样性与生产力的关系具有十分重要的意义。该研究在内蒙古短花针茅草原区设置了202个样地进行群落调查, 在干旱区及半干旱区两种资源供给下, 分析了物种丰富度、功能群丰富度与生产力的关系, 旨在解决两个科学问题: 1)物种多样性和功能群多样性中, 哪一种与生产力关系更为密切?2)资源供给对多样性和生产力关系的影响。结果表明: 1)物种丰富度、群落生产力与年降水量呈正相关关系, 而功能群丰富度与年降水量之间不存在显著相关性; 2)群落生产力随物种丰富度的增加而增加, 且两者间呈正线性关系, 功能群丰富度与生产力之间不存在显著相关关系; 3)资源供给会影响多样性与生产力之间的关系, 资源供给低时, 多样性对生产力贡献较低, 资源供给高时, 多样性对生产力的贡献较高。该研究丰富了多样性与生产力关系的研究, 同时, 考虑到植物功能性状的研究在近几年受到生态学家的重视, 且多数研究集中于小尺度的人工控制实验, 因此, 在大尺度自然生态系统中开展功能性状多样性与生态系统功能关系的研究将十分必要。     

Global patterns of plant diversity

Summary Using 94 data sets from across the globe, we explored patterns of mean community species richness, landscape species richness, mean similarity among communities and mosaic diversity. Climate affected community species richness primarily through productivity while other climatic factors were secondary. Climatic equability affected species richness only in temperate regions where richness was greatest at high levels of temperature variability and low levels of precipitation variability. Landscape species richness correlated positively with community species richness. A global gradient in mean similarity existed but was uncorrelated with community species richness. Mean similarity was least and mosaic diversity was greatest between 25 and 30° latitude. The most diverse landscapes (low mean similarity) correlated with warm temperatures, high elevations, large areas and large seasonal temperature fluctuations. The most complex landscapes (high mosaic diversity) correlated with large areas, high productivity and warm winters. We compared diversity measures among continents and found only one significant difference: Australian landscapes have greater mosaic diversity than African landscapes. Based on our analyses we propose two hypotheses: (1) for plants, biotic interactions are more important in structuring landscapes in warmer climates and (2) longer isolated landscapes have more clearly differentiated ecological subunits.     

Helminth parasite diversity discloses age and sex differences in the foraging behaviour of southern lapwings (Vanellus chilensis)

Parasites with heteroxen cycles are important sources of information on the trophic relations of hosts. This is particularly instructive for species whose age‐based or sex‐based differences are hardly detected by behavioural observations. Here, we describe the helminth community of the omnivorous southern lapwing (Vanellus chilensis) and evaluate whether it is affected by the host's sex, age and body size. The species is sexually monomorphic in body length, but males are slightly heavier than females. We analysed 112 individuals collected in Curitiba, Brazil, in March 2010. All hosts were parasitized. The helminth community was composed of 10 species (the digeneans Leucochloridium parcum and Athesmia sp., the cestode Infula macrophallus, the acantocephalans Plagiorhynchus sp., Centrorhynchus sp., Mediorhynchus sp., and an unidentified Gigantorhynchida, and the nematodes Heterakis psophiae, Dispharynx nasuta and an unidentified Capillariidae), seven of which were novel reports for this host species. Prevalence ranged from <1% to 99%. Whereas I. macrophallus was the most prevalent species, D. nasuta showed the highest mean intensity and abundance of infection. The former was found in most hosts as single male–female pairs, suggesting the occurrence of intrasexual competition. The infracommunities of juvenile birds showed a higher parasite species richness than those of adult males and females, suggesting the exploitation of a wider array of prey. However, the three classes harboured seven parasite species. Differences in parasite diversity (lower in juveniles, intermediate in adult males and higher in adult females) reflect the evenness in the distribution of parasite specimens among taxa in each age–sex class and are compatible with differences in their foraging strategy. Finally, we conclude based on the cycles of the heteroxen species that southern lapwings preyed upon molluscs, coleopterans, woodlice and earthworms.     

Human impacts decouple a fundamental ecological relationship—The positive association between host diversity and parasite diversity

Human impacts on ecosystems can decouple the fundamental ecological relationships that create patterns of diversity in free‐living species. Despite the abundance, ubiquity, and ecological importance of parasites, it is unknown whether the same decoupling effects occur for parasitic species. We investigated the influence of fishing on the relationship between host diversity and parasite diversity for parasites of coral reef fishes on three fished and three unfished islands in the central equatorial Pacific. Fishing was associated with a shallowing of the positive host‐diversity–parasite‐diversity relationship. This occurred primarily through negative impacts of fishing on the presence of complex life‐cycle parasites, which created a biologically impoverished parasite fauna of directly transmitted parasites resilient to changes in host biodiversity. Parasite diversity appears to be decoupled from host diversity by fishing impacts in this coral reef ecosystem, which suggests that such decoupling might also occur for parasites in other ecosystems affected by environmental change.     

Antimicrobial defences increase with sociality in bees

Evidence for the antiquity and importance of microbial pathogens as selective agents is found in the proliferation of antimicrobial defences throughout the animal kingdom. Social insects, typified by crowding and often by low genetic variation, have high probabilities of disease transmission and eusocial Hymenoptera may be particularly vulnerable because of haplodiploidy. Mechanisms they employ to reduce the risk of disease include antimicrobial secretions which are particularly important primary barriers to infection. However, until now, whether or not there is selection for stronger antimicrobial secretions when the risk of disease increases because of sociality has not been tested. Here, we present evidence that the production of progressively stronger antimicrobial compounds was critical to the evolution of sociality in bees. We found that increases in group size and genetic relatedness were strongly correlated with increasing antimicrobial strength. The antimicrobials of even the most primitive semi-social species were an order of magnitude stronger that those of solitary species, suggesting a point of no return, beyond which disease control was essential. Our results suggest that selection by microbial pathogens was critical to the evolution of sociality and required the production of strong, front-line antimicrobial defences.