Abiotic and biotic determinants of the seasonal dynamics of the tick Rhipicephalus appendiculatus in South Africa

Abstract. The creation of a generic population model for the tick Rhipicephalus appendiculatus requires a detailed, quantified understanding of the interactions of these ticks with their biotic and abiotic environment in the different parts of their range, from the tropical regions of East Africa to the temperate regions of South Africa. The much greater seasonal variation in climatic conditions, particularly temperature, further from the equator introduces variable development rates and diapause into the life cycle. Estimates of natural temperature-dependent interstadial development periods, derived from a combination of published laboratory and field data, were applied to published data on the seasonal abundance of R.appendiculatus on three farms in the Eastern Cape Province of South Africa. This enabled an assessment of which ticks of one stage give rise to which ticks of the next stage, from which (a) the onset and duration of diapause in unfed adults, and (b) seasonal interstadial mortality indices in the form of fc-values, could be estimated. The contribution of biotic (tick density) and abiotic (climatic) factors as predictors of mortality at each life stage was investigated by step-wise multiple regression. Density-independent mortality at the female-to-larval stage is correlated with geographically variable climatic factors, minimum temperature at two farms and minimum relative humidity at the third. The other two stages are governed by density-dependent mortality, which, it is argued, may be caused largely by the hosts' acquired resistance to ticks. As expected on theoretical grounds, this density dependence is weaker nearer to the edge of the tick's range and for the more vulnerable immature stages.     

Climate,satellite imagery and the seasonal abundance of the tick Rhipicephalus appendiculatus in southern Africa: a new perspective

Abstract. Recent predictive models for the distribution of the African tick Rhipicephalus appendiculatus Neumann, based on the computer packages CLIMEX and BIOCLIM and data derived from meteorological satellites, and for the seasonal dynamics of the same tick using the computer simulation models ECFXPERT and T3HOST, all have their limitations. Statistical analysis of the relationships between the seasonal abundance of all three life stages of this tick and climatic and satellite-derived data from five sites in southern Africa, taken from the literature, supports a new perspective that it is the timing of the questing activity of the desiccation-vulnerable larvae that determines the pattern of the tick's seasonal dynamics. The timing of the activity of nymphs and adults is determined by temperature-dependent development rates plus the delaying phenomenon of photoperiod-sensitive diapause, the timing and duration of which have evolved to achieve maximum generation survival by ensuring the occurrence of eggs and larvae during periods of optimal climatic conditions. The most important environmental factor appears to be night-time minimum temperature, determining condensation and saturation deficit and thus the tick's ability to replenish moisture lost during the daytime and so to survive while questing for hosts. It is the larvae whose numbers are correlated most closely with these factors, consistent with earlier experimental results showing larvae to be most susceptible to desiccating conditions. There is a statistical linkage between larval tick numbers and satellite imagery, arising from the correlation between larval numbers and minimum temperature and saturation deficit conditions, and in turn the relationship between these climatic conditions and the subsequent vegetational changes monitored by the satellites. Moisture availability to larvae is likely to be the critical factor throughout the geographical range of R.appendiculatus, but the precise combination of climatic conditions that optimize moisture availability and questing tick survival can be expected to vary geographically. The relationships between ticks, temperatures and satellite data in parts of equatorial Africa have yet to be established. These correlative patterns highlight both the critical life stage and environmental factors when trying to understand temporal, and ultimately spatial, variations in tick abundance.     

Dynamic and temporal structure of the troglobitic beetle Speonomus hydrophilus (Coleoptera: Bathyscimae)

The population ecology of the beetle Speonomus hydrophilus , occurring both in caves (reduced fluctuations in many abiotic parameters) and under the deepest layer of soil in mountains (MSS, more exposed to climatic variations), was studied in four habitats in the French central Pyrenees We have assessed some of the characteristics of the environment where these populations occur c g physical data (altitude and exposure), geologic data (nature of the parent-rock) and abiotic parameters (temperature with its seasonal fluctuations) and we investigated the relative importance of environmental structure and ecological characteristics on the temporal organization of S hydrophilus and the troglobitic fauna which cohabits The climatic study shows the existence of an annual thermal cycle which is regular and well marked for the MSS habitats but slightly out of phase with the surface cycle These periodic variations however slight may be stressful for troglobitic species In the MSS populations, the phenology of the entire community is reflected in the pattern seen in Speonomus The analysis of faunal profiles shows that samples follow the same seasonal succession during the annual cycle A potential seasonal rhythm of emergence may reflect a seasonal rhythm of vitellogenesis which produces a rhythm of egg-laying     

Comparative off‐host survival of larval Rocky Mountain Wood Ticks (Dermacentor andersoni) collected from ecologically distinct field populations

Dermacentor andersoni (Ixodida: Ixodidae) Stiles, also known as the Rocky Mountain Wood Tick (RMWT), is found throughout the western United States and transmits pathogens of importance to human and animal health. The distributions and activity patterns of RMWTs are shaped by regional climatic variation. However, it is unknown if responses to climatic variation differ across the tick's geographical range. The objective of this narrow study was to test the hypothesis that the responses of RMWTs to abiotic conditions [e.g. temperature and RH (relative humidity)] vary among populations. We collected RMWTs from ecologically distinct field sites in the states of Montana and Oregon (USA). In the laboratory, we tracked weekly survival of tick larvae under four combinations of RH (75% and 98%) and temperature (26 and 32 °C) that reflected the range of conditions observed in the source habitats during spring‐summer. For both populations, larval survival time decreased at the higher ambient temperature (50% mortality 1–2 weeks earlier). Differences in RH did not affect the survival time of larvae from Oregon. By contrast, the survival time of larvae from Montana decreased at the lower RH (50% mortality 1 week earlier). These data suggest that the tolerance limits for water stress may differ among populations of D. andersoni.     

Occurrence and distribution of zoosporic organisms in water bodies from Brazilian Cerrado

Zoosporic organisms are commonly found in different aquatic ecosystems; however little is known about the influence of environmental factors and seasonal fluctuations on their occurrence and distribution. This study investigated patterns of abundance, frequency and diversity of these organisms and their relationship with some abiotic factors and seasonality in a cerrado remnant in Sao Paulo, Brazil. Water samples were collected at four dates in two areas of this remnant during the rainy and dry seasons, and 13 abiotic factors were analyzed. From 48 samples collected, 32 taxa were isolated with the multiple baiting technique. The community structure of the zoosporic organisms showed some changes in response to different spatial distribution and seasonal fluctuations, probably influenced by distinct abiotic characteristics of each area or climatic period. Still, the similarity between areas and seasons indicated by S?rensen index and diversity (richness, evenness, Shannon and Simpson indices) were high.     

Intraspecific differentiation of social behavior and shelter selection in Mesobuthus gibbosus (Brullé, 1832) (Scorpiones: Buthidae)

We compared seasonal shelter selection and social behavior of Mesobuthus gibbosus from autumn to mid-summer in two similar phryganic ecosystems, in continental Greece (near Volos city) and in insular Greece (eastern Crete), and in the laboratory under simulated abiotic conditions. Our results showed that shelter selection is a critical indicator of the seasonal social behavior of the species. The abrupt climatic changes in spring caused a differentiation in shelter selection between the cold period (November–February) and the warm period (March–June) at both sites. Sociality was exhibited only during winter in the field and was more extensive under cold conditions in the laboratory. Co-occurrence of scorpions proved to be age-specific, facilitated by population density and by harsh abiotic conditions during winter, and negatively influenced by intraspecific competition, which was higher in continental Greece. The response of scorpions to changes of abiotic factors reveals synchronization of seasonal shelter selection with climatic changes.     

Assessing the use of camera-based indices for characterizing canopy phenology in relation to gross primary production in a deciduous broad-leaved and an evergreen coniferous forest in Japan

Recent studies have reported that seasonal variation in camera-based indices that are calculated from the digital numbers of the red, green, and blue bands (RGB_DN) recorded by digital cameras agrees well with the seasonal change in gross primary production (GPP) observed by tower flux measurements. These findings suggest that it may be possible to use camera-based indices to estimate the temporal and spatial distributions of photosynthetic productivity from the relationship between RGB_DN and GPP. To examine this possibility, we need to investigate the characteristics of seasonal variation in three camera-based indices (green excess index [GE], green chromatic coordinate [rG], and HUE) and the robustness of the relationship between these indices and tower flux-based GPP and how it differs among ecosystems. Here, at a daily time step over multiple years in a deciduous broad-leaved and an evergreen coniferous forest, we examined the relationships between canopy phenology assessed by using the three indices and GPP determined from tower CO₂ flux observations, and we compared the camera-based indices with the corresponding spectra-based indices estimated by a spectroradiometer system. We found that (1) the three camera-based indices and GPP showed clear seasonal patterns in both forests; (2) the amplitude of the seasonal variation in the three camera-based indices was smaller in the evergreen coniferous forest than in the deciduous broad-leaved forest; (3) the seasonal variation in the three camera-based indices corresponded well to seasonal changes in potential photosynthetic activity (GPP on sunny days); (4) the relationship between the three camera-based indices and GPP appeared to have different characteristics at different phenological stages; and (5) the camera-based and spectra-based HUE indices showed a clear relationship under sunny conditions in both forests. Our results suggest that it might be feasible for ecologists to establish comprehensive networks for long-term monitoring of potential photosynthetic capacity from regional to global scales by linking satellite-based, in situ spectra-based, and in situ camera-based indices.     

Projected effects of climate change on tick phenology and fitness of pathogens transmitted by the North American tick Ixodes scapularis

Ixodes scapularis is the principal tick vector of the Lyme borreliosis agent Borrelia burgdorferi and other tick-borne zoonoses in northeastern North America. The degree of seasonal synchrony of nymphal and larval ticks may be important in influencing the basic reproductive number of the pathogens transmitted by I. scapularis. Because the seasonal phenology of tick vectors is partly controlled by ambient temperature, climate and climate change could shape the population biology of tick-borne pathogens. We used projected monthly normal temperatures, obtained from the second version of the Canadian Coupled Global Climate Model (CGCM2) under emissions scenario A2 of the Intergovernmental Panel on Climate Change for a site in southern Ontario, Canada, to simulate the phenology of I. scapularis in a mathematical model. The simulated seasonal abundance of ticks then determined transmission of three candidate pathogens amongst a population of white-footed mice (Peromyscus leucopus) using a susceptible-infected-recovered (SIR) model. Fitness of the different pathogens, in terms of resilience to changes in tick and rodent mortality, minima for infection duration, transmission efficiency and particularly any additional mortality of rodents specifically associated with infection, varied according to the seasonal pattern of immature tick activity, which was different under the temperature conditions projected for the 2020s, 2050s and 2080s. In each case, pathogens that were long-lived, highly transmissible and had little impact on rodent mortality rates were the fittest. However, under the seasonal tick activity patterns projected for the 2020s and 2050s, the fitness of pathogens that are shorter-lived, less efficiently transmitted, and more pathogenic to their natural hosts, increased. Therefore, climate change may affect the frequency and distribution of I. scapularis-borne pathogens and alter their evolutionary trajectories.     

An empirical quantitative framework for the seasonal population dynamics of the tick Ixodes ricinus

The wide geographic and climatic range of the tick Ixodes ricinus, and the consequent marked variation in its seasonal population dynamics, have a direct impact on the transmission dynamics of the many pathogens vectored by this tick species. We use long-term observations on the seasonal abundance and fat contents (a marker of physiological ageing) of ticks, and contemporaneous microclimate at three field sites in the UK, to establish a simple quantitative framework for the phenology (i.e. seasonal cycle of development) of I. ricinus as a foundation for a generic population model. An hour-degree tick inter-stadial development model, driven by soil temperature and including diapause, predicts the recruitment (i.e. emergence from the previous stage) of a single cohort of each stage of ticks each year in the autumn. The timing of predicted emergence coincides exactly with the new appearance of high-fat nymphs and adults in the autumn. Thereafter, fat contents declined steadily until unfed ticks with very low energy reserves disappeared from the questing population within about 1 year from their recruitment. Very few newly emerged ticks were counted on the vegetation in the autumn, but they appeared in increasing numbers through the following spring. Larger ticks became active and subsequently left the questing population before smaller ones. Questing tick population dynamics are determined by seasonal patterns of tick behaviour, host-contact rates and mortality rates, superimposed on a basal phenology that is much less complex than has hitherto been portrayed.     

Understanding the Relationships between Landscape Connectivity and Abundance of Ixodes ricinus Ticks

The density of Ixodes ricinus ticks in a heterogeneous landscape of northern Spain was determined and associated with some aspects of habitat topology. The habitat mosaic was used to quantify connectivity between patches of different tick density. The analysis revealed that patches with high tick abundance are ‘stepping-stone’ territories that, when removed from the landscape, cause large changes in connectivity. Sites with medium tick abundance do not cause such a critical transition in connectivity. Patches with low tick abundance, but optimal abiotic conditions for survival are located within the minimum cost corridors network joining the patches, while those sites where the tick has been intermittently collected are located at variable distances from this network. These results suggest that tick distribution in a zone is highly affected not only by abiotic variables (vegetation and weather) but also by host movements. Whether these high-density ‘stepping-stone’ patches occur in other tick species needs to be evaluated because of the potential implications of these foci for human health. This revised version was published online in July 2006 with corrections to the Cover Date.     

SEASONAL MORTALITY PATTERNS IN NON‐HUMAN PRIMATES: IMPLICATIONS FOR VARIATION IN SELECTION PRESSURES ACROSS ENVIRONMENTS

Examining seasonal mortality patterns can yield insights into the drivers of mortality and thus potential selection pressures acting on individuals in different environments. We compiled adult and juvenile mortality data from nine wild non‐human primate taxa to investigate the role of seasonality in patterns of mortality and address the following questions: Is mortality highly seasonal across species? Does greater environmental seasonality lead to more seasonal mortality patterns? If mortality is seasonal, is it higher during wet seasons or during periods of food scarcity? and Do folivores show less seasonal mortality than frugivores? We found seasonal mortality patterns in five of nine taxa, and mortality was more often tied to wet seasons than food‐scarce periods, a relationship that may be driven by disease. Controlling for phylogeny, we found a positive relationship between the degree of environmental seasonality and mortality, with folivores exhibiting more seasonal mortality than frugivores. These results suggest that mortality patterns are influenced both by diet and degree of environmental seasonality. Applied to a wider array of taxa, analyses of seasonal mortality patterns may aid understanding of life‐history evolution and selection pressures acting across a broad spectrum of environments and spatial and temporal scales.     

Impact of biodiversity and seasonality on Lyme-pathogen transmission

Lyme disease imposes increasing global public health challenges. To better understand the joint effects of seasonal temperature variation and host community composition on the pathogen transmission, a stage-structured periodic model is proposed by integrating seasonal tick development and activity, multiple host species and complex pathogen transmission routes between ticks and reservoirs. Two thresholds, one for tick population dynamics and the other for Lyme-pathogen transmission dynamics, are identified and shown to fully classify the long-term outcomes of the tick invasion and disease persistence. Seeding with the realistic parameters, the tick reproduction threshold and Lyme disease spread threshold are estimated to illustrate the joint effects of the climate change and host community diversity on the pattern of Lyme disease risk. It is shown that climate warming can amplify the disease risk and slightly change the seasonality of disease risk. Both the “dilution effect” and “amplification effect” are observed by feeding the model with different possible alternative hosts. Therefore, the relationship between the host community biodiversity and disease risk varies, calling for more accurate measurements on the local environment, both biotic and abiotic such as the temperature and the host community composition.     

Abiotic mosaics affect seasonal variation of plant resources and influence the performance and mortality of a leaf-miner in Gambel’s oak (Quercus gambelii, Nutt.)

Abstract Conceptual models of terrestrial trophic dynamics have emphasized the potential influence of various abiotic factors, though empirical studies have found generalities to be scarce. Progress may result through an increased use of experimental gradients in tandem with existing, natural gradients of abiotic factors that are more difficult to manipulate. Along an elevation/climatic gradient, a fertilization experiment was conducted to examine the impact of environmentally induced variation in foliar nitrogen on tri-trophic interactions for Phyllonorycter sp., a leaf-miner (Lepidoptera: Gracillariidae), in Rocky Mountain white oak, Quercus gambelii. Microclimate determined the relative effect of host-plant and natural-enemy effects for Phyllonorycter sp. Cooler, more humid microclimates resulted in a higher biomass payoff (total biomass/density). Further, increased seasonal variation in foliar nitrogen content at warmer, drier sites significantly increased parasitism rates. Fertilization treatments increased foliar nitrogen content and resulted in increased early-instar mortality. Fertilization treatment also acted in a non-linear fashion with microclimate to influence spatial patterns in parasitism rates. Overall, microclimate was found to affect characteristics of each trophic level, including host-plant nitrogen dynamics, Phyllonorycter sp. performance, and parasitism rates. Additionally, nutrient availability altered patterns of parasitoid-related mortality in Phyllonorycter sp. within microclimates. These results suggest that local topographic variation in combinations of abiotic factors, or abiotic mosaics, has important effects for spatial patterns of tri-trophic interactions.     

Epidemiology of canine babesiosis in relation to the activity ofDermacentor reticulatus in southern Jura (France)

Dates of onset of canine babesiosis within a hyperendemic focus in France (Rhone Valley, south of Jura and north of the Alps) were compared to the seasonal population level of the adult stage ofDermacentor reticulatus over a 12-month period (December 1982–November 1983). Cases of babesiosis occurred in spring and fall when adultD. reticulatus were active. The fluctuations of the vector tick population and the onsets of canine babesiosis were also correlated with climatic changes: no tick activity or clinical cases of disease were detected in winter (low temperature) or in summer. The slight disparities observed between both distributions might be explained by various factors such as the development of immunity against the parasites, the intervals between tick bites and appearance of symptoms of the disease, or the asynchrony between different biotopes.     

The relationships between habitat topology, critical scales of connectivity and tick abundance Ixodes ricinus in a heterogeneous landscape in northern Spain

The habitat mosaic was used to quantify connectivity between patches of different tick density of the notorious tick species Ixodes ricinus in an attempt to determine the cause of variations in tick abundance among apparently homogeneous sites in northern Spain. The analysis revealed that patches with high tick abundance are "stepping-stone" territories that, when removed from the landscape, cause large changes in connectivity. Sites with medium tick abundance do not cause such a critical transition in connectivity. Patches with low tick abundance, but optimal abiotic conditions for survival, are located within the minimum cost corridors network joining the patches, while those sites where the tick has been intermittently collected are located at variable distances from this network. Sites where the tick is consistently absent, but where the habitat is predicted to be suitable (old, heterogeneous forests of Quercus spp.) for the tick, are very separated from this main network of connections. These results suggest that tick distribution in a zone is highly affected not only by abiotic variables (vegetation and weather) but also by host movements. Dispersal of the tick is a function of how the hosts perceive the habitat, and the habitat's permeability to host movement. Permanent tick populations seem to be supported by the existence of these critical, high density patches, located at significant places within the habitat network.     

Evaluating the economic damage threshold for bont tick (Amblyomma hebraeum) control in Zimbabwe

Controlling ticks and tick-borne diseases by frequent applications of acaricides (e.g., dipping) is costly, and can leave treated livestock vulnerable to epizootics of tick-borne diseases should the system of applying acaricides break down. The concept of only applying acaricides on an infrequent (strategic) basis often relies on the target tick population displaying a seasonal cycle. However, as adult bont tick (Amblyomma hebraeum) infestations in Zimbabwe's lowveld do not have a strictly seasonal pattern of occurrence, it is recommended that tick control only be applied when bont tick infestations are equal to, or greater than, their economic damage threshold. The economic damage threshold is the minimum average weekly standard female tick burden sufficient to cause damage equal in dollar value to the costs of applying tick control. Assuming that each standard female tick represents a 10 gram weight loss, the economic damage threshold (standard female ticks/week) is equivalent to the ratio of the producer price of beef (liveweight equivalent): per head cost of dipping (Eqn (3)). To illustrate the application of the threshold methodology, it was assumed that the producer price of beef was Z$1.63/kg (U.S.$0.33/kg) and that tick control cost Z$0.29/hd/dip (U.S.$0.06/hd/dip). This gave a threshold of 18 standard female ticks/head/week. Using tick counts obtained from 20 Brahman cattle held at Mbizi in southeastern Zimbabwe, it was shown that for the 1988 calendar year there were only 32 weeks when the economic damage threshold was met or exceeded. This is substantially less that the 44 dippings per year that have been, until very recently, legally required in Zimbabwe. Sensitivity analysis showed that a 10% rise in the cost of dipping reduced to 23 (a 28% decrease) the number of weeks when tick burdens exceeded the economic damage threshold. By further assuming that an acaricide application and residual effects will cause a 3–5 week interval before the next application may be required, the number of weeks when the tick burden was equal to or greater than the threshold of 18 standard females/week fell to just 9–12 weeks. Three factors may cause an alteration in the economic damage threshold: i) tick burdens may cause damage to the udders; ii) secondary infestations (e.g., screw-worm) may cause economic damage; and iii) nutritional stress of the cattle may reduce the actual average per tick weight loss. Until further data becomes available, it is recommended that the economic damage threshold methodology be used as described here, and that farmers closely observe their herds during the suggested weekly tick sampling for udder damage and secondary infestations.     

The use of time-series analysis to forecast bont tick (Amblyomma hebraeum) infestations in Zimbabwe

Studying the dynamics of tick infestations on cattle is an essential step in developing optimal strategies for tick control. Successful strategic tick control requires accurate predictions of when tick infestations will reach predetermined threshold levels. In the case ofAmblyomma hebraeum, earlier work has shown that there is no consistent pattern of seasonal activity. This means that a statistical model for predictingA. hebraeum infestations cannot reliably use climatic factors as the only independent variables. An alternative method is to apply time-series, or auto-regressive moving-average (ARMA), analysis which uses only the past population patterns to predict future trends. This technique was applied to a data set consisting of 108 weekly tick counts ofA. hebraeum (adult males, standard females, flat females and standard nymphs), conducted at an experimental station in southeastern Zimbabwe. The ability of the ARMA models to fit and predict actual tick infestations was judged using two sets of criteria. The first set focused on the goodness-of-fit, and used the adjustedR ² values,Q statistic and the Akaike Information Criteria. The second set of criteria measured the forecasting accuracy of an estimated equation, and consisted of regressing a 9-period forecast against an actual out-of-sample data set not used in the estimation process. The root mean square error of the forecast was also considered when comparing several models for the same data set. Using these criteria, the models estimated using the ARMA technique were judged to both fit and forecast with sufficient accuracy to warrant their use in strategic tick control. Although the success of using ARMA to forecastA. hebraeum is partly due to the non-seasonal behaviour of the species, the results presented here suggest that it is worthwhile exploring the use of ARMA techniques to model the dynamics of other tick species. Where independent variables exert considerable influence on the dynamics of a tick species, these variables can be incorporated into an ARMA-style model.     

Impact of Nesting Mortality on Avian Breeding Phenology: A Case Study on the Red-Backed Shrike (Lanius collurio)

The seasonal timing of avian reproduction is supposed primarily to coincide with favourable feeding conditions. Long-term changes in avian breeding phenology are thus mostly scrutinized in relation to climatic factors and matching of the food supplies, while the role of nesting mortality is largely unexplored. Here we show that higher seasonal mean daily mortality rate leads to a shift in the distribution of breeding times of the successful nests to later dates in an an open-nesting passerine bird, the red-backed shrike Lanius collurio. The effect appeared to be strong enough to enhance or counteract the influence of climatic factors and breeding density on the inter-annual variation in mean hatching dates. Moreover, the seasonal distribution of reproductive output was shifted to larger, or smaller, broods early in the season when the nesting mortality increased, or decreased, respectively, during the season. We suggest that population level changes in timing of breeding caused by a general advancement of spring and of the food supplies might be altered by the seasonality in nesting mortality. Hence, we argue that consideration of nesting mortality is of major importance for understanding long-term trends in avian phenology, particularly in species capable of renesting.     

Demographics in an alpine reindeer herd: effects of density and winter weather

We examined how population density, winter weather, snow conditions, and 2 large-scale climatic indices (North Atlantic Oscillation, NAO, and Arctic Oscillation, AO) influenced demography (reproduction and mortality) in an alpine herd of semi-domesticated reindeer Rangifer tarandus between 1959 and 2000 in Finnish Lapland. The herd lived on heavily grazed lichen pastures, with winter densities between 0.8 and 3.9 individuals km⁻². Icing conditions occurred every 7th yr, on an average, and decreased reproductive rate (calves/females) by 49%. In general linear models icing remarkably increased the fit of snow models to reproductive rate. Incorporation of an interaction term between icing and the snow depth index provided better fit than a model without interaction. Delayed snowmelt decreased reproductive rate. For the day of snowmelt, however, the model without interaction was better than the interaction model. These 3 models provided the best fit to the data and accounted for 51–54% of the variation in reproductive rate. Winter mortality was related to density and large-scale climatic indices, but not to local winter weather except a slight increase in mortality during an icing winter. The best model for winter mortality, including reindeer density and NAO, accounted for 26% of variation in mortality. Three factors may be involved explaining weak density dependence or the lack of such dependence; climate change scenarios that predict higher winter temperature, more frequent thawing-freezing periods, and deeper snow would be expected to decrease reproductive rate and increase winter mortality of reindeer and thus to reduce profitability of reindeer husbandry. In contrast, early springs would be advantageous for reindeer in the short term.     

Climate extremes promote fatal co-infections during canine distemper epidemics in African lions

Extreme climatic conditions may alter historic host-pathogen relationships and synchronize the temporal and spatial convergence of multiple infectious agents, triggering epidemics with far greater mortality than those due to single pathogens. Here we present the first data to clearly illustrate how climate extremes can promote a complex interplay between epidemic and endemic pathogens that are normally tolerated in isolation, but with co-infection, result in catastrophic mortality. A 1994 canine distemper virus (CDV) epidemic in Serengeti lions (Panthera leo) coincided with the death of a third of the population, and a second high-mortality CDV epidemic struck the nearby Ngorongoro Crater lion population in 2001. The extent of adult mortalities was unusual for CDV and prompted an investigation into contributing factors. Serological analyses indicated that at least five "silent" CDV epidemics swept through the same two lion populations between 1976 and 2006 without clinical signs or measurable mortality, indicating that CDV was not necessarily fatal. Clinical and pathology findings suggested that hemoparsitism was a major contributing factor during fatal epidemics. Using quantitative real-time PCR, we measured the magnitude of hemoparasite infections in these populations over 22 years and demonstrated significantly higher levels of Babesia during the 1994 and 2001 epidemics. Babesia levels correlated with mortalities and extent of CDV exposure within prides. The common event preceding the two high mortality CDV outbreaks was extreme drought conditions with wide-spread herbivore die-offs, most notably of Cape buffalo (Syncerus caffer). As a consequence of high tick numbers after the resumption of rains and heavy tick infestations of starving buffalo, the lions were infected by unusually high numbers of Babesia, infections that were magnified by the immunosuppressive effects of coincident CDV, leading to unprecedented mortality. Such mass mortality events may become increasingly common if climate extremes disrupt historic stable relationships between co-existing pathogens and their susceptible hosts.