Social living mitigates the costs of a chronic illness in a cooperative carnivore.

Infection risk is assumed to increase with social group size, and thus be a cost of group living. We assess infection risk and costs with respect to group size using data from an epidemic of sarcoptic mange (Sarcoptes scabiei) among grey wolves (Canis lupus). We demonstrate that group size does not predict infection risk and that individual costs of infection, in terms of reduced survival, can be entirely offset by having sufficient numbers of pack-mates. Infected individuals experience increased mortality hazards with increasing proportions of infected pack-mates, but healthy individuals remain unaffected. The social support of group hunting and territory defence are two possible mechanisms mediating infection costs. This is likely a common phenomenon among other social species and chronic infections, but difficult to detect in systems where infection status cannot be measured continuously over time.

Evidence of trade-offs shaping virulence evolution in an emerging wildlife pathogen.

In the mid-1990s, the common poultry pathogen Mycoplasma gallisepticum (MG) made a successful species jump to the eastern North American house finch Haemorhous mexicanus (HM). Subsequent strain diversification allows us to directly quantify, in an experimental setting, the transmission dynamics of three sequentially emergent geographic isolates of MG, which differ in the levels of pathogen load they induce. We find significant among-strain variation in rates of transmission as well as recovery. Pathogen strains also differ in their induction of host morbidity, measured as the severity of eye lesions due to infection. Relationships between pathogen traits are also investigated, with transmission and recovery rates being significantly negatively correlated, whereas transmission and virulence, measured as average eye lesion score over the course of infection, are positively correlated. By quantifying these disease-relevant parameters and their relationships, we provide the first analysis of the trade-offs that shape the evolution of this important emerging pathogen.

Zoonotic schistosomiasis in non-human primates: past, present and future activities at the human-wildlife interface in Africa.

Schistosomiasis is one of the world's most widely distributed and prevalent parasitic diseases. Less widely recognized is that some species of Schistosoma, including several that commonly affect humans, also cause disease in other mammalian species; in particular, infections in non-human primates are known. With interest increasing in emerging zoonotic diseases, the status of schistosomiasis as a zoonotic infection is in need of re-appraisal, especially in light of advances in application of molecular screening and epidemiological tools where newly reported infections raise general animal welfare and conservation concerns. Focusing on Africa, this review provides a summary of the occurrence of schistosomiasis in non-human primates and discusses new ways in which surveillance for schistosomiasis should be integrated into more effective conservation management and disease control strategies. Emphasis is on the more common forms of human schistosomiasis, their clinical manifestations and epidemiological significance in terms of infection reservoir potential.

Common garden experiment reveals pathogen isolate but no host genetic diversity effect on the dynamics of an emerging wildlife disease.

Host genetic diversity can mediate pathogen resistance within and among populations. Here we test whether the lower prevalence of Mycoplasmal conjunctivitis in native North American house finch populations results from greater resistance to the causative agent, Mycoplasma gallisepticum (MG), than introduced, recently-bottlenecked populations that lack genetic diversity. In a common garden experiment, we challenged wild-caught western (native) and eastern (introduced) North American finches with a representative eastern or western MG isolate. Although introduced finches in our study had lower neutral genetic diversity than native finches, we found no support for a population-level genetic diversity effect on host resistance. Instead we detected strong support for isolate differences: the MG isolate circulating in western house finch populations produced lower virulence, but higher pathogen loads, in both native and introduced hosts. Our results indicate that contemporary differences in host genetic diversity likely do not explain the lower conjunctivitis prevalence in native house finches, but isolate-level differences in virulence may play an important role.

Probable causes of increasing brucellosis in free-ranging elk of the Greater Yellowstone Ecosystem.

While many wildlife species are threatened, some populations have recovered from previous overexploitation, and data linking these population increases with disease dynamics are limited. We present data suggesting that free-ranging elk (Cervus elaphus) are a maintenance host for Brucella abortus in new areas of the Greater Yellowstone Ecosystem (GYE). Brucellosis seroprevalence in free-ranging elk increased from 0-7% in 1991-1992 to 8-20% in 2006-2007 in four of six herd units around the GYE. These levels of brucellosis are comparable to some herd units where elk are artificially aggregated on supplemental feeding grounds. There are several possible mechanisms for this increase that we evaluated using statistical and population modeling approaches. Simulations of an age-structured population model suggest that the observed levels of seroprevalence are unlikely to be sustained by dispersal from supplemental feeding areas with relatively high seroprevalence or an older age structure. Increases in brucellosis seroprevalence and the total elk population size in areas with feeding grounds have not been statistically detectable. Meanwhile, the rate of seroprevalence increase outside the feeding grounds was related to the population size and density of each herd unit. Therefore, the data suggest that enhanced elk-to-elk transmission in free-ranging populations may be occurring due to larger winter elk aggregations. Elk populations inside and outside of the GYE that traditionally did not maintain brucellosis may now be at risk due to recent population increases. In particular, some neighboring populations of Montana elk were 5-9 times larger in 2007 than in the 1970s, with some aggregations comparable to the Wyoming feeding-ground populations. Addressing the unintended consequences of these increasing populations is complicated by limited hunter access to private lands, which places many ungulate populations out of administrative control. Agency-landowner hunting access partnerships and the protection of large predators are two management strategies that may be used to target high ungulate densities in private refuges and reduce the current and future burden of disease.

Dynamics of a multihost pathogen in a carnivore community.

1. We provide the first theoretical analysis of multihost disease dynamics to incorporate social behaviour and contrasting rates of within- and between-group disease transmission. 2. A stochastic susceptible-infected-recovered (SIR) model of disease transmission involving one to three sympatric species was built to mimic the 1994 Serengeti canine distemper virus outbreak, which infected a variety of carnivores with widely ranging social structures. The model successfully mimicked the erratic and discontinuous spatial pattern of lion deaths observed in the Serengeti lions under a reasonable range of parameter values, but only when one to two other species repeatedly transmitted the virus to the lion population. 3. The outputs from our model suggest several principles that will apply to most directly transmitted multihost pathogens: (i) differences in social structure can significantly influence the size, velocity and spatial pattern of a multihost epidemic; and (ii) social structures that permit higher intraspecific neighbour-to-neighbour transmission are the most likely to transmit disease to other species; whereas (iii) species with low neighbour-to-neighbour intraspecific transmission suffer the greatest costs from interspecific transmission.

Climate disruption and parasite-host dynamics: patterns and processes associated with warming and the frequency of extreme climatic events.

Levels of parasitism and the dynamics of helminth systems is subject to the impact of environmental conditions such that we may expect long term increases in temperature will increase the force of infection and the parasite's basic reproduction number, R0. We postulate that an increase in the force of infection will only lead to an increase in mean intensity of adults when adult parasite mortality is not determined by acquired immunity. Preliminary examination of long term trends of parasites of rabbits and grouse confirm these predictions. Parasite development rate increases with temperature and while laboratory studies indicate this is linear some recent studies indicate that this may be non-linear and would have an important impact on R0. Warming would also reduce the selective pressure for the development of arrestment and this would increase R0 so that in systems like the grouse and Trichostrongylus tenuis this would increase the instability and lead to larger disease outbreaks. Extreme climatic events that act across populations appear important in synchronizing transmission and disease outbreaks, so it is speculated that climate disruption will lead to increased frequency and intensity of disease outbreaks in parasite populations not regulated by acquired immunity.

Sexually transmitted diseases in polygynous mating systems: prevalence and impact on reproductive success.

Studies of disease in relation to animal mating systems have focused on sexual selection and the evolution of sexual reproduction. Relatively little work has examined other aspects of ecological and evolutionary relationships between host social and sexual behaviour, and dynamics and prevalence of infectious diseases; this is particularly evident with respect to sexually transmitted diseases (STDs). Here, we use a simulation approach to investigate rates of STD spread in host mating systems ranging from permanent monogamy to serial polygyny or polyandry and complete promiscuity. The model assumes that one sex (female) is differentially attracted to the other, such that groups of varying size are formed within which mating and disease transmission occur. The results show that equilibrium disease levels are generally higher in females than males and are a function of variance in male mating success and the likelihood of a female switching groups between mating seasons. Moreover, initial rates of disease spread (determining whether an STD establishes in a population) depend on patterns of host movement between groups, variance in male mating success and host life history (e.g. mortality rates). Male reproductive success can be reduced substantially by a sterilizing STD and this reduction is greater in males that are more 'attractive' to females. In contrast, females that associate with more attractive males have lower absolute fitness than females associating with less attractive males. Thus, the potential for STDs to act as a constraint on directional selection processes leading to polygyny (or polyandry) is likely to depend on the details of mate choice and group dynamics.

Effective time to satisfactory intubation conditions after administration of rocuronium in adults. Comparison of propofol and thiopentone for rapid sequence induction of anaesthesia.

We determined the effective time to satisfactory intubation conditions after the administration of rocuronium 0.6 mg.kg-1 to 120 unpremedicated adult patients anaesthetised with propofol 2.5 mg.kg-1 or thiopentone 5 mg.kg-1. Intubation conditions were assessed in 10 subgroups of 12 patients at 30, 40, 50, 60 and 70 s. The effective times to satisfactory intubation conditions in 50 and 90% of patients were obtained by the method of maximum likelihood after log time-probit response transformations. Intubation conditions after induction of anaesthesia with propofol were satisfactory in 5/12 patients at 30 s, 7/12 at 40 s, 10/12 at 50 s, 11/12 at 60 s and 11/12 at 70 s compared with 1/12 patients at 30 s, 2/12 at 40 s, 5/12 at 50 s, 7/12 at 60 s and 8/12 at 70 s after induction with thiopentone. The effective times to satisfactory intubation conditions in 50% and 90% (95% confidence intervals) of patients after rocuronium 0.6 mg.kg-1 were 34 (26-40) s and 61 (50-81) s in patients given propofol compared with 57 (48-69) s and 101 (79-167) s in patients given thiopentone. We conclude that rocuronium 0.6 mg.kg-1 may be a suitable alternative to suxamethonium during rapid sequence induction of anaesthesia with propofol in situations where suxamethonium is contraindicated.

Prevention of population cycles by parasite removal.

The regular cyclic fluctuations in vertebrate numbers have intrigued scientists for more than 70 years, and yet the cause of such cycles has not been clearly demonstrated. Red grouse populations in Britain exhibit cyclic fluctuations in abundance, with periodic crashes. The hypothesis that these fluctuations are caused by the impact of a nematode parasite on host fecundity was tested by experimentally reducing parasite burdens in grouse. Treatment of the grouse population prevented population crashes, demonstrating that parasites were the cause of the cyclic fluctuations.

Patterns of macroparasite aggregation in wildlife host populations.

Frequency distributions from 49 published wildlife host-macroparasite systems were analysed by maximum likelihood for goodness of fit to the negative binomial distribution. In 45 of the 49 (90%) data-sets, the negative binomial distribution provided a statistically satisfactory fit. In the other 4 data-sets the negative binomial distribution still provided a better fit than the Poisson distribution, and only 1 of the data-sets fitted the Poisson distribution. The degree of aggregation was large, with 43 of the 49 data-sets having an estimated k of less than 1. From these 49 data-sets, 22 subsets of host data were available (i.e. host data could be divided by either host sex, age, where or when hosts were sampled). In 11 of these 22 subsets there was significant variation in the degree of aggregation between host subsets of the same host-parasite system. A common k estimate was always larger than that obtained with all the host data considered together. These results indicate that lumping host data can hide important variations in aggregation between hosts and can exaggerate the true degree of aggregation. Wherever possible common k estimates should be used to estimate the degree of aggregation. In addition, significant differences in the degree of aggregation between subgroups of host data, were generally associated with significant differences in both mean parasite burdens and the prevalence of infection.

Transmission dynamics and host-parasite interactions of Trichostrongylus tenuis in red grouse (Lagopus lagopus scoticus).

Two components of the transmission dynamics of Trichostrongylus tenuis in red grouse are examined and quantified, namely parasite transmission rate and density-dependent reductions in egg production. Age-intensity data for birds of known age suggest that the rate of parasite uptake increases during the first 6 mo of a bird's life and this increase reflects an increase in feeding rate with age and exhibits no signs of self-cure. Analysis of these age-intensity curves permits us to estimate the transmission rate of the free-living infective stages. Reinfection rates of adults treated to reduce parasite intensities were not significantly different from infection rates of naive immature grouse. Secondary infections continued to rise over a period of 18 mo and this suggests that there is no strong host-mediated response against the parasite. Any density-dependent reduction in parasite fecundity is probably very weak and would act through interspecific competition between parasites. Initial analysis of worm egg production in relation to the intensity of worm infection found weak evidence of density-dependent suppression of egg production at high worm intensities. However, a more rigorous analysis found that such a relationship suffered from Type I errors and was a consequence of the aggregated distribution of the parasites. Any density-dependent suppression of parasite egg production is too weak to be detected and would only occur at high worm intensities. The potential density-dependent reductions in fecundity on the population dynamics of T. tenuis and red grouse are examined using a mathematical model. The model suggests that the presence of density-dependent reductions in worm fecundity could produce significant reductions in the propensity of the grouse-nematode system to exhibit population cycles. The sustained cycles observed in the long-term dynamics of the grouse populations in the study area suggest that density-dependent reductions in worm fecundity and establishment are either absent or only operating at levels that are not detectable in field studies.

Allometry and simple epidemic models for microparasites.

Simple mathematical models for microparasites offer a useful way to examine the population dynamics of different viral and bacterial pathogens. One constraint in applying these models in free-living host populations is the paucity of data with which to estimate transmission rates. Here we recast a standard epidemiological model by setting the birth and death rates of the host population and its density as simple allometric functions of host body weight. We then use standard threshold theorems for the model in order to estimate the minimum rate of transmission for the parasite to establish itself in a mammalian host population. Transmission rates that produce different comparable values of the parasites' basic reproductive number, RO, are themselves allometric functions of host body size. We have extended the model to show that hosts having different body sizes suffer epidemic outbreaks whose frequency scales with body size. The expected epidemic periods for pathogens in different mammalian populations correspond to cycles observed in free-living populations.

The population dynamics of communities of parasitic helminths.

This paper considers the dynamics of a host (animal) species that would grow exponentially in the absence of parasitism, and a community of parasite species that may regulate this growth. The model consists of a single differential equation for the host and one for each of the parasite species. This level of simplicity is achieved by assuming that each parasite species has a negative binomial distribution within the host population, with either zero covariance between the species (exploitation competition), or a specified covariance structure (interference competition). Conditions on the model parameters that determine the abundance of the different species are formulated, as are conditions that determine when a parasite species can invade a community and when a species is likely to be squeezed out. The results show that highly aggregated parasite species are more likely to coexist, but are less able to regulate their host population. A negative correlation between the distributions of the parasite species enhances both their ability to coexist and their ability to regulate the host population. The results of this analysis apply more generally to other systems where communities of exploiter species coexist on discretely distributed hosts, for example, insects on plants.

Behavioural constraints and conservation biology: Conspecific attraction and recruitment.

Wildlife managers have devoted considerable time and research to determine how animals judge the suitability and quality of their habitat. These analyses typically center on habitat characteristics and often produce equivocal results. An additional method of habitat assessment is to examine the extent to which animals, and particularly birds, use conspecifics as cues to establishing breeding and feeding territories. Conspecifics can be indicators of habitat quality, or they might intrinsically affect reproductive success. Here we discuss the implications of conspecific attraction for biologists who wish to conserve endangered species that are living in fragmented habitats.