Search

The spatial availability of social resources is speculated to structure animal movement decisions, but the effects of social resources on animal movements are difficult to identify because social resources are rarely measured. Here, we assessed whether varying availability of a key social resource—access to receptive mates—produces predictable changes in movement decisions among bighorn sheep in Nevada, the United States. We compared the probability that males made long-distance 'foray' movements, a critical driver of connectivity, across three ecoregions with varying temporal duration of a socially mediated factor, breeding season. We used a hidden Markov model to identify foray events and then quantified the effects of social covariates on the probability of foray using a discrete choice model. We found that males engaged in forays at higher rates when the breeding season was short, suggesting that males were most responsive to the social resource when its existence was short lived. During the breeding season, males altered their response to social covariates, relative to the non-breeding season, though patterns varied, and age was associated with increased foray probability. Our results suggest that animals respond to the temporal availability of social resources when making the long-distance movements that drive connectivity. This article is part of the theme issue 'The spatial–social interface: a theoretical and empirical integration'. [ABSTRACT FROM AUTHOR]

Epizootic events of pneumonia, presumably caused by Mycoplasma ovipneumoniae, in bighorn sheep (Ovis canadensis) have been observed in the western United States and Canada. Until recently, it was thought that populations of Mexican (O. c. mexicana) and Nelson's (O. c. nelsoni) desert bighorn sheep in Texas, USA, had not been exposed to Mycoplasma. Evidence of disease and potential population decline from outbreaks of M. ovipneumoniae are now known from several populations across the Trans‐Pecos Ecoregion with documented instances of pneumonia and bluetongue in desert bighorn sheep from the Van Horn Mountains and Black Gap Wildlife Management Area. These disease events, especially those in 2019–2021, may be a result of increasing populations of aoudad (Ammotragus lervia), an introduced and invasive ungulate, in the region. With large population sizes and similar movement patterns as desert bighorn sheep, aoudad potentially are the reservoirs for bacterial and viral diseases, such as pneumonia and bluetongue, and are possibly contributing to the decline of desert bighorn sheep. Herein, we optimized the multi‐locus sequence typing (MLST) with modifications in the Taq polymerase and annealing temperatures to determine the genetic identity of Mycoplasma strains or species within the nasal passages of desert bighorn sheep and aoudad in the Trans‐Pecos Ecoregion of Texas. Four loci (small ribosomal unit, 16S; 16S‐23S intergenic spacer region, IGS; RNA polymerase B, rpoB; gyrase B, gyrB) were characterized using MLST. Based on results from the modified MLST technique, we identified 9 desert bighorn sheep and 5 aoudad with M. ovipneumoniae, 9 aoudad with bacterial sequences genetically similar to M. conjunctivae, and 10 aoudad with bacterial sequences genetically similar M. hyopneumoniae. Of these, 9 aoudad possessed bacterial sequences genetically similar to both M. conjunctivae and M. hyopneumoniae. Among the 4 diagnostic loci, genetic divergence of M. ovipneumoniae ranged from 0.00–0.90% among desert bighorn sheep and aoudad. Future sampling efforts of seemingly asymptomatic aoudad, and asymptomatic, visibly sick, or deceased desert bighorn sheep, are important to monitor the spread of disease in desert bighorn sheep populations across mountain ranges in western Texas. It is imperative that aoudad removal plans are implemented to reduce and eliminate current infections and putative transmission of M. ovipneumoniae, prevent future disease outbreaks of pneumonia, and ultimately conserve desert bighorn sheep for future generations. [ABSTRACT FROM AUTHOR]

Low lamb recruitment can be an obstacle to bighorn sheep (Ovis canadensis) conservation and restoration. Causes of abortion and neonate loss in bighorn sheep, which may affect recruitment, are poorly understood. Toxoplasma gondii is a major cause of abortion and stillbirth in domestic small ruminants worldwide, but no reports exist documenting abortion or neonatal death in bighorn sheep attributable to toxoplasmosis. Between March 2019 and May 2021, eight fetal and neonatal bighorn lamb cadavers from four western US states (Idaho, Montana, Nebraska, and Washington) were submitted to the Washington Animal Disease Diagnostic Laboratory for postmortem examination, histologic examination, and ancillary testing to determine the cause of abortion or neonatal death. Necrotizing encephalitis characteristic of toxoplasmosis was identified histologically in six of eight cases, and T. gondii infection was confirmed by PCR in five cases with characteristic lesions. Other lesions attributable to toxoplasmosis were pneumonia (3/5 cases) and myocarditis (2/5 cases). Protozoal cysts were identified histologically within brain, lung, heart, skeletal muscle, adipose tissue, or a combination of samples in all five sheep with PCR-confirmed T. gondii infections. Seroprevalence of T. gondii ranged from 40–81% of adult females sampled in the Washington population in October and November 2018–2021, confirming high rates of exposure before detection of Toxoplasma abortions in this study. Of 1,149 bighorn sheep postmortem samples submitted to Washington Animal Disease Diagnostic Laboratory between January 2000 and May 2021, 21 of which were from fetuses or neonates, a single case of chronic toxoplasmosis was diagnosed in one adult ewe. Recent identification of Toxoplasma abortions in bighorn sheep suggests that toxoplasmosis is an underappreciated cause of reproductive loss. Abortions and neonatal mortalities should be investigated through postmortem and histologic examination, particularly in herds that are chronically small, demographically stagnant, or exhibit reproductive rates lower than expected. [ABSTRACT FROM AUTHOR]

Ecological context—the biotic and abiotic environment, along with its influence on population mixing dynamics and individual susceptibility—is thought to have major bearing on epidemic outcomes. However, direct comparisons of wildlife disease events in contrasting ecological contexts are often confounded by concurrent differences in host genetics, exposure histories, or pathogen strains. Here, we compare disease dynamics of a Mycoplasma ovipneumoniae spillover event that affected bighorn sheep populations in two contrasting ecological contexts. One event occurred on the herd's home range near the Rio Grande Gorge in New Mexico, while the other occurred in a captive facility at Hardware Ranch in Utah. While data collection regimens varied, general patterns of antibody signal strength and symptom emergence were conserved between the two sites. Symptoms appeared in the captive setting an average of 12.9 days postexposure, average time to seroconversion was 24.9 days, and clinical signs peaked at approximately 36 days postinfection. These patterns were consistent with serological testing and subsequent declines in symptom intensity in the free‐ranging herd. At the captive site, older animals exhibited more severe declines in body condition and loin thickness, higher symptom burdens, and slower antibody response to the pathogen than younger animals. Younger animals were more likely than older animals to clear infection by the time of sampling at both sites. The patterns presented here suggest that environment may not be a major determinant of epidemiological outcomes in the bighorn sheep—M. ovipneumoniae system, elevating the possibility that host‐ or pathogen‐factors may be responsible for observed variation. [ABSTRACT FROM AUTHOR]

Controlling persistent infectious disease in wildlife populations is an ongoing challenge for wildlife managers and conservationists worldwide, and chronic diseases in particular remain a pernicious problem.Here, we develop a dynamic pathogen transmission model capturing key features of Mycoplasma ovipneumoniae infection, a major cause of population declines in North American bighorn sheep Ovis canadensis. We explore the effects of model assumptions and parameter values on disease dynamics, including density‐ versus frequency‐dependent transmission, the inclusion of a carrier class versus a longer infectious period, host survival rates, disease‐induced mortality and recovery rates and the epidemic growth rate. Along the way, we estimate the basic reproductive ratio, R0, for M. ovipneumoniae in bighorn sheep to fall between approximately 1.36 and 1.74.We apply the model to compare efficacies across a suite of management actions following an epidemic, including test‐and‐remove, depopulation‐and‐reintroduction, range expansion, herd augmentation and density reduction.Our results suggest that test‐and‐remove, depopulation‐and‐reintroduction and range expansion could help persistently infected bighorn sheep herds recovery following an epidemic. By contrast, augmentation could lead to worse outcomes than those expected in the absence of management. Other management actions that improve host survival or reduce disease‐induced mortality are also likely to improve population size and persistence of chronically infected herds.Synthesis and applications. Dynamic transmission models like the one employed here offer a structured, logical approach for exploring hypotheses, planning field experiments and designing adaptive management. We find that management strategies that removed infected animals or isolated them within a structured metapopulation were most successful at facilitating herd recovery from a low‐prevalence, chronic pathogen. Ideally, models like ours should operate iteratively with field experiments to triangulate on better approaches for managing wildlife diseases. [ABSTRACT FROM AUTHOR]

Secondary sexual traits (e.g., horns and antlers) have ecological and evolutionary importance and are of management interest for game species. Yet, how these traits respond to emerging threats like infectious disease remains underexplored. Infectious pneumonia threatens bighorn sheep (Ovis canadensis) populations across North America and we hypothesized it may also reduce horn growth in male sheep. We assess the effect of pneumonia on horn size in male bighorn sheep using 12 herd datasets from across the western United States that had horn growth and disease data. Disease resulted in 12–35% reduction in increment (yearly) length and 3–13% reduction in total horn length in exposed individuals. The disease effect was prolonged when pathogens continued to circulate in sheep populations. Further, disease likely delays the age at which horns reach ¾‐curl and prevents achievement of full‐curl. This is further evidenced with 6 of the 12 herds experiencing an increase in average age at harvest following die‐off events. [ABSTRACT FROM AUTHOR]

A respiratory disease epizootic at the National Bison Range (NBR) in Montana in 2016–2017 caused an 85% decline in the bighorn sheep population, documented by observations of its unmarked but individually identifiable members, the subjects of an ongoing long‐term study. The index case was likely one of a small group of young bighorn sheep on a short‐term exploratory foray in early summer of 2016. Disease subsequently spread through the population, with peak mortality in September and October and continuing signs of respiratory disease and sporadic mortality of all age classes through early July 2017. Body condition scores and clinical signs suggested that the disease affected ewe groups before rams, although by the end of the epizootic, ram mortality (90% of 71) exceeded ewe mortality (79% of 84). Microbiological sampling 10 years to 3 months prior to the epizootic had documented no evidence of infection or exposure to Mycoplasma ovipneumoniae at NBR, but during the epizootic, a single genetic strain of M. ovipneumoniae was detected in affected animals. Retrospective screening of domestic sheep flocks near the NBR identified the same genetic strain in one flock, presumptively the source of the epizootic infection. Evidence of fatal lamb pneumonia was observed during the first two lambing seasons following the epizootic but was absent during the third season following the death of the last identified M. ovipneumoniae carrier ewe. Monitoring of life‐history traits prior to the epizootic provided no evidence that environmentally and/or demographically induced nutritional or other stress contributed to the epizootic. Furthermore, the epizootic occurred despite proactive management actions undertaken to reduce risk of disease and increase resilience in this population. This closely observed bighorn sheep epizootic uniquely illustrates the natural history of the disease including the (presumptive) source of spillover, course, severity, and eventual pathogen clearance. [ABSTRACT FROM AUTHOR]

ABSTRACT Infectious disease contributed to historical declines and extirpations of bighorn sheep ( Ovis canadensis) in North America and continues to impede population restoration and management. Reports of pneumonia outbreaks in free-ranging bighorn sheep following contact with domestic sheep have been validated by the results of 13 captive commingling experiments. However, ecological and etiological complexities still hinder our understanding and control of respiratory disease in wild sheep. In this paper, we review the literature and summarize recent data to present an overview of the biology and management of pneumonia in bighorn sheep. Many factors contribute to this population-limiting disease, but a bacterium ( Mycoplasma ovipneumoniae) host-specific to Caprinae and commonly carried by healthy domestic sheep and goats, appears to be a primary agent necessary for initiating epizootics. All-age epizootics are usually associated with significant population declines, but mortality rates vary widely and factors influencing disease severity are not well understood. Once introduced, M. ovipneumoniae can persist in bighorn sheep populations for decades. Carrier females may transmit the pathogen to their susceptible lambs, triggering fatal pneumonia outbreaks in nursery groups, which limit recruitment and slow or prevent population recovery. The demographic costs of disease persistence can be equal to or greater than the impacts of the initial epizootic. Strain typing suggests that spillover of M. ovipneumoniae into bighorn sheep populations from domestic small ruminants is ongoing and that consequences of spillover are amplified by movements of infected bighorn sheep across populations. Therefore, current disease management strategies focus on reducing risk of spillover from reservoir populations of domestic sheep and goats and on limiting transmission among bighorn sheep. A variety of techniques are employed to prevent contacts that could lead to transmission, including limiting the numbers and distribution of both wild and domestic species. No vaccine or antibiotic treatment has controlled infection in domestic or wild sheep and to date, management actions have been unsuccessful at reducing morbidity, mortality, or disease spread once a bighorn sheep population has been exposed. More effective strategies are needed to prevent pathogen introduction, induce disease fadeout in persistently infected populations, and promote population resilience across the diverse landscapes bighorn sheep inhabit. A comprehensive examination of disease dynamics across populations could help elucidate how disease sometimes fades out naturally and whether population resilience can be increased in the face of infection. Cross-jurisdictional adaptive management experiments and transdisciplinary collaboration, including partnerships with members of the domestic sheep and goat community, are needed to speed progress toward sustainable solutions to protect and restore bighorn sheep populations. © 2017 The Wildlife Society. [ABSTRACT FROM AUTHOR]

Superspreading, the phenomenon where a small proportion of individuals contribute disproportionately to new infections, has profound effects on disease dynamics. Superspreading can arise through variation in contacts, infectiousness or infectious periods. The latter has received little attention, yet it drives the dynamics of many diseases of critical public health, livestock health and conservation concern. Here, we present rare evidence of variation in infectious periods underlying a superspreading phenomenon in a free-ranging wildlife system. We detected persistent infections of Mycoplasma ovipneumoniae, the primary causative agent of pneumonia in bighorn sheep ( Ovis canadensis), in a small number of older individuals that were homozygous at an immunologically relevant genetic locus. Interactions among age-structure, genetic composition and infectious periods may drive feedbacks in disease dynamics that determine the magnitude of population response to infection. Accordingly, variation in initial conditions may explain divergent population responses to infection that range from recovery to catastrophic decline and extirpation. [ABSTRACT FROM AUTHOR]

Understanding both contact and probability of transmission given contact are key to managing wildlife disease. However, wildlife disease research tends to focus on contact heterogeneity, in part because the probability of transmission given contact is notoriously difficult to measure. Here, we present a first step towards empirically investigating the probability of transmission given contact in free-ranging wildlife., We used measured contact networks to test whether bighorn sheep demographic states vary systematically in infectiousness or susceptibility to Mycoplasma ovipneumoniae, an agent responsible for bighorn sheep pneumonia., We built covariates using contact network metrics, demographic information and infection status, and used logistic regression to relate those covariates to lamb survival. The covariate set contained degree, a classic network metric describing node centrality, but also included covariates breaking the network metrics into subsets that differentiated between contacts with yearlings, ewes with lambs, and ewes without lambs, and animals with and without active infections., Yearlings, ewes with lambs, and ewes without lambs showed similar group membership patterns, but direct interactions involving touch occurred at a rate two orders of magnitude higher between lambs and reproductive ewes than between any classes of adults or yearlings, and one order of magnitude higher than direct interactions between multiple lambs., Although yearlings and non-reproductive bighorn ewes regularly carried M. ovipneumoniae, our models suggest that a contact with an infected reproductive ewe had approximately five times the odds of producing a lamb mortality event of an identical contact with an infected dry ewe or yearling. Consequently, management actions targeting infected animals might lead to unnecessary removal of young animals that carry pathogens but rarely transmit., This analysis demonstrates a simple logistic regression approach for testing a priori hypotheses about variation in the odds of transmission given contact for free-ranging hosts, and may be broadly applicable for investigations in wildlife disease ecology. [ABSTRACT FROM AUTHOR]

ABSTRACT Transmission of pathogens commonly carried by domestic sheep and goats poses a serious threat to bighorn sheep ( Ovis canadensis) populations. All-age pneumonia die-offs usually ensue, followed by asymptomatic carriage of Mycoplasma ovipneumoniae by some of the survivors. Lambs born into these chronically infected populations often succumb to pneumonia, but adults are usually healthy. Surprisingly, we found that introduction of a new genotype (strain) of M. ovipneumoniae into a chronically infected bighorn sheep population in the Hells Canyon region of Washington and Oregon was accompanied by adult morbidity (100%) and pneumonia-induced mortality (33%) similar to that reported in epizootics following exposure of naïve bighorn sheep. This suggests an immune mismatch occurred that led to ineffective cross-strain protection. To understand the broader context surrounding this event, we conducted a retrospective analysis of M. ovipneumoniae strains detected in 14 interconnected populations in Hells Canyon over nearly 3 decades. We used multi-locus sequence typing of DNA extracts from 123 upper respiratory tract and fresh, frozen, and formalin-fixed lung samples to identify 5 distinct strains of M. ovipneumoniae associated with all-age disease outbreaks between 1986 and 2014, a pattern consistent with repeated transmission events (spillover) from reservoir hosts. Phylogenetic analysis showed that the strain associated with the outbreak observed in this study was likely of domestic goat origin, whereas strains from other recent disease outbreaks probably originated in domestic sheep. Some strains persisted and spread across populations, whereas others faded out or were replaced. Lack of cross-strain immunity in the face of recurrent spillovers from reservoir hosts may account for a significant proportion of the disease outbreaks in bighorn sheep that continue to happen regularly despite a century of exposure to domestic sheep and goats. Strain-specific immunity could also complicate efforts to develop vaccines. The results of our study support existing management direction to prevent contacts that could lead to pathogen transmission from domestic small ruminants to wild sheep, even if the wild sheep have previously been exposed. Our data also show that under current management, spillover is continuing to occur, suggesting that enhanced efforts are indicated to avoid introducing new strains of M. ovipneumoniae into wild sheep populations. We recommend looking for new management approaches, such as clearing M. ovipneumoniae infection from domestic animal reservoirs in bighorn sheep range, and placing greater emphasis on existing strategies to elicit more active cooperation by the public and to increase vigilance on the part of resource managers. © 2016 The Wildlife Society. [ABSTRACT FROM AUTHOR]

Bighorn sheep mortality related to pneumonia is a primary factor limiting population recovery across western North America, but management has been constrained by an incomplete understanding of the disease. We analysed patterns of pneumonia-caused mortality over 14 years in 16 interconnected bighorn sheep populations to gain insights into underlying disease processes., We observed four age-structured classes of annual pneumonia mortality patterns: all-age, lamb-only, secondary all-age and adult-only. Although there was considerable variability within classes, overall they differed in persistence within and impact on populations. Years with pneumonia-induced mortality occurring simultaneously across age classes (i.e. all-age) appeared to be a consequence of pathogen invasion into a naïve population and resulted in immediate population declines. Subsequently, low recruitment due to frequent high mortality outbreaks in lambs, probably due to association with chronically infected ewes, posed a significant obstacle to population recovery. Secondary all-age events occurred in previously exposed populations when outbreaks in lambs were followed by lower rates of pneumonia-induced mortality in adults. Infrequent pneumonia events restricted to adults were usually of short duration with low mortality., Acute pneumonia-induced mortality in adults was concentrated in fall and early winter around the breeding season when rams are more mobile and the sexes commingle. In contrast, mortality restricted to lambs peaked in summer when ewes and lambs were concentrated in nursery groups., We detected weak synchrony in adult pneumonia between adjacent populations, but found no evidence for landscape-scale extrinsic variables as drivers of disease., We demonstrate that there was a >60% probability of a disease event each year following pneumonia invasion into bighorn sheep populations. Healthy years also occurred periodically, and understanding the factors driving these apparent fade-out events may be the key to managing this disease. Our data and modelling indicate that pneumonia can have greater impacts on bighorn sheep populations than previously reported, and we present hypotheses about processes involved for testing in future investigations and management. [ABSTRACT FROM AUTHOR]

Group living facilitates pathogen transmission among social hosts, yet temporally stable host social organizations can actually limit transmission of some pathogens. When there are few between-subpopulation contacts for the duration of a disease event, transmission becomes localized to subpopulations. The number of per capita infectious contacts approaches the subpopulation size as pathogen infectiousness increases. Here, we illustrate that this is the case during epidemics of highly infectious pneumonia in bighorn lambs (Ovis canadensis). We classified individually marked bighorn ewes into disjoint seasonal subpopulations, and decomposed the variance in lamb survival to weaning into components associated with individual ewes, subpopulations, populations and years. During epidemics, lamb survival varied substantially more between ewe-subpopulations than across populations or years, suggesting localized pathogen transmission. This pattern of lamb survival was not observed during years when disease was absent. Additionally, group sizes in ewe-subpopulations were independent of population size, but the number of ewe-subpopulations increased with population size. Consequently, although one might reasonably assume that force of infection for this highly communicable disease scales with population size, in fact, host social behaviour modulates transmission such that disease is frequency-dependent within populations, and some groups remain protected during epidemic events. [ABSTRACT FROM AUTHOR]