Animal movements among habitat patches or populations are important for maintaining long-term genetic and demographic viability, but connectivity may also facilitate disease spread and persistence. Understanding factors that influence animal movements is critical to understanding potential transmission risk and persistence of communicable disease in spatially structured systems. We evaluated effects of sex, age and Mycoplasma ovipneumoniae infection status at capture on intermountain movements and seasonal movement rates observed in desert bighorn sheep ( Ovis canadensis nelsoni ) using global positioning system collar data from 135 individuals (27 males, 108 females) in 14 populations between 2013 and 2018, following a pneumonia outbreak linked to the pathogen M. ovipneumoniae in the Mojave Desert, California, USA. Based on logistic regression analysis, intermountain movements were influenced by sex, age and most notably, infection status at capture: males, older animals and uninfected individuals were most likely to make such movements. Based on multiple linear regression analysis, females that tested positive for M. ovipneumoniae at capture also had lower mean daily movement rates that were further influenced by season. Our study provides empirical evidence of a pathogenic infection decreasing an individual's future mobility, presumably limiting that pathogen's ability to spread, and ultimately influencing transmission risk within a spatially structured system.
Habitat fragmentation is an important driver of biodiversity loss and can be remediated through management actions aimed at maintenance of natural connectivity in metapopulations. Connectivity may protect populations from infectious diseases by preserving immunogenetic diversity and disease resistance. However, connectivity could exacerbate the risk of infectious disease spread across vulnerable populations. We tracked the spread of a novel strain of Mycoplasma ovipneumoniae in a metapopulation of desert bighorn sheep Ovis canadensis nelsoni in the Mojave Desert to investigate how variation in connectivity among populations influenced disease outcomes. M. ovipneumoniae was detected throughout the metapopulation, indicating that the relative isolation of many of these populations did not protect them from pathogen invasion. However, we show that connectivity among bighorn sheep populations was correlated with higher immunogenetic diversity, a protective immune response and lower disease prevalence. Variation in protective immunity predicted infection risk in individual bighorn sheep and was associated with heterozygosity at genetic loci linked to adaptive and innate immune signalling. Together, these findings may indicate that population connectivity maintains immunogenetic diversity in bighorn sheep populations in this system and has direct effects on immune responses in individual bighorn sheep and their susceptibility to infection by a deadly pathogen. Our study suggests that the genetic benefits of population connectivity could outweigh the risk of infectious disease spread and supports conservation management that maintains natural connectivity in metapopulations.
Peninsular bighorn sheep (Ovis canadensis nelsoni) are found exclusively in Southern California and Baja Mexico. They are federally endangered due to multiple threats, including introduced infectious disease. From 1981 - 2017, we conducted surveillance for 16 pathogens and estimated population sizes, adult survival, and lamb survival. We used mixed effects regression models to assess disease patterns at the individual and population levels. Pathogen infection/exposure prevalence varied both spatially and temporally. Our findings indicate that the primary predictor of individual pathogen infection/exposure was the region in which an animal was captured, implying that transmission is driven by local ecological or behavioral factors. Higher Mycoplasma ovipneumoniae seropositivity was associated with lower lamb survival, consistent with lambs having high rates of pneumonia-associated mortality, which may be slowing population recovery. There was no association between M. ovipneumoniae and adult survival. Adult survival was positively associated with population size and parainfluenza-3 virus seroprevalence in the same year, and orf virus seroprevalence in the previous year. Peninsular bighorn sheep are recovering from small population sizes in a habitat of environmental extremes, compounded by infectious disease. Our research can help inform future pathogen surveillance and population monitoring for the long-term conservation of this population.
A 2013 outbreak of respiratory disease in bighorn sheep from California's Mojave Desert metapopulation caused high mortality in at least one population. Subsequent PCR and strain-typing indicate widespread infection of a single strain of Mycoplasma ovipneumoniae throughout this region. Serosurvey of archived samples showed that some populations have had antibodies to M. ovipneumoniae since at least 1986, although pre-2013 strain-type data are unavailable.
ABSTRACTBeginning in the early 1900s, poly‐factorial, poly‐microbial pneumonia was identified as a disease affecting bighorn sheep (Ovis canadensis) and it continues to threaten bighorn populations, posing an ongoing management challenge. In May and June 2013, a pneumonia outbreak linked to the pathogen Mycoplasma ovipneumoniae led to an all‐age die‐off of desert bighorn sheep (O. c. nelsoni) at Old Dad Peak in the Kelso Mountains of the Mojave Desert in California, USA. Subsequently, we observed clinical signs of respiratory disease among bighorn sheep in multiple neighboring ranges. Our objective was to investigate post‐outbreak survival of adult female bighorn across 9 populations from 2014 to 2017 in the Mojave Desert and evaluate the relationship between M. ovipneumoniae infection and survival, while testing effects of range factors that could potentially influence differences in adult female survival (i.e., forage quality, winter precipitation, population abundance). We fitted adult females with radio‐collars following the outbreak and collected serum and nasal swab samples for competitive enzyme‐linked immunosorbent assay (cELISA) and polymerase chain reaction (PCR) testing to determine exposure and infection status at time of capture. We tracked survival of 115 adult females with radio‐collars and used the known‐fate model in Program MARK to evaluate effects and estimate survival from November 2013 to March 2017. Annual survival was negatively correlated with positive infection status at capture but varied across populations with respect to differences in range conditions. Summer and autumn forage quality, as represented by mean normalized difference vegetation index (NDVI) values for summer and autumn, was positively correlated with overwinter survival, whereas winter precipitation (a proxy for winter severity) was negatively correlated with overwinter survival. Population abundance was negatively correlated with annual survival, suggesting a potential density‐dependent effect. Model‐averaged annual survival estimates ranged from 0.700 ± 0.07 (SE) to 0.945 ± 0.026 for infected individuals and 0.896 ± 0.03 to 0.983 ± 0.011 for uninfected individuals. We conclude that summer and autumn forage quality, indexed by NDVI, may partially offset the negative effect associated with M. ovipneumoniae infection on host survival. Our survival modeling results suggest that chronic infection may have afflicted adult females that were PCR‐positive (i.e., infected with M. ovipneumoniae) at time of capture. We propose programmatic re‐testing of infected individuals to assess pathogen persistence at the individual level and evaluate whether selective culling might potentially help to reduce prevalence and transmission within populations. © 2020 The Wildlife Society.
Babesiosis is a potentially fatal tick-borne zoonotic disease caused by a species complex of blood parasites that can infect a variety of vertebrates, particularly dogs, cattle, and humans. In the United States, human babesiosis is caused by two distinct parasites, Babesia microti and Babesia duncani. The enzootic cycle of B. microti, endemic in the northeastern and upper midwestern regions, has been well characterised. In the western United States, however, the natural reservoir host and tick vector have not been identified for B. duncani, greatly impeding efforts to understand and manage this zoonotic disease. Two and a half decades after B. duncani was first described in a human patient in Washington State, USA, we provide evidence that the enzootic tick vector is the winter tick, Dermacentor albipictus, and the reservoir host is likely the mule deer, Odocoileus hemionus. The broad, overlapping ranges of these two species covers a large portion of far-western North America, and is consistent with confirmed cases of B. duncani in the far-western United States.
Ticks (Acari: Ixodidae) were collected from 44 desert bighorn sheep (Ovis canadensis) and 10 mule deer (Odocoileus hemionus) in southern California, US during health inspections in 2015-16. Specimens were identified and screened by PCR analysis to determine the presence and prevalence of Bartonella, Borrelia, and Rickettsia species in ticks associated with these wild ruminants. None of the 60 Dermacentor hunteri and 15 Dermacentor albipictus ticks tested yielded positive PCR results. Additional tick specimens should be collected and tested to determine the prevalence of these confirmed or suspected tickborne pathogens within ruminant populations.
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.
OBJECTIVE To present a case series of idiopathic lipoidal corneal degeneration in falcons. ANIMALS STUDIED Five falcons including three peregrine falcons (Falco peregrinus), one prairie falcon (Falco mexicanus), and one red-naped shaheen (Falco peregrinus babylonicus) were observed to develop slowly progressive corneal opacification that began at the temporal limbus and extended centripetally across the cornea over a period of years. Four of the birds were over 20 years old. PROCEDURES All animals underwent complete ophthalmic examinations. A red-naped shaheen underwent ocular imaging via spectral-domain optical coherence tomography. Two peregrine falcons were euthanized due to declining health, and their eyes were examined histologically. RESULTS The opacities were pale and granular, with frequent vascularization associated perilimbally. Diffuse neutral lipid was observed in stromal cells throughout the corneal stroma of both clear and opaque areas of the cornea, sparing only the acellular anterior limiting lamina. Clusters of cholesterol crystals surrounded by macrophages were present in the mid-stroma. Fibrosis was evident in a subepithelial location, which separated the epithelium from the anterior limiting lamina. Ultrastructurally, diffuse vacuolization of the keratocytes was observed. No other ophthalmic or systemic abnormalities were noted. CONCLUSIONS Results suggest that lipid degeneration occurs rarely in captive falcons of advanced age. The underlying cause is unclear. Though unsubstantiated, possible contributing factors include dyslipoproteinemia, corneal trauma, diet, and age-related alterations in corneal metabolism. The initiation of pathology at the temporal limbus, as well as slow progression, suggests that exposure contributes to the onset and progression of this unique keratopathy.
The authors captured bighorn sheep (Ovis canadensis) comprising a small population in the San Bernardino Mountains of California and evaluated the degree of infestation by mites of the genus Psoroptes for each individual. The animals were treated with two novel methods: amitraz-impregnated collars and cyfluthrin-impregnated ear tags and recaptured the following year to evaluate the effect of treatment. The authors compared data on degree of infestation for animals recaptured in the posttreatment year, detected no significant interyear differences in infestation severity scores among animals treated with amitraz or cyfluthrin, and could not detect any differences between treatment types. However, a significant (P<0.10) decreased pattern in severity scores from the beginning to the end of treatments was detected, suggesting a cumulative therapeutic value in repeated annual treatments across the 3-yr period. Additionally, the authors detected a lower median mite severity score between 2000 and a later capture in 2006. These positive outcomes may be the result of previous treatments during 2000-2002, but environmental covariates not accounted for could have been contributing factors. Avermectin drugs with longer release profiles may be a more effective treatment option in this and other small bighorn sheep populations that are compromised with mite infestations.
Vernon C. Bleich, Jeffrey T. Villepique, James H. Davis, Steven G. Torres, and Ben J. Gonzales California Department of Fish and Wildlife, Sierra Nevada Bighorn Sheep Recovery Program, 407 W. Line St., Bishop, CA 93514 USA California Department of Fish and Wildlife, P.O. Box 3222, Big Bear City, CA 92314 USA California Department of Fish and Wildlife, Wildlife Investigations Laboratory, 1701 Nimbus Rd., Rancho Cordova, CA 95670 USA
Whole blood and serum mineral concentrations were measured in diverse bighorn sheep (Ovis canadensis) metapopulations in California, and 90% reference intervals were determined. While there were some statistical differences between median concentrations among the different metapopulations, detected values were generally in good agreement with concentrations reported for other bighorn sheep populations and with reference ranges widely accepted for domestic sheep (Ovis aries). Although median whole blood selenium and serum copper concentrations were within adequate ranges reported for domestic sheep, some metapopulations had substantial numbers of individuals whose concentrations would be considered suboptimal for domestic sheep. There are a number of factors that can influence mineral concentrations in wildlife species such as bighorn sheep and that make the establishment of reference ranges challenging. However, the establishment of mineral reference ranges is important for such species, as their health and productivity are increasingly scrutinized and actively managed.
Brucella abortus has been an important wildlife disease issue for most of the last century, especially because wildlife species are considered to be important disease reservoirs for cattle. Diagnostic uncertainty, caused in part by cross-reactions of antibodies to environmental pathogens such as Yersinia enterocolitica O:9 on standard Brucella serology, has exacerbated the challenges of managing the disease and has highlighted the need for test validation in wildlife species. The western immunoblot was evaluated for use in detecting B. abortus exposure in elk (Cervus elaphus) and for ruling out exposure to cross-reacting bacteria. Samples collected from 2003 to 2006, including 54 female and immature elk from four different elk herds, were tested using standard Brucella serologic methods (card, rapid automated presumptive [RAP], and rivanol tests), as well as the western immunoblot. Samples (n=28) from animals known to be naturally infected with B. abortus biovar 1 served as positive controls. For presumed negative samples, sera (n=26) were collected from two elk herds in which negative serologic tests, and the absence of clinical signs of disease such as abortions, supported Brucella-negative classification. In addition to these study samples, serologic data from 12 tule elk (Cervus elaphus nannodes) were provided from the California Department of Fish and Game in order to illustrate a field application of the western blot. The western immunoblot had the highest sensitivity (1.0; % 0.899-1.0) and specificity (1.0; 0.891-1.0) among all tests used in the study. The Kappa statistic for agreement between the western blot and the card, rivanol, and RAP tests were 0.701, 0.808, and 0.921, respectively, showing good to excellent agreement with the standard diagnostic tests currently in use. Although the western immunoblot is more expensive and time intensive than other tests, in this limited study, it was shown to be reliable for establishing and confirming B. abortus disease status in elk. In addition to this study, subsequent applications of the western blot assay have been successful in detecting Yersinia sp. exposure in elk after their antibodies cross-reacted on standard Brucella serology.
The prevalence and phenotypic variability of Pasteurella and Mannheimia isolates from Sierra Nevada bighorn sheep (Ovis canadensis sierrae), White Mountain bighorn sheep (Ovis canadensis nelsoni), and domestic sheep (Ovis aries) from California, USA, were compared. The White Mountain bighorn sheep population had a recent history of pneumonia-associated mortality, whereas the Sierra Nevada bighorn sheep population had no recent history of pneumonia-associated mortality. The domestic sheep flocks were pastured in areas geographically near both populations but were not known to have direct contact with either bighorn sheep population. Oropharyngeal swab samples were collected from healthy domestic and bighorn sheep and cultured to characterize bacterial species, hemolysis, biogroups, and biovariants. Pasteurella trehalosi and Mannheimia haemolytica were detected in all of the study populations, but the relative proportion of each bacterial species differed among sheep populations. Pasteurella trehalosi was more common than M. haemolytica in the bighorn sheep populations, whereas the opposite was true in domestic sheep. Mannheimia haemolytica was separated into 11 biogroups, and P. trehalosi was characterized into two biogroups. Biogroup distributions for M. haemolytica and P. trehalosi differed among the three populations; however, no difference was detected for the distribution of P. trehalosi biogroups between the Sierra Nevada bighorn sheep and domestic sheep. The prevalence odds ratios (pOR) for the distribution of M. haemolytica biogroups suggested little difference between White Mountain bighorn sheep and domestic sheep compared with Sierra Nevada bighorn sheep and domestic sheep, although these comparisons had relatively large confidence intervals for the point estimates. Hemolytic activity of the isolates was not different among the sheep populations for M. haemolytica but was different for P. trehalosi. No clear evidence of association was found in the Pasteurellaceae distribution between the White Mountains bighorn sheep and domestic sheep.
We studied mountain sheep in the vicinity of three high-wall limestone mines in San Bernardino County, CA, USA to evaluate factors that influenced habitat use and, specifically, to investigate the influence of mining activity on distribution of those specialized ungulates. We used aerial telemetry data to estimate a resource selection function by fitting a logistic regression model and then comparing environmental characteristics at observed sheep locations to those at random locations. Distribution of mountain sheep was most influenced by a fire in 1999 that resulted in an area they avoided. Mountain sheep used steeper slopes, areas of lower terrain roughness, higher elevations, and areas closer to escape terrain than were random points. In contrast, sheep avoided areas near roads (federal and state highways, local roads, and off-road vehicle trails) but used areas near hiking trails and a railway. Water sources had the smallest effect of the factors considered, with sheep being associated with areas further from water points than were random locations. The disturbed area associated with the mines had a moderate influence on distribution, with sheep being associated with areas closer to the mine than were random points. Mining activities can alter terrain features and vegetation structure or composition in a way that promotes occupancy by sheep if they create steep slopes and rugged terrain (escape terrain) or reduce vegetation density or height (i.e., improve visibility). Whether increased occupancy reflects a benefit depends on the demographic responses of those sheep to the resources and conditions available on mine sites; information about those responses remains lacking.
Despite evidence that domestic sheep diseases threaten the persistence of bighorn sheep populations, the economic consequences of restricting domestic sheep grazing has polarized the debate, with some arguing that disease risk posed by domestic sheep has been exaggerated and grazing restrictions should be eased. We constructed a model to assess how different management strategies (grazing allotment closures, grazing time reductions, and reduced probability of stray domestic sheep) affect the risk of respiratory disease transmission from domestic sheep to endangered Sierra Nevada bighorn sheep, and to predict population-level impacts of an outbreak. Even when management strategies reduced risk of interspecies contact to less than 2% per year, our model predicted a 50% probability of a catastrophic respiratory disease outbreak during the next 10 bighorn sheep generations. If an outbreak occurs in the near future, the model predicts that the smallest Sierra Nevada bighorn sheep population would have a 33% probability of quasi-extinction. To eliminate all risk of contact and potential disease transmission, domestic sheep cannot be grazed on allotments that overlap with areas utilized by Sierra Nevada bighorn sheep. Where wildlife and domestic animal populations share limited habitat, and there is documented evidence of a substantial disease threat and extinction risk, stakeholders must recognize that the only way to eliminate contact and risk of disease transmission is to give priority to one species or the other. If conservation is the priority, difficult decisions will need to be made to balance trade-offs between economic livelihoods and species conservation.
Between 2 August and 22 September 2000, 37 hunter-killed tule elk (Cervus elaphus nannodes) were evaluated at the Grizzly Island Wildlife Area, California, USA, for evidence of paratuberculosis. Elk were examined post-mortem, and tissue and fecal samples were submitted for radiometric mycobacterial culture. Acid-fast isolates were identified by a multiplex polymerase chain reaction (PCR) that discriminates among members of the Mycobacterium avium complex (MAC). Histopathologic evaluations were completed, and animals were tested for antibodies using a Johne's enzyme–linked immunosorbent assay (ELISA) and agar gel immunodiffusion. In addition, 104 fecal samples from tule elk remaining in the herd were collected from the ground and submitted for radiometric mycobacterial culture. No gross lesions were detected in any of the hunter-killed animals. Mycobacterium avium subsp. paratuberculosis (MAP) was cultured once from ileocecal tissue of one adult elk and was determined to be a strain (A18) found commonly in infected cattle. One or more isolates of Mycobacterium avium subsp. avium (MAA) were isolated from tissues of five additional adult elk. Gastrointestinal tract and lymph node tissues from 17 of the 37 elk (46%) examined had histopathologic lesions commonly seen with mycobacterial infection; however, acid-fast bacteria were not observed. All MAC infections were detected from adult elk (P = 0.023). In adult elk, a statistically significant association was found between MAA infection and ELISA sample-to-positive ratio (S/P)≥0.25 (P=0.021); four of five MAA culture–positive elk tested positive by ELISA. Sensitivity and specificity of ELISA S/P≥0.25 for detection of MAA in adult elk were 50% and 93%, respectively. No significant associations were found between MAC infection and sex or histopathologic lesions. Bacteriologic culture confirmed infection with MAP and MAA in this asymptomatic tule elk herd. The Johne's ELISA was useful in signaling mycobacterial infection on a population basis but could not discriminate between MAA and MAP antibodies. The multiplex PCR was useful in discriminating among the closely related species belonging to MAC.
A portable modified steel mesh panel trap was developed to reduce injuries that wild pigs (Sus scrofa) suffered when captured in simple square panel traps. We avoided use of physical restraint for immobilizations by using a drug combination of Telazol(R) and xylazine hydrochloride (HCl) that was injected with blow darts. Heart rates, respiration rates, and body temperatures were all lower for animals immobilized with Telazol and xylazine HCl compared to relatively few animals that were restrained for immobilization with ketamine HCl and xylazine HCl. We captured an average of 2.5 pigs per trap night, and in 343 captures only 11 animals were injured. There were no capture-related mortalities. Also, the trap was efficient at capturing multiple-animal groups, and there was no difference in the mean size of pig groups captured using a rep-hinged gale ((x) over bar = 4.7, SE = 0.6) or a side-hinged squeeze gate ((x) over bar = 4.2, SE = 0.4).
We used a combination of Telazol® (3.3 mg/kg) and xylazine hydrochloride (1.6 mg/kg) to immobilize 144 wild pigs (Sus scrofa) with blow darts. This drug combination was safe and effective for rapidly immobilizing animals ranging in size from 34 to >170 kg and avoided difficulties associated with hand injections. For 123 single injection immobilizations, mean (±SD) induction times and effective handling periods averaged 5 (±2.5) and 52 (±18) min, respectively, and animals generally recovered for release within 120 min of initial injections. Animals that required two injections to immobilize (n = 21) received lower initial doses of Telazol® and xylazine hydrochloride than those immobilized with a single injection because of errors in estimating body sizes; we found that there was a threshold dose required to immobilize wild pigs from 2.8 to 3.3 mg/kg Telazol® and 1.4 to 1.6 mg/kg xylazine. Although neither age or sex influenced immobilization parameters, animals in good condition required longer to recover than those in poor condition. However, animals immobilized with two injections recovered as rapidly as those immobilized with a single injection. Heart rates and body temperatures declined slightly during the immobilization period, but respiration rates and blood oxygen saturation levels remained stable. In general, single injection immobilizations were preferable because they minimized problems associated with injecting partially immobilized animals. Because it was difficult to accurately estimate the sizes of large wild pigs (≥90 kg), and because wild pigs that were partially immobilized were difficult to handle, we recommend increasing the drug doses to 4 mg/kg Telazol® and 2 mg/kg xylazine hydrochloride when injecting relatively large animals to assure single injection immobilizations. Although recovery periods may be prolonged, higher doses of Telazol® and xylazine should be safe based on data from domestic pigs.