Weaning Conflict in Desert and Mountain Bighorn Sheep (Ovis canadensis): An Ecological Interpretation

Field data on weaning behavior in bighorn sheep (Ovis canadensis) were collected from populations in British Columbia, southern California, and one transplanted from the original B.C. study site to eastern Oregon. These areas were designated mountain, desert, and transplant, respectively. Seasons that were energetically stressful to lactating ewes were predicted. Summers were implicated for desert ewes; winters for ewes in colder and more seasonal northern environments. Although the temporal distribution of milk available to lambs varied between the three study populations, ewes generally weaned their lambs prior to the onset of the predicted stressful periods. In the desert, weaning occurred more abruptly than it did in the two more northern populations. The evolution of weaning strategies is discussed in subspecies of bighorn sheep based on proximate and ultimate factors.     

Epizootic Pneumonia of Bighorn Sheep following Experimental Exposure to Mycoplasma ovipneumoniae

Background

Bronchopneumonia is a population limiting disease of bighorn sheep (Ovis canadensis). The cause of this disease has been a subject of debate. Leukotoxin expressing Mannheimia haemolytica and Bibersteinia trehalosi produce acute pneumonia after experimental challenge but are infrequently isolated from animals in natural outbreaks. Mycoplasma ovipneumoniae, epidemiologically implicated in naturally occurring outbreaks, has received little experimental evaluation as a primary agent of bighorn sheep pneumonia.

Methodology/Principal Findings

In two experiments, bighorn sheep housed in multiple pens 7.6 to 12 m apart were exposed to M. ovipneumoniae by introduction of a single infected or challenged animal to a single pen. Respiratory disease was monitored by observation of clinical signs and confirmed by necropsy. Bacterial involvement in the pneumonic lungs was evaluated by conventional aerobic bacteriology and by culture-independent methods. In both experiments the challenge strain of M. ovipneumoniae was transmitted to all animals both within and between pens and all infected bighorn sheep developed bronchopneumonia. In six bighorn sheep in which the disease was allowed to run its course, three died with bronchopneumonia 34, 65, and 109 days after M. ovipneumoniae introduction. Diverse bacterial populations, predominantly including multiple obligate anaerobic species, were present in pneumonic lung tissues at necropsy.

Conclusions/Significance

Exposure to a single M. ovipneumoniae infected animal resulted in transmission of infection to all bighorn sheep both within the pen and in adjacent pens, and all infected sheep developed bronchopneumonia. The epidemiologic, pathologic and microbiologic findings in these experimental animals resembled those seen in naturally occurring pneumonia outbreaks in free ranging bighorn sheep.     

The ecology, structure and functions of social play in Bighorn sheep (Ovis canadensis)

Social play was studied in Bighorn sheep ( Ovis canadensis ) by examining its structure in two natural populations and by inferring functions through sexual differences in behaviour. Lambs discriminated the size of lambs with which they played. During contact play they responded most often to similarly sized individuals, whereas in locomotor play they chased and followed all sizes. Males engaged in more contact-oriented play than females. This evidence supports the idea that contact-oriented play developed in part to provide training for combatant skills in males and locomotor play as part of a general anitpredator strategy in both sexes. Social play was severely restricted in a desert environment due to risks and personal injury. The implications of lack of play in natural populations are discussed.     

Dynamics of Pneumonia in a Bighorn Sheep Metapopulation

Abstract: We investigated the dynamics of 8 populations of a bighorn sheep (Ovis canadensis) metapopulation in Hells Canyon, USA from 1997 to 2003. Pneumonia was the most common cause (43%) of adult mortality and the primary factor limiting population growth. Cougar (Puma concolor) predation was the second most-frequent source (27%) of adult mortality but did not reduce the rate of population growth significantly. Most pneumonia-caused mortality occurred in fall and early winter and most cougar predation occurred in late winter and early spring. Average annual survival of adult males (0.84) was lower than females (0.91). Pneumonia was the most common known cause of lamb mortality (86%), and pneumonia-related mortality was detected whenever summer lamb survival was <50%. Pneumonia-caused mortality rates in lambs were high from 21 days to 91 days of age and peaked at 42 days to 70 days. Summer pneumonia epizootics in lambs were independent of pneumonia-caused mortality in adults. Pneumonia-caused mortality occurred at the population level and was not synchronized geographically or temporally among populations. Although catastrophic all-age pneumonia-epizootics have previously been described in bighorn sheep, we found that chronic, although sporadic, pneumonia-caused mortality in adults and lambs can also have important effects on the dynamics of bighorn sheep populations.     

Survival of Adult Female Bighorn Sheep Following a Pneumonia Epizootic

Beginning 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.     

Exposure to Psoroptes sp. mites is common among bighorn sheep (Ovis canadensis) populations in California.

Sera (n = 806) from 50 populations of bighorn sheep (Ovis canadensis) in California (USA) were evaluated for antibodies to Psoroptes sp. mites using a kinetic enzyme-linked immunosorbent assay (ELISA). Test values for each sample were determined to be either positive or negative at each of two ELISA cutoff values that provided either 100% sensitivity (low cutoff) or 100% specificity (high cutoff), respectively. One hundred sixty-eight (20.8%) sera were seropositive at the low cutoff value, and 87 (10.8%) of these sera also were seropositive at the high cutoff value. Eleven populations were designated as scabies-suspect and 25 populations were designated as scabies-positive because they had at least one seropositive animal at the low and the high cutoff values, respectively. Based on these results, exposure to Psoroptes sp. mites appeared to be widely distributed among bighorn sheep populations from 1980 to 1990 and infested animals may have been present prior to 1980.     

Psoroptic scabies in Rocky Mountain bighorn sheep (Ovis canadensis canadensis) from Wyoming

Thirteen Rocky Mountain bighorn sheep (Ovis canadensis canadensis) with clinical signs of psoroptic scabies were captured in Wyoming. Signs included droopy ears, depilation on the head and neck, and exudate in the ears. Mites were identified as either Psoroptes cervinus or P. equi. Two ewes with scabies at the time of original capture had no clinical signs of mite infection 1 and 2 yr later.     

Birthdate and survival in bighorn lambs (Ovis canadensis)

The relationship between parturition date and lamb survival was investigated among individually marked bighorn ewes in south-western Alberta, Canada. Lambs were born from 17 May to 21 July. Most births occurred in the first two weeks of the lambing period. Lambs born in May enjoyed greater survival than lambs born in June and July. The viability of lambs born after 10 June was extremely low. Inadequate nutrition is suggested as the cause of greater mortality of lateborn lambs. Ewes that give birth late are at an earlier stage of lactation when forage quality declines and may not produce sufficient milk to ensure lamb survival. Their lambs have access to high-quality forage for a shortened period. The proportion of late lambs appeared to be correlated with ewe density in the winter range.     

Bighorn Sheep Abundance Following a Suspected Pneumonia Epidemic in Rocky Mountain National Park

Abstract: Anecdotal evidence of a pneumonia epizootic among bighorn sheep (Ovis canadensis canadensis) in Rocky Mountain National Park (RMNP), Colorado, USA, during the mid-1990s prompted park officials to examine the current condition of the herds. Here we present a mark—resight study design to estimate population abundance that, in many circumstances, is a reliable and cost-effective alternative to traditional mark—recapture or to indices of population abundance. We captured 59 adult females and radiocollared them via helicopter net-gunning during winter 2002–2003. From ground resighting surveys conducted May—September, we estimated the total RMNP bighorn population at 389.9 (SE = 34.9, CI = 327.2–464.6) in 2003 and 366.4 (SE = 34.7, CI = 304.4–441.0) in 2004. Previous abundance estimates suggest a park-wide decline has occurred between the late 1980s and the suspected pneumonia epidemic of the mid-1990s. Although the 2 years of data from our study are not enough to predict whether the herds are capable of recovering to previous levels, they provide park officials the tools necessary to make the most informed decisions for future monitoring and management of this fragile species.     

Response of vaccinated and unvaccinated bighorn sheep (Ovis canadensis canadensis) to experimental respiratory syncytial virus challenge

Five Rocky Mountain bighorn sheep (Ovis canadensis canadensis), approximately 5 mo old and without detectable antibody titers to respiratory syncytial virus (RSV), were assigned to two groups to study the effects of RSV challenge inoculation in vaccinated (n = 3) and unvaccinated (n = 2) bighorns. The three lambs vaccinated with a modified live bovine RSV vaccine developed a detectable antibody response to the vaccine. Vaccinated and unvaccinated lambs challenged with an ovine isolate of RSV developed increased levels of neutralizing antibody, but clinical signs of disease were not observed. Neutralizing antibody titers to RSV remained higher (2-4-fold) in vaccinated lambs over time when compared to unvaccinated lambs.     

Comparison of pulmonary defense mechanisms in Rocky Mountain bighorn (Ovis canadensis canadensis) and domestic sheep

Alveolar macrophages were obtained from Rocky Mountain bighorn sheep (Ovis canadensis canadensis) and domestic sheep for the purpose of comparing pulmonary host defense mechanisms in the two species. Specific variables studied included (1) characterization of the cell types present in the lung, (2) alveolar macrophage phagocytic and bactericidal functions, (3) measurement of protein levels in lavage fluid, and (4) measurement of cortisol levels in lavage fluid. While phagocytic cell populations were similar between bighorn and domestic sheep, a significantly higher percentage of lymphocytes were present in bighorns than domestics (20% in bighorn versus 6% in domestic sheep). Significant differences were not observed in the phagocytic or bactericidal functions of macrophages between the two species. Significant differences were not observed in either lavage fluid protein levels or in cortisol levels.     

Osteoporosis and cranial asymmetry in a mountain sheep (Ovis canadensis)

The skull of a mountain sheep (Ovis canadensis cremnobates) exhibiting osteoporosis was recovered from San Diego County, California. This specimen also exhibited lateral asymmetry of the cranium and rostrum, malocclusion of molars, and mandibular asymmetry. Investigators are cautioned about potentially confusing lesions associated with chronic frontal sinusitis with lesions of osteoporosis in mountain sheep. To our knowledge this is the first report of osteoporosis in this species.     

Genomic association with pathogen carriage in bighorn sheep (Ovis canadensis)

Genetic composition can influence host susceptibility to, and transmission of, pathogens, with potential population‐level consequences. In bighorn sheep (Ovis canadensis), pneumonia epidemics caused by Mycoplasma ovipneumoniae have been associated with severe population declines and limited recovery across North America. Adult survivors either clear the infection or act as carriers that continually shed M. ovipneumoniae and expose their susceptible offspring, resulting in high rates of lamb mortality for years following the outbreak event. Here, we investigated the influence of genomic composition on persistent carriage of M. ovipneumoniae in a well‐studied bighorn sheep herd in the Wallowa Mountains of Oregon, USA. Using 10,605 SNPs generated using RADseq technology for 25 female bighorn sheep, we assessed genomic diversity metrics and employed family‐based genome‐wide association methodologies to understand variant association and genetic architecture underlying chronic carriage. We observed no differences among genome‐wide diversity metrics (heterozygosity and allelic richness) between groups. However, we identified two variant loci of interest and seven associated candidate genes, which may influence carriage status. Further, we found that the SNP panel explained ~55% of the phenotypic variance (SNP‐based heritability) for M. ovipneumoniae carriage, though there was considerable uncertainty in these estimates. While small sample sizes limit conclusions drawn here, our study represents one of the first to assess the genomic factors influencing chronic carriage of a pathogen in a wild population and lays a foundation for understanding genomic influence on pathogen persistence in bighorn sheep and other wildlife populations. Future research should incorporate additional individuals as well as distinct herds to further explore the genomic basis of chronic carriage.     

Seroprevalence of Toxoplasma gondii in Rocky Mountain bighorn sheep (Ovis canadensis)

Serum samples from 697 Rocky Mountain bighorn sheep (Ovis canadensis) from North America were examined for antibodies to Toxoplasma gondii by the modified agglutination test incorporating mercaptoethanol and formalin-fixed tachyzoites. Antibodies to T. gondii were found in 25 of 697 (3.6%) sheep in titers of 1:25 (8 sheep), 1:50 (4 sheep), 1:100 (7 sheep), 1:200 (1 sheep), 1:400 (1 sheep), 1:800 (1 sheep), and 1:1,600 (3 sheep). This is the first record of T. gondii exposure in bighorn sheep.