Muellerius capillaris (Mueller, 1889) (Nematoda: Protostrongylidae): an unusual finding in Rocky Mountain bighorn sheep (Ovis canadensis canadensis Shaw) in South Dakota

Lungs and fecal samples from nine hunter-killed Rocky Mountain bighorn sheep were examined for lungworms. All samples contained adults and/or larvae of Muellerius capillaris (Mueller, 1889). Protostrongylus spp., the lungworms commonly reported from bighorn sheep, were not present in any samples. Larvae of M. capillaris bear a spine on the dorsal side of the posterior end and are shorter than dorsal-spined larvae of other lungworms recorded from North American ungulates. Larvae similar in shape but longer than those of Muellerius were found in free-ranging bighorn sheep in Alberta and British Columbia. In addition, dorsal-spined larvae have been found in bighorn sheep in Montana, North Dakota, and Washington. The identity of the dorsal-spined larvae is known only from sheep in South Dakota. Thus, caution must be taken when diagnosing lungworm infections in Rocky Mountain bighorn sheep.     

Relationships among fecal lungworm loads, fecal glucocorticoid metabolites, and lamb recruitment in free-ranging Rocky Mountain bighorn sheep

Most wild Rocky Mountain big-horn sheep (Ovis canadensis canadensis) in northern latitudes are infected with lungworms. Indirect effects of lungworms on bighorn sheep are unknown, but high pulmonary burdens might increase stress (i.e., elevated glucocorticoid levels), and chronic stress could in turn decrease fitness. We hypothesized that high lungworm burdens in Rocky Mountain bighorn ewes increase stress, thereby increasing lamb mortality. To test our hypothesis, one subherd of bighorn sheep in Custer State Park, South Dakota was provided a free-choice loose mineral mix containing the anthelmintic fenbendazole every six weeks from March 1999 to August 2000 to eliminate lungworms; another subherd served as the control. Daily, individually marked ewes were located telemetrically from the ground and uniquely marked animals were observed until they defecated. After the herd moved from the area, fecal samples were collected and stored at -23 C. A consistent number of samples per season per herd (x-=16.56+/-3.99 samples) were collected. Fecal larval lungworm levels (LPG) in the treatment subherd were lower than levels in the control subherd; however, there was no difference in fecal glucocorticoid metabolite (FGM) levels between the two subherds. Fecal glucocorticoid metabolite levels varied by season in both subherds, with levels in winter lower than during the other three seasons. Lamb:ewe ratios were not different between the control and treatment subherds at the end of summer 1999. In contrast, the treatment group had a lower lamb:ewe ratio at the end of summer 2000 despite having lower LPG. However, this result was attributed to lower ewe production, not lower lamb survival. The LPG levels were not correlated with FGM concentrations; instead, FGM levels might reflect normal seasonal patterns. Other factors, including contagious ecthyma, were more important for determining lamb mortality than LPG and FGM levels during our study. We suggest further experimental work over a longer duration to address these relationships.     

Fecal counts of lungworm larvae and reproductive effort in bighorn sheep, Ovis canadensis

Because parasite resistance and reproduction require metabolic resources, life-history models predict a tradeoff between current reproduction and parasite load. These tradeoffs have been widely studied in birds, but few studies have been conducted on mammals. We monitored lungworm ( Protostrongylus spp.) larvae counts in bighorn sheep ( Ovis canadensis ) over four years to examine how individual differences in fecal output of lungworm larvae (LPG) by yearlings and adults were affected by season, sex, body mass, age and reproductive effort. We also compared lamb mass at six months and LPG. Overall, we found that LPG varies seasonally, peaking in females prior to lambing and in males during the rut. Age had no effect on LPG for either sex. During autumn, we found no effect of age or mass on LPG for sheep one year and older. Lamb body size or sex did not affect LPG. Females that weaned a lamb had higher counts than females that did not produce a lamb or females whose lamb died during summer. For rams, social rank and testosterone levels were not related to LPG but LPG increased with time spent searching for estrous ewes during the rut. Our results suggest a tradeoff between parasite resistance and reproductive effort in bighorn sheep of both sexes.     

Infectious keratoconjunctivitis in bighorn sheep, Silver Bell Mountains, Arizona, USA

An infectious keratoconjunctivitis (IKC) epizootic in bighorn sheep (Ovis canadensis) occurred in the Silver Bell Mountains, Arizona, USA, from 1 December 2003 to 31 March 2004. We used standard culture methods and polymerase chain reaction (PCR) amplification of the 16S rRNA gene to test for the causative agents of IKC and other diseases reported to be associated with bighorn sheep populations. All bighorn sheep and domestic goat test results were negative except for Mycoplasma spp. and Branhamella spp. The culture and PCR results differed. Conjunctival swabs from four of 19 IKC-affected bighorn sheep tested by culture were positive for Mycoplasma spp., whereas 22 of 22 bighorn sheep samples tested by PCR were positive for Mycoplasma spp. None of 13 domestic goats tested positive by culture for Mycoplasma spp., whereas five of 16 tested positive by PCR. Three of 16 domestic goats and seven of 24 IKC-affected bighorn sheep tested positive for Branhamella spp. by culture. Bighorn sheep began showing clinical signs of IKC between 21 and 28 days following initial detection of domestic goats in bighorn sheep habitat. The IKC epizootic lasted 122 days, and individual bighorn sheep were blind for an average of 38.4 days. Given the clear potential for disease transmission to bighorn sheep, we recommend that land managers not allow the pasturing of domestic goats near occupied bighorn sheep habitat.     

Evaluating apparent competition in limiting the recovery of an endangered ungulate

Predation can disproportionately affect endangered prey populations when generalist predators are numerically linked to more abundant primary prey. Apparent competition, the term for this phenomenon, has been increasingly implicated in the declines of endangered prey populations. We examined the potential for apparent competition to limit the recovery of Sierra Nevada bighorn sheep (Ovis canadensis sierrae), an endangered subspecies under the US Endangered Species Act. Using a combination of location, demographic, and habitat data, we assessed whether cougar (Puma concolor) predation on endangered bighorn sheep was a consequence of their winter range overlap with abundant mule deer (Odocoileus hemionus). Consistent with the apparent competition hypothesis, bighorn sheep populations with higher spatial overlap with deer exhibited higher rates of cougar predation which had additive effects on adult survival. Bighorn sheep killed by cougars were primarily located within deer winter ranges, even though those areas constituted only a portion of the bighorn sheep winter ranges. We suspect that variation in sympatry between bighorn sheep and deer populations was largely driven by differences in habitat selection among bighorn sheep herds. Indeed, bighorn sheep herds that experienced the highest rates of predation and the greatest spatial overlap with deer also exhibited the strongest selection for low elevation habitat. Although predator-mediated apparent competition may limit some populations of bighorn sheep, it is not the primary factor limiting all populations, suggesting that the dynamics of different herds are highly idiosyncratic. Management plans for endangered species should consider the spatial distributions of key competitors and predators to reduce the potential for apparent competition to hijack conservation success.     

Influence of rainfall on lungworm infections in bighorn sheep.

From 1959 through 1968, lungs from 124 Rocky Mountain bighorn sheep (Ovis c. canadensis) from the Sun River herd in western Montana were examined for lungworm infections. All lungs were infected with Protostrongylus stilesi and 104 (84%) contained concurrent infections of P. stilesi and P. rushi. Significant correlations were observed between levels of lungworm infection and total rainfall during April, May, and June of each year. An explanation of this in terms of terrestrial snail (intermediate host) populations and a suggestion for the possible use of these data in developing a predictive model for forecasting lungworm levels for use in in bighorn sheep management are given.     

Sharing of Pasteurella spp. between free-ranging bighorn sheep and feral goats

Pasteurella spp. were isolated from feral goats and free-ranging bighorn sheep (Ovis canadensis canadensis) in the Hells Canyon National Recreation Area bordering Idaho, Oregon, and Washington (USA). Biovariant 1 Pasteurella haemolytica organisms were isolated from one goat and one of two bighorn sheep found in close association. Both isolates produced leukotoxin and had identical electrophoretic patterns of DNA fragments following cutting with restriction endonuclease HaeIII. Similarly Pasteurella multocida multocida a isolates cultured from the goat and one of the bighorn sheep had D type capsules, serotype 4 somatic antigens, produced dermonecrotoxin and had identical HaeIII electrophoretic profiles. A biovariant U(beta) P.haemolytica strain isolated from two other feral goats, not known to have been closely associated with bighorn sheep, did not produce leukotoxin but had biochemical utilization and HaeIII electrophoretic profiles identical to those of isolates from bighorn sheep. It was concluded that identical Pasteurella strains were shared by the goats and bighorn sheep. Although the direction of transmission could not be established, evidence suggests transmission of strains from goats to bighorn sheep. Goats may serve as a reservoir of Pasteurella strains that may be virulent in bighorn sheep; therefore, goats in bighorn sheep habitat should be managed to prevent contact with bighorn sheep. Bighorn sheep which have nose-to-nose contact with goats should be removed from the habitat.     

Restoration Potential of Bighorn Sheep in a Prairie Region

Efforts to recover Rocky Mountain bighorn sheep (Ovis canadensis canadensis) throughout western North America have had limited success with the majority of current populations remaining in small and isolated areas on a fraction of their historical range. Prairie environments with rugged topography throughout the Northern Great Plains ecoregion were historically occupied by relatively robust bighorn sheep populations. We predicted there is likely unrealized potential habitat for restoring bighorn sheep to these areas; however, relatively little attention has been devoted to identifying habitat in unoccupied prairie regions. We used global positioning system (GPS)-collar data collected from 43 female bighorn sheep in 2 populations located in the eastern Montana, USA, portion of the Northern Great Plains during 2014–2018 to estimate a population-level annual resource selection model and identify the important factors affecting bighorn sheep resource selection. We extrapolated model predictions across eastern Montana's prairie region and identified potential habitat to understand restoration potential and assist with future translocations of bighorn sheep. Resource selection of bighorn sheep was most strongly associated with terrain slope and ruggedness, tree canopy cover, and a normalized difference vegetation index metric. Within currently unoccupied areas of the historical range, the model extrapolation predicted 7,211 km² of habitat, with most owned and managed by private landowners (44%), Bureau of Land Management (33%), and the United States Fish and Wildlife Service (15%). Our results provide an empirical evaluation of landscape covariates influencing resource selection of bighorn sheep occupying prairie environments and provide a habitat model that may be generalizable to other areas in the Northern Great Plains ecoregion. We demonstrate substantial potential for restoration opportunities of bighorn sheep in the Northern Great Plains ecoregion. Broad restoration of bighorn sheep across the ecoregion would likely require strong collaboration among and between public resource managers, private landowners, and livestock producers given the heterogeneous land ownership patterns, management strategies, and domestic sheep distributions. © 2020 The Wildlife Society.     

Desert bighorn sheep habitat selection,group size,and mountain lion predation risk

Bighorn sheep (Ovis canadensis) evolved for thousands of years in the presence of numerous predators, including mountain lions (Puma concolor). Bighorn sheep have presumably developed predator avoidance strategies; however, the effectiveness of these strategies in reducing risk of mountain lion predation is not well understood. These strategies are of increasing interest because mountain lion predation on bighorn sheep has been identified as a leading cause of mortality in some sheep populations. Therefore, we investigated how mountain lions affect both bighorn sheep habitat selection and risk of mortality in Arizona, USA. We used 2 approaches to investigate the predator-prey relationship between mountain lions and bighorn sheep. We fit 103 bighorn sheep (81 females and 22 males) with global positioning system radio-collars in 2 Arizona populations from 2013 to 2017, and used a negative binomial resource selection probability function to evaluate whether bighorn sheep selected for habitat features in accordance with presumed predator avoidance strategies, including terrain ruggedness, slope, topographic position, and horizontal obstruction, in 2 seasons (winter and summer). We then estimated how habitat features such as terrain ruggedness, slope, horizontal obstruction, and group size, influence the risk of mortality due to mountain lion predation using an Andersen-Gill proportional hazards model. Generally, both sexes selected areas with lower horizontal obstruction and intermediate ruggedness and slope, but selection patterns differed between seasons and sexes. The use of more rugged areas and steeper slopes decreased the risk of mortality due to mountain lion predation, consistent with presumed predator avoidance strategies. Increased group size decreased risk of bighorn sheep mortality due to mountain lion predation but this effect became marginal at approximately 10 individuals/group. We did not identify a relationship between horizontal obstruction and bighorn sheep mortality risk. Our findings can be used in habitat and population management decisions such as the prioritization of habitat restoration sites or selection of translocation sites. In addition, we suggest that augmentation of low-density bighorn sheep populations may reduce mountain lion predation risk by increasing group size, and that releasing large groups of bighorn sheep in population augmentation and reintroduction efforts may help to reduce mountain lion predation.     

Comparison of Endocrine Response to Stress Between Captive-Raised and Wild-Caught Bighorn Sheep

ABSTRACT Stress hormones in Rocky Mountain bighorn sheep (Ovis canadensis canadensis), produced in response to environmental changes, road development, or high population density, may impact their immune systems to a threshold level that predisposes them to periodic, large-scale mortality. We compared the stress response to a novel environmental situation and repeated handling between bighorn sheep born and raised in captivity (CR) and bighorn sheep born in the wild (WC) and brought into captivity. We measured plasma epinephrine, norepinephrine, cortisol, and fecal glucocorticoid metabolites (FGM). Three weeks after each group's arrival we used a one-time drop-net event to elicit an acute stress response, and we collected blood samples from each sheep over 35 minutes, as well as one fecal sample. We collected blood and fecal samples from both groups on 7 other occasions over the subsequent 6 months. We also collected fecal samples from the pen at approximately 24-hour intervals for 3 days following every handling event to monitor the stress response to handling. We found that CR sheep had a stronger autonomic nervous system response than WC sheep, as measured by epinephrine and norepinephrine levels, but we found a very similar hypothalamic-pituitary–adrenal axis (HPA) response, measured by cortisol levels, to the acute stress event of a drop-net restraint. We also found that once the WC sheep had acclimated, as indicated by the return to the initial baseline FGM levels within 12 weeks, the CR and WC groups' HPA responses to sampling events were not significantly different from one another. Fecal samples can provide a noninvasive mechanism for managers to monitor baseline FGM for a given herd. Using long-term monitoring of FGM rather than values from a single point in time may allow managers to correlate these levels to outside influences on the herd and better understand the impacts of management changes, population density, or increased human developments on the health of the sheep population.     

Occurrence, diagnosis, and strain typing of Mycobacterium avium subspecies paratuberculosis infection in Rocky Mountain bighorn sheep (Ovis canadensis canadensis) in southwestern Alberta

The role that wildlife may play in the transmission of Mycobacterium avium subspecies paratuberculosis (Map), the causative agent of Johne's disease (JD), and the potential consequences of infection in these populations are being given increasing consideration. A yearling male Rocky Mountain bighorn sheep (Ovis canadensis canadensis) from southwestern Alberta, Canada, was found infected with Map in August 2009. Clinical signs of emaciation and diarrhea and histologic findings of diffuse granulomatous enteritis of the distal ileum, lymphadenitis of the mesenteric lymph nodes, and lymphangitis of the ileum were similar to previously described cases of JD in bighorn sheep. Infection with Map was confirmed by bacterial isolation through fecal culture, acid-fast staining, and polymerase chain reaction (PCR) of IS900. The Map1506 gene was sequenced, and the isolate was identified as a Cattle (Type II) strain. In a follow-up herd-level survey, three of 44 fecal samples (7%) from individual bighorn sheep from the same herd as the index case were PCR-positive and identified as Type II Map strains. Twenty-five samples from a distant bighorn population were negative. Additional strain typing of the isolates from the index case and the positive fecal samples was done by sequencing three discriminatory short sequence repeat (SSR) regions. All four SSR profiles differed from one another, suggesting multiple introductions or a long-existing circulation of Map within this bighorn population. Detailed molecular analyses are essential for understanding and managing diseases at the wildlife-livestock interface.     

Comparison of three fecal steroid metabolites for pregnancy detection used with single sampling in bighorn sheep (Ovis canadensis)

We conpared three fecal steroid metabolite assays for their usefulness in detecting pregnalcy among free-ranging Rocky Mountain bighorn sheep (Ovis canadensis canadensis) from Bighorn Canyon National Recreation Area, Wyoming and Montana (USA) and captive bighorn ewes at ZooMontana in Billings, Montana. Fecal samples were collected from 11 free-ranging, radio-collared bighorn ewes in late January-May 2001 and from 20 free-ranging, radio-collared ewes in late March to mid-May 2002. Free-ranging ewes were monitored the following spring to determine whether or not they lambed. In addition, two captive ewes were studied at ZooMontana. With three exceptions, free-ranging bighorn ewes that produced lambs had nonspecific progesterone metabolite (iPdG) levels of >1800 ng/g feces and iPdG levels >7000 ng/gm feces when samples were collected between early March and mid-May. Samples collected earlier in the year were inconclusive. One false negative was suspected to be the result of sample collection error. Of the captive ewes, nonspecific pregnanediol-3alpha-glucuronide (PdG) and iPdG followed a predictable curve over the course of the 180-day pregnancies. We conclude that estrone conjugates are not useful in diagnosing pregnancy; however, fecal steroid analysis of PdG and iPdG can be used to accurately determine pregnancy and reproductive function in bighorn sheep. This holds great potential as a noninvasive technique for understanding the role of reproductive disease in wild bighom sheep.     

Elaeophorosis in bighorn sheep in New Mexico

Two bighorn sheep (Ovis canadensis) in New Mexico (USA) were found to be naturally infected with Elaeophora schneideri. An adult ram examined in 1997 in the Fra Cristobal Mountains had 26 nematodes in the carotid and iliac arteries, and microfilariae were present in the skin, nasal mucosa, brain, and lungs. This ram was markedly debilitated prior to euthanasia and extensive crusty, scabby lesions were observed on its head. In 1998, a yearling ewe found dead adjacent to Watson Mountain near the Gila Wilderness area was found to have 13 nematodes present in its heart. This is the first report of E. schneideri in bighorn sheep, and we suggest that bighorn sheep are susceptible to E. schneideri infection wherever they coexist with mule deer (Odocoileus hemionus hemionus) and appropriate tabanid vectors.     

Quantifying Landscape Ruggedness for Animal Habitat Analysis: A Case Study Using Bighorn Sheep in the Mojave Desert

Abstract: Terrain ruggedness is often an important variable in wildlife habitat models. Most methods used to quantify ruggedness are indices derived from measures of slope and, as a result, are strongly correlated with slope. Using a Geographic Information System, we developed a vector ruggedness measure (VRM) of terrain based on a geomorphological method for measuring vector dispersion that is less correlated with slope. We examined the relationship of VRM to slope and to 2 commonly used indices of ruggedness in 3 physiographically different mountain ranges within the Mojave Desert of the southwestern United States. We used VRM, slope, distance to water, and springtime bighorn sheep (Ovis canadensis nelsoni) adult female locations to model sheep habitat in the 3 ranges. Using logistic regression, we determined that the importance of ruggedness in habitat selection remained consistent across mountain ranges, whereas the relative importance of slope varied according to the characteristic physiography of each range. Our results indicate that the VRM quantifies local variation in terrain more independently of slope than other methods tested, and that VRM and slope distinguish 2 different components of bighorn sheep habitat.     

Is Habitat Constraining Bighorn Sheep Restoration? A Case Study

Rocky Mountain bighorn sheep (Ovis canadensis) restoration continues to be a challenge throughout western North America despite nearly a century of efforts dedicated to the species' recovery. Though bighorn sheep restoration may be constrained by several environmental factors and behavioral tendencies, areas with unrealized restoration potential may exist if novel restoration strategies are considered. We used global positioning system (GPS) location data from 27 female bighorn sheep within the southern portion of the Madison Range in southwest Montana, USA, 2015–2017, to develop and validate winter and summer habitat models, which we extrapolated throughout the entire Madison Range to identify potential seasonal habitat. We estimated potential bighorn sheep minimum population estimates within the extrapolation area by linking our top-ranked winter habitat model to population count data. During summer, female bighorn sheep selected areas characterized by rugged and steep terrain, reduced canopy cover, southwestern aspects, and ridgelines. During winter, female bighorn sheep selected areas characterized by low elevations, southwestern aspects, steep slopes, reduced canopy cover, ridgelines, high normalized difference vegetation index amplitude, and areas close to steep terrain. Predicted summer habitat was concentrated along the high-elevation ridgelines associated with steep slopes and reduced canopy cover. Predicted winter habitat occurred in a non-contiguous distribution primarily along the low-elevation, southwest-facing aspects along the western slopes of the Madison Range. Our results suggest that the Madison Range may be capable of supporting 780–1,730 animals, which is 2–4 times the number of bighorn sheep currently observed within the range. Further, our findings provide managers with a quantification of female bighorn sheep habitat and suggests that a strategy focused on establishing a metapopulation through a series of within-range translocations may enhance bighorn sheep restoration. We suggest that similar restoration opportunities may be common in other unoccupied areas of bighorn sheep historical range. © 2020 The Wildlife Society.     

Morphometric analysis of Psoroptes spp. mites from bighorn sheep, mule deer, cattle, and rabbits

A morphometric study of Psoroptes spp. mites was conducted to address difficulties encountered in species identification and to provide insights into the phylogenetic relationships between mites found on different hosts. A discriminant analysis employing 9 morphologic characters revealed that the lengths of the outer opisthosomal setae and the lateral margins of the opisthosomal knobs were the 2 most important characters for grouping mites according to host species. This analysis clearly separated mites collected from allopatric populations of bighorn sheep, rabbits, and cattle into discrete groups. However, differences were not detected between mites collected from sympatric populations of infested mule deer and bighorn sheep, suggesting that these mites were not host specific and represented a single interbreeding population. Differences also were not detected among mites collected from the ears and body of bighorn sheep and rabbits, demonstrating that the location of mites on a given host should not be used as a primary criterion in species identification.     

Bighorn sheep, a new host record for Parelaphostrongylus odocoilei (Nematoda: Protostrongylidae)

Larval nematodes with a dorsal spine on the tail were recovered from fecal samples of California bighorn sheep (Ovis canadensis californiana) in northeastern Washington State, USA. The identity of these dorsal-spined larvae (DSL) was established by single-strand conformation polymorphism (SSCP) analyses of a partial fragment of the first internal transcribed spacer of the ribosomal DNA. The SSCP profiles of individual DSL from bighorn sheep were compared with those of DSL of five protostrongylid species (Parelaphostrongylus andersoni, P odocoilei, P. tenuis, Elaphostrongylus rangiferi, and Muellerius capillaris) but were identical to only those of P. odocoilei. This study represents the first confirmed identification of P. odocoilei in bighorn sheep.     

Assessment of Predictive Habitat Models for Bighorn Sheep in California's Peninsular Ranges

ABSTRACT We developed predictive habitat models for a bighorn sheep (Ovis Canadensis) population in the Peninsular Ranges of southern California, USA, using 2 Geographic Information System modeling techniques, Ecological Niche Factor Analysis (ENFA) and Genetic Algorithm for Rule-set Production (GARP). We used >16,000 Global Positioning System locations from 34 animals in 5 subpopulations to develop and test ENFA and GARP models, and we then compared these models to each other and to the expert-based model presented in the United States Fish and Wildlife Service's Recovery Plan for this population. Based on a suite of evaluation methods, we found both ENFA and GARP to provide useful predictions of habitat; however, models developed with GARP appeared to have higher predictive power. Habitat delineations resulting from GARP models were similar to the expert-based model, affirming that the expert-based model provided a useful delineation of bighorn sheep habitat in the Peninsular Ranges. In addition, all 3 models identified continuous bighorn sheep habitat from the northern to southern extent of our study area, indicating that the Recovery Plan's recommendation of maintaining habitat connectivity throughout the range is an appropriate goal.     

Insects feeding on desert bighorn sheep, domestic rabbits, and Japanese quail in the Santa Rosa mountains of southern California.

Desert bighorn sheep (Ovis canadensis cremnobates), a domestic rabbit (Oryctolagus cuniculus), and Japanese quail (Coturnix japonica) were used as bait animals to collect blood-feeding flies in an area of active blue-tongue and epizootic hemorrhagic disease virus transmission. Precipitin tests were used to confirm the blood source where feasible. Eight species of Culicoides, members of the Leptoconops kerteszi group, Simulium spp., Anopheles franciscanus, and Stomoxys calcitrans were collected from the bighorn sheep. Feeding on the bighorn sheep by Culicoides brookmani (n = 25), C. variipennis (n = 6), C. cacticola (n = 1), and Simulium spp. (n = 3) was confirmed by precipitin testing. Primary species attacking the rabbit were C. brookmani, C. variipennis, and the L. kerteszi group. The quail were attacked primarily by members of the C. copiosus group and the L. kerteszi group.