Genetic variation and population structure in desert bighorn sheep: implications for conservation

Bighorn sheep populations experienced a drastic reduction in both distribution and abundance until the advent of modern wildlife management, where improving viability of extant populations and translocating animals into historical habitat range have been the most important management policies. The fact that subspecies relationships among bighorn are ambiguous,together with the importance of selecting appropriate source stock and the expense of translocation projects, makes an understanding of subspecies relationships and genetic variation, within and between populations, important for the management and conservation of this species. In this study, genetic variation in 279 bighorn sheep from 13 study sites in Arizona, California, New Mexico and Alberta, Canada were examined by analyzing ten microsatellite loci to determine interpopulation differentiation and relationships between closely related taxa. All populations contained a substantial amount of genetic variation. Genetic differences between populations were large and roughly proportional to geographic distance. The significance of this to desert subspecies relationships and management is discussed.     

Comparing contemporary models to traditional indices to estimate abundance of desert bighorn sheep

Aerial surveys for large ungulates produce count data that often underrepresent the number of animals. Errors in count data can lead to erroneous estimates of abundance if they are not addressed. Our objective was to address imperfect detection probability by developing a framework that produces realistic and defensible estimates of bighorn sheep (Ovis canadensis) abundance. We applied our framework to a population of desert bighorn sheep (O. c. nelsoni) in the Great Basin, Nevada, USA. We captured and marked 24 desert bighorn sheep with global positioning system (GPS)-collars and then conducted helicopter surveys naïve to the locations of collared animals. We developed a Bayesian integrated data model to leverage information from telemetry data, helicopter survey counts, and habitat characteristics to estimate abundance while accounting for availability and perception probability (i.e., detection given availability). Distance to ridgeline, terrain ruggedness, tree cover, and slope influenced perception probability of sheep given they were viewable from the helicopter. There was also annual variation in perception probability (2018: median = 0.64, credible interval [CrI] = 0.37–0.87; 2019: median = 0.81, CrI = 0.49–0.97). The abundance estimates from the integrated data model decreased from 2018 (594; 95% CrI = 537–656) to 2019 (487; 95% CrI = 436–551). In addition, accounting for availability and imperfect perception resulted in greater estimates of abundance compared to traditional directed search methods, which were 340 for 2018 and 320 for 2019. Our modeling framework can be used to generate more defensible population estimates of bighorn sheep and other large mammals that have been surveyed in a similar manner.     

The rise and fall of psoroptic scabies in bighorn sheep in the San Andres Mountains, New Mexico

Between 1978 and 1997, a combination of psoroptic scabies (Psoroptes spp.), mountain lion (Puma concolor) predation, and periodic drought reduced a population of native desert bighorn sheep (Ovis canadensis) in the San Andres Mountains (SAM), New Mexico, from >200 individuals to a single ewe. In 1999, this ewe was captured, ensured to be Psoroptes-free, and released back into the SAM. Eleven radio-collared rams were translocated from the Red Rock Wildlife Area (RRWA) in New Mexico into the SAM range and monitored through 2002 to determine whether Psoroptes spp. mites were still in the environment. None of these sentinel rams acquired scabies during this period, and no additional native sheep were found to be present in the range. In 2002, 51 desert bighorn sheep were translocated into the SAM from the Kofa National Wildlife Refuge in Arizona (n = 20) and the RRWA in New Mexico (n = 31). Twenty-one bighorn sheep have died in the SAM since that time, but Psoroptes spp. mites have not been detected on any of these animals, nor have they been found on mule deer (Odocoileus hemionus) sampled since 2000. We conclude that psoroptic scabies is no longer present in the San Andres bighorn sheep population and that psoroptic scabies poses a minimal to nonexistent threat to the persistence of this population at this time.     

Survival of captive-born Ovis canadensis in North American zoos

The survival rate of North American bighorn sheep, Ovis canadensis, housed in several zoological gardens was analyzed. Complete herd histories, including birth and death data as well as causes of mortality, were collected from seven institutions. Lambs were divided into inbred and noninbred animals, with lambs being considered inbred if they had an inbreeding coefficient greater than zero. The rate of survival of inbred and noninbred lambs was compared using “survival equalling one year” and “survival equalling six months.” Another analysis compared the survival rate of male and female inbred lambs and male and female noninbred lambs. Age at death was also compared in inbred and noninbred lambs. The analysis of the data for the seven collections, located in various geographic areas and housing various subspecies, indicates that inbreeding depression is a mortality factor in the captive management of North American bighorn sheep. Therefore, long-term survival of captive or isolated wild populations will depend on maintaining genetic diversity within the herds through careful selection of breeding stock in captive populations and introduction of nonrelated animals into isolated wild populations.     

Hemorrhagic disease in bighorn sheep in Arizona

Two bighorn sheep from Arizona (USA) were submitted for necropsy. One was a Rocky Mountain bighorn (Ovis canadensis canadensis) and the other was a desert bighorn (Ovis canadensis mexicana). Both had lesions consistent with those of hemorrhagic disease (HD). Epizootic hemorrhagic disease virus (EHDV) type-2 and bluetongue virus (BTV) type-17, respectively, were isolated from the sheep tissues. To our knowledge, HD caused by either EHDV or BTV infection has not been documented previously in Arizona bighorn sheep.     

Toxoplasmic encephalitis in a free-ranging Rocky Mountain bighorn sheep from Washington

A 4-mo-old free-ranging Rocky Mountain bighorn sheep (Ovis canadensis canadensis) from the Hells Canyon area (Washington, USA) was diagnosed with encephalitis associated with Toxoplasma gondii infection. The sheep had concurrent pneumonic pasteurellosis and resided in a geographic area with endemic Pasteurella-associated pneumonia and mortality in bighorn sheep. The brain had multifocal necrotizing and nonsuppurative encephalitis with intralesional protozoa. The protozoa were identified as T. gondii by immunohistochemistry. To our knowledge, this is the first report of T. gondii infection in a Rocky Mountain bighorn sheep.     

Desert bighorn sheep mortality due to presumptive type C botulism in California

During a routine telemetry flight of the Mojave Desert (California, USA) in August 1995, mortality signals were detected from two of 12 radio-collared female desert bighorn sheep (Ovis canadensis) in the vicinity of Old Dad Peak in San Bernardino County (California). A series of field investigations determined that at least 45 bighorn sheep had died near two artificial water catchments (guzzlers), including 13 bighorn sheep which had presumably drowned in a guzzler tank. Samples from water contaminated by decomposing bighorn sheep carcasses and hemolyzed blood from a fresh bighorn sheep carcass were tested for the presence of pesticides, heavy metals, strychnine, blue-green algae, Clostridium botulinum toxin, ethylene glycol, nitrates, nitrites, sodium, and salts. Mouse bioassay and enzyme-linked immunosorbent assay detected type C botulinum toxin in the hemolyzed blood and in fly larvae and pupae. This, coupled with negative results from other analyses, led us to conclude that type C botulinum poisoning was most likely responsible for the mortality of bighorn sheep outside the guzzler tank.     

Potential Foraging Decisions by a Desert Ungulate to Balance Water and Nutrient Intake in a Water-Stressed Environment

Arid climates have unpredictable precipitation patterns, and wildlife managers often provide supplemental water to help desert ungulates endure the hottest, driest periods. When surface water is unavailable, the only source of water for ungulates comes from the forage they consume, and they must make resourceful foraging decisions to meet their requirements. We compared two desert bighorn sheep (Ovis canadensis nelsoni) populations in Arizona, USA: a treatment population with supplemental water removed during treatment, and a control population. We examined whether sheep altered their seasonal diets without supplemental water. We calculated water and nutrient intake and metabolic water production from dry matter intake and forage moisture and nitrogen content, to determine whether sheep could meet their seasonal daily water and nutrient requirements solely from forage. Diets of sheep were higher in protein (all seasons) and moisture (autumn and winter) during treatment compared to pretreatment. During treatment, sheep diet composition was similar between the treatment and control populations, which suggests, under the climatic conditions of this study, water removal did not influence sheep diets. We estimated that under drought conditions, without any surface water available (although small ephemeral potholes would contain water after rains), female and male sheep would be unable to meet their daily water requirements in all seasons, except winter, when reproductive females had a nitrogen deficit. We determined that sheep could achieve water and nutrient balances in all seasons by shifting their total diet proportions by 8–55% from lower to higher moisture and nitrogen forage species. We elucidate how seasonal forage quality and foraging decisions by desert ungulates allow them to cope with their xeric and uncertain environment, and suggest that, with the forage conditions observed in our study area during this study period, providing supplemental water during water-stressed periods may not be necessary for desert bighorn sheep.     

Behavioural reactions of desert bighorn sheep to avian scavengers

I report previously undocumented encounters between avian scavengers and desert bighorn sheep ( Ovis canadensis nelsoni ). These encounters are of interest because they occurred only during the nursery season when lamb mortality was high. Desert bighorn sheep responded to the presence of avian scavengers with typical anti-predator behaviours, which may indicate that avian scavengers are potential predators or that desert bighorn sheep are unable to distinguish between avian scavengers and avian predators.     

Immunologic responses of domestic and bighorn sheep to a multivalent Pasteurella haemolytica vaccine

The efficacy of a Pasteurella haemolytica vaccine (serotypes A1, A2, and T10) to induce humoral antibodies and alter colonization of the upper respiratory tract by related P. haemolytica spp. strains was evaluated in 10 bighorn (Ovis canadensis canadensis) and 10 domestic (Ovis aries) sheep. Sheep of each species were divided into five pairs based on age and history of respiratory disease. One sheep in each pair was vaccinated twice 2 wk apart with 2 ml of vaccine (VAC group) and the remaining animals (NV group) were injected with 2 ml of sterile saline. Mild, transient lameness was the only observed adverse effect. Blood sera from the sheep were tested for agglutinating antibodies against whole cells of A1, A2, and T10 and for leukotoxin neutralizing antibodies. Antibody titers were expressed as the reciprocal log2 of the highest reactive dilutions. Domestic sheep > 1-yr-old and two bighorn sheep with a history of A1 infection had higher titers throughout the study against A1 cells than domestic sheep < 1-yr-old and bighorns without a history of A1 infection. Both domestic and bighorn sheep had log2 titers of 8 to 12 against A2 cells and 6 to 12 against T10 cells during this time. Bighorn sheep in the VAC group had 2 to 32 fold titer increases for A1 cells by 2 wk post-vaccination (PV) compared to 0 to 2 fold increases in VAC domestic sheep. Two to 16 and 0 to 8 fold increases in antibodies titers to A2 and T10 cells, respectively, were detected in sera of both VAC groups. Sera of bighorn sheep with a history of respiratory disease and all domestic sheep had log2 leukotoxin neutralizing antibody titers of 4 to 14 in contrast to < or = 2 in sera of bighorn sheep without a history of respiratory disease. Neutralizing antibody titers of two bighorns without a history of respiratory disease in the VAC group increased from log2 0 to 5 in one and from 0 to 9 in the other 2 wk PV. Antibody increases in these animals were no longer evident at 16 wk PV while titers of animals with histories of disease remained relatively stable. The types and numbers of Pasteurella spp. isolated from nasal and pharyngeal swabs varied throughout the study without conclusive evidence of suppression of colonization. Although the animals were not experimentally challenged to determine the efficacy of the vaccine, one VAC and one NV bighorn sheep died following introduction of an A2 P. haemolytica strain when leukotoxin neutralizing antibodies had returned to pre-vaccination levels. This vaccine appeared to be safe for use in bighorn sheep and stimulated moderate but transient increases in antibody levels which should provide some protection against naturally occurring disease. A vaccine which would induce production of high and maintained antibodies against multiple strains of P. haemolytica would be valuable for use in bighorn sheep maintained in captivity or when captured for relocation.     

Hematologic and blood chemistry values of the Masai ostrich (Struthio camelus)

Normal mean values for hematocrit, hemoglobin concentration, erythrocyte and leukocyte counts, hematimetric indices, erythrocyte dimensions, glucose, urea, uric acid, cholesterol, creatinine, total bilirubin, serum aspartate aminotransferase, serum alanine aminotransferase, alkaline phosphatase, creatinine phosphokinase, lactic dehydrogenase, inorganic phosphorus, chloride, total plasma protein, sodium, potassium, calcium, and magnesium were obtained from the blood or plasma of four Masai ostriches (Struthio camelus) when juveniles at 5 mo of age and as adults 1 yr later in the Barcelona Zoo (Spain). Young ostriches had significantly lower concentrations of hematocrit, hemoglobin concentration, calcium, and magnesium, and higher levels of total protein and potassium, than the adult individuals. The rest of the parameters were not significantly different between the two age groups. The data obtained provide reference values for Masai ostriches.     

New ruminant hosts and wider geographic range identified for Babesia odocoilei (Emerson and Wright 1970)

Babesia odocoilei was found to infect two previously unknown host species, desert bighorn sheep (Ovis canadensis nelsoni) and musk oxen (Ovibos moschatus), both of which are members of the family Bovidae. Previously, B. odocoilei has been reported in only Cervidae hosts. New geographic regions where B. odocoilei infections have not been reported previously include Pennsylvania and New York, where fatal babesiosis has occurred in reindeer (Rangifer tarandus tarandus); New Hampshire, where elk (Cervus elaphus canadensis) have been affected; and California, home of the infected desert bighorn sheep. Infection with B. odocoilei in these hosts was confirmed by parasite small subunit ribosomal RNA gene sequence analysis. A serosurvey for B. odocoilei antibody activity in New Hampshire showed prevalence rates of 100% at two elk farms and 12% at another farm. Control of potential vector ticks, Ixodes scapularis, especially when translocating livestock, is imperative to prevent outbreaks of babesiosis in managed herds of potential host species.     

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.     

Habitat use by desert bighorn sheep during lambing

Habitat used for predator escape may be a factor limiting restoration of desert bighorn sheep (Ovis canadensis mexicana) in small mountain ranges in New Mexico. Female bighorn sheep seek isolation in discrete areas for parturition. Although parturition sites are used only for <3 days they play an important role in neonate survival. We compared habitat characteristics at pre-parturition (n = 21), parturition (n = 38), random (n = 38), and post-parturition sites (n = 21). At each site we calculated distance to steep terrain, elevation, ruggedness, slope, and visibility. Parturition and post-parturition sites were higher in elevation and more rugged than sites used during pre-parturition. Post-parturition sites were closer to terrain with 100% slope than the pre-parturition or parturition sites. Post-parturition sites had higher visibility and steeper slopes than the pre-parturition sites. Parturition sites were steeper, higher in elevation, more rugged, and had lower visibility than the random sites. Fidelity to parturition sites was observed on 1 of 38 occasions. Because lambing sites are used for short periods they are poorly understood and often ignored by land managers. Additional research is recommended in other desert bighorn sheep populations to understand better and predict habitat use during the lambing period.     

Effects of capture on biological parameters in free-ranging bighorn sheep (Ovis canadensis): evaluation of normal, stressed and mortality outcomes and documentation of postcapture survival

Blood samples and physiological data were collected from 634 bighorn sheep (Ovis canadensis) captured by four different methods between 1980 and 1986 in the western United States. These parameters were evaluated for selected physiological, biochemical and hematological values. Postcapture biological parameters were compared among bighorn sheep according to four different outcomes; normal, stressed or compromised, capture myopathy (CM) mortality, and accidental mortality. Significant differences (P less than 0.05) were noted between outcome groups relative to certain parameters: temperature, respiration, creatinine phosphokinase (CPK), lactic dehydrogenase (LDH), serum glutamic oxaloacetic transaminase (SGOT), blood urea nitrogen (BUN), glucose, white blood cell count (WBC) and plasma pH. Such differences between groups may help in evaluating the clinical status of bighorn sheep at capture, enabling one to predict those animals that might develop CM at a later date, indicate candidates for preventive medical treatment prior to release, and/or which should be followed closely to determine long-term survival. Evaluation of follow-up data (n = 77) related to outcome status and long-term survival of bighorn sheep indicated that less than 4% (3 of 77) were dead within 1 mo of capture (one of these had been classified as normal and two as stressed or compromised at capture); less than 3% (3 of 77) were dead greater than 1 mo, and less than 6 mo after capture two were classified in the stressed outcome and one as diseased. Eighty-eight percent (68 of 77) were alive from 1 mo to 5 yr after capture (53 were classified as normal, 12 as stressed or compromised and 3 as diseased), and 2% (1 of 77) had chronic CM but was still alive (this animal had been classified as normal). Of 77 sheep in the follow-up group, less than 3% (2 of 77) were not observed following capture (one was classified as normal and one as stressed and diseased). Of the fatalities, less than 3% (2 of 40) had been captured by the net-gun and less than 4% (1 of 27) by drive-net. Those two unobserved in the follow-up group also had been caught with the net-gun, 5% (2 of 40). The single surviving CM case had been captured by the net-gun. Although the net-gun appears to be one of the safest methods of capturing individual bighorn sheep, based on evaluation of capture data and biological parameters, it may not be associated with the best long-term survival in some bighorn sheep.(ABSTRACT TRUNCATED AT 400 WORDS)     

Population Dynamics and Behavior of Bighorn Sheep with Infectious Keratoconjunctivitis

ABSTRACT Introduced disease is a major mortality factor in some populations of bighorn sheep (Ovis canadensis). Epizootics of infectious keratoconjunctivitis (IKC) and contagious ecthyma occurred in bighorn sheep in the Silver Bell Mountains of south-central Arizona, USA, from 1 December 2003 to 31 March 2004. Our objectives were to 1) investigate the influence of the epizootic on abundance and demographics and 2) examine how IKC affected the mortality, behavior, and movements of clinically affected animals. Morbidity was 39%, and all sex and age classes were affected. The population declined 23%, with most mortality in the adult female (1 M, 11 F) segment of the population. Of the diseased animals that were marked (n = 27), 44% recovered and 44% died. Predation (50%) and starvation (33%) were the primary causes of mortality of diseased bighorn sheep. Bighorn sheep that were infected spent less time feeding and moved less than noninfected animals during the epizootic. Managers might be able to minimize losses of infected animals through predator control. To minimize losses to starvation, managers should refrain from any activity that disturbs infected animals (including treatment) because disturbances increase energy expenditures and expose infected animals to injury.     

Brucellosis in captive Rocky Mountain bighorn sheep (Ovis canadensis) caused by Brucella abortus biovar 4

Nine (four female, five male) captive adult Rocky Mountain bighorn sheep (Ovis canadensis) contracted brucellosis caused by Brucella abortus biovar 4 as a result of natural exposure to an aborted elk (Cervus elaphus) fetus. Clinical signs of infection were orchitis and epididymitis in males and lymphadenitis and placentitis with abortion in females. Gross pathologic findings included enlargement of the testes or epididymides, or both, and yellow caseous abscesses and pyogranulomas of the same. Brucella abortus biovar 4 was cultured in all bighorn sheep from a variety of tissues, including testes/epididymides, mammary gland, and lymph nodes. All bighorn sheep tested were positive on a variety of standard Brucella serologic tests. This is the first report of brucellosis caused by B. abortus in Rocky Mountain bighorn sheep. It also provides evidence that bighorn sheep develop many of the manifestations ascribed to this disease and that infection can occur from natural exposure to an aborted fetus from another species. Wildlife managers responsible for bighorn sheep populations sympatric with Brucella-infected elk or bison (Bison bison) should be cognizant of the possibility of this disease in bighorn sheep.     

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.