Evaluating habitat selection models to predict critical habitat for mountain goats in northwest British Columbia

Anthropogenic activity imposes increasing pressure on wildlife populations globally; these pressures can affect habitat suitability and function, modify wildlife space use, and influence population viability. Native mountain goat (Oreamnos americanus) populations can be negatively affected by anthropogenic disturbance and modify their space use in response to land development and recreational activity. From 2018 to 2020, we studied space use of mountain goats northeast of Smithers, British Columbia, Canada, an area that is subject to increasing anthropogenic development and yearlong recreational activities. We aimed to generate models that would improve our ability to identify habitat for mountain goats relative to existing survey data and established ungulate winter ranges. Using resource selection function (RSF) analyses generated from global positioning system (GPS) collar data, we identified influential habitat covariates and compared these covariates and RSF values to existing habitat models. Additionally, we compared the extent to which our models were congruent with existing resource selection probability functions, were congruent with aerial survey data, and overlapped existing ungulate winter ranges previously derived from predictive models inside and outside of the study area. Overall, our models noted higher RSF values among GPS data relative to aerial survey data for winter months, while results for summer habitats were comparable. In extending our RSFs outside of the study area and evaluating the overlap with ungulate winter ranges in adjacent areas, values were similar, albeit lower, as is expected given that the models were developed elsewhere. Ultimately, these models, combined with existing methods, improve the accuracy and reliability of identified, important areas of habitat for mountain goats. We recommend that the RSF models generated here be used in conjunction with aerial survey data and existing methods to delineate ungulate winter ranges for mountain goats in similar eco-regions in British Columbia. The models developed here support existing methods that have been used to delineate or validate ungulate winter ranges for mountain goats in British Columbia and help facilitate mitigation measures to support the continued use of important winter habitat and significant landscape features that play a role in ensuring population viability and resilience through time.     

Genome-wide cross-amplification of domestic sheep microsatellites in bighorn sheep and mountain goats

We tested for cross‐species amplification of microsatellite loci located throughout the domestic sheep (Ovis aries) genome in two north American mountain ungulates (bighorn sheep, Ovis canadensis, and mountain goats, Oreamnos americanus). We identified 247 new polymorphic markers in bighorn sheep (≥ 3 alleles in one of two study populations) and 149 in mountain goats (≥ 2 alleles in a single study population) using 648 and 576 primer pairs, respectively. Our efforts increased the number of available polymorphic microsatellite markers to 327 for bighorn sheep and 180 for mountain goats. The average distance between successive polymorphic bighorn sheep and mountain goat markers inferred from the Australian domestic sheep genome linkage map (mean ± 1 SD) was 11.9 ± 9.2 and 15.8 ± 13.8 centimorgans, respectively. The development of genomic resources in these wildlife species enables future studies of the genetic architecture of trait variation.     

A Sightability Model for Mountain Goats

ABSTRACT Unbiased estimates of mountain goat (Oreamnos americanus) populations are key to meeting diverse harvest management and conservation objectives. We developed logistic regression models of factors influencing sightability of mountain goat groups during helicopter surveys throughout the Cascades and Olympic Ranges in western Washington during summers, 2004–2007. We conducted 205 trials of the ability of aerial survey crews to detect groups of mountain goats whose presence was known based on simultaneous direct observation from the ground (n = 84), Global Positioning System (GPS) telemetry (n = 115), or both (n = 6). Aerial survey crews detected 77% and 79% of all groups known to be present based on ground observers and GPS collars, respectively. The best models indicated that sightability of mountain goat groups was a function of the number of mountain goats in a group, presence of terrain obstruction, and extent of overstory vegetation. Aerial counts of mountain goats within groups did not differ greatly from known group sizes, indicating that under-counting bias within detected groups of mountain goats was small. We applied Horvitz-Thompson-like sightability adjustments to 1,139 groups of mountain goats observed in the Cascade and Olympic ranges, Washington, USA, from 2004 to 2007. Estimated mean sightability of individual animals was 85% but ranged 0.75–0.91 in areas with low and high sightability, respectively. Simulations of mountain goat surveys indicated that precision of population estimates adjusted for sightability biases increased with population size and number of replicate surveys, providing general guidance for the design of future surveys. Because survey conditions, group sizes, and habitat occupied by goats vary among surveys, we recommend using sightability correction methods to decrease bias in population estimates from aerial surveys of mountain goats.     

Estimating mountain goat abundance using DNA from fecal pellets

Non-invasive collection of tissue samples to obtain DNA for microsatellite genotyping required to estimate population size has been used for many wildlife species but rarely for ungulates. We estimated mountain goat (Oreamnos americanus) population size on a mountain complex in southwestern British Columbia by identification of individuals using DNA obtained from fecal pellet and hair samples collected during 3 sampling sessions. We identified 55 individuals from 170 samples that were successfully genotyped, and estimated a population of 77 mountain goats (SE = 7.4). Mean capture probability was 0.38 (SE = 0.037) per session. Our technique provides one of the first statistically rigorous estimates of abundance of an ungulate species using DNA derived primarily from fecal pellets. Our technique enables managers to obtain minimum counts or population estimates of ungulates in areas of low sightability that can be used for conservation and management. © 2011 The Wildlife Society.     

Survival of adult mountain goats in Washington: effects of season,translocation, snow,and precipitation

Elucidating patterns of adult survival rates is key to understanding population dynamics of large mammals. We used data from 7 separate studies of mountain goats (Oreamnos americanus) conducted from 2002 through 2022 in western Washington, USA, to quantify survival rates in relation to key biotic and abiotic factors using known fate models implemented in program MARK. We monitored 324 (206 females, 118 males) radio-marked mountain goats for 178,339 days. A substantial number of mountain goats in our sample (n = 217) had been translocated from the Olympic Peninsula to the Cascade Mountains on Washington's mainland, providing an added opportunity to examine translocation effects. We adopted a sequential modeling approach, first building a set of models to examine fundamental survival patterns by age, sex, season, study area, and translocated status. We used variables retained from the top model in a second set to investigate relationships between annual survival and local weather covariates hypothesized by previous studies to influence mountain goat behavior, habitat selection, and vital rates. Survival among adult females in spring was slightly lower than other sex and age categories, but seasonal patterns were otherwise not evident. There were significant negative relationships between survival and winter snow depth, an index of the previous year's drought, and mean daily temperature during the previous May, and a positive relationship with previous year's precipitation. Weather effects were similar among resident and translocated animals. By the end of the study period, weather effects had evidently contributed to reductions in mountain goat survival to a level low enough that population stability was unlikely. Because the frequency of droughts and warm spring temperatures are expected to increase with climate change, mountain goat populations in Washington will likely be increasingly challenged as the atmosphere warms.     

The Origin and Genetic Status of Insular Caprines in the Eastern Mediterranean: A Case Study of Free-Ranging Goats (Capra aegagrus cretica) on Crete

The history and species status of free-ranging goats inhabiting the Eastern Mediterranean islands is discussed with reference to morphometric, archaeological and genetic findings. A case study on the free-ranging goats on Crete (Capra aegagrus cretica) is presented. The phenotype of the Cretan goat resembles that of the wild bezoar goat (C. aegagrus). However, the mitochondrial DNA of cytochrome b and d-loop sequences shows affinity with domestic goats. It has been suggested that the Cretan goat represents a feral animal that was introduced onto the island during the 6th millennium b.c. as a primitive domestic, and has retained the wild morphotype but has undergone significant genetic change. An alternative explanation, and the one that is proposed here, is that the goat was introduced onto the island in wild form and released as a food source. Subsequent introgressions with domestic animals, especially ewes, have influenced its genotype. These conclusions are applicable to other free-living goats and sheep which inhabit islands in the Eastern Mediterranean.     

Mountain goat survival in coastal Alaska: Effects of age,sex, and climate

Ecological theory predicts that individual survival should vary between sex and age categories due to differences in allocation of nutritional resources for growth and reproductive activities. During periods of environmental stress, such relationships may be exacerbated, and affect sex and age classes differently. We evaluated support for hypotheses about the relative roles of sex, age, and winter and summer climate on the probability of mountain goat (Oreamnos americanus) survival in coastal Alaska. Specifically, we used known-fates analyses (Program MARK) to model the effects of age, sex, and climatic variation on survival using data collected from 279 radio-marked mountain goats (118 M, 161 F) in 9 separate study areas during 1977–2008. Models including age, sex, winter snowfall, and average daily summer temperature (during Jul–Aug) best explained variation in survival probability of mountain goats. Specifically, our findings revealed that old animals (9+ yr) have lower survival than younger animals. In addition, males tended to have lower survival than females, though differences only existed among prime-aged adult (5–8 yr) and old (9+ yr) age classes. Winter climate exerted the strongest effects on mountain goat survival; summer climate, however, was significant and principally influenced survival during the following winter via indirect effects. Furthermore, old animals were more sensitive to the effects of winter conditions than young or prime-aged animals. These findings detail how climate interacts with sex and age characteristics to affect mountain goat survival. Critically, we provide baseline survival rate statistics across various age, sex, and climate scenarios. These data will assist conservation and management of mountain goats by enabling detailed, model-based demographic forecasting of human and/or climate-based population impacts. © 2011 The Wildlife Society.     

Evidence for Introgressive Hybridization of Captive Markhor (Capra falconeri) with Domestic Goat: Cautions for Reintroduction

Markhors (Capra falconeri) are among the most endangered mammal species, and several conservation measures, including ex situ breeding, are implemented to prevent their extinction. We studied sequence diversity and differentiation of the first hypervariable segment of the mitochondrial DNA control region among C. f. heptneri and C. f. megaceros kept in four zoos in relationship to lineages of other wild and domestic goats, to assess for the first time the level of molecular distinctness and variability among those subspecies, and to check for possible introgression by related Capra taxa, such as domestic goats. Levels of differentiation between some Capra falconeri lineages and modern domestic goats were similar to levels between other wild goat species (i.e., Capra aegagrus, Capra ibex) and domestic goats. Among pure markhor lineages, paraphyly was observed for C. f. heptneri, suggesting occurrence of shared ancestral polymorphism among markhor subspecies and/or ancient or recent gene exchange between subspecies. Interestingly, 35.7% of all studied markhors from three zoos are introgressed by the domestic goat. Furthermore, despite relatively small breeding group sizes, markhors have maintained a relatively high proportion of mtDNA variation within zoo groups. In any case, the existence of markhors introgressed with domestic goat DNA in zoos should be considered when selecting markhors for ex situ breeding programs with the aim of building up a stock for later reintroduction into the wild.     

Habitat shift and time budget of the Tibetan argali: the influence of livestock grazing

Livestock production is the primary source of livelihood and income in most of the high steppe and alpine regions of the Indian Trans-Himalaya. In some areas, especially those established or proposed for biodiversity conservation, recent increases in populations of domestic livestock, primarily sheep and goats, have raised concern about domestic animals competitively excluding wild herbivores from the rangelands. We evaluated the influence of domestic sheep and goat grazing on the habitat use and time budget of the endangered Tibetan argali Ovis ammon hodgsoni in the proposed Gya-Miru Wildlife Sanctuary, Ladakh, India. We asked if the domestic sheep and goat grazing and collateral human activities relegate the argali to sub-optimal habitats, and alter their foraging time budgets. Data were collected on habitat use and time budget of a population of c. 50 argalis before and after c. 2,000 sheep and goats moved onto their winter pasture in the Tsabra catchment of the aforementioned reserve. Following the introduction of domestic sheep and goats, argalis continued to use the same catchment but shifted to steeper habitats, closer to cliffs, with lower vegetation cover, thus abandoning previously used plant communities with denser cover. Argalis’ active time spent foraging also decreased by 10% in response to the presence of livestock. These results suggest a clear disturbance effect of livestock on argalis, and indicate a potential for competition, conceivably a significant disadvantage for argalis in winter when forage availability is minimal.     

Wildlife Interactions within Highway Underpasses

Highway underpasses are a common management tool used to lessen wildlife-vehicle collisions on roadways. Despite their widespread use, the effects of predator-prey interactions and human disturbances on wildlife within underpasses have not been well studied. To understand the effect of species interactions and human disturbances on wildlife traveling through underpasses, we analyzed camera data from 3 underpasses in Hallelujah Junction Wildlife Area, Sierra County, California, USA, from June 2017 to December 2018. We recorded 3,589 detections, which were predominately mule deer (Odocoileus hemionus), rodents, lagomorphs, California quail (Callipepla californica), bobcats (Lynx rufus), mountain lions (Puma concolor), and coyotes (Canis latrans). We used occupancy modeling and daily activity estimates to analyze species' spatial and temporal activity within the underpasses. Predator-prey interactions and human disturbances were among the most important factors that influenced wildlife travel through the underpasses. Mule deer avoided underpasses highly used by mountain lions, and mountain lions followed mule deer daily temporal activity patterns and seasonal activity patterns. These results indicate that predator-prey interactions influenced deer and mountain lion use of the underpasses. Coyotes favored underpasses and seasons with higher rodent and lagomorph presence, suggesting that the presence of prey was also important to coyote use of the underpasses. Coyotes, mountain lions, and bobcats all exhibited either temporal or spatial avoidance of human activity within the underpasses. California quail avoided predators within the underpasses and favored underpasses and times with high human activity. Our study suggests that underpass managers need to closely monitor the effect of predator-prey interactions and human activity on wildlife within underpasses to ensure these interactions do not discourage wildlife from using them. © 2019 The Wildlife Society.     

Deciphering translocations from relicts in Baranof Island mountain goats: is an endemic genetic lineage at risk?

Human-mediated movement of wildlife is a common practice in North America. Some translocations have occurred where local populations were thought to be extinct or simply not present. In Alaska’s Alexander Archipelago, mountain goats (Oreamnos americanus) were not considered indigenous and were introduced to Baranof Island in 1923. However, a range-wide survey using microsatellites and mitochondrial DNA revealed a distinct genetic subpopulation endemic to the island. In this study, we attempted to clarify the evolutionary history of the mountain goats on Baranof Island by examining sequence variation in the Y chromosome. We first screened five regions of the Y chromosome in a subset of mountain goats from across their native range. We detected a single polymorphic site in the SRY promoter, and subsequently sequenced this gene in 100 mountain goats. A unique Y chromosome polymorphism was restricted to Baranof Island and an area near Haines, Alaska, and not detected in the presumed source population. An island-to-mainland dispersal scenario from a cryptic refugial population during the retreat of the Cordilleran ice-sheet would account for this distribution. Overall, these data support the hypothesis that a glacial relict population of mountain goats was present on the island prior to introduction. Based on a combination of mitochondrial, microsatellite, and Y chromosome data, we recommend recognizing Baranof Island mountain goats as an evolutionary significant unit.     

Does wild boar rooting affect livestock grazing areas in alpine grasslands?

Interactions between traditional livestock management practices and wildlife activities are important in the conservation of many mountain ecosystems including the summer rangelands in the Spanish Central Pyrenees, where rooting by wild boar (Sus scrofa) is a large disturbance that can reduce the amount of area available to grazing livestock. This study explored the likely impact of wild boar rooting on Pyrenean grasslands. It quantified the extent of wild boar rooting in livestock grazing areas and determined whether wild boars selected or avoided areas depending on the type of livestock and stocking rates. Wild boar rooting affected 16% of livestock grazing area and occurred in sites that were grazed by cattle, rather than by sheep. In addition, a preference for areas that had intermediate stocking rates was found. The relationship between the increase in the number of wild boars and trends in livestock management suggests that the extent of wild boar rooting will increase especially in cattle grazing areas, and therefore, the area available for cattle grazing in Pyrenean mountain rangelands would decrease significantly.     

Long-term data reveal effects of climate,road access,and latitude on mountain goat horn size

Potential negative artificial selection on horn size is a concern for many harvested ungulates. The mountain goat (Oreamnos americanus) has distinct black horns, but targeting animals based on horn size in the field can be challenging. We analyzed over 23,000 horn records that included base circumference and total length, from which we also derived horn volume, from mountain goats harvested in Alaska, British Columbia, and the Northwest Territories from 1980 to 2016. We tested 3 potential drivers of horn size variation: geographical location, environmental conditions, and artificial selection. We found no support for a latitudinal effect with surprisingly little variation across the sampling distribution. The Pacific Decadal Oscillation had the largest effect outside age in the model, suggesting a role of climate in shaping variation. Mountain goats harvested closer to roads had larger horns, indicating that ease of access might allow hunters to be more selective, though the effect size was small. Our findings reinforce the value of accurate and complete record keeping on horn size, age, and sex of harvested animals, and highlight the importance of explicitly considering climate and accessibility when devising management strategies for the mountain goat.     

Restoration of Insular Ecosystems: Control of Feral Goats on Aldabra Atoll, Republic of Seychelles

The control of feral goats (Capra hircus) was studied on Aldabra Atoll, Republic of Seychelles, from October 1993 to May 1994 and November 1994 to May 1995. A total of 832 goats was killed on Aldabra using the Judas goat technique and traditional hunting methods. Twenty-eight goats equipped with radio transmitters (Judas goats) were used to locate and kill other feral goats. The remnant goat populations on Ile Picard (n=13) and Ile Malabar (n=19) were eradicated during the first season. On Grande Terre, a total of 798 (374 M : 424 F) goats were killed. Mean group size was 3.2 with a range of 1–20. Judas goat hunting became increasingly important over time with 18.0% (n=85) of goats killed in the presence of Judas goats in the first season compared to 42.3% (n=126) of goats killed during the second season. The overall kill rate for the project was almost 2 times greater for Judas goat hunting (0.61 goats killed/h) than traditional hunting (0.32 goats killed/h). The home range size of each Judas goat and the number of goats killed in association with it was significantly related. Using the Leslie–Davis removal method of population estimation, 84 goats were estimated to remain on all of Grande Terre at the end of thebreak project. This revised version was published online in July 2006 with corrections to the Cover Date.     

Heading for the hills? Evaluating spatial distribution of woodland caribou in response to a growing anthropogenic disturbance footprint

Anthropogenic landscape change (i.e., disturbance) is recognized as an important factor in the decline and extirpation of wildlife populations. Understanding and monitoring the relationship between wildlife distribution and disturbance is necessary for effective conservation planning. Many studies consider disturbance as a covariate explaining wildlife behavior. However, we propose that there are several advantages to considering the spatial relationship between disturbance and wildlife directly using utilization distributions (UDs), including objective assessment of the spatially explicit overlap between wildlife and disturbance, and the ability to track trends in this relationship over time. Here, we examined how central mountain woodland caribou (Rangifer tarandus caribou) distribution changed over time in relation to (i) anthropogenic disturbance, baseline range (defined using telemetry data from 1998 to 2005), and alpine habitat; and (ii) interannual climate variation (North Pacific Index; NPI). We developed seasonal UDs for caribou in west‐central Alberta and east‐central British Columbia, Canada, monitored with GPS collars between 1998 and 2013. We mapped the cumulative annual density of disturbance features within caribou range and used indices of overlap to determine the spatial relationship and trend between caribou UDs, anthropogenic disturbance, baseline range, alpine habitat, and the NPI. Anthropogenic disturbance increased over time, but the overlap between caribou UDs and disturbance did not. Caribou use of alpine habitat during spring, fall, and late winter increased over time, concurrent with a decrease in use of baseline range. Overlap between caribou UDs and disturbance increased during spring and fall following relatively cold, snowy winters (high NPI), but overall, climate did not explain changes in caribou distribution over time. We provide evidence supporting the hypothesis that caribou populations adjust their spatial distribution in relation to anthropogenic landscape change. Our findings could have implications for population persistence if distributional shifts result in greater use of alpine habitat during winter. Monitoring long‐term changes in the distribution of populations is a valuable component of conservation planning for species at risk in disturbed landscapes.     

Genetic Variations of 13 Indigenous Chinese Goat Breeds Based on Cytochrome b Gene Sequences

Phylogenetic relationships among and genetic variability within 13 Chinese indigenous goat breeds and Boer goat were analyzed using cytochrome b gene sequences. There were 44 variable sites found in a 642 bp sequence, and 46 Cyt b haplotypes were subsequently defined. The phylogeny analysis of haplotypes in combination with goat Cyt b sequences from GenBank shows that Chinese goats are obviously separated from wild goats and might come from Capra aegagrus. Further analysis indicated that indigenous Chinese goats might descend from at least two lineages; most of the individuals analyzed could be classified into lineage A as defined by Luikart, but five other goats were of uncertain lineage. The Tibet plateau is a possible place of origin for Chinese goats. The neighbor-joining tree based on pairwise differences among populations shows that most Tibetan goats, except the Middle Tibet type, cluster closely with North China goats, and then with South China goats. This result confirms that differences in genetic structure exist among goats in different geographic locations. Nucleotide diversity varied among populations. Tibet and North China goats had higher genetic diversity than South China goats. The fixation index (F _st=87.72%) suggested that most of the total genetic variation was due to variation within populations. In addition, the results indicate that Cyt b gene sequence information alone might not be enough for phylogeny analysis among breeds within species, as shown by fewer polymorphic sites and lower bootstrap values on the neighbor-joining tree.     

Low MHC DRB class II diversity in the mountain goat: past bottlenecks and possible role of pathogens and parasites

Major histocompatibility complex (MHC) genes are the most polymorphic in vertebrates and code for molecules playing a central role in pathogen resistance. We studied levels of MHC DRB class II diversity in a long-term study population of mountain goats (Oreamnos americanus) at Caw Ridge, Alberta, and two other populations from British Columbia, Canada. Only two alleles were found among the three populations sampled. The Caw Ridge population was fixed for one of the two MHC DRB alleles, but this lack of variation did not appear to have affected it negatively because the population doubled over two decades and had no history of any apparent infectious diseases. Past population bottlenecks during Pleistocene glaciations are thought to have been the main factor contributing to the low levels of MHC diversity in mountain goats, a hypothesis supported by our previous work reporting low polymorphism at neutral loci. Additionally, the limited MHC variability in mountain goats may be related to its northern distribution as we found that allelic diversity at MHC DRB class II in wild ungulates decreases with increasing latitude, possibly as a result of low parasite diversity at high latitudes. The low MHC variation in mountain goats and other northern ungulates such as muskoxen (Ovibos moschatus) may expose these species to population outbreaks that could be generated by introduced pathogens or northward shifts in the distribution of pathogens with global climate warming.     

The wild goats Capra aegagrus Erxleben, 1777 of the Mediterranean Sea and the Eastern Atlantic Ocean islands

• 1 Populations of wild goats that can be referred to as phenotypes of the pasang, or Bezoar goat, or wild goat Capra aegagrus Erxleben, 1777, still occur on several of the Mediterranean and Eastern Atlantic islands. Other populations became extinct not many decades ago.
• 2 Fossil evidence for the natural spread of the wild goat to any of these islands has not been found. Originating in the Near East, the region of its natural range and its earliest domestication, the species was introduced by humans onto the Mediterranean islands starting as early as the Pre‐Pottery Neolithic.
• 3 The islands of the Eastern Atlantic were colonized artificially by animals of ‘pre‐Hispanic’ origin. The principal phenotypes expressed by all these populations of goats can be matched with the characteristics of both C. a. aegagrus Erxleben, 1777 (aegagrus phenotype) and C. a. dorcas Reichenow, 1888 (dorcas phenotype).
• 4 Most of these animals currently survive on the islands without being fed by humans, and should be protected.

     

A standard set of polymorphic microsatellites for threatened mountain ungulates (Caprini,Artiodactyla)

Nearly 70% of the world's mountain ungulate taxa are endangered. The availability of a standard set of DNA markers for forensic and molecular ecology studies would help to establish conservation programs and detect poaching activities of these endangered taxa. We tested 60 published microsatellite primer pairs from bovids (cattle, sheep and goat) on 49 individuals from 11 taxa including six wild goat‐like species (Capra spp.), three divergent wild sheep (Ovis spp.), and two chamois (Rupicapra spp.) species. Approximately 30 microsatellites amplified a microsatellite‐like PCR product in all three genera, and with the exception of ILST097, nearly all the loci were polymorphic within most of the 11 species.     

Characterization of 29 polymorphic artiodactyl microsatellite markers for the mountain goat (Oreamnos americanus)

We report the results of a cross‐species amplification test of 156 bovine, ovine and cervid microsatellite markers in a wild population of mountain goats, Oreamnos americanus, inhabiting Caw Ridge, Alberta, Canada. Twenty‐nine markers were found to be low to moderately polymorphic with between two to nine alleles per locus. Observed heterozygosity ranged from 0.14 to 0.85 for a sample of 215 mountain goats. This set of markers will be used in parentage analyses to construct the pedigree of the long‐term studied population and to investigate the effects of individual genetic variability on life‐history traits.