Swift Fox (Vulpes velox): COSEWIC assessment and status report 2021

Official title: COSEWIC assessment and status report on the Swift Fox (Vulpes velox) in Canada

Threatened 2021

COSEWIC assessment summary

Assessment Summary – April 2021

Common name: Swift Fox

Scientific name: Vulpes velox

Status: Threatened

Reason for designation: This small prairie canid was extirpated from Canada in the 1930s. Following reintroduction programs initiated in 1983, it has re-established in southern Alberta and Saskatchewan as well as in Northern Montana. Regular monitoring suggests that the population reached a peak in 2005 but had subsequently declined when surveyed again in 2014/15. The reason for the decline is unknown but suspected to be related to severe winter conditions in 2010/11. Occupancy surveys in 2015 and 2018 suggest the population has remained stable since 2010/11. The species persists at very low numbers. Threats include accidental or intentional poisoning, disease, habitat loss, habitat fragmentation, and severe winters.

Occurrence: Alberta, Saskatchewan

Status history: Last seen in Saskatchewan in 1928. Designated Extirpated in April 1978. Status re-examined and designated Endangered in April 1998 after successful re-introductions. Status re-examined and confirmed in May 2000. Status re-examined and designated Threatened in November 2009. Status re-examined and confirmed in May 2021.

COSEWIC executive summary

Swift Fox

Vulpes velox

Wildlife species description and significance

Swift Fox (Vulpes velox) is one of the smallest canids and one of the fastest mammals in North America, capable of reaching 60 km/h. It has an adult body length of 68–88 cm, a shoulder height of 30–32 cm and a body mass of 1.5–3.0 kg, with males being slightly larger than females. Pelage on the upper body is dark buffy-grey, with sides, legs, and beneath the tail orange-tan, and the undersides buff to pure white. Swift Fox is a grassland specialist and considered an important indicator of the integrity of the short-grass and mixed-grass prairies. Swift Fox have a strong spiritual and cultural significance for the Niitsitapi (Blackfoot Confederacy) and specifically for the Kainai (Blood) Tribe in Alberta, Canada.

Distribution

The historical range of Swift Fox extended from central Alberta, south to central Texas, and from North Dakota west to central Colorado. In Canada, it originally occurred from the Pembina Hills in Manitoba, across southern Saskatchewan, and west to the foothills of the Rocky Mountains in Alberta. Swift Fox were extirpated from Canada by the 1930s and reintroductions from 1983 to 1997 led to the current distribution. Swift Fox currently occur in the southernmost portion of the prairies on both sides of the Alberta-Saskatchewan border and in and around Grasslands National Park, Saskatchewan. The Canadian and adjacent Montana population is geographically and genetically isolated from the contiguous Swift Fox range in the United States.

Habitat

Swift Fox inhabit grasslands on level terrain or gently rolling hills, with short, sparse herbaceous vegetation. They prefer relatively dry homogeneous areas and avoid cropland and fragmented habitats. The range of Swift Fox encompasses some of the most modified landscapes in North America, and conversion of at least 70% native prairie to cropland agriculture has been implicated as a primary reason for the historical range contraction of this species. Current estimates indicate that only 19% of the Canadian Prairies ecozone remains as grassland habitat.

Biology

Swift Fox are opportunistic foragers that eat mammals, birds, insects, plants, and carrion. Swift Fox either dig their own dens or modify those of other species such as American Badger and ground squirrels. They are one of the most burrow-dependent canids and use burrows throughout the year as refugia from predators, protection from extreme weather, shelter to prevent excess water loss, periodic resting cover, and as places to raise young. Females have litters of four and up to eight pups. Pups disperse between 9.5 and 18 months of age over distances typically less than 15 km from the natal area. Swift Fox that survive their first year usually live for 3–7 years. They are territorial, with home ranges in Canada averaging 32 km² in size. Predation by Coyote is the primary known natural mortality factor in Canada, but Golden Eagles, American Badger, and possibly Red Fox can kill Swift Fox.

Population sizes and trends

The current population of Swift Fox in Canada and adjacent Montana arose from reintroductions. Four systematic winter surveys were conducted between 1996/1997 and 2014/2015 to monitor the population with estimates reaching a high of 1163 foxes (adults and juveniles combined) in 2005/2006. The most recent (2014/2015) population estimate was 523 foxes across a 14,402 km² survey area in Canada. The number of reproductively mature individuals in Canada was estimated to be 445. An inferred population decline of 45% took place between surveys conducted in 2005/2006 and 2014/2015. Camera trapping in 2015 and 2018 suggested that the population has remained stable at the lower abundance.

Threats and limiting factors

Accidental poisoning from the misuse of toxicants (rodenticides, predacides), road mortality from collision with vehicles, and interspecific competition with and predation by other sympatric canids are currently among the most important threats and limiting factors to Swift Fox in Canada. Diseases are ongoing threats to small canids around the world and this could impact Swift Fox although no significant disease outbreaks have been documented in Swift Fox yet. Limiting factors include predation by Coyote and severe winters. These threats and factors may act singly or synergistically to affect Swift Fox survival, reproductive success, and distribution.

Protection, status and ranks

Swift Fox is listed as Threatened within Schedule 1 of the Species at Risk Act, and as Endangered under the Alberta Wildlife Act and the Saskatchewan Wildlife Act (and associated Wild Species at Risk Regulations). The International Union for the Conservation of Nature (IUCN) lists the species as “Least Concern”. Under the General Status of Wild Species 2015, Swift Fox was listed as Imperilled in Canada, Critically Imperilled to Imperilled in Alberta, Vulnerable in Saskatchewan, and Presumed Extirpated in Manitoba. NatureServe ranks Swift Fox as Vulnerable across its global distribution and as Vulnerable to Critically Imperilled in Canada. The Alberta Conservation Information Management System also ranks Swift Fox as Critically Imperilled to Imperilled, the Saskatchewan Conservation Data Centre as Vulnerable, and the Manitoba Conservation Data Centre as Presumed Extirpated. A recovery strategy for Swift Fox is available and Critical Habitat has been identified in southwestern Saskatchewan and in Grasslands National Park under two multi-species action plans.

Technical summary

Vulpes velox

Swift Fox

Renard véloce

Range of occurrence in Canada (province/territory/ocean): Alberta, Saskatchewan

Quantitative Analysis

Is the probability of extinction in the wild at least [20% within 20 years or 5 generations, or 10% within 100 years]? N/A

Threats (direct, from highest impact to least, as per IUCN Threats Calculator)

Was a threats calculator completed for this species? Yes, June 2019 (Appendix 1)

Overall threat impact: High

Key threats were identified as:

What additional limiting factors are relevant?

Intraguild predation and competition (Red Fox, Coyote), and prey availability which may interact synergistically with some of the above threats.

⁺ See Table 3 (Guidelines for modifying status assessment based on rescue effect)

Data sensitive species

Is this a data sensitive species? No

Status history

COSEWIC: Last seen in Saskatchewan in 1928. Designated Extirpated in April 1978. Status re-examined and designated Endangered in April 1998 after successful re-introductions. Status re-examined and confirmed in May 2000. Status re-examined and designated Threatened in November 2009. Status re-examined and confirmed in May 2021.

Status and reasons for designation

Status: Threatened

Alpha-numeric codes: D1

Reasons for designation: This small prairie canid was extirpated from Canada in the 1930s. Following reintroduction programs initiated in 1983, it has re-established in southern Alberta and Saskatchewan as well as in Northern Montana. Regular monitoring suggests that the population reached a peak in 2005 but had subsequently declined when surveyed again in 2014/15. The reason for the decline is unknown but suspected to be related to severe winter conditions in 2010/11. Occupancy surveys in 2015 and 2018 suggest the population has remained stable since 2010/11. The species persists at very low numbers. Threats include accidental or intentional poisoning, disease, habitat loss, habitat fragmentation, and severe winters.

Applicability of Criteria

Criterion A (Decline in Total Number of Mature Individuals):

Not Applicable. May meet Threatened A2b but camera trapping surveys between 2015 and 2018 and data from adjacent Montana suggest that the population has not continued to decline since the suspected harsh-winter die-off in 2010/11.

Criterion B (Small Distribution Range and Decline or Fluctuation):

Not Applicable. Below the threshold for Threatened B1 with an EOO of 10,998 km² but no continuing decline and no identified threat that could rapidly affect the entire Canadian population, thus more than 10 locations.

Criterion C (Small and Declining Number of Mature Individuals):

Not Applicable. All individuals are part of a single population with fewer than 1000 mature individuals, but camera trapping surveys between 2015 and 2018 and data from adjacent Montana suggest that the population has not continued to decline since the suspected harsh winter die-off in 2010.

Criterion D (Very Small or Restricted Population):

Meets Threatened, D1, Number of mature individuals estimated to be 445.

Criterion E (Quantitative Analysis):

Not applicable. No quantitative analysis was performed.

Preface

Since the last assessment by COSEWIC in 2009, there has been one intensive capture-mark-recapture session in the winter of 2014/15 and two camera trapping sessions in the summers of 2015 and 2018. The 2014/15 session identified a significant decline in Swift Fox abundance which was thought to be due to a severe winter in 2010/11. The camera trapping surveys suggested that the population remained low but stable between 2015 and 2018.

Road mortality, accidental poisoning from the misuse of toxicants (rodenticides and predacides), and changes in the dynamics of interspecific competitors brought about by anthropogenic factors are thought to be important threats to the Swift Fox. Changes in federal and provincial government policies regarding community pastures since 2009 have brought uncertainty about the future of large tracts of native grassland in Swift Fox range that were previously maintained as compatible pasture land. Many of these tracts of land are now part of 15-year leases. While oil and gas development has previously been extensive in the Canadian Swift Fox range, it has largely subsided during this period, due in part to low commodity prices.

As part of the Recovery Strategy for the Swift Fox, critical habitat has been identified in Grasslands National Park, Saskatchewan, and in an area known as “South-of-the-Divide”, representing the Saskatchewan portion of the Milk River drainage basin. In addition, multiple species action plans have been developed for those areas to conserve the Swift Fox and other species at risk, and their supporting habitats, through collaboration with land owners and other stakeholders. Critical habitat is awaiting identification in Alberta.

The contiguous Canada-Montana Swift Fox population was found to exist as two clusters (‘eastern’, ‘western’) based on genetic differences. These clusters reflect the geographical distribution of the original reintroduction sites around Grasslands National Park and the Alberta-Saskatchewan border. They are indicative of reduced gene flow between the two areas, possibly due to habitat characteristics that limit dispersal.

COSEWIC history

The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) was created in 1977 as a result of a recommendation at the Federal-Provincial Wildlife Conference held in 1976. It arose from the need for a single, official, scientifically sound, national listing of wildlife species at risk. In 1978, COSEWIC designated its first species and produced its first list of Canadian species at risk. Species designated at meetings of the full committee are added to the list. On June 5, 2003, the Species at Risk Act (SARA) was proclaimed. SARA establishes COSEWIC as an advisory body ensuring that species will continue to be assessed under a rigorous and independent scientific process.

COSEWIC mandate

The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) assesses the national status of wild species, subspecies, varieties, or other designatable units that are considered to be at risk in Canada. Designations are made on native species for the following taxonomic groups: mammals, birds, reptiles, amphibians, fishes, arthropods, molluscs, vascular plants, mosses, and lichens.

COSEWIC membership

COSEWIC comprises members from each provincial and territorial government wildlife agency, four federal entities (Canadian Wildlife Service, Parks Canada Agency, Department of Fisheries and Oceans, and the Federal Biodiversity Information Partnership, chaired by the Canadian Museum of Nature), three non-government science members and the co-chairs of the species specialist subcommittees and the Aboriginal Traditional Knowledge subcommittee. The Committee meets to consider status reports on candidate species.

Definitions (2021)

Wildlife Species

A species, subspecies, variety, or geographically or genetically distinct population of animal, plant or other organism, other than a bacterium or virus, that is wild by nature and is either native to Canada or has extended its range into Canada without human intervention and has been present in Canada for at least 50 years.

Extinct (X)

A wildlife species that no longer exists.

Extirpated (XT)

A wildlife species no longer existing in the wild in Canada, but occurring elsewhere.

Endangered (E)

A wildlife species facing imminent extirpation or extinction.

Threatened (T)

A wildlife species likely to become endangered if limiting factors are not reversed.

Special Concern (SC)*

A wildlife species that may become a threatened or an endangered species because of a combination of biological characteristics and identified threats.

Not at Risk (NAR)**

A wildlife species that has been evaluated and found to be not at risk of extinction given the current circumstances.

Data Deficient (DD)***

A category that applies when the available information is insufficient (a) to resolve a species’ eligibility for assessment or (b) to permit an assessment of the species’ risk of extinction.

* Formerly described as “Vulnerable” from 1990 to 1999, or “Rare” prior to 1990.

** Formerly described as “Not In Any Category”, or “No Designation Required.”

*** Formerly described as “Indeterminate” from 1994 to 1999 or “ISIBD” (insufficient scientific information on which to base a designation) prior to 1994. Definition of the (DD) category revised in 2006.

The Canadian Wildlife Service, Environment and Climate Change Canada, provides full administrative and financial support to the COSEWIC Secretariat

Wildlife species description and significance

Name and classification

Scientific name: Vulpes velox, Say 1823

English name: Swift Fox

French name: Renard véloce

Aboriginal names: Senopah (Aamsskaapi Piikani [Blackfeet Nation]), Sinopaa (Kainai [Blood Tribe])

Classification: Class – Mammalia; Order – Carnivora; Family – Canidae; Genus – Vulpes

First described by Say (1823) as Canis velox (see Thwaites 1905), Swift Fox was re-assigned to the genus Vulpes by Audubon and Bachman (1851), and their status as a full species has been validated by genetic work (Mercure et al. 1993). Merriam (1902) gave subspecific designations of V. v. hebes (northern Swift Fox; Canada and northern United States [US]) and V. v. velox (southern Swift Fox; southern range in US). Despite significant geographic variation in morphology within the species, these subspecific assignments are not considered valid (Stromberg and Boyce 1986; Dragoo et al. 1990).

Morphological description

Swift Fox are one of the smallest North American canids, measuring 68–88 cm in length (including the tail), standing 30–32 cm at the shoulder, and with a body mass of 1.5–3.0 kg. On average, males are 8% larger than females. Winter pelage is long and dense, with dark buffy-grey upper parts, orange-tan sides, legs, and ventral surface of the tail, with buff to pure white fur underneath; summer pelage is shorter and more reddish. Swift Fox have black patches on either side of the muzzle and a black-tipped tail, which along with their small size and light colouration, distinguish them from all other foxes in Canada (Naughton 2012). Swift Fox in Canada are not known to exhibit distinct colour phases (Cypher 2003).

Population spatial structure and variability

Results of analyses from genotyped hair samples collected during live-trap surveys of Swift Fox in 2001/2002 and 2005/2006 showed the presence of two genetic clusters in the Alberta, Saskatchewan, and adjacent Montana area (Cullingham and Moehrenschlager 2013; 2019). The east and west genetic clusters generally match the geographic locations of original Swift Fox release sites in Canada even though efforts were made during the reintroductions to ensure both locations were founded from similar mixed sources (Carbyn 1998). Agriculture (crop production) was a factor believed to limit gene flow between some populations in the US (Schwalm et al. 2014). However, Cullingham and Moehrenschlager (2019) reported that the distribution of cropland did not explain the two genetic clusters, but there was a relationship between gene flow and terrain ruggedness and isolation by distance. Genetic analyses measured with 18 microsatellite loci over the two survey periods indicated that the Canada-Montana population was genetically diverse (i.e., a relatively high heterozygosity H_o: 0.476-0.858) and showed a growing total effective population in the absence of ongoing released animals (Cullingham and Moehrenschlager 2013; 2019).

Designatable units

The species is considered to be a single designatable unit in Canada. No subspecies designation is currently recognized under V. velox (Mercure et al. 1993). Temporal genetic analysis of the Canada-northern Montana population three years after the reintroduction program ended, showed high genetic diversity, a growing effective population size, and evidence of two genetic clusters (see Population Spatial Structure and Variability section above). The observed structure is thought to have arisen from founder effects of the reintroduction locations and/or the presence of a dispersal barrier between the two clusters and limited genetic mixing (Cullingham and Moehrenschlager 2019). Long-distance dispersal up to nearly 200 km has been observed (Ausband and Moehrenschlager 2009) but is thought to be rare for the species. Swift Fox typically disperse within about 15 km (Moehrenschlager 2000; Sovada et al. 2003), which would limit gene flow between groups separated by unsuitable habitat. East and west subpopulations are still highly related, and differences are thought to have arisen in part from human-related habitat fragmentation; therefore, the two clusters are not considered “discrete”.

Special significance

Swift Fox are one of the smallest canids and one of the fastest mammals on the North American prairies, running up to 60 km/h (Moehrenschlager and Sovada 2004). It is also one of the few canids in North America to use dens throughout the year. It is a specialist of short-stature grasslands (Egoscue 1979; Gese and Thompson 2014) and as such, it is considered an important indicator species of the former extent of short-grass and mixed-grass prairies (Sovada et al. 2009). As a wide-ranging meso-predator in Canada negatively impacted by habitat fragmentation (Moehrenschlager et al. 2007a), its continued presence attests to the intactness and the functioning of the remaining short and mixed grassland ecosystems.

Swift Fox has strong spiritual and cultural significance for the Kainai (Blood Tribe) in Alberta (Pruss et al. 2008) and the Aamsskaapi Piikani (Blackfeet Nation) in Montana, USA (Waters et al. 2007).

Distribution

Global range

Historically, Swift Fox occurred in mixed- and short-grass prairies from southern Alberta, Saskatchewan, and Manitoba, south to New Mexico and Texas, and from North Dakota west to Montana (Figure 1; Moehrenschlager and Sovada 2016) – an estimated range of approximately 1.5 million km² (Sovada et al. 2009).

Figure 1. Estimated historical (from USGS and IUCN 2016) and current Swift Fox distribution in North America (from Figure 2, Harrison et al. 2004; NENHP 2011; Schwalm et al. 2012; USGS and IUCN 2016; Bly pers. comm. 2018; Bjornlie pers. comm. 2018; Dowd Stukel pers. comm. 2018; CONHP 2018; Schmidt et al. 2018; Montana Field Guide 2019; NDGFD 2019). Map prepared by F. Boulin.

The most dramatic reductions in the range of Swift Fox occurred in the mid-1800s to the early-1900s due to the activities of European settlers on the Great Plains. By the late 1950s Swift Fox started to show signs of recovery in some core areas of its historical range. By 2006, Swift Fox was estimated to occupy about 30% of its historical North American range (Sovada et al. 2009).

Figure 2. Map of current Swift Fox distribution and extent of occurrence (EOO) in Canada and northern Montana. EOO boundaries are based on the 2018 camera trap survey data of Swift Fox detections (Moehrenschlager et al. 2020). Map prepared by S. Allen.

Figure 3. Evidence of Swift Fox on townships that were replicated in 2014/2015 (from release-trapping or camera trapping) and in 2018 (from camera trapping only) (from Moehrenschlager et al. 2020).

Canadian range

Prior to 1900, the Canadian distribution of Swift Fox likely approximated what is now recognized as the Mixed and Moist Mixed Grassland Ecoregions of the Prairies Ecozone (ESTR Secretariat 2014) in southern Alberta and Saskatchewan, and possibly the southwestern corner of Manitoba, an area that extends from the Pembina Hills in Manitoba west to the foothills of the Rocky Mountains in Alberta (Figure 1; Carbyn et al. 1994). The northern distributional limit in Canada was originally the 53^rd parallel in Alberta. Sovada et al. (2009) estimated the historical Canadian range to be 362 436 km², or 25% of its North American range. Swift Fox was extirpated from Canada and the last specimen was collected in 1928 near Govenlock, Saskatchewan, 14 km east of the Alberta border and 28 km north of the US border (Carbyn 1998). A sighting near Manyberries, Alberta in 1938 was reported in a 1950 newspaper article (Pied Piper 1950).

From 1983 to 1997 reintroduction efforts through captive breeding and translocation from the US allowed a Swift Fox population to become established in two areas of southwestern Saskatchewan and southeastern Alberta. As the population grew, it expanded into adjacent north-central Montana (Moehrenschlager and Sovada 2016). Observations outside the core range in Canada have been made in the Yorkton, Hudson Bay, Saskatoon, Battlefords, and Wynyard areas in Saskatchewan and in Banff National Park, Alberta (Bilyk pers. comm. 2017; Keith pers. comm. 2017). The current Canadian and adjacent Montana population is geographically and genetically isolated from the rest of the Swift Fox range in the United States (Figure 1; Schwalm et al. 2014; Alexander et al. 2016).

Search effort

Data on the distribution of Swift Fox in Canada and adjacent northern Montana come from various sources. They come from targeted surveys or incidental encounters, pre-industrial development surveys, academic or other research work added to the Alberta Fish and Wildlife Management Information System, the Saskatchewan Conservation Data Centre database, and Parks Canada Agency’s database for Grasslands National Park. Four standardized capture-mark-recapture (hereafter mark-recapture) and sign surveys have been conducted between 1996/1997 and 2014/2015 during winter months in Alberta, Saskatchewan, and adjacent northeastern Montana to assess population status and distribution (Moehrenschlager and Moehrenschlager 2018). The original Canadian Swift Fox population’s suspected range was determined by the National Swift Fox Recovery Team in 1996 and comprised 108 townships in Canada, of which about 75% were randomly selected for subsequent sampling (Cotterill 1997). An additional 80 townships (75% were randomly sampled) were added in Montana in 2000/2001, 49 in 2005/2006, and another 19 in 2014/2015, for a combined total of 256 townships (Moehrenschlager and Moehrenschlager 2018). In 2014/2015, 189 townships were sampled in Canada and Montana (102 in Canada), including sites in 170 townships that had been surveyed in 2005/2006 and 19 new townships. Both mark-recapture and camera-trapping were used for the survey, with some townships surveyed using both techniques for comparison. Mark-recapture trapping was conducted in 94 townships (42 in Canada) and camera-trapping in 133 townships (98 in Canada) in a minimum convex polygon area of 32,608.5 km² (Moehrenschlager and Moehrenschlager 2018). An additional round of camera trapping following the methods and placement of the 2014/2015 survey was conducted in the summer of 2015 and 2018 (Moehrenschlager et al. 2020).

Extent of occurrence and index of area of occupancy

Canada

The extent of occurrence (EOO) of the Canadian segment of the Swift Fox population as calculated from the 2014/2015 survey data, covered 10,998 km² (Figure 2). This is a 22% decrease from 14,038 km² calculated in 2005/2006. The EOO for sample locations with Swift Fox detections in 2018 is estimated at 6,405 km² (Moehrenschlager 2021 pers. comm.). These estimates are not directly comparable, but the number of townships with Swift Fox presence detected in the 2015 and 2018 can serve as an index of change in distribution. In both surveys, 67 townships were replicated and 17 and 16 townships had detections in 2015 and 2018, respectively (Moehrenschlager et al. 2020).

The index of area of occupancy (IAO) for the Canada portion of the range remained relatively stable between 2005/2006 (4,655 km²) and 2014/2015 (4,411 km²) (Moehrenschlager and Moehrenschlager 2018). The IAO was calculated by Moehrenschlager and Moehrenschlager (2018) for 2014/2015 (and recalculated for 2005/2006) as follows: 1) initial captures from mark-recapture and camera-trapping were plotted; 2) all trap sites on transects that had captures were buffered by 3.19 km (approximate radius of 32 km² Swift Fox home range – inferred occupation) and the number of 2 x 2-km grid cells overlapped by occupied buffer zones were identified; 3) the number of inferred occupied grid cells was adjusted using a probability of detection of 0.865, which is the average of estimated mark-recapture and camera-trapping samples; 4) the proportion of occupied grid squares was determined by dividing the number of squares inferred in step #3 by those determined in step #2; 5) the total number of grid squares overlapping the sampled population was counted; 6) the proportion of occupied grid squares from step #4 was multiplied by the total number of grid squares comprising the Swift Fox study area; and 7) the number of grid squares inferred to have Swift Fox presence was multiplied by 4 km² to obtain the IAO.

Montana

The EOO of the Montana segment of the Swift Fox population from the 2014/2015 survey data covered 6,916 km². This is a 40% decrease from 11,495 km² calculated in 2005/2006. The IAO in Montana also showed a decrease of 54% between the two survey periods, from 4,314 km² in 2005/2006 to 1,999 km² in 2014/2015, largely due to fewer occupied townships in 2014/2015 (Moehrenschlager and Moehrenschlager 2018).

Canada/Montana

The EOO of the entire Canada-Montana population of Swift Fox as calculated from the last survey (2014/2015) covered 19,779 km². This is a decrease of 10% from the 21,954 km² calculated in 2005/2006. The EOO for the area sampled in 2018 is 14,713 km² (Moehrenschlager 2021 pers. comm.). However, the 2014/2015 and 2018 EOOs cannot be directly compared as the 2018 EOO does not include off transect effort that was previously included. The IAO for the entire Canada-Montana range decreased by 29% between 2005/2006 and 2014/2015 from 8,969 km² to 6,410 km², largely due to a reduced number of occupied areas in Montana in 2014/2015 (Moehrenschlager and Moehrenschlager 2018). The IAO from sample locations with Swift Fox detection in 2018 is 2,928 km², although this estimate cannot be directly compared to previous estimates (Moehrenschlager 2021 pers. comm.).

Habitat

Habitat requirements

Primary habitat for Swift Fox is short- or mixed-grass prairie on level or gently rolling terrain (Moehrenschlager and Sovada 2016). Individuals usually select areas where vegetation is short (30 cm or less) and sparse, and with minimal elevational changes or topographic features such as canyons, steep hills, or coulees (Whitaker 1997; Russell 2006; Moehrenschlager et al. 2007a) but will sometimes select slightly raised sites within those areas to den (Pruss 1999; Tannerfeldt et al. 2003). Those areas offer greater visibility and mobility, facilitating prey hunting and detection and evasion of predators (Sovada et al. 2009).

In Canada and adjacent Montana, Swift Fox prefer native grassland over cropland (Carbyn 1998). In other regions of the range, Swift Fox have adapted to atypical habitats [e.g., mixed agricultural areas (Moehrenschlager and Sovada 2016), sagebrush steppe and short-grass prairie transition (Olson and Lindzey 2002), and piñon-juniper habitat (Covell 1992)]. In Kansas, for example, Swift Fox use fallow wheat or milo fields, winter wheat fields, and dryland crop fields (most fields fallowed every other year) and exhibited similar survival rates and home range sizes to those living in native prairie habitat (Sovada et al. 2003). Farmlands within the range of Swift Fox in Canada do not appear to be similarly usable. Annual crop production and associated tillage, or other potential ecological factors, such as the abundance of competing Red Fox (Moehrenschlager et al. 2004) may preclude their use by Swift Fox (Moehrenschlager et al. 2007a). Availability of mixed-grass prairie in Alberta and Saskatchewan therefore appears critical to sustain this reintroduced population (Carbyn 1998).

Dens are a fundamental element of habitat for this canid, and several are used throughout the year (Allardyce and Sovada 2003). Den distribution and density is important in terms of providing shelter for rearing young, providing protection against extreme climate conditions and for avoiding Coyote (Canis latrans) or Golden Eagle (Aquila chrysaetos) predation (Tannerfeldt et al. 2003; Moehrenschlager et al. 2007b). Two types of dens are recognized: natal dens, where young are born and raised and shelter dens that provide protection from weather extremes and escape refugia from predators (Tannerfeldt et al. 2003). Natal dens typically have a more complex structure with several entrance holes (four in top 75% quartile in Canada; Pruss 1999), while shelter dens may have only one opening (Tannerfeldt et al. 2003). Use of up to 14 dens in a season has been recorded (Kitchen et al. 1999). The highest number of dens is used during the pup-rearing season (Schauster et al. 2002). Pairs in Canada have been documented to use up to eight different den sites during the pup whelping/rearing period (Pruss 1994; Tannerfeldt et al. 2003). Den shifts, typically less than 500 m in Canada, are frequent (Pruss 1994; Tannerfeldt et al. 2003). However, a pair in the Alberta-Saskatchewan border area was observed moving a litter of seven pups across a highway to a new den 1.9 km from their original one (Tannerfeldt et al. 2003).

The presence of fossorial species, like American Badger (Taxidea taxus), ground squirrels (Urocitellus spp.), and prairie dogs (Cynomys spp.) in Swift Fox habitat is therefore desirable, providing burrows, which they may readily modify (Tannerfeldt et al. 2003). However, they may use unmodified badger or similar sized holes as shelter dens (Moehrenschlager et al. 2007b) or excavate their own dens (Hillman and Sharps 1978). Anthropogenic habitats have also been used for denning such as roadsides, culvert pipes, cemeteries, and farm buildings (Kilgore 1969; Hillman and Sharps 1978; Tannerfeldt et al. 2003). In Canada, badger excavations appear to be their main source of burrows (Pruss 1999; Moehrenschlager et al. 2007b) with badger burrow density being directly related to the density of ground-squirrel openings (Proulx et al. 2011).

Moehrenschlager et al. (2007a) analyzed population survey data to assess critical habitat characteristics that predict Swift Fox occurrence. They developed model parameters from winter live-trap survey data collected in Alberta, Saskatchewan, and Montana between 15 October 2005 and 15 February 2006. They also used satellite image data from August 1999 and September/October 2000 to assess 15 environmental variables in the trapping area that might be useful in predicting Swift Fox distribution. Four habitat parameters successfully explained variation in Swift Fox presence: moisture, topography, cropland, and habitat fragmentation. Swift Fox primarily occurred in relatively dry areas and tended to avoid areas with large elevation changes, preferring habitats with gently sloping terrain. Cropland and the presence of Swift Fox were negatively associated. Swift Fox also avoided fragmented landscapes, edge habitats, and roads. Habitat parameters affected the occurrence of Swift Fox more than 5 km from their capture site, likely owing to home range size (Moehrenschlager et al. 2007a). From October 2008 to April 2009, A. Moehrenschlager conducted surveys with scent-post camera-traps in 32 previously sampled and 28 unsampled townships in the western half of the Canadian Swift Fox range. Results from this effort suggest that the model is not only predictive of fox presence/absence among surveys, but also highly predictive of Swift Fox presence in previously unsurveyed regions (A. Moehrenschlager in COSEWIC 2009).

Home ranges of Canadian and Montana Swift Fox are among the largest in the species’ range. Average home range size of 36 Swift Fox in the Alberta-Saskatchewan border region was 40.8 ± 6.1 km², based on the adaptive kernel (ADK) method (Worton 1989) (31.9 ± 4.8 km² based on the fixed kernel method [FK; Seaman and Powell 1996]; Moehrenschlager et al. 2007b). In north-central Montana, Zimmerman et al. (2003) estimated ADK home ranges between 8.7 km² and 20.3 km² for five individuals. In north-eastern Montana, Swift Fox home ranges were similar to those in Canada at 42.0 ± 4.7 km² (FK; Butler et al. 2020). In western Kansas, Sovada et al. (2003) estimated an average home range of 15.9 ±1.6 km² (ADK) for 21 adult Swift Fox, with similar estimates in both cropland and rangeland.

Habitat trends

The substantial conversion of native prairie habitats to farmland was one of the main contributors to the decline in Swift Fox across the Great Plains of North America (Egoscue 1979). Sovada et al. (2009) estimated the historical range of Swift Fox largely based on the extent of shortgrass and mixed-grass prairie ecosystems but also based on published accounts and historical records of Swift Fox and expert opinion. They determined that about 39% of high-quality (short to medium height perennial grass without shrubs) and 10% of less suitable (medium height perennial grass with sparse shrubs or tall grass) grassland habitats were still available from the estimated historical range of 1.5 million km². They considered an additional 25% of agricultural lands predominantly under dryland cropping practices as potentially suitable habitats. They estimated that in the remaining 39% of high-quality grassland habitat, 41% (sampling units: counties in the United States, townships in Canada) of those were occupied by Swift Fox. In Canada alone, they estimated that 39% of the former 362,436 km² Swift Fox range still contained high-quality grassland habitats and about 5% was occupied by the species (Sovada et al. 2009).

The Ecosystem Status and Trends Report (ESTR) Secretariat (2014) reported that an estimated 70% of the native prairie vegetation in the 465,094 km² Prairies Ecozone was lost prior to the 1990s (most of it prior to the 1930s, due to conversion to cropland) and that less than 25% of the ecozone remained as native grassland. Using agriculture census data from Statistics Canada, the ESTR Secretariat (2014) considered that the percentage of “natural land for pasture” decreased from 27% in 1971 to 24% in 1986 and remained around that level until at least 2006. However, grassland loss continues; an overall 10% loss of native grasslands was recorded on transects in some parts of the Prairies Ecozone between 1985 to 2001 (ESTR Secretariat 2014).

A recent predictive model of Swift Fox occurrence was developed by Parks Canada and A. Moehrenschlager using the preliminary approach of Moehrenschlager et al. (2007a) to identify critical habitat for the species. Fourteen summer landscape-scale habitat variables were analysed within 3 km of Swift Fox captures using data from the 2005/2006 winter mark-recapture survey. Results tested using previous and subsequent winter survey data (1996/1997, 2000/2001, 2008/2009) indicated that 53% (approx. 8765 km²) of the total area of the species’ range provided habitat attributes that were suitable to contain 89% of Swift Fox occurrences (Parks Canada Agency 2016), which is about 2.4% of the former Swift Fox range, as estimated in Sovada et al. (2009).

Biology

Life cycle and reproduction

The life cycle of Swift Fox varies with latitude. In the southern portion of their range, they breed from December to early January (Kilgore 1969) and as late as mid-February to March in Canada (Asa and Valdespino 2003). Females are monoestrous (Allardyce and Sovada 2003) and gestation is about 51 days (Schroeder 1985). Litter size is usually about four pups (Olson and Lindzey 2002) but can be as high as eight (Carbyn et al. 1994; Moehrenschlager 2000). In Canada, litter size was correlated to female’s body weight prior to the mating season (Moehrenschlager 2000). Pups emerge from the natal den at about 3-4 weeks of age and are weaned at 6–7 weeks (Kilgore 1969; Hines 1980). In Canada, emergence dates were May 25 - June 9 and timing appeared related to weather conditions during the months leading up to, and including, the breeding season (Pruss 1994; Moehrenschlager 2000). Both males and females are sexually mature and can breed in their first year, but not all actually do. In Colorado, New Mexico, Texas, and northern Montana, 50% of juvenile females with known fate that survived to breeding age reproduced as yearlings (Kamler et al. 2004; Ausband and Foresman 2007). In Canada, 85% of females (including 1 year olds and likely males; Moehrenschlager pers. comm. 2018) are reproductively active (Moehrenschlager et al. 2004). Using inferred (to account for pup [0-6 months-old] mortality) juvenile survival rate to 12 months and actual average adult survival rate of 0.45 for Swift Fox in Canada (Moehrenschlager et al. 2007a,b), average fertility of 3.8 pups per reproductive pairs (Moehrenschlager et al. 2007a), and assuming post-first year fertility and survival rates to remain constant over an assumed 9 years (maximum age of reproduction for females; Moehrenschlager et al. 2007a), the generation length in Canada was calculated to be 2 years using the formula in option 1 of the IUCN Standards and Petitions Subcommittee (2017).

Swift Fox appear to be highly plastic in their social grouping strategies. A typical social group consists of a mated pair with pups but breeding groups of various composition exist. They generally maintain stable pair bonds (Kilgore 1969) but do not necessarily mate for life (Kitchen et al. 2006). Extra-pair mating occurs frequently (Kitchen et al. 2006). In Alberta, a male was observed with the litters of two different females on the same day (Moehrenschlager 2000). Trios (two females and one male or two males and one female) and even quartets (one male and three females and two males and two females) occasionally form, with some individuals of the social unit sometimes being related (Covell 1992; Schauster et al. 2002; Kitchen et al. 2006; Poessel and Gese 2013). Kamler et al. (2004) assessed the relationship between Swift Fox density, mating systems and group structure in northwestern Texas. Polygynous groups, communal denning, and non-breeding females occurred in the high Swift Fox density area with lower mortality rate from Coyote predation, whereas only monogamous pairs occurred at the low-density area where mortality from Coyote predation was significantly higher (Z = 2.55, P = 0.01). Non-breeding individuals may act as helpers to raise pups (Tannerfeldt et al. 2003).

Swift Fox generally live 3 - 7 years in the wild (Mamo 1994; Reid 2006). In the Alberta-Saskatchewan border region, annual adult Kaplan-Meier survival rates ranged from 0.38 (95% CI = 0.15-0.60) to 0.52 (95% CI = 0.29-0.72) in 1995-1998 (Moehrenschlager et al. 2007b), similar to estimates from the core range in the US (Schauster et al. 2002; Zimmerman et al. 2003). Juvenile (collared at 6 months of age) survival rates for the same period were 0 – 0.63 (95% CI =0.23-0.86) (Moehrenschlager et al. 2007b) and was similar to the US (i.e., 0.13 – 0.69, Covell 1992; Sovada et al. 1998; Kitchen et al. 1999; Schauster et al. 2002).

Diet and predation

Swift Fox are opportunistic omnivorous meso-predators with diets that vary geographically and seasonally, depending on the diversity and abundance of local prey species (Moehrenschlager et al. 2004). In the Alberta-Saskatchewan border region, scat analysis revealed that mammals are the most frequent prey (68.3%; composed of 59.5% rodents, 7.1% lagomorphs, and 1.7% large mammals [presumably carrion]), followed by insects (23.8%), and birds (8.0%) (Moehrenschlager et al. 2007b). Reynolds et al. (1991) found that small rodents and carrion from ungulates were most important with a small portion from lagomorphs and ground squirrels. In northwestern Texas, mammals and insects dominated the diet of Swift Fox. Mammals (rodents) were most important in all seasons, except autumn when insects became more important (Kamler et al. 2007). Murid rodents were the most frequent prey of Swift Fox in Kansas and Nebraska (Hines and Case 1991; Sovada et al. 2001), although hare (Lepus spp.) were also common (Cutter 1958; Cameron 1984; Zumbaugh et al. 1985). Black-tailed Prairie Dogs (Cynomys ludovicianus) are most important in South Dakota (Uresk and Sharps 1986), but unlikely to be important in Canada because of their rarity and limited distribution to only Grasslands National Park and neighbouring Masefield and Dixon community pastures (Tuckwell and Everest 2009). In general, prey items are not larger than White-tailed Jackrabbits (Lepus townsendii; 3 – 4 kg). In winter, Swift Fox in Canada rely heavily on voles (Moehrenschlager et al. 2007b). During 9360 trap-nights from November 1995 to March 1996 in the same study area as Moehrenschlager et al. (2007b), Klausz (1997) only caught 163 small mammals. This small abundance, along with low species diversity (96% were Deermouse [Peromyscus maniculatus] and 4% were shrews [Sorex spp.]), led to the hypothesis that low hunting success in the late winter period could lead to high mortalities of Swift Fox, as they would presumably have exhausted their fat reserve (Moehrenschlager et al. 2007b).

Mortality is primarily attributable to natural predation or human causes. There is no evidence that disease is currently an important source of mortality (Moehrenschlager et al. 2004). Direct human-caused mortality is caused by poison, shooting, collisions with vehicles, and trapping (Allardyce and Sovada 2003). Predation by Coyotes is the principal cause of natural mortality across the species’ range but Golden Eagle predation may also be important in some years in Canada (Allardyce and Sovada 2003, Moehrenschlager et al. 2007b).

Physiology and adaptability

Swift Fox are well adapted morphologically and behaviourally for life in an open semi-arid environment where water may be scarce, climate can be extreme, and shelters are limited. Morphologically, their slender skeleton and long legs are adapted for running (Egoscue 1979), which helps them evade predators and hunt fast prey such as lagomorphs. Swift Fox grow a longer and denser pelage for winter and the foot pads are almost completely covered with fur, thus improving body insulation (Egoscue 1979; Geffen and Girard 2003). In summer, a thin coat and a small body with a large surface to volume ratio dissipates heat effectively (Geffen and Girard 2003). Swift Fox can maintain water balance with food alone (Geffen and Girard 2003). When freestanding water is available, a Swift Fox requires about 210 g of food per day; in the absence of freestanding water, moisture requirements can be satisfied by eating about 330 g per day (Flaherty and Plaake 1986).

Dispersal and migration

Swift Fox dispersal takes place primarily as juveniles but can take place as adults as well. Pups break close associations with their parents at 4 – 6 months (Covell 1992) and, in Canada, begin dispersal in August with all pups dispersed by 18 months of age (Pruss 1994; Moehrenschlager 2000). Juvenile dispersal in northeastern Colorado involved three strategies: some juveniles remained in their natal territory helping the mated pair to raise their litter, while others dispersed to neighbouring or to distant territories (Schauster et al. 2002). Juvenile dispersal distances average 10 - 15 km and are generally similar for males and females in northern populations (Moehrenschlager 2000; Schauster et al. 2002; Sovada et al. 2003; Ausband and Foresman 2007), but distances > 50 km and up to 191 km have been reported from a reintroduced northern population in Montana (Ausband and Moehrenschlager 2009). In northeastern Colorado, monitored adult foxes dispersed 12 km (on average), largely during the breeding/gestation season (7 - 12%) but also during the pup-rearing and dispersal seasons (3% each; Schauster et al. 2002).

Interspecific interactions

Coyotes are important predators of Swift Fox in Canada (Moehrenschlager et al. 2007b) and, along with Red Fox, compete with Swift Fox (Moehrenschlager and Sovada 2016; see the Threats and Limiting Factors section).

Population sizes and trends

Reintroductions

Canada

The current population of Swift Fox in Canada is derived from reintroductions into Canada and adjacent Montana. From 1983 to 1996, 479 Swift Fox were released in the Alberta-Saskatchewan border area, and from 1990 to 1997, 420 individuals were released into the Grasslands National Park and Wood Mountain areas (Carbyn 1998). About 90% of the Swift Fox released in Alberta and Saskatchewan came from captivity (Carbyn 1998). These were supplied by Cochrane Ecological Institute (formerly the Wildlife Reserve of Western Canada) of west-central Alberta which began a Swift Fox breeding program in 1971 using animals sourced from Colorado (Moehrenschlager and Moehrenschlager 2001; Smeeton and Weagle 2000). Other facilities that contributed include: Calgary Zoo (Alberta; 1983–1994), Moose Jaw Wild Animal Park (Saskatchewan; 1984–1995), and Edmonton Valley Zoo (Alberta; 1989–1997) (Carbyn 1998). From 1973 to 1986, 151 Swift Fox were translocated from the US to support either the release or captive-breeding programs, with 99 from Wyoming, 40 from Colorado, and 12 from South Dakota (Carbyn 1998). Breeding records were kept by the breeding facilities and pairing was done to ensure maximum genetic heterozygosity with an inbreeding coefficient of <0.05 (Smeeton et al. 2003). Animals carrying the same bloodlines were released into different reintroduction areas. The last releases occurred in 1996 and 1997 into the border and Grasslands National Park areas, respectively. By 2001, the Swift Fox population had significantly increased in abundance and distribution and nearly all captures during Swift Fox surveys were of wild-born individuals that included juvenile and older animals (Moehrenschlager and Moehrenschlager 2001). Details about the reintroduction program are found in Carbyn (1998).

In 1989, a reintroduction was also attempted in the Milk River Ridge area of south-central Alberta with the release of 61 individuals (Brechtel et al. 1993). This program was discontinued after only one year due to high predator abundance and intensive predator and rabies control programs (Brechtel et al. 1993). Since that time, there has not been any compelling evidence of Swift Fox presence in the region (COSEWIC 2009).

In 2004, 15 Swift Fox were released onto Kainai (Blood Tribe) lands in southwestern Alberta in a 1,424-km² area of fescue prairie and foothills (Smeeton 2006). This area, to the west of current Swift Fox range, was within dispersal distance of the Aamsskaapi Piikani (Blackfeet Nation) in Montana where reintroductions during 1998−2002 had established a small population of foxes (Ausband and Foresman 2007). A verbal report to the Swift Fox Recovery Team from Blood Tribe land management staff in 2005 (J. Nicholson in COSEWIC 2009) indicated that the five radio-collared individuals from this effort either went missing (3), shed their collar (1) or were found dead (1), and there has been no additional documented follow-up.

In the US

Swift Fox were reintroduced into the Blackfeet Nation (1998-2002) and Fort Peck Tribal Lands (2006) in Montana, and Bad River Ranches (2007), Lower Brule Tribal Lands (2006), and Badlands National Park (2007) in South Dakota (Sovada et al. 2009). One female captured as juvenile in 2003 on the Aamsskaapi Piikani lands east of Glacier National Park in north-central Montana, dispersed a straight-line distance of 191 km across a highly fragmented landscape to southeastern Alberta. She was re-captured within 3 km of Swift Fox males in February 2006 and potentially represents gene flow to the Canadian Swift Fox population (Ausband and Moehrenschlager 2009). This is the only record of a Swift Fox from another reintroduced population having dispersed into the Canadian population. In the autumn of 2020, 27 Swift Fox were introduced on to Tribal lands in northern Montana (Fox 2020). If they establish, they will represent the first new genetics introduced into the linked Canada-US population in a decade (A. Moehrenschlager pers. comm. 2021).

Sampling effort and methods

Population surveys have been conducted four times since reintroductions stopped; from 31 October until mid-February in 1996/1997, 2000/2001, 2005/2006, and 2014/2015 in Alberta, Saskatchewan, and adjacent northeastern Montana (see Search Effort). The four surveys have used standardized field methodology using catch and release of live animals using baited box traps, combined with incidental sightings. These surveys were conducted at the scale of townships in Alberta, Saskatchewan, and Montana. However, number and distribution of the townships that were sampled in each survey varied over time. Several townships remained constant between each survey and were termed “replicated townships” throughout this document. The 2014/2015 survey also tested camera-trapping as an alternative method to catch-and-release for occupancy objectives and this was followed up in the summer of 2015 and 2018 with surveys using only camera-trapping (Moehrenschlager and Moehrenschlager 2018; Moehrenschlager et al. 2020). Although camera trapping alone can not provide abundance information, as individual foxes are not recognizable, it can be used to infer trend.

During the 2014/2015 survey, Moehrenschlager and Moehrenschlager (2018) sampled 189 townships in Canada and Montana. Sampling was conducted in 102 townships in Canada and 87 in Montana. Of these, replicated sites were surveyed in 170 of 191 townships that had been sampled in 2005/2006 plus an additional 19 townships. In 2018, 147 townships in Canada (n = 67) and Montana (n = 80), which had been sampled in 2014/2015, were sampled using camera traps alone (Moehrenschlager et al. 2020).

Individual release or camera-traps were placed at 1-km intervals along 5 km continuous sections of trail that were closest to the centre of respective townships. Distance adjustments, by up to 100 m, were made to allow trap placement along fences or on top of hills. Each township was surveyed with six camera or release traps for three nights. Release-trapping was conducted on consecutive nights when possible, but this was dependent on weather conditions. Both mark-recapture and camera-trapping were tested at 38 townships in Canada to compare the relative efficiency of each method with similar detection efficiencies (Moehrenschlager and Moehrenschlager 2018).

Abundance

Using the same methods and effort as in 2005/2006, Moehrenschlager and Moehrenschlager (2018) conducted mark-recapture trapping and captured a total of 63 individual Swift Fox distributed across the Canada-Montana survey range. Another 15 foxes were inferred to occur from the photographs taken with the camera-trapping method. Using Spatially Explicit Capture-Recapture (SECR) methods (Efford 2017; R Core Team 2015), they estimated Swift Fox abundance for the 2,148.96 km² SECR-generated effective sampling area (ESA). By dividing the estimated abundance by the ESA, they obtained a Swift Fox density of 0.0036 ± SE 0.008 foxes / km². By extrapolating across the entire Canada-Montana survey range, they estimated that 522.9 ± SE 115.2 foxes occurred across the 14,402 km² of survey range in Canada, 346.9 ± SE 79.5 foxes across the 9,562 km² in Montana, and 870.1 ± SE 191.7 foxes across the entire 23,964 km² Canada-Montana sample range (Moehrenschlager and Moehrenschlager 2018).

Male and female Swift Fox are reproductively mature at one year of age (Moehrenschlager 2000) and 85% of females (likely similar for males; Moehrenschlager pers. comm. 2018) are reproductively active in the population (Moehrenschlager et al. 2004), therefore the number of mature individuals in the Canadian Swift Fox population was estimated as 0.85 X 522.9 = 445 individuals at the end of the 2014/2015 survey.

Fluctuations and trends

The 2014/2015 Swift Fox survey is the first one to indicate a decline in the post-reintroduction Swift Fox population since surveys began in 1996/1997. Abundance estimates were derived from slightly different methods and thus cannot be directly compared with previous ones. However, Swift Fox abundance is thought to have decreased between 2005/2006 and 2014/2015 (Moehrenschlager and Moehrenschlager 2018). Alternatively, percent change in replicated townships with evidence of Swift Fox (from release-trap or camera-traps) between survey years can be used as an index of population change. Replicated townships with evidence of Swift Fox went from 39.7% in 2000/2001 and climbed to 52.1% in 2005/2006, for an increase of 31% between the two survey years (Moehrenschlager and Moehrenschlager 2006). Between the 2005/2006 and 2014/2015 surveys, replicated townships with evidence of Swift Fox declined from 44.7% to 24.7%, respectively, indicating a decrease of about 45% (Moehrenschlager and Moehrenschlager 2018). Camera trapping in 147 townships across Canada and Montana that were replicated in 2015 and 2018 detected Swift Fox in 36 and 40 townships, respectively, suggesting the population has remained stable (Moehrenschlager et al. 2020). Within Canada, foxes were detected in 17 and 16 townships in 2015 and 2018, respectively (Moehrenschlager et al. 2020).

Accounting for respective capture and recapture probabilities, mark-recapture abundance estimates at the 94 sampling sites where release-trapping was replicated were 68.9 ± SE 15.4 (95% C.I. 59.3 - 90.0) for 2014/2015 compared to 243.4 ± SE 15.4 (95% C.I. 218.8 - 280.1) for 2005/2006. This represents a decrease of 75.4% between those two most recent survey years using the IUCN Criterion A model for exponential decline calculated with two data points. However, 12 replicated townships that had evidence of foxes in 2014/2015 did not have evidence in 2005/2006, which slightly offset the decrease in replicated townships with captures and/or fox abundance at sampling sites (Moehrenschlager and Moehrenschlager 2018).

Incidental evidence from camera trap surveys conducted in 2009 in a sample of previously surveyed townships did not indicate any drastic declines, suggesting that the population decline happened after 2009 but before 2014/2015 (Moehrenschlager and Moehrenschlager 2018). Because of the sampling interval, it is difficult to pinpoint what might have caused the decline. The 2005/2006 Canadian Swift Fox population reached a high during the time of a sustained high abundance of Richardson’s Ground Squirrel in the prairies between 2000 and 2009 (Proulx 2010). Winter severity is thought to have been a contributing factor to the decline and likely affected the Swift Fox population uniformly. The winter of 2010/2011 was particularly severe and energetic constraints could have acted synergistically with increased mortality from predation and competition to drive down the population (Moehrenschlager and Moehrenschlager 2018).

Rescue effect

Repopulation of the Canadian segment of the Swift Fox population in the event of a decline or extirpation may be possible with foxes dispersing from adjacent Montana. With the reintroduction efforts that took place in Canada between 1983 and 1997 (Carbyn 1998), the growing Canadian Swift Fox population has expanded into adjacent land in Montana (Moehrenschlager and Moehrenschlager 2006). Although the range has expanded southward since the reintroductions it has not reconnected with the Swift Fox range in the south which creates northern (contiguous Canada-Montana population) and southern (core) segments to the global Swift Fox range (Figure 1). With the northern segment occupying parts of Alberta and Saskatchewan in Canada and parts of Montana in the United States. Results of genetic analyses from hair samples taken during two survey years (2000/2001 and 2005/2006) across the contiguous Canada-Montana population confirmed genetic exchange across the US border (Cullingham and Moehrenschlager 2013). The north-central Montana Swift Fox would therefore represent an important rescue source providing that 1) surplus individuals were available in Montana to disperse into Canada, 2) the factor(s) that have caused the demise of the Canadian segment of the population were not affecting the Montana segment, and 3) suitable habitat was still available in Canada. As the Canadian and Montana populations are contiguous and responding to threats and limiting factors in a similar way, rescue is likely to be possible but limited.

On a broader scale, evidence from field surveys and genetic analyses indicate that the Canada-Montana population is geographically and genetically isolated from the core Swift Fox range to the south in the US (Schwalm et al. 2014; Alexander et al. 2016, Butler et al. 2020). Despite the nearly 200 km dispersal of a juvenile Swift Fox between the Blackfeet Reservation in northern Montana and the contiguous Canada-Montana population, long-distance dispersals across fragmented habitat are uncommon (Ausband and Moehrenschlager 2009), limiting the rescue potential from the core Swift Fox range in the US.

Threats and limiting factors

Canadian Swift Fox faces multiple threats and limiting factors in its changing habitat, which may act singly or cumulatively to affect the population. Important threats include accidental poisoning from the misuse of rodenticides or predacides and disease. Important limiting factors include predation and interspecific competition by and with other canids and food availability, particularly during severe winters.

Threats that have been identified are categorized below, following the IUCN-CMP (International Union for the Conservation of Nature – Conservation Measures Partnership) unified threats classification system (based on Salafsky et al. 2008). They are listed in order of decreasing severity of impact, ending with those for which scope or severity is unknown. The overall threat impact is considered to be High (see Appendix 1 for details).

Threats

IUCN 9. Pollution (high-medium threat impact)

Agricultural and forestry effluents - accidental poisoning

Historically, poisoning for predator control had serious impacts on Swift Fox populations, likely contributing to their decline in the early 20th century (Scott-Brown et al. 1987). Direct poisoning of Swift Fox is now illegal in Canada and therefore has declined as a threat. However, as a carnivore, Swift Fox is at risk of secondary poisoning from consuming prey items poisoned by rodenticide or anticoagulants or from poisoned baits (toxicants) used for control of other predators (Proulx 2011).

Following a Richardson’s Ground Squirrel population outbreak on the prairies, the Pest Management Regulatory Agency (PMRA) of Canada approved an Emergency Registration program which allowed the restricted use of 2% liquid strychnine concentrate (LSC) for severe Richardson’s Ground Squirrel infestations starting in 2007 in Saskatchewan and in 2008 in Alberta. The label provides a precautionary warning to the user to “not apply the product if these (Swift Fox or Burrowing Owl) or other species at risk that may feed on strychnine bait or ground squirrels are present in your area” (PMRA 2012).

Zinc phosphide is another acute poison registered for control of ground squirrels, Northern Pocket Gopher (Thomomys talpoides), and other rodents in Canada. Because it rapidly converts to phosphine gas and is not retained in toxicologically significant amounts in body tissue of the primary consumer, zinc phosphide is considered of low secondary hazard, except to raptors or other scavengers that may consume the gastrointestinal tract which may still contain undigested bait (Erickson and Urban 2004). As per the label, users must ensure that the bait is applied only to active burrows and ensure that there is no evidence of species at risk activity or presence in burrows (PMRA 2007).

It is municipal staff (Saskatchewan) or their appointed Agriculture Service Board (Alberta) that are responsible for the sale of rodenticides or anticoagulants to landowners. Staff must approve the sales and landowners must sign a form at the time of purchase acknowledging that they do not have Swift Fox on their property (Matz pers. comm. 2017; Storch pers. comm. 2017; Merrill pers. comm. 2017; Wilkins pers. comm. 2017a). Merrill (pers. comm. 2017) and Storch (pers. comm. 2017) mentioned that producers and landowners in the range of Swift Fox are very conscientious of Swift Fox re-establishment and are not interested in compromising these efforts or the ability of landowners in Alberta to obtain 2% LSC to control ground squirrel populations.

Toxicants (or predacides) used to control Coyotes or wolves that are depredating livestock or potentially carrying rabies may be lethal for the Swift Fox if used within the Swift Fox range and if not used according to the label. Sodium cyanide is a restricted toxicant registered to the Government of Alberta for Coyote control (PMRA 2011) and sodium monofluoroacetate (known as Compound 1080) is another restricted toxicant registered to the Governments of Alberta and Saskatchewan for the control of wolves and Coyotes (PMRA 2014). The Alberta Ministry of Agriculture and Forestry no longer issues toxicants for Coyote control in Swift Fox areas (Merrill pers. comm. 2017). Similarly, in Saskatchewan, the only registered toxicant, Compound 1080, is no longer used within the primary Swift Fox range (defined as Townships 1-7 and Ranges 1-30 west of the 3^rd meridian) and is used only after three days of scent-post tests have revealed no sign of Swift Fox in a restricted buffer area south of the South Saskatchewan River and west from Tugaske, Moose Jaw, Milestone, and Minton (Saskatchewan Fur Program 2012).

In Alberta, there have been no Swift Fox carcasses handed over to the provincial Fish and Wildlife Laboratory (Pybus pers. comm. 2017) nor reports of Swift Fox poisoning (Nicholson pers. comm. 2017) since the emergency registration of 2% LSC was approved in 2008. Similarly, in Saskatchewan, there has been no incidence of Swift Fox poisoning reported to Saskatchewan Agriculture (Wilkins pers. comm. 2017a) nor to the Saskatchewan Fish and Wildlife authorities since the emergency registration in 2007 (Prieto pers. comm. 2017).

Restrictions for use, labeling, laws, regulations, and policies that have been put in place by government ministries and agencies in Canada, Alberta, and Saskatchewan to minimize the risk of primary or secondary poisoning of Swift Fox may in part be effective. However, it is often difficult to find carnivore carcasses in this landscape and establish a cause of death (Proulx 2011). The potential for direct or secondary poisoning still exists if regulatory measures are not observed, instructions on toxicants are not followed, or if Swift Fox venture onto farmland outside their core area (Proulx 2011). In addition, limited to no follow-up is done on application sites, especially in cropland bordering Swift Fox areas or at the interface between cropland and native grass. As part of Health Canada’s National Pesticide Compliance Program in Fiscal Year 2014-2015, 107 on-farm inspections of liquid strychnine concentrate users in Saskatchewan and Alberta resulted in five violations related to not following label use directions (Health Canada 2016). Secondary poisoning may affect Swift Fox the same way as Red Fox (73% fewer adult Red Foxes per km of road in area with high (90%) poisoning than in area with low (20%) poisoning; Proulx and MacKenzie 2012). This threat is ongoing as toxicants continue to be used on the prairies for the control of rodents and Coyote populations.

IUCN 8. Invasive and other problematic species and genes (medium threat impact)

Invasive non-native/alien species

Canine diseases are growing threats to species of conservation concern around the world (Marino et al. 2017). Most pathogens are either foreign in origin or variants that have been introduced to North America with European immigration. Domestic Dogs (C. familiaris) are known to pose a significant risk as reservoirs for infectious diseases, especially for wild canids (Aguirre 2009). The most important diseases that may affect the Swift Fox include those caused by viruses such as 1) Rabies (type Rabies lyssavirus; endemic to North America but canine variants brought through European immigration; Velasco-Villa et al. 2017), 2) Canine distemper (Canine Distemper Virus; became distributed worldwide along with domestic dogs as human populations expanded; CABI 2018), 3) Canine parvovirus (CPV; originating from Europe or Eurasia and identified in dogs in the late 1970s; Knobel et al. 2014), and those caused by bacteria, such as the sylvatic plague: a flea-transmitted disease caused by the bacterium Yersinia pestis (Olsen 1981) and introduced to western North America from Asia through non-native rats within the last 125 years (Touchman et al. 2007).

Swift Fox are not known to be reservoirs for rabies. Antibodies to rabies virus were found in a Swift Fox in Wyoming with no evidence of infection (Miller et al. 2000). However, a sudden outbreak of rabies transmitted by skunks was the suspected cause of a dramatic five-fold reduction in the closely related San Joaquin Kit Fox (V. macrotis mutica) over a three-year period in California (White et al. 2000). Skunks are one of the most important rabies vector species in Canada and the US (Brown et al. 2014) and their range overlaps that of Swift Fox (Helgen and Reid 2016). Distemper has been responsible for population declines in Black-footed Ferrets (Mustela nigripes), Catalina Island foxes (Urocyon littoralis catalinae), Gray Wolves, Coyotes, and many other wild carnivores worldwide (Kapil and Yeary 2011). Two Swift Fox have died of canine distemper virus in Wyoming (Olson and Lindzey 2002). In wild canids, CPV has been confirmed only in Coyotes and Gray Wolves with the virus associated with mortality and poor pup survival (Knobel et al. 2014). However, high prevalence of antibodies has been detected in many other canid species, including the Swift Fox (Miller et al. 2000). A limited serological survey of 21 Swift Fox in Canada found 52% to have antibodies to distemper, with high titres indicative of previous natural exposure to the pathogen, and 100% to have antibodies to CPV, with 71% having high titres (Miller et al. 2000; COSEWIC 2009). Exposure to infectious pathogens in Swift Fox is likely common but infection and illness may be rare (Pybus and Williams 2003).

Sylvatic plague entered the native rodent population in 1908 to spread rapidly to its current distribution by 1950. It results in large scale epizootics among its most susceptible rodent host groups: prairie dogs (Cynomys spp.) and ground squirrels (Spermophilus spp., now Ictidomys spp., Poliocitellus spp., Callospermophilus spp., and Urocitellus spp. for genera occurring in Canada; Helgen et al. 2009) (Touchman et al. 2007). Canids are infected by flea bites or by consuming infected rodents (Thomas et al. 1989) and could serve to transmit plague. Sylvatic plague is endemic (Leighton et al. 2001) to areas coincident with Swift Fox range. Antibody prevalence in Swift Fox was 100% in northwestern Texas (n=12; McGee et al. 2006), 51% in southeastern Colorado (Gese et al. 2004), and 6% in New Mexico (n=16; Harrison 2003). In a study in Colorado, 24% of the 61 captured foxes were seropositive for plague antibodies in an area with epizootic plague activity in prairie dog colonies. However, none of the fleas found on the foxes were positive for Y. pestis and Swift Fox were not confirmed as reservoirs for plague (Salkeld et al. 2007). Nevison (2017) came to a similar conclusion for a small, reintroduced population of Swift Fox in South Dakota where 69.6% of Swift Fox were seropositive for plague. In Canada, sylvatic plague was detected in Grasslands National Park in one Black-tailed Prairie Dog in 2010 and in one prairie dog and two Richardson’s Ground-squirrels in 2017 (Antonation et al. 2014; Shury pers. comm. 2017), substantiating its presence in Canadian Swift Fox range. Pybus and Williams (2003) indicated that, with a few exceptions, most predators are resistant to sylvatic plague and Swift Fox are also likely resistant to developing clinical or infectious plague.

Swift Fox exposure to canine diseases has not been well studied in Canada and the United States and the sources and effects of known diseases in Swift Fox populations are not fully understood (Moehrenschlager et al. 2004; Moehrenschlager and Sovada 2016). Regardless, diseases are well-documented in other endangered wild canids (Laurenson et al. 2004), including the closely related San Joaquin Kit Fox (White et al. 2000), and Swift Fox coexist with Domestic Dogs, Coyotes, Red Foxes, and Striped Skunks (Mephitis mephitis) in Canada. Although the two population clusters in Canada suggest some barrier to dispersal (Cullingham and Moehrenschlager 2013; 2019), disease could still threaten all Swift Fox if other canids that move freely through their range act as vectors. The western cluster during the 2005/2006 peak abundance year represented about 79% of the estimated Swift Fox population (Moehrenschlager and Moehrenschlager 2006). The severity of the threat may be slight as no significant disease outbreaks have been documented in Swift Fox to date (Moehrenschlager et al. 2004).

IUCN 2. Agriculture and Aquaculture (low threat impact)

Annual and perennial non-timber crops

Native grassland conversion into farmland (row crops and hay) is thought to have been the main factor contributing to the Swift Fox range constriction through habitat loss and fragmentation (Moehrenschlager and Sovada 2016). Although nearly 70% of native vegetation conversion in the Prairies Ecozone took place prior to the 1990s (ESTR 2014) the rate of conversion has recently decreased as less native prairie land with adequate soil and moisture is available to grow crops or tame forage. However, a substantial increase in prices of commodity crops or the development of crop varieties better adapted to marginal land varieties could stimulate further native prairie conversion to cropland (ECCC 2017). In addition, the threat of conversion of large blocks of native prairie to agriculture and other land uses in Swift Fox range has re-emerged because of changes in government policies.

Changes in government policies have added uncertainty to the conservation of some remaining native grasslands. In 2012, the Government of Canada announced that it would cease operating the Prairie Farm Rehabilitation Administration (PFRA) Community Pastures Program and would turn over management and administration of the reversionary pasture lands to the respective provincial governments over the five-year period of 2014-2018 (Phillips 2015). In Saskatchewan, there are 62 former PFRA Community Pastures totaling 714,918 ha that have been or will be transitioned. Of those, eight are within newly identified Swift Fox critical habitat in Saskatchewan (AAFC 2009; and see ECCC 2017). They include Lone Tree (transitioned in 2013), Masefield (transitioned in 2017), Val Marie, Reno 1 and Reno 2 (transitioned in 2018), Battle Creek, Nashlyn, and Govenlock (McInnis pers. comm. 2017), for a total area of 166,428 ha, of which 85% (140,741 ha) was in native cover as of 2005 (Phillips 2015). Various scenarios have been proposed for ownership (sale of land with native prairie with “no-break no-drain” easements) or management of those pastures, but at the time of this report, leasing (15 years) from the Government of Saskatchewan by organized groups of former pasture patrons was the approach taken for all the pastures that had been transferred to the province (McInnis pers. comm. 2017, 2018). However, other options were also being explored for Battle Creek, Nashlyn, and Govenlock, which were still under control of the Government of Canada up until 2019 (McInnis pers. comm. 2018).

In Alberta, a single 41,000 ha PFRA-administered complex of three community pastures was located within the Canadian Forces Base (CFB) Suffield. It was found within the historical Swift Fox range, but outside of the current range. The pasture’s land control reverted to the Department of National Defence (DND) in 2014 and a memorandum of understanding was developed between DND and the former pasture patrons for continued grazing. In addition, the federally managed Onefour Agricultural Research Substation, which totaled 17,300 ha within the Swift Fox range, was also administered by PFRA. Most of the land (16,098 ha) that was originally deeded to Alberta, was returned under provincial administration and management in 2013. In December 2016, a memorandum of understanding was signed between the Government of Alberta and the University of Alberta to allow the university long-term and stable access to conduct academic agricultural research on the property. Under its new status, the Onefour University Research Ranch continues to be grazed by a local grazing association through a grazing management plan that includes provisions for the management and protection of species at risk habitats and is protected from development through Disposition Reservation (DRS) with a Protective Notation (PNT) (Ehlert pers. comm. 2017). The ranch also includes 3,885 ha designated as “Heritage Rangeland Natural Area” and protected under the Alberta Wilderness Areas, Ecological Reserves, Natural Areas and Heritage Rangelands Act and some areas falling under the federal Emergency Order for the Protection of the Greater Sage Grouse (Centrocercus urophasianus urophasianus), which provide additional protection to the grasslands.

In March 2017, the Government of Saskatchewan officially announced that its Saskatchewan Pastures Program was to be phased out over a 3-year period. The program includes 50 pastures operating on 315,655 ha (780,000 acres) of land, with 1,300 current patrons in Saskatchewan (Government of Saskatchewan 2016). Three of these pastures, Arena, Dixon, and Mankota, are within recently identified Swift Fox Critical Habitat in Saskatchewan (Saskatchewan Ministry of Agriculture 2016; and see ECCC 2017). They cover a total of 57,314 ha, of which 92% (52,923 ha) is native pasture (Parsons pers. comm. 2017). The last grazing season for these three pastures as part of the program will be 2019, after which they will switch over organized pasture-patrons control under a 15-year lease with the Government of Saskatchewan to ensure continued grazing and environmental stewardship of the land. Current and potential non-agricultural uses (e.g., oil and gas activity, sand and gravel reserves, etc.) will be allowed to continue on these lands with setback distances and restrictions on timing where species at risk occur (McInnis pers. comm. 2018).

With regard to the transfer of PFRA community pastures, there are two aspects to this threat: 1) the potential outright loss of habitat from a possible sale of the lands by the Government of Saskatchewan, and 2) the loss and fragmentation of habitat from a change in policies and regulations with change in jurisdictions, leading to a potential increase in industrial activities in those areas. From a habitat loss perspective, this threat is large in scope because 38 of 84 (40.4 %) foxes caught in the Alberta-Saskatchewan border region were on townships that contained PFRA lands in the 2005/2006 peak Swift Fox abundance year (Moehrenschlager and Moehrenschlager 2006). This area includes the three PFRA community pastures (totalling 83,694 ha) that remain to be transferred to Saskatchewan as of the end of 2018. It is unknown what the fate of the remaining PFRA community pastures will be after being transferred, but so far, all former pastures’ patron groups have entered into a 15-year lease with the Government of Saskatchewan and there was no interest by any of these groups to purchase the land (McInnis pers. comm. 2018). With PFRA pastures transitioning from federal to provincial Crown land, the immediate prohibitions associated with designated species at risk under SARA no longer apply and species, residence, and critical habitat protection fall under provincial responsibility. In Saskatchewan, the Wild Species at Risk Regulations of the Wildlife Act and the Activity Restriction Guidelines for Sensitive Species (Government of Saskatchewan 2017a) provide for protection of the Swift Fox and its residence, while the environmental assessment process accounts for species residence, critical habitat, and native grasslands (Government of Saskatchewan 2018). It is unknown how the Saskatchewan acts, policies, and regulations, and the enforcement associated with them compare to those associated with SARA with respect to effectively protecting Swift Fox, its residence, and its critical habitat, but these were the tools in place for the protection of species at risk on all provincial Crown land prior to this transfer.

IUCN 3. Energy production and mining (low threat impact)

Renewable and non-renewable energy production may affect Swift Fox directly through increased mortality related to increased road network and/or increased traffic, or indirectly through habitat loss, fragmentation or degradation, increased anthropogenic disturbance, change in density of predators or competitors and/or decrease in food availability (Cypher et al. 2003; Moehrenschlager and Moehrenschlager 2006; Moehrenschlager et al. 2007a). Mortality associated with roads and increased traffic is dealt with in the Transportation and Service Corridors section below.

Oil and gas production

In Wyoming, the probability of Swift Fox extirpation was positively correlated with the number of years energy development was observed within 31 km² grids (Van Fleet et al. 2015). Conversely, in southwestern Saskatchewan, oil and gas sites with baited scent posts were visited regularly by Swift Fox regardless of the time since initial development (Hockaday 2011). Swift Fox den use and survival were not adversely affected by pipeline construction activities, as long as dens were not destroyed, although there was some evidence of negative effects on reproduction due to disturbance during a pipeline construction (Moehrenschlager 2000). At the Naval Petroleum Reserves in California, there was no evidence that oil fields affected survival, reproduction, dispersal space use, den use, food habits, or food availability of Kit Foxes (V. macrotis) and population trends were similar between the developed and the undeveloped areas but abundance in the developed area was half that in the undeveloped area (Cypher et al. 2003). A reduction in carrying capacity associated with 70% habitat disturbance is believed to have caused the decline. In the Milk River watershed, which encompasses much of the current Swift Fox range, the Milk River Watershed Council Canada (MRWCC) reported a total of 2,856 wells associated with oil and gas activity in Alberta, of which 35% (992) were still active in 2012 (MRWCC 2013). In the Montana portion of the watershed, the number of wells reported for 2012 was 9,586, of which 9,465 were active) (MRWCC 2013). In the South of the Divide (SoD) area, which represents the Saskatchewan portion of the Milk River watershed, ECCC (2017) reported 1,350 oil and gas wells capable of production, and 750 wells actively producing as of 2013. They indicated that oil wells and oil reserves were concentrated in a relatively small area, east of the town of Eastend while natural gas wells and estimated remaining gas reserves were concentrated in the western third of the SoD area. The latter area would have the greatest impact on Swift Fox habitat. While oil and gas activity has been relatively important in the Swift Fox range between 1997 and 2008, with peak numbers of new wells in 2005 for Saskatchewan and in 2007 for Alberta (Moehrenschlager and Moehrenschlager 2006), the recession of 2008, which saw the price of oil plummeting (Depersio 2018), has since nearly stalled new well drilling on the Alberta side and greatly reduced it in Saskatchewan.

A large segment of the Swift Fox population (especially the western cluster on the Saskatchewan side and adjacent Montana where much oil and gas activity is taking place) could be affected by energy development. There are still large reserves of oil and especially natural gas that remain undeveloped in the South of the Divide region of Saskatchewan (ECCC 2017). While traffic associated with producing oil and gas wells is still present, pressure from new oil and gas infrastructure development is currently much reduced in the Swift Fox range with low oil and gas prices and is likely to shift only with increasing prices.

IUCN 4. Transportation and service corridors (low threat impact)

Roads and railroads

Roads may cause direct Swift Fox mortality through collisions with vehicles, cause habitat loss and degradation, fragment the landscape, lead to avoidance (loss of functional habitat), and may impede gene flow (Moehrenschlager et al. 2007a). Increasing energy development activities on the Canadian prairies is associated with increasing road development and traffic, increasing the risk of collisions, especially with juveniles (Allardyce and Sovada 2003; Pruss et al. 2008). However, roadways may also be attractive to Swift Fox through avoidance of core area of Coyote ranges, by providing additional food sources from carrion and increased abundance of small mammals in taller disturbed roadside grass, and as travel corridors (Pruss 1999; Harrison and Whitaker-Hoagland 2003). It is unknown whether roads act as ecological traps for Swift Fox; however, they can be an important source of mortality in some areas. In a fragmented landscape of northwestern Texas, vehicle collisions were the primary (42%) cause of Swift Fox mortality due to the presence of a two-lane highway (Kamler et al. 2003b). In a predominantly ranching area of Canada, Moehrenschlager et al. (2007b) found that 7.7% of 39 Swift Fox with known cause of mortality between 1995 and 1998 were road kills. Through logistic regression modeling using three years of winter survey data from 1996/1997 to 2005/2006, Moehrenschlager et al. (2007a) showed that habitat fragmentation from linear landscape disturbance such as roads had a negative effect on Swift Fox presence that extended up to at least 5 km and suggested a loss of functional habitat. In addition, Red Foxes are known to benefit from the presence of roads and associated infrastructure on the landscape, which provide them with food sources and corridors into remote areas where they could compete with other smaller foxes (Cypher et al. 2003; Selås et al. 2010).

Overall, Swift Fox range is in an area of low road density in southwestern Saskatchewan and southeastern Alberta where the development of new roads is largely associated with energy development activity, which is currently low. The majority of roads tend to have low traffic volume but there are ongoing talks about extending the border crossing hours at Wild Horse, AB to 24, which would lead to an increased volume of traffic on Highway #41 during the hours of high Swift Fox activity (Nicholson, pers. comm. 2019).

IUCN 11. Climate change and severe weather (low threat impact)

Droughts

Based on recent climate change predictions, droughts are likely to increase in prevalence and severity in the Canadian Prairie Ecozone (Thorpe 2011). Swift Fox may not be directly impacted by drought as they can meet their water requirements from food alone (Flaherty and Plaake 1986). However, severe, or prolonged droughts may impact their body condition and survival (Herrero et al. 1991) and, as with San Joaquin Kit Fox, decrease their reproductive success because of prey scarcity (White and Ralls 1993; Moehrenschlager et al. 2007a). Interspecific competition can exacerbate the problem. It is unknown to what extent reduced food availability may impact the Swift Fox population but in the San Joaquin Kit Fox, the numerical response was dramatic and rapid, leading to a >50% decrease in Kit Fox captures over two consecutive years, with lower reproduction believed to be the main driver (Cypher et al. 2003). Multiple drought years may occur over a 10-year period in the Canadian Swift Fox range, potentially affecting the entire population. A recent study on the Black-tailed Prairie Dogs near Val Marie, Saskatchewan recognized three drought years during an eight-year span between 2007 and 2014 (Stephens et al. 2018).

Temperature extremes

Swift Fox are behaviourly and morphologically adapted to cope with seasonal environmental changes within their range (Carbyn 1998; Geffen and Girard 2003). However, at the northern extent of their range, increased energetic demand coupled with decreased prey availability during severe winters may lead to mortality from starvation and/or increased vulnerability to killing by predators or intraguild competitors (Moehrenschlager et al. 2007b; Moehrenschlager and Moehrenschlager 2018). Moehrenschlager and Moehrenschlager (2018) speculated that the severe winter of 2010-2011 might have been a possible cause for a 45% decline in population between the 2005/2006 and 2014/2015 survey years.

Limiting factors

Predation

Coyote on the prairies have expanded in distribution and abundance with the extripation of the Gray Wolf (Canis lupus) and the collapse of the prairie ecosystem as it existed prior to European settlement (Herrero et al. 1991). Predation by Coyote is the principal natural cause of Swift Fox mortality across the range (Allardyce and Sovada 2003) and this has hindered Swift Fox reintroduction efforts in Canada (Carbyn et al. 1994). Of the 89 Alberta and Saskatchewan Swift Fox carcasses examined by Carbyn et al. (1994) between 1983 and 1992, 38% were thought to be killed by Coyotes, 8% were likely killed by avian predators (of which 71% were Golden Eagles), and 7% were known or suspected American Badger kills. Moehrenschlager et al. (2007b) found that 31% of radio-collared Swift Fox were killed by Coyotes and 33% by Golden Eagles (n = 39, value for one of the three years of the study), while all closely related Kit Foxes in Mexico survived the duration of the study. Swift Fox home ranges were about three times as large as the Kit Fox’s and had significantly (Kruskal–Wallis Χ² = 35.8, d.f. = 1, P < 0.001) fewer escape holes within them, which made Swift Fox more susceptible to density-dependent encounters with sympatric Coyotes and depredation by Coyotes and Golden Eagles. In Texas, increases in Swift Fox survival rates and recruitment occurred following Coyote control, although the long term effect was unknown (Kamler et al. 2003a). In southern Colorado, Coyote population reduction temporarily increased survival rates of both juvenile and adult Swift Fox, although effects on adults were highly dependent on the timing of Coyote control (Karki et al. 2007). Robinson (1961) and Linhart and Robinson (1972) found no increase in Swift Fox following Coyote control in New Mexico, Colorado, and Wyoming and Swift Fox survival was not enhanced in Canada after Coyote harvesting, owing to increases in kills by Golden Eagle and Red Fox expansion into core Swift Fox habitat (Moehrenschlager 2000). Coyote populations were decimated in southwest Saskatchewan in 2010 due to a bounty control program (Proulx and MacKenzie 2012) but it is unknown what impact this had on the Swift Fox population.

There are no reports of direct Swift Fox predation by the larger Red Fox, which lives normally in more peripheral farmland and other fragmented anthropogenic areas (Larivière and Pasitschniak-Arts 1996). However, Red Fox have been observed depredating the closely related Arctic Fox (Vulpes lagopus; Elmhagen et al. 2017) and the San Joaquin Kit Fox (Ralls and White 1995). San Joaquin Kit Fox carcasses were partially eaten and completely buried (Ralls and White 1995); a behaviour typical to Red Fox that would result in undetected Swift Fox carcasses where the two coexisted without Coyote.

Interspecific competition

Interspecific competition, particularly with other canids may limit Swift Fox. Competitive interaction strength is likely related to prey abundance (Creel et al. 2001). Given that Coyotes seldom consume Swift Fox after killing them is evidence for interference competition (Moehrenschlager et al. 2007b). Kitchen et al. (1999) suggested that den use and some dietary partitioning allowed Coyote and Swift Fox to cohabit. However, Moehrenschlager et al. (2007b) found that Swift Fox in Canada and Kit Fox in Mexico did not and likely could not partition their habitat relative to Coyotes and that killing of Swift or Kit Foxes was likely from density-dependent chance encounters with Coyotes. However, Mitchell (2018) found that Swift Fox in the Dakotas avoided areas of high probability of occupancy by sympatric Coyote and Red Fox. Similarly, Red Fox avoided areas of high probability of occupancy by Coyote. Red Fox have recently expanded their range on the Canadian prairies, stimulated at least in part by habitat fragmentation, human presence and intervention, and access to anthropogenic food sources and shelters (Kamler and Ballard 2002; Mueller et al. 2018). This relatively novel competitor with Swift Fox may be even more significant than Coyote because sympatric Red Fox can exclude Swift Fox from their range through a combination of interference and exploitative competition. In disturbed agricultural landscapes, Coyotes typically select areas of natural cover and avoid denning in proximity to high human activity (farmsteads and residential areas), whereas Red Fox avoid habitats used by sympatric Coyote and use peripheral human-modified habitats for denning and as refugia where anthropogenic food sources and shelters exist (Gosselink et al. 2003; Tannerfeldt et al. 2003; Mueller et al. 2018). Red Fox are known to take over Swift Fox dens and could use these refugia to invade Swift Fox areas in the absence of Coyote (Moehrenschlager et al. 2004). While there is considerable dietary overlap between Swift Fox and Coyote that coexist in Canadian native prairies, the overlap is even greater between sympatric foxes creating greater potential for exploitative competition (Moehrenschlager et al. 2004). Red Fox tend to occur at higher densities than Coyote, which means that the likelihood of Red Fox encounters in Swift Fox home ranges would be greater than that of Coyote encounters (Moehrenschlager et al. 2004). The interaction between Coyote and Red Fox may affect the strength of competition between Red Fox and Swift Fox (Tannerfeldt et al. 2003). Moehrenschlager et al. (2004), indicated that Swift Fox persistence, in areas of sympatry, would require a sufficiently large population of Coyote to keep Red Fox numbers low, yet a sufficiently small population to minimize Swift Fox mortalities. The dynamic of ecological interactions between these three sympatric canids in a changing human-altered environment is poorly understood and may have important ramifications for the sustainability of a Swift Fox population in Canada (Pruss et al. 2008).

Food availability

Food availability has likely been reduced for the Swift Fox since European settlement of the Canadian prairies and the extirpation of Bison (Bison bison). Previously, Bison carrion, either killed by a larger predator, or dying of natural causes, would likely have periodically supplemented their diet, especially during severe winters (Herrero et al. 1991; Sovada et al. 2009). Coyote abundance would have been kept in check by wolves (Berger and Gese 2007), which probably would have had more tolerance for and less dietary overlap (Kitchen et al. 1999) with Swift Fox (Herrero et al. 1991). In addition, the loss of mixed-grass prairie to crop production and changes in grazing patterns and intensity brought about by the replacement of native grazers with domestic cattle, have significantly transformed the prairie ecosystem and likely contributed to a decrease in prey populations for the Swift Fox (Sovada et al. 2009). This maybe be particularly important in the winter when insects and hibernating Richardson’s Ground Squirrels (Urocitellus richardsonii) are unavailable (Michener 1983). They must consequently rely on other small mammals (Moehrenshchlager et al. 2007b), which may be rare in some years and decline over winter (Klausz 1997). As a result, late winter is likely the most critical period for generalist Swift Fox and lack of prey availability may lead to starvation (Klausz 1997). Moehrenschlager et al. (2007b) observed one case of late winter starvation in their study area and noted that foxes were more food-stressed in Canada than closely related Kit Fox in Mexico, which likely led to home-range sizes three times the size of those in Mexico, with associated increase in vulnerability to predators such as Golden Eagle and Coyote (Moehrenschlager et al. 2007b).

Intraguild competition for prey may also add pressure on Swift Fox during times of low prey abundance. Swift Fox diet overlaps considerably with that of Coyote (Kitchen et al. 1999) and the two may compete for food. However, in Canada, Swift Fox diet overlaps even more with that of Red Fox (Moehrenschlager et al. 2004). As a result, the larger Red Fox may exclude Swift Fox from some areas or prevent them from establishing in unoccupied suitable areas (Allardyce and Sovada 2003; Moehrenschlager et al. 2004).

The predator-prey dynamic is complex for the Swift Fox as it is confounded by interspecific and intraguild competition, which in turn is further linked to additional factors related to habitat loss, degradation, and fragmentation, and weather extremes that may act singly or cumulatively. Moreover, prey availability may not only affect Swift Fox survival, but it may also impact productivity, as Swift Fox litter sizes in Canada are correlated to body weights prior to and during the breeding season (late February to March; Moehrenschlager 2000).

Number of locations

Currently, the most serious plausible threat to the Swift Fox in Canada is the use of toxicants and the potential for secondary poisoning. However, it is unlikely that toxicant poisoning would act as a single event that would rapidly affect the majority of individuals in the Canadian population. Thus, the number of locations was considered as > 10.

Protection, status and ranks

Legal protection and status

Swift Fox is protected nationally under Schedule 1 (Threatened Species) of the Canadian Species at Risk Act - SARA (Government of Canada 2018a). SARA contains immediate prohibitions on federal lands against the killing, harming, harassing, capturing, taking, possessing, collecting, buying, selling, or trading of individuals of endangered, threatened, and extirpated species listed in Schedule 1 of the Act. The Act also contains a prohibition against the damage or destruction of their residences (e.g., nest or den) (Government of Canada 2018b). A recovery strategy has been developed for the species (Pruss et al. 2008).

In Alberta, Swift Fox is listed as Endangered under the Alberta Wildlife Act, which confers protection of individuals through prohibitions on hunting and trapping, as well as protecting den sites. The legislation does not protect habitat apart from the den. This is accomplished through Standards and Conditions that apply to formal disposition applications approved under the Public Lands Act (primarily relating to oil and gas development on Crown land; Government of Alberta 2017c), as well as Directives for wind energy and solar energy development (Government of Alberta 2017a,b). On private lands, guidelines for dates of activity restrictions and setback distances have been developed to “help land users minimize, or avoid, potential adverse effects on selected wildlife and wildlife resources when conducting activities.” The guidelines for Swift Fox suggest that all levels of land use activity should be prohibited within 500 m of a den during breeding and pup-rearing season (16 February to 31 July). Outside this period, setbacks of 50 m, 100 m, and 500 m are recommended for low, medium, and high impact disturbances, respectively (Government of Alberta 2011).

Swift Fox is listed as Endangered under the Wild Species at Risk Regulations of the Saskatchewan Wildlife Act. This designation confers enhanced protection of individuals and their dens. Provincial guidelines for sensitive species provide additional habitat protection around den sites. During the breeding and pup-rearing period (15 February to 31 August), low-level (e.g., foot traffic, small vehicles) and medium-level (e.g., large vehicles, small pipeline construction, operation of compressor stations) disturbances are prohibited within 500 m, whereas high levels of disturbance activity (e.g., construction, seismic exploration, forest harvest) are not permitted within 2 km (MEFWLB 2017). During the rest of the year, the setback distances from Swift Fox dens are 100 m, 500 m, and 2 km for low, medium, and high levels of disturbance activity, respectively (MEFWLB 2017).

In the United States, Swift Fox was petitioned in 1992 for listing under the Endangered Species Act. Following an initial review, the US Fish and Wildlife Service (USFWS) concluded that listing of the Swift Fox was “warranted but precluded by higher listing priorities” (USFWS 1995). In 2001, because of new information, the USFWS concluded that “originally identified threats were no longer applicable” and the Swift Fox was removed from the candidate list (USFWS 2001). In response to the initial petition, state and federal agencies, tribal and non-governmental organizations, and representatives from Canada formed the Swift Fox Conservation Team in 1994 and developed the Conservation Assessment and Conservation Strategy of Swift Fox (Vulpes velox) in the United States to “assemble existing information, collect new biological data, implement swift fox monitoring and management programs, and advance Swift Fox conservation and restoration and avoid future listing under the Endangered Species Act” (Dowd Stukel 2017).

The Swift Fox is not currently listed under CITES (CITES 2017).

Non-legal status and ranks

Swift Fox was assessed as Extirpated by COSEWIC in 1978. They were re-assessed as Endangered in May 2000 after initial reintroduction success. The status was re-examined and assessed as Threatened in November 2009 (COSEWIC 2009).

The IUCN Red List of Threatened Species ranking for Swift Fox is Least Concern (Moehrenschlager and Sovada 2016). Under the General Status of Wild Species 2015, Swift Fox was listed as N2 (Imperilled) in Canada, S1S2 (Critically Imperilled to Imperilled) in Alberta, S3 (Vulnerable) in Saskatchewan and SX (Presumed Extirpated) in Manitoba (CESCC 2016). NatureServe (2021) ranks Swift Fox globally as G3 (Vulnerable; last reviewed April 2016) and nationally (Canada) as N3 (Vulnerable; last reviewed January 2018). The Alberta Conservation Information Management System (ACIMS 2017) ranks Swift Fox as S1S2 (Critically Imperilled to Imperilled; updated July 2017), the Saskatchewan Conservation Data Centre (SKCDC 2018) ranks it as S3 (Vulnerable; updated February 2018), and the Manitoba Conservation Data Centre ranks it as SX (Presumed Extirpated; MBCDC 2016).

Habitat protection and ownership

Section 49 (1)(a) of SARA requires that Action Plans include an identification of the species’ critical habitat (to the extent possible) unless such critical habitat was fully identified in a recovery strategy (Pruss et al. 2008). New critical habitat was partially identified as part of action plans for multiple species at risk in the South of the Divide region of southwest Saskatchewan and in Grasslands National Park (ECCC 2017; Parks Canada Agency 2016). Critical habitat was identified using modifications and refinements of the habitat suitability model developed by Moehrenschlager et al. (2007a) as well as other scientific information about habitat requirements of the species, and field records from provinces, universities, non-profit organizations, and federal departments (Parks Canada Agency 2016). Critical habitat was identified as “those areas within the current range of the species in which the combination of habitat attributes is at least as favourable for Swift Fox as the majority of locations where Swift Fox occurrences were documented” (Parks Canada Agency 2016). The biophysical attributes that went into the development of the refined model were assessed within a 3-km radius of Swift Fox captures from the 2005/2006 winter mark-recapture survey and included:

• large tracts of intact (i.e., native) prairie
• short (< 25 cm high), sparse, and relatively homogeneous vegetation
• level or low variation in terrain roughness (gently sloping terrain or few. topographic features such as canyons, steep hills, or coulees)
• dry, well-drained soils
• high density of burrows created by fossorial mammals
• limited cropland
• limited invasive species
• adequate availability of prey items (small mammals and insects)

Critical habitat identified on federal lands in these action plans was protected from destruction under section 58(1) of SARA or through Orders made under subsections 58(4) and 58(5) (Parks Canada Agency 2016). Notices were published in the Canada Gazette on September 24^th, 2016, for Grasslands National Park (Queen’s Printer for Canada 2016), and on February 17^th, 2018, for Prairie National Wildlife Area (unit number 11) (Queen’s Printer for Canada 2018), advising of the coming into force of these Swift Fox critical habitat areas 90 days ensuing the respective notice publication dates.

The portions of critical habitat identified in the above action plans that are on non-federal lands will be assessed by Environment and Climate Change Canada for the protection they currently offer. Provincial laws and legal instruments in Saskatchewan will be reviewed in collaboration with the Government of Saskatchewan to determine if they prevent destruction of critical habitat. If there are gaps, other federal provisions under SARA, legislation or conservation measures will be reviewed to determine whether they prevent destruction of critical habitat (ECCC 2017).

Critical habitat has not been identified yet but will be identified for Alberta (ECCC 2017). Currently, Swift Fox habitat in Alberta benefits from protection conferred by the Greater Sage-grouse Emergency Order (Government of Canada 2013) as well as other regulatory requirements within the Greater Sage Grouse Range on provincial Crown lands (Downey pers. comm. 2017).

About one-sixth of the Canadian Swift Fox population resides within the boundaries of Grasslands National Park (Moehrenschlager and Sovada 2016), making their habitat federally protected.

Acknowledgements and authorities contacted

The writer of this updated report would like to thank the writers of the previous two reports, Jay Gedir and Lu Carbyn, whose earlier work is the foundation of this report. Axel Moehrenschlager provided recent survey data and valuable insight on the status of the Swift Fox population in Canada and adjacent Montana, and access to published and unpublished documents. Lonnie Bilyk, Andrea Benville, and Shelley Pruss provided Swift Fox occurrence data from the Alberta Fish and Wildlife Management System, the Saskatchewan Conservation Data Centre and Parks Canada, respectively. Andre Breault, Andrea Benville, Daniel Buell, Lu Carbyn, Nishanthy Chitravelu, Sue Cotterill, Brandy Downey, Pat Fargey, Chris Friesen, Mike Gollop, Susan Goods, Jeff Keith, Kamal Khidas, Dave Matz, Rachel McDonald, Brent McInnis, Phil Merrill, Amanda J Miller, Gerry Ehlert, Joel Nicholson, Sara Olimb, Rod Parsons, Beatriz Prieto, Gilbert Proulx, Shelley Pruss, Margo Pybus, Doni Schwalm, Todd Shury, Jason Storch, Bill Watkins, Richard Wilkins, and Greg Wilson contributed or facilitated access to valuable information for the report. Graphic images for figures were provided by Axel Moehrenschlager, the Milk River Watershed Council Canada, the Canadian Museum of Nature, Kristy Bly, Eileen Dowd Stukel, and Nichole Bjornlie. Dana Blouin assisted with editing the report and locating some information. Graham Forbes and Stephen Petersen (Co-chairs, COSEWIC Terrestrial Mammals Specialist Subcommittee), Sydney Allen, Angèle Cyr, Karen Timm, and Jenny Wu (Scientific Project Officers, COSEWIC Secretariat) and Neil Jones and Gloria Goulet (ATK Subcommittee, COSEWIC Secretariat) facilitated this status report update. We acknowledge Gordon Court who generously provided his Swift Fox photo. The many contributors of Swift Fox records in the Alberta Fish and Wildlife Management Information System are acknowledged in Appendix 2 as required by Alberta Environment and Parks (Bilyk pers. comm. 2017).

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Biographical summary of report writer(s)

François Blouin is a wildlife biology consultant with over 22 years of experience on species at risk management and conservation. He obtained his B.Sc. in Biology from Bishop’s University in Sherbrooke, Quebec, a graduate diploma in Wildlife Management from Université du Québec à Rimouski, and his M.Sc. in Biology from Université du Québec à Montréal. He has since worked for the Manitoba Conservation Data Centre, Operation Grassland Community (Alberta), the MULTISAR program with Alberta Sustainable Resource Development (now Alberta Environment and Parks), and with the Alberta Prairie Conservation Program. He has completed multiple wildlife field surveys, habitat conservation plans, species assessment reports as well as developed species at risk Beneficial Management Practices for agricultural producers.

Appendix 1. Threat calculator for the Swift Fox

Species or ecosystem scientific name: Swift Fox, Vulpes velox

Element ID: Not applicable

Elcode: Not applicable

Date: 10/06/2019

Assessor(s): Stephen Petersen (Terrestrial Mammals Co-chair), Karen Timm (COSEWIC Secretariat), Rosie Nobre-Soares (COSEWIC Secretariat), Jennifer Heron (facilitator), Francois Blouin, Gord Court, Lu Carbyn.

References: N/A

Assigned overall threat impact: B = High

Impact adjustment reasons: N/A

Overall threat comments: Overall calculated and assigned threat impact of Very High - High from draft calculator decreased to High based on COSEWIC discussion at the April 2021 COSEWIC species assessment meeting. These changes are shown in red font.

Classification of Threats adopted from IUCN-CMP, Salafsky et al. (2008).

Appendix 2. Contributors of Swift Fox records to the Alberta Fish and Wildlife Management System as of June 2017

Angela Chen, Eric Brownrigg, Jennifer Carpenter, Krissy Bush, Leah Darling, Maria Olsen, Melissa Mcintosh, Mike Swystun

CWS - Edmonton

Jon Groves, Steve Swartz

Axel Moehrenschlager

Dale Kastelen

Julie Landry-Deboer, Megan Jensen

Axys Environmental Consulting Ltd.

Danielle Cross

Karsten Heuer

Brandy Downey

Danielle Cross, Len Lupyczuk

Kelly Sturgess

Brent Cowan

Darryl Jarina

Ken Pitcher, Tanner Broadbent

Brian Vanderlinden

Dave Scobie

Kristen Rumbolt-Miller

Brian Wanner

Deanna White, Lee Moltzahn

Kristi Anderson

Bruce Cairns

Dennis Milner

Leah Daoust, Maria Didkowsky

Calgary Zoological Society

Francois Blouin

Lee Finstad

Cameron Aldridge

Geoff Holroyd, Helen Trefry

Mike Swystun

Cameron Jackson

Gerry Ehlert

Orrin Dell

Cameron Lockerbie

Ghostpine Environmental Services Ltd.

Peter Balagus

Canadian Wildlife Service - Edmonton

Helen Trefry

Rob Morrison

Chad Lyttle

Jason Allison

Robert Wapple

Charles Mamo

Jason Lewis

Ron Fode

Christina Devlin

Jennifer Hemsing

Sandy Barrett

Christy Sikina

Joe Harty

Susan Cotterill

Cleve Wershler

Joel Nicholson

Syl Pompu

Cliff Thesen

Joel Nicholson, Kim Morton, Mike Grue

Terry Hood

Colin Starkevich

Joel Nicholson, Mike Grue, Mike Verhage

Troy Wellicome

Corey Scobie

John Taggart

Valerie Toft

Corey Skiftun

Johnathan Cumming, Megan Wright

Wildlife Management