scholarly journals How apparent competition and predator responses led to the decline of Arctic ground squirrels in the boreal forests of the southwest Yukon

Author(s):  
Jeffery R Werner ◽  
Elizabeth A Gillis ◽  
Rudy Boonstra ◽  
Charles J Krebs

Throughout much of North America’s boreal forest, the cyclical fluctuations of snowshoe hare populations (Lepus americanus) may cause other herbivores to become entrained in similar cycles. Alternating apparent competition and indirect mutualism via prey switching are the mechanisms behind this interaction. Our purpose is to document a change in the role of indirect interactions between sympatric populations of hares and arctic ground squirrels (Urocitellus parryii plesius), and to emphasize the influence of predation for controlling ground squirrel numbers. We used mark-recapture to estimate the population densities of both species over a 25-year period that covered two snowshoe hare cycles. We analysed the strength of association between snowshoe hare and ground squirrel numbers and changes in the seasonal and annual population growth rates of ground squirrels over time. A hyperbolic curve best describes the per capita rate of increase of ground squirrels relative to their population size, with a single stable equilibrium and a lower critical threshold below which populations drift to extinction. The crossing of this unstable boundary resulted in the subsequent uncoupling of ground squirrel and hare populations following the decline phase of their cycles in 1998. The implications are that this sustained Type II predator response led to the local extinction of ground squirrels. Arctic ground squirrels may also have exhibited an Allee effect caused by the disruption of social signalling of approaching predators when few individuals are left in a colony.

PeerJ ◽  
2016 ◽  
Vol 4 ◽  
pp. e2303 ◽  
Author(s):  
Jeffery R. Werner ◽  
Elizabeth A. Gillis ◽  
Rudy Boonstra ◽  
Charles J. Krebs

Throughout much of North America’s boreal forest, the cyclical fluctuations of snowshoe hare populations (Lepus americanus) may cause other herbivores to become entrained in similar cycles. Alternating apparent competition via prey switching followed by positive indirect effects are the mechanisms behind this interaction. Our purpose is to document a change in the role of indirect interactions between sympatric populations of hares and arctic ground squirrels (Urocitellus parryii plesius), and to emphasize the influence of predation for controlling ground squirrel numbers. We used mark-recapture to estimate the population densities of both species over a 25-year period that covered two snowshoe hare cycles. We analysed the strength of association between snowshoe hare and ground squirrel numbers, and the changes to the seasonal and annual population growth rates of ground squirrels over time. A hyperbolic curve best describes the per capita rate of increase of ground squirrels relative to their population size, with a single stable equilibrium and a lower critical threshold below which populations drift to extinction. The crossing of this unstable boundary resulted in the subsequent uncoupling of ground squirrel and hare populations following the decline phase of their cycles in 1998. The implications are that this sustained Type II predator response led to the local extinction of ground squirrels. When few individuals are left in a colony, arctic ground squirrels may also have exhibited an Allee effect caused by the disruption of social signalling of approaching predators.


2016 ◽  
Author(s):  
Jeffery R Werner ◽  
Elizabeth A Gillis ◽  
Rudy Boonstra ◽  
Charles J Krebs

Throughout much of North America’s boreal forest, the cyclical fluctuations of snowshoe hare populations (Lepus americanus) may cause other herbivores to become entrained in similar cycles. Alternating apparent competition and indirect mutualism via prey switching are the mechanisms behind this interaction. Our purpose is to document a change in the role of indirect interactions between sympatric populations of hares and arctic ground squirrels (Urocitellus parryii plesius), and to emphasize the influence of predation for controlling ground squirrel numbers. We used mark-recapture to estimate the population densities of both species over a 25-year period that covered two snowshoe hare cycles. We analysed the strength of association between snowshoe hare and ground squirrel numbers and changes in the seasonal and annual population growth rates of ground squirrels over time. A hyperbolic curve best describes the per capita rate of increase of ground squirrels relative to their population size, with a single stable equilibrium and a lower critical threshold below which populations drift to extinction. The crossing of this unstable boundary resulted in the subsequent uncoupling of ground squirrel and hare populations following the decline phase of their cycles in 1998. The implications are that this sustained Type II predator response led to the local extinction of ground squirrels. Arctic ground squirrels may also have exhibited an Allee effect caused by the disruption of social signalling of approaching predators when few individuals are left in a colony.


2000 ◽  
Vol 78 (8) ◽  
pp. 1309-1319 ◽  
Author(s):  
Andrea E Byrom ◽  
Tim J Karels ◽  
Charles J Krebs ◽  
Rudy Boonstra

We examined whether arctic ground squirrel (Spermophilus parryii plesius) populations in northern boreal forest in the Yukon Territory, Canada, were limited by food, predators, or a combination of both, during the decline and low phases of a snowshoe hare cycle. From 1990 to 1995, populations were monitored in large-scale (1 km2) experimental manipulations. Squirrels were studied on eight 9-ha grids: four unmanipulated control grids, two food-supplemented grids, a predator-exclosure grid, and a predator-exclosure + food-supplemented grid. Population density was measured on all grids by livetrapping and active-season survival was measured using radiotelemetry. Population densities were lowest in 1992 and 1993 (2 years after the snowshoe hare population decline). Rates of population change were negative from 1991 to 1993, when predation pressure was most intense after the snowshoe hare decline, and positive from 1993 to 1995, when hares and predators were at low densities. Predation accounted for 125 of 130 mortalities (96%) of radio-collared squirrels. Adult survival was significantly lower in 1992 and 1993 than in 1994 and 1995, and was a strong predictor of annual rates of population change in arctic ground squirrels. Treatments were ranked as follows in their effect on adult survival: predator exclosure + food-supplemented > food-supplemented > predator exclosure > controls. Juvenile survival was lowest in 1992, and food addition and predator removal separately increased juvenile survival. On average, predator exclusion increased population densities twofold, food supplementation increased densities fourfold, and food supplementation and predator removal together increased densities 10-fold. We conclude that food and predation interact to limit arctic ground squirrel populations in the boreal forest during the decline and low phases of the snowshoe hare cycle. The snowshoe hare cycle may indirectly create a lagged secondary fluctuation in arctic ground squirrel populations through shared cyclic predators.


1984 ◽  
Vol 247 (4) ◽  
pp. R722-R727 ◽  
Author(s):  
D. R. Caprette ◽  
J. B. Senturia

The effects of low temperature on mechanical performance of the isolated left ventricles of the 13-lined ground squirrel (a hibernator) and the rat (a nonhibernator) were studied. In addition, low-temperature performance of hearts from summer-active, winter-hibernating, and winter-active ground squirrels were compared. By measuring pressure (P) generated against a balloon inserted into the left ventricle, maximum developed pressure (DP) and maximum rate of increase of P (peak dP/dt) were determined over a temperature range of 5–20 degrees C. The DP and dP/dt of the rat ventricle exhibited significantly greater reduction in magnitude at reduced temperature, compared with those of ground squirrel ventricle. Rat, but not ground squirrel, hearts exhibited arrhythmias of various kinds, including extra-systoles, tachycardia, pulsus alternans, and periods of asystole. Hearts from winter-active ground squirrels developed greater pressures than those from winter-hibernating and summer-active animals. This evidence suggests that disruption of cell communication in the nonhibernator ventricular myocardium plays an important role in the failure of the nonhibernator heart at low body temperatures. Contractility of the seasonal hibernator's heart is influenced by both season and hibernation itself, possibly through shifts in myocardial metabolism. However, seasonal adaptations appear not to be required to confer the special resistance of the seasonal hibernator's heart to the deleterious effects of low temperature.


1998 ◽  
Vol 76 (3) ◽  
pp. 592-596 ◽  
Author(s):  
Anne H Hubbs ◽  
Rudy Boonstra

We used radiotelemetry to study the effects of food addition and predator reduction on the home-range sizes of adult Arctic ground squirrels (Spermophilus parryii) on large-scale experimental grids in the boreal forest of the southwestern Yukon Territory. Home ranges were 2-7 times smaller on food-supplemented grids than on nonsupplemented grids, regardless of whether large mammalian predators were present. Similarly, core areas (where 50% of activities occur) were 8-11 times smaller on food-supplemented grids. Food availability rather than predator presence primarily determined the sizes of home ranges and core areas of Arctic ground squirrels.


2015 ◽  
Vol 42 (2) ◽  
pp. 176 ◽  
Author(s):  
Jeffery R. Werner ◽  
Charles J. Krebs ◽  
Scott A. Donker ◽  
Rudy Boonstra ◽  
Michael J. Sheriff

Context The arctic ground squirrel (Urocitellus parryii) comprised 17% of the biomass of herbivores in the Yukon boreal forest during the summer months from 1987 to 1996 and was responsible for 23% of the energy flow at the herbivore level. By 2000, ground squirrel populations in this region collapsed to nearly zero and have remained there. Aims We summarise the population monitoring (since 1975) and recent experimental work that has been done on this key herbivore in the Kluane area of the southern Yukon to test one mechanistic hypothesis as the possible explanation for this population collapse and subsequent lack of recovery: predation. Methods Ground squirrels are the preferred summer prey of bird and mammal predators when snowshoe hare (Lepus americanus) populations are declining. We used translocations into formerly occupied habitat and radiotelemetry to determine movements and causes of death from 2009 to 2014. We surveyed 158 sites between 2008 and 2013 to measure the disappearance of colonies in alpine and forest habitats over 25 000 km2. Key results Ground squirrels from 2000 to 2013 comprised a small fraction of the herbivore biomass in the boreal forest zone, down from 17% earlier. Most forest populations (~95%) are currently extinct, whereas just over half (65%) of low-elevation meadow populations are locally extinct. One hypothesis is that ground squirrels in the forest have been driven into a predator pit from which they cannot recover. They remain abundant in alpine tundra (93% occupancy rate) and around airport runways and human habitations (97% occupancy), but there is no apparent dispersal from alpine areas down into the boreal forest. Conclusion The predator pit hypothesis is a likely explanation for the initial collapse and sustained decline in population size from 2000 to 2013. Recent attenuation of the hare cycle and milder winter climate have allowed shrubs to expand throughout the forest, thereby reducing visibility and increasing predation risk. This conclusion will be tested in further research using reintroductions to formerly occupied sites. Implication If the loss of this herbivore from the boreal forest is not reversed, predator pressure on the other major herbivores of the montane forest zone is likely to change significantly.


1975 ◽  
Vol 228 (1) ◽  
pp. 325-330 ◽  
Author(s):  
W Galster ◽  
PR Morrison

The hibernation season in the arctic ground squirrel (Citellus undulatus) is broken into 8- to 18- day cycles by short homeothermal periods during which the carboydrate reserves depleted during hibernation are replenished. This study follows a number of metabolities in tissues and body fluids to assess the sources for reconstitution of the glucose reserves: lactate, urea, ammonia, free fatty acid, glycerol, triglyceride, and glucose in plasma; glycogen in liver and muscle; and urea and ammonia nitrogen in urine. Fat is the major energy source during both homeothermal and heterothermal periods, the contribution from glucose being limited to glycolysis. Reconstitution of glycogen is accomplished prior to reentry through maximal use of substrates from all sources including glycolysis, fat, and protein metabolism. Of the new gluconeogenic substrate, one-fourth is supplied from protein and three-fourths from fat.


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