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75<br />

Beschta 2004). Predation rates were high on porcupines <strong>du</strong> ring the two years of study and<br />

predation on a<strong>du</strong>lts was linked to the presence of snow covering the ground (chap. 3). We<br />

observed porcupines re<strong>du</strong>cing the time spent outsi<strong>de</strong> of the <strong>de</strong>n and the <strong>du</strong>ration of activity<br />

bouts when snow pen<strong>et</strong>rability increased. We suggest porcupines lowered expo<strong>sur</strong>e to<br />

predators by re<strong>du</strong>cing the time spent outsi<strong>de</strong> of the <strong>de</strong>n when snow pen<strong>et</strong>rability, and<br />

therefore predation risk, was high.<br />

Besi<strong>de</strong>s re<strong>du</strong>cing the time spent outsi<strong>de</strong> of the <strong>de</strong>n, porcupines also became more diurnal<br />

with <strong>de</strong>creasing Tc. North American porcupines and ro<strong>de</strong>nts from the Hystricidae family are<br />

known to be essentially nocturnal (Roze 1989, Roll <strong>et</strong> al. 2006). However, only 47% of<br />

porcupines' active time was <strong>du</strong>ring the night and porcupines were more diurnal at cold<br />

temperatures. Ro<strong>de</strong>nts living in cold environments are more 1ikely to be diurnal, possibly<br />

because they exploit the walmer hours of the diel cycle (Roll <strong>et</strong> al. 2006). Still, diel activity<br />

patterns are phylogen<strong>et</strong>ically constrained (Roll <strong>et</strong> al. 2006). Hence, being cathemeral (i .e.<br />

being active <strong>du</strong>ring both day and night, Tattersall 1987, Tattersall 2006) may represent the<br />

best compromise nocturnal animais like porcupines can make to avoid expo<strong>sur</strong>e to cold night<br />

temperatures. Porcupines also became more diurnal when snow pen<strong>et</strong>rability increased and<br />

one hypothesis is that being active <strong>du</strong>ring the daytime re<strong>du</strong>ced the likelihood of encountering<br />

predators (Bakker <strong>et</strong> al. 2005).<br />

As we could expect from our thermal map, using a <strong>de</strong>n was at the basis of porcupines'<br />

behavioural thermoregu<strong>la</strong>tory strategy. Porcupines modified the time spent outsi<strong>de</strong> of the <strong>de</strong>n<br />

and the timing of activity according to Tc, and this may allow huge energy savings (this<br />

study, Humphries <strong>et</strong> al. 2005).<br />

Microhabitats used outsi<strong>de</strong> of the <strong>de</strong>n<br />

Outsi<strong>de</strong> their <strong>de</strong>ns, porcupines mainly used coniferous trees. Operative temperature did<br />

not <strong>influence</strong> microhabitat use once porcupines were insi<strong>de</strong> coniferous trees. Rather, the<br />

microhabitat choice varied with the behaviour of the animal: porcupines preferentially<br />

exploited open microhabitats and tops of trees for feeding and covered microhabitats and<br />

bottoms of trees for resting. According to our thennal map, energy expenditure was generally<br />

higher in open and arboreal than in other microhabitats. Feeding sites therefore did not appear<br />

to be chosen based on the protection they offered against the thennal environment. We<br />

generate the testable hypothesis that porcupines fed on tops of trees because they offered the

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