Indirect impacts of a highway on movement behavioral states of a threatened tortoise and implications for landscape connectivity.

Roads have often been identified as barriers to the movement of free-ranging animals. However, whether restoration of landscape connectivity across roadways can mitigate barriers to movement is insufficiently understood in light of indirect effects of roads on wildlife movement. We GPS-tagged free-ranging Mojave desert tortoises (Gopherus agassizii) to quantify movement behavioral states using hidden Markov models in relation to a major highway and to document use of existing, permeable culverts. We then used the observed movement behaviors to parameterize simulations of tortoise movement to evaluate alternative culvert designs and placements for enhancing connectivity across the roadway. Tortoises were most active during mid-day, in warm temperatures, and when close to the highway. The highway affected transition probabilities between movement states, as females were more likely than males to switch to an energy-demanding traveling movement state, remain in that state, and move farther than usual within that state. In contrast, males were more likely than females to continue in the low-energy resting state when close to the highway, but if traveling, to travel farther than usual. We observed two highway crossings by a tagged tortoise, which was a higher rate of crossing than in simulated tortoises. Simulated crossing rates increased with culvert size and culvert density, and size and density appeared more important for crossing than if culverts were placed singly or in pairs. Existing culvert densities across the region appeared potentially sufficient for long-term genetic connectivity, but only if retrofitted to allow for tortoise access and passing. We concluded that existing highway traffic may indirectly depress tortoise populations adjacent to the highway, particularly via negative impacts to female movements, and that existing culverts in washes should be retrofitted to allow for periodic tortoise crossings to improve structural connectivity for occasional passage.