Competitive Exclusion as a Factor Influencing the Distributions of Two Species of Terrestrial Salamanders

The salamander Plethodon richmondi shenandoah is restricted to areas of talus on Hawksbill Mountain, Shenandoah National Park, Virginia, while P. cinereus inhabits the soil outside the talus. To test the hypothesis that the former species is excluded from the soil by the latter species, three enclosures were constructed in each of four habitats: (i) in talus consisting only of bare rocks, (ii) in talus where soil has infiltrated between but not under rocks, (iii) in isolated pockets of shallow soil within the rocky confines of the talus, and (iv) in deep soil outside the talus. In each habitat one enclosure contained an isolated sample of shenandoah, another an isolated sample of cinereus, and the third a mixture of the two species. The above testes indicated that: (i) neither species survived the extremely dry conditions of the bard rocks for more than a week, (ii) shenandoah expressed a higher survivorship than did cinereus in this relatively dry habitat, (iii) in the shallow soil shenandoah had lower survivorship in the presence of cinereus than it did in isolation, and (iv) in the deep soil shenandoah had poorer survivorship than cinereus in the mixed species enclosure and poorer survivorship than its isolated control. P. cinereus apparently inhibits the presence of shenandoah in areas of soil, while shenandoah survives better under rocky conditions. However, the talus is a suboptimal habitat for shenandoah, since its survivorship in isolated enclosures is significantly better in the presence of soil. It seems unlikely that salamanders other than cinereus could exclude shenandoah from the soil or that other animals exert a strong influence. Predation, diseases, and parasites also do not appear to exclude shenandoah from areas of deep soil. Competitive exclusion by cinereus has probably restricted the distribution of shenandoah to the suboptimal talus refugium.