Evolution of viviparity in squamate reptiles (Sceloporus spp.): a variant of the cold-climate model

An alternative to the cold-climate model for the evolution of viviparity is that the impetus for the initial transition from oviparity to viviparity is not an increase in the duration of egg retention but a shift in the location of nests to more superficial and thus warmer locations in the soil profile as temperature declines with increasing altitude or latitude. Shallower nest placement, however, would lead to increased egg mortality as a result of physiological stress or predation, and enhanced egg mortality would thus provide the initial benefit of extended egg retention. To test this hypothesis, I examined the thermal biology of three species of Sceloporus lizards living at high altitudes: S. virgatus (1800 m), S. aeneus (2800 m), and S. bicanthalis (3200 m). The oviparous S. virgatus and S. aeneus females laid eggs at depths of 6 and 2 cm, and mean body/nest temperatures were 24.6/25.2 degree C and 19.9/20.8 degree C, respectively. Because the initial increment in the duration of egg retention is presumably short, females of these oviparous species cannot initially keep their eggs appreciably warmer than they would be in nests. In contrast, mean temperatures of simulated nests (17.6 degree C) of the viviparous S. bicanthalis were similar to the 17 degree C low temperature threshold for development, and temperatures in some nests fell below freezing at night and rose to lethal levels during the day. Mean body temperatures of S. bicanthalis females were 20.1 degree C; eggs retained during the entire developmental period would hatch 17 days sooner than they would if laid in a nest. Extended egg retention at the highest elevation site would thus provide two benefits: reduced mortality of eggs and accelerated development relative to that in a nest. These benefits, however, accrue because of the initial shift of nests from deeper to more superficial depths in the soil profile.