Thermal biology of sea snakes and sea kraits.

Temperature probably had no direct effect on the evolution of sea kraits within their center of origin, a geologically stable thermal zone straddling the equator, but may have indirectly affected expansions and contractions in distributions beyond that zone through global fluctuations that caused alternation of higher and lower sea levels. The northern limit of the Laticauda colubrina complex seems to be the 20°C isotherm; in the south, the range does not reach that isotherm because there is no land (also a habitat requirement of sea kraits) within the zone of suitable temperature. The relationship of temperature to the pattern of geographic variation in morphology supports either the hypothesis of peripheral convergence or the developmental hypothesis but does not distinguish between them. Quadratic surfaces relating cumulative scores for coloration and morphological characters to global position showed a strong latitudinal component and an even stronger longitudinal one in which the direction of the latitudinal effect was reversed between east and west. A multivariate analysis revealed that while morphological characters vary significantly by location and climate when tested separately, when the influence of location on morphology is taken into account, no residual relationship between climate and morphology remains. Most marine snakes have mean upper temperature tolerances between 39°C and 40°C and operate at temperatures much nearer their upper thermal limits than their lower limits but still avoid deleterious extremes by diving from excessively hot water to deeper, cooler strata, and by surfacing when water is cold. At the surface in still water in sunlight, Pelamis can maintain its body temperature slightly above that of the water, but whether this is significant in nature is questionable. As temperature falls below 18-20°C, survival time is progressively reduced, accompanied by the successive occurrence of cessation of feeding, cessation of swimming, and failure to orient. Acclimation does not seem to be in this species' repertoire. In the water column, marine snakes track water temperature; on land, sea kraits can thermoregulate by basking, selecting favorable locations, and by kleptothermy. Laticauda colubrina adjusts its reproductive cycle geographically in ways that avoid breeding in the coldest months. Mean voluntary diving time is not temperature-dependent within the normal range of temperatures experienced by marine snakes in the field, but is reduced in water colder than 20°C. On land, much as while diving in the sea, sea kraits maintain long periods of apnea; intervals between breaths are inversely related to temperature.