Stable isotopes (δ18O and δ13C) in Spirula spirula shells from three major oceans indicate developmental changes paralleling depth distributions

Stable isotopes (delta O-18 and delta C-13) were measured in successive chambers of the aragonitic shells of the small deep-sea squid Spirula spirula (Linnaeus 1758) (class Cephalopoda, subclass Coleoidea, order Sepioidea, family Spirulidae) to determine whether their depth distributions change with age. The spiral shells, ranging in diameter from 18 to 23 mm (30-38 chambers), were collected between 2000 and 2006 from beaches in six widely separated locations in three oceans, the Atlantic (Tobago and Canary Islands), Indian (Madagascar, Maldives, and Perth, Australia), and Pacific Oceans (Ulladulla, Australia). The patterns for both isotopes were highly correlated in specimens from all six sites. The delta O-18 data suggest that after hatching at depths > 1,000 m at temperatures of 4-6 degrees C, the squid migrate into shallower, warmer waters at 12-14 degrees C at depths of 400-600 m. Subsequently, the increasing delta O-18 values suggest a migration back into somewhat cooler, deeper habitats. The delta C-13 values also revealed three ontogenetic stages in all six specimens, including a major shift from positive to negative values, which probably corresponds to sexual maturation, the initiation of reproduction, and concomitant changes in diet. In three of the six specimens (from Tobago, Canary Islands, and Maldives) a fourth embryonic stage (not detected in the oxygen data) was accompanied by markedly less positive delta C-13 values in the first few chambers. These data, combined with the scanty life history information from previous studies of S. spirula, can be used to compare the habitat requirements of related extant and fossil cephalopod genera.