Embryonic staging series for the beach spawning, terrestrially incubating California grunion Leuresthes tenuiswith comparisons to other Atherinomorpha

California grunion Leuresthes tenuis synchronize spawning with tidal cycles, so the embryos incubate in a terrestrial environment, delay hatching until cued by a specific environmental trigger, and may extend incubation for up to an additional four weeks. These adaptations, however, do not appear to alter the morphology or sequence of early developmental stages as compared to other Atherinomorph fishes in the Orders Beloniformes and Cyprinodontiformes. Embryonic development is described in a series of 30 stages based on morphology observed by light microscopy. Stages are placed in five periods: zygote and cleavage, blastula, gastrula, segmentation and organogenesis, and hatching competence. Embryos from a southern population of L. tenuis in Los Angeles are compared with embryos found >560 km north in San Francisco Bay. Northern L. tenuis embryos developed more slowly at several stages than southern embryos and reached hatching competence later, but both locations maintained synchrony with the tidal cycle for both spawning and hatching. The variation in rates of development and stage at hatching readiness are forms of developmental heterochrony that may be associated with evolutionary adaptation or morphological plasticity within this highly successful clade.