Nocturnal and crepuscular behavior in elasmobranchs: a review of movement, habitat use, foraging, and reproduction in the dark

It is commonly assumed that elasmobranchs (sharks, skates, and rays) are most active during dark periods (dawn, dusk, night). However, this assertion has not been critically evaluated. It is also unclear whether dark periods are primarily utilized for the performance of important life-history events, such as mating. If this were the case, low-light periods would be of significance to elasmobranch conservation as some anthropogenic activities (night fishing, lighting) could disproportionately impact fitness of species that are more active in the dark. Here, we review and summarize previous studies on elasmobranch behavior during nocturnal and crepuscular periods focusing on patterns of movement, habitat use, foraging, and reproduction. A review of 166 studies provided mixed results for widely-assumed increased elasmobranch activity when dark. Frequency of foraging and horizontal movement (distance travelled, activity space) were reported as greater only during crepuscular periods in the majority (>50%) of reviewed studies (28 of 43 and 78 of 125 studies, respectively), a pattern not evident during night. No pervasive patterns emerged for increased habitat use or reproductive behaviors during dark. We did not find any particular habitat type consistently supporting increased activity during dark, nor did we find evidence that higher trophic level elasmobranchs were more active when dark. Thus, generalizations about increased elasmobranch activity during dark periods are currently not supported. While research on the behavior of elasmobranchs during dark periods has been increasing, many knowledge gaps remain and we present a set of research priorities to assist in the development of future investigations.