The environmental conditions in the ocean have long been considered relatively more stable through time compared to the conditions on land. Advances in sensing technologies, however, are increasingly revealing substantial fluctuations in abiotic factors over ecologically and evolutionarily relevant timescales in the ocean, leading to a growing recognition of the dynamism of the marine environment as well as new questions about how this dynamism may influence species' vulnerability to global environmental change. In some instances, the diurnal or seasonal variability in major environmental change drivers, such as temperature, pH and seawater carbonate chemistry, and dissolved oxygen, can exceed the changes expected with continued anthropogenic global change. While ocean global change biologists have begun to experimentally test how variability in environmental conditions mediates species' responses to changes in the mean, the extensive literature on species' adaptations to temporal variability in their environment and the implications of this variabilityfor their evolutionary responses has not been well integrated into the field. Here, we review the physiological mechanisms underlying species' responses to changes in temperature, pCO 2 /pH (and other carbonate parameters), and dissolved oxygen, and discuss what is known about behavioral, plastic, and evolutionary strategies for dealing with variable environments. In addition, we discuss how exposure to variability may influence species' responses to changes in the mean conditions and highlight key research needs for ocean global change biology. K E Y W O R D Sadaptive tracking, bioenergetic/behavioral strategies, manipulative experiments, physiological adaptations, variable environments KROEKER Et al.

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SUMMARYWe studied the locomotion and behavior of Dosidicus gigas using pop-up archival transmitting (PAT) tags to record environmental parameters (depth, temperature and light) and an animal-borne video package (AVP) to log these parameters plus acceleration along three axes and record forward-directed video under natural lighting. A basic cycle of locomotor behavior in D. gigas involves an active climb of a few meters followed by a passive (with respect to jetting) downward glide carried out in a finsfirst direction. Temporal summation of such climb-and-glide events underlies a rich assortment of vertical movements that can reach vertical velocities of 3ms , characteristic of the midwater oxygen minimum zone (OMZ), can influence the daytime depth of squid, but this depends on location and season, and squid can ʻdecoupleʼ from this environmental feature. Light is also an important factor in determining daytime depth, and temperature can limit nighttime depth. Vertical velocities were compared over specific depth ranges characterized by large differences in dissolved oxygen. Velocities were generally reduced under OMZ conditions, with faster jetting being most strongly affected. These data are discussed in terms of increased efficiency of climband-glide swimming and the potential for foraging at hypoxic depths.

Dosidicus gigas (Humboldt or jumbo flying squid) is an economically and ecologically influential species, yet little is known about its natural behaviors because of difficulties in studying this active predator in its oceanic environment. By using an animal-borne video package, National Geographic's Crittercam, we were able to observe natural behaviors in free-swimming D. gigas in the Gulf of California with a focus on color-generating (chromogenic) behaviors. We documented two dynamic displays without artificial lighting at depths of up to 70 m. One dynamic pattern, termed 'flashing' is characterized by a global oscillation (2-4 Hz) of body color between white and red. Flashing was almost always observed when other squid were visible in the video frame, and this behavior presumably represents intraspecific signaling. Amplitude and frequency of flashing can be modulated, and the phase relationship with another squid can also be rapidly altered. Another dynamic display termed 'flickering' was observed whenever flashing was not occurring. This behavior is characterized by irregular wave-like activity in neighboring patches of chromatophores, and the resulting patterns mimic reflections of down-welled light in the water column, suggesting that this behavior may provide a dynamic type of camouflage. Rapid and global pauses in flickering, often before a flashing episode, indicate that flickering is under inhibitory neural control. Although flashing and flickering have not been described in other squid, functional similarities are evident with other species.