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Visual signals rapidly relay information, facilitating behaviors and ecological interactions that shape ecosystems. However, most known signaling systems can be restricted by low light levels-a pervasive condition in the deep ocean, the largest inhabitable space on the planet. Resident visually cued animals have therefore been hypothesized to have simple signals with limited information-carrying capacity. We used cameras mounted on remotely operated vehicles to study the behavior of the Humboldt squid, Dosidicus gigas , in its natural deep-sea habitat. We show that specific pigmentation patterns from its diverse repertoire are selectively displayed during foraging and in social scenarios, and we investigate how these behaviors may be used syntactically for communication. We additionally identify the probable mechanism by which D. gigas , and related squids, illuminate these patterns to create visual signals that can be readily perceived in the deep, dark ocean. Numerous small subcutaneous (s.c.) photophores (bioluminescent organs) embedded throughout the muscle tissue make the entire body glow, thereby backlighting the pigmentation patterns. Equipped with a mechanism by which complex information can be rapidly relayed through a visual pathway under low-light conditions, our data suggest that the visual signals displayed by D. gigas could share design features with advanced forms of animal communication. Visual signaling by deep-living cephalopods will likely be critical in understanding how, and how much, information can be shared in one of the planet's most challenging environments for visual communication., Competing Interests: The authors declare no competing interest.

The light environment of the mesopelagic realm of the ocean changes with both depth and viewer orientation, and this has probably driven the high diversity of visual adaptations found among its inhabitants. The mesopelagic 'cockeyed' squids of family Histioteuthidae have unusual eyes, as the left and right eyes are dimorphic in size, shape and sometimes lens pigmentation. This dimorphism may be an adaptation to the two different sources of light in the mesopelagic realm, with the large eye oriented upward to view objects silhouetted against the dim, downwelling sunlight and the small eye oriented slightly downward to view bioluminescent point sources. We used in situ video footage from remotely operated vehicles in the Monterey Submarine Canyon to observe the orientation behaviour of 152 Histioteuthis heteropsis and nine Stigmatoteuthis dofleini We found evidence for upward orientation in the large eye and slightly downward orientation in the small eye, which was facilitated by a tail-up oblique body orientation. We also found that 65% of adult H. heteropsis ( n = 69) had yellow pigmentation in the lens of the larger left eye, which may be used to break the counterillumination camouflage of their prey. Finally, we used visual modelling to show that the visual returns provided by increasing eye size are much higher for an upward-oriented eye than for a downward-oriented eye, which may explain the development of this unique visual strategy.This article is part of the themed issue 'Vision in dim light'., (© 2017 The Author(s).)

Climate-driven range shifts are ongoing in pelagic marine environments, and ecosystems must respond to combined effects of altered species distributions and environmental drivers. Hypoxic oxygen minimum zones (OMZs) in midwater environments are shoaling globally; this can affect distributions of species both geographically and vertically along with predator-prey dynamics. Humboldt (jumbo) squid (Dosidicus gigas) are highly migratory predators adapted to hypoxic conditions that may be deleterious to their competitors and predators. Consequently, OMZ shoaling may preferentially facilitate foraging opportunities for Humboldt squid. With two separate modeling approaches using unique, long-term data based on in situ observations of predator, prey, and environmental variables, our analyses suggest that Humboldt squid are indirectly affected by OMZ shoaling through effects on a primary food source, myctophid fishes. Our results suggest that this indirect linkage between hypoxia and foraging is an important driver of the ongoing range expansion of Humboldt squid in the northeastern Pacific Ocean., (© 2014 John Wiley & Sons Ltd.)

The deep-sea squid Grimalditeuthis bonplandi has tentacles unique among known squids. The elastic stalk is extremely thin and fragile, whereas the clubs bear no suckers, hooks or photophores. It is unknown whether and how these tentacles are used in prey capture and handling. We present, to our knowledge, the first in situ observations of this species obtained by remotely operated vehicles (ROVs) in the Atlantic and North Pacific. Unexpectedly, G. bonplandi is unable to rapidly extend and retract the tentacle stalk as do other squids, but instead manoeuvres the tentacles by undulation and flapping of the clubs' trabecular protective membranes. These tentacle club movements superficially resemble the movements of small marine organisms and suggest the possibility that G. bonplandi uses aggressive mimicry by the tentacle clubs to lure prey, which we find to consist of crustaceans and cephalopods. In the darkness of the meso- and bathypelagic zones the flapping and undulatory movements of the tentacle may: (i) stimulate bioluminescence in the surrounding water, (ii) create low-frequency vibrations and/or (iii) produce a hydrodynamic wake. Potential prey of G. bonplandi may be attracted to one or more of these as signals. This singular use of the tentacle adds to the diverse foraging and feeding strategies known in deep-sea cephalopods.

Little is known about the reproductive habits of deep-living squids. Using remotely operated vehicles in the deep waters of the Monterey Submarine Canyon, we have found evidence of mating, i.e. implanted sperm packages, on similar body locations in males and females of the rarely seen mesopelagic squid Octopoteuthis deletron. Equivalent numbers of both sexes were found to have mated, indicating that male squid routinely and indiscriminately mate with both males and females. Most squid species are short-lived, semelparous (i.e. with a single, brief reproductive period) and promiscuous. In the deep, dark habitat where O. deletron lives, potential mates are few and far between. We suggest that same-sex mating behaviour by O. deletron is part of a reproductive strategy that maximizes success by inducing males to indiscriminately and swiftly inseminate every conspecific that they encounter.

Visual behaviors are prominent components of intra- and interspecific communication in shallow-water cephalopods. Meso- and bathypelagic cephalopods were believed to have limited visual communication, other than bioluminescence, due to the reduced illumination at depth. To explore potential visual behaviors in mesopelagic squid, we used undersea vehicles to observe 76 individuals of Octopoteuthis deletron. In contrast to predictions, we found this species capable of a variety of visually linked behaviors not previously reported for a deep-ocean cephalopod. The resultant ethogram describes numerous chromatic, postural, locomotor, and bioluminescent behavioral components. A few common body patterns-the whole appearance of the individual involving multiple components-are characterized. The behaviors observed from individual squid were compared using a Non-metric Multi-Dimensional Scaling (NMDS) ordination, onto which hydrographic and observation parameters were mapped. Observation length, specimen collection, and contact with the vehicle affected which behaviors were performed. A separate NMDS, analyzing the body patterns, indicated that these sets of behavioral components could be visualized as groups within the NMDS ordination. While the functional roles of the behaviors described are not yet known, our findings of numerous behaviors in O. deletron clearly indicate that bioluminescence is not the sole method of visual communication by deep-sea squid.

A unique 16-year time series of deep video surveys in Monterey Bay reveals that the Humboldt squid, Dosidicus gigas, has substantially expanded its perennial geographic range in the eastern North Pacific by invading the waters off central California. This sustained range expansion coincides with changes in climate-linked oceanographic conditions and a reduction in competing top predators. It is also coincident with a decline in the abundance of Pacific hake, the most important commercial groundfish species off western North America. Recognizing the interactive effects of multiple changes in the environment is an issue of growing concern in ocean conservation and sustainability research.

Gonatus onyx is one of the most abundant cephalopods in the Pacific and Atlantic Oceans and is an important prey species for a variety of vertebrate predators, but a full understanding of its life history has been hampered because spawning occurs at great depths, where observation is difficult. Here we describe post-spawning egg care, or brooding, in this deep-sea squid. Our finding is unexpected because this behaviour differs from the reproductive habits of all other known squid species.