RESUMO
BACKGROUND: Prey can alter their behavior when detecting predator cues. Little is known about which sensory channel, number of channels, or the interaction among channels that shrimp species use to evaluate the threat from predators. The amphidromous shrimp Xiphocaris elongata has an induced defense, an elongated rostrum, where predatory fishes are present. We sought to test if kairomones or visual cues when presented singly from fish either eating flakes or shrimp, had more effect on altering the temporal feeding and refuge use patterns of long-rostrum (LR) X. elongata. We were also interested in elucidating potential interactions among cues when presented simultaneously in different combinations (kairomones + visual + mechanosensory, kairomones + alarm + visual, kairomones + alarm, kairomones + visual) on the same response variables. We expected that when presented alone kairomones will significantly increase refuge use and decrease foraging, particularly late at night, in comparison to visual cues alone, and that multiple cues when presented simultaneously will further increase refuge use and decrease foraging at night. METHODS: We exposed shrimp to individual or multiple cues from the predatory fish mountain mullet, Augonostomus monticola. We examined shrimp behavior with respect to refuge use and foraging activity during four time periods (after sunset, nighttime, sunrise, and sunset) in a 24-hour period. RESULTS: Shrimp presented fish visual and chemical cues singly did not differ from one another but differed from control shrimp (no cues) with respect to refuge use or foraging. The number of shrimp using refuge in the treatment with most cues (KVM: kairomones+ visual + mechanosensory) was higher than in all the treatments with less cues. A significant decline in foraging was observed when multiple cues were presented simultaneously. The highest number of shrimp foraged one hour after sunset and at nighttime. A significant interaction was observed between cue treatments and time periods, with shrimp in the KVM treatment foraging less and using more refuge late at night and at sunrise than shrimp in other treatments or time periods. CONCLUSIONS: The observation that fish chemical and visual cues when presented singly produced similar refuge use and foraging patterns was contrary to expectation and suggests that visual and chemical cues, when presented alone, provide redundant information to X. elongata with regards to predation threat. The significant increase in refuge use and reduction in foraging observed in the KVM treatment suggest multimodal signal enhancement in the perception of threat. This makes evolutionary sense in "noisy" environments, such as streams, where detection, localization, and intention of predators is much improved when cues are received through multiple sensory channels.
RESUMO
Aposematic coloration is traditionally considered to signal unpalatability or toxicity. In mammals, most research has focused on just one form of defense, namely, noxious anal secretions, and its black-and-white advertisement as exemplified by skunks. The original formulation of aposematism, however, encompassed a broader range of morphological, physiological, and behavioral defenses, and there are many mammal species with black-and-white contrasting patterns that do not have noxious adaptations. Here, using Bayesian phylogenetic models and data from 1726 terrestrial nonvolant mammals we find that two aspects of conspicuous coloration, black-and-white coloration patterns on the head and body, advertise defenses that are morphological (spines, large body size), behavioral (pugnacity), and physiological (anal secretions), as well as being involved with sexual signaling and environmental factors linked to crypsis. Within Carnivora, defensive anal secretions are associated with complex black-and-white head patterns and longitudinal black-and-white body striping; in primates, larger bodied species exhibit irregular patches of black-and-white pelage; and in rodents, pugnacity is linked to sharp countershading and irregular blocks of white and black pelage. We show that black-and-white coloration in mammals is multifunctional, that it serves to warn predators of several defenses other than noxious anal secretions, and that aposematism in mammals is not restricted to carnivores.