Abstract

In mammals, lactation can be the most energetically expensive part of the reproductive cycle. Thus, when energy needs are compromised due to predation risk, environmental disturbance, or resource scarcity, future reproductive success can be impacted. In marine and terrestrial environments, foraging behavior is inextricably linked to predation risk. But quantification of foraging energetics for lactating animals under predation risk is less understood. In this study, we used a spatially explicit individual‐based model to study how changes in physiology (lactating or not) and the environment (predation risk) affect optimal behavior in dolphins. Specifically, we predicted that an adult dolphin without calf would incur lower relative energetic costs compared to a lactating dolphin with calf regardless of predation risk severity, antipredator behavior, or prey quality consumed. Under this state‐dependent analysis of risk approach, we found predation risk to be a stronger driver in affecting total energetic costs (foraging plus locomotor costs) than food quality for both dolphin types. Further, contrary to our hypothesis, after accounting for raised energy demands, a lactating dolphin with calf does not necessarily have higher relative‐to‐baseline costs than a dolphin without calf. Our results indicate that both a lactating (with calf) and non‐lactating dolphin incur lowered energetic costs under a risk‐averse behavioral scheme, but consequently suffer from lost foraging calories. A lactating dolphin with calf could be particularly worse off in lost foraging calories under elevated predation risk, heightened vigilance, and increased hiding time relative to an adult dolphin without calf. Further, hiding time in refuge could be more consequential than detection distance for both dolphin types in estimated costs and losses incurred. In conclusion, our study found that reproductive status is an important consideration in analyzing risk effects in mammals, especially in animals with lengthy lactation periods and those exposed to both biological and nonbiological stressors.

Highlights

  • State-­based decision-­making under predation risk entails a trade-­off between feeding and vigilance (Brown & Kotler, 2004; Lima, 1998)

  • We added a bioenergetics module to the model developed by Srinivasan, Grant, Swannack, and Rajan (2010), which was created to explore the evolution of antipredator behavior in dusky dolphins in the Kaikoura Canyon

  • Detection: This strategy maximized time spent in the refuge by the duskies We calculated caloric costs of nightly foraging behavior for both LD and AD, under each of these five scenarios, under each of six predation risk levels, and under each of four dolphin prey capture costs represented as a percentage caloric content of prey item consumed (5%, 10%, 15%, and 20%)

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Summary

| INTRODUCTION

State-­based decision-­making under predation risk entails a trade-­off between feeding and vigilance (Brown & Kotler, 2004; Lima, 1998). Smaller-­bodied species lack these metabolic stores and are unable to fast for extended periods (Costa, 2009; Mann, 2009; Oftedal, 1997) They reduce activity, which can serve as an ancillary strategy to overcome energy deficits, as in non-human primates (Barrett, Halliday, & Henzi, 2006; Dufour & Sauther, 2002). Direct observations of predation events are rare (Constantine, Visser, Buurman, Buurman, & McFadden, 1998; Visser, 1999), but there are multiple accounts of duskies fleeing at top speed when killer whales are detected, leaving the area entirely until the threat has subsided, or seeking refuge in shallow waters

| METHODS
| DISCUSSION
Findings
CONFLICT OF INTEREST
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