Foraging strategies are of great ecological interest, as they have a strong impact on the fitness of an individual and can affect its ability to cope with a changing environment. Recent studies on foraging strategies show a higher complexity than previously thought due to intraspecific variability. To reliably identify foraging strategies and describe the different foraging niches they allow individual animals to realize, high-resolution multivariate approaches which consider individual variation are required. Here we dive into the foraging strategies of Galápagos sea lions (Zalophus wollebaeki), a tropical predator confronted with substantial annual variation in sea surface temperature. This affects prey abundance, and El Niño events, expected to become more frequent and severe with climate change, are known to have dramatic effects on sea lions. This study used high-resolution measures of depth, GPS position and acceleration collected from 39 lactating sea lion females to analyze their foraging strategies at an unprecedented level of detail using a novel combination of automated broken stick algorithm, hierarchical cluster analysis and individually fitted multivariate hidden Markov models. We found three distinct foraging strategies (pelagic, benthic, and night divers), which differed in their horizontal, vertical and temporal distribution, most likely corresponding to different prey species, and allowed us to formulate hypotheses with regard to adaptive values under different environmental scenarios. We demonstrate the advantages of our multivariate approach and inclusion of individual variation to reliably gain a deeper understanding of the adaptive value and ecological relevance of foraging strategies of marine predators in dynamic environments.

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Amphibious mammal species alternate between foraging at sea and attendance on land. Due to thermoregulatory requirements, they often haul out during the night, making social interactions difficult to observe. We tested the suitability of UHF-proximity loggers for assessing social relationships among Galápagos sea lions Zalophus wollebaeki. To survive periods at sea, proximity loggers were embedded in epoxy. Automatic downloads to receiving stations rendered logger recovery unnecessary. Encounters were logged within a range of 10 m. Logs provided information about interacting individuals as well as time and duration of encounters. 'Received signal strength indicator' values correlated with distance, but were influenced by antenna angle and environmental factors. Laboratory tests and validation in the field demonstrated that the spatial resolution corresponded to 2 m. Data needed to be corrected, as single encounters were recorded as multiple logs and reciprocity of loggings between animals was not always achieved. Digital data correlated with observational data, but associations were 4 times more likely to be detected using proximity loggers. Data revealed that non-territorial males frequently associated for extended periods of time, especially during the night and even outside the study colony. The modified proximity logger system proves an excellent tool to determine social structure in situations where direct observation is limited. KEY WORDS: Amphibious species · Proximity logger · Social structureResale or republication not permitted without written consent of the publisher AQUATIC BIOLOGY

Population monitoring is vital for conservation and management. However, simple counts of animals can be misleading and this problem is exacerbated in seals (pinnipeds) where individuals spend much time foraging away from colonies. We analyzed a 13-year-series of census data of Galapagos sea lions (Zalophus wollebaeki) from the colony of Caamaño, an islet in the center of the Galapagos archipelago where a large proportion of animals was individually marked. Based on regular resighting efforts during the cold, reproductive (cold-R; August to January) and the warm, non-reproductive (warm-nR; February to May) season, we document changes in numbers for different sex and age classes. During the cold-R season the number of adults increased as the number of newborn pups increased. Numbers were larger in the morning and evening than around mid-day and not significantly influenced by tide levels. More adults frequented the colony during the warm-nR season than the cold-R season. Raw counts suggested a decline in numbers over the 13 years, but Lincoln-Petersen (LP-) estimates (assuming a closed population) did not support that conclusion. Raw counts and LP estimates were not significantly correlated, demonstrating the overwhelming importance of variability in attendance patterns of individuals. The probability of observing a given adult in the colony varied between 16% (mean for cold-R season) and 23% (warm-nR season) and may be much less for independent 2 to 4 year olds. Dependent juveniles (up to the age of about 2 years) are observed much more frequently ashore (35% during the cold-R and 50% during the warm-nR seasons). Simple counts underestimate real population size by a factor of 4–6 and may lead to erroneous conclusions about trends in population size.