Abstract

Rapid development of a successful foraging strategy is critical for juvenile survival, especially for naïve animals that receive no parental guidance. However, this process is poorly understood for many species. Although observation of early‐life movements is increasingly possible with miniaturisation of animal‐borne telemetry devices, analytical limitations remain. Here, we tracked 29 recently‐weaned grey seal Halichoerus grypus pups from colonies in two geographically distinct regions of the United Kingdom. We analysed at‐sea movements of pups throughout their initial months of nutritional independence to investigate the ontogeny of behaviour‐specific (foraging and travelling) movement patterns. Using generalized hidden Markov models (HMMs), we extended the conventional HMM framework to account for temporal changes in putative foraging and travelling movement characteristics, and investigate the effects of intrinsic (sex) and extrinsic (environment) factors on this process. Putative foraging behaviour became more tortuous with time, and travelling became faster and more directed, suggesting a reduction in search scale and an increase in travel efficiency as pups shifted from exploration to an adult‐like repeatable foraging strategy. Sex differences in movement characteristics were evident from colony departure, but sex‐specific activity budgets were only detected in one region. We show that sex‐specific behavioural strategies emerge before sexual size dimorphism in grey seals, and suggest that this phenomenon may occur in other long‐lived species. Our results also indicate that environmental variation may affect the emergence of sex‐specific foraging behaviour, highlighting the need to consider interacting intrinsic and extrinsic factors in shaping movement strategies of long‐lived vertebrates. Moreover, comparing the behavioural state estimations to those of a conventional HMM (no variation in state‐specific movement parameters) revealed differences in the amount and location of foraging activity, with implications for spatial conservation management. Overlooking intrinsic and extrinsic variation in movement processes could distort our understanding of foraging ecology, population dynamics and conservation requirements.

Highlights

  • The period immediately following nutritional independence is one of the most vulnerable times in the life of an animal (Lindström 1999)

  • Using location and dive data from recently-weaned pups tagged on breeding colonies in two distinct regions of the United Kingdom (UK; Northeast Scotland, West Wales), we examine the movement characteristics of putative foraging and travelling throughout the first four months of nutritional independence at sea

  • Predation is a limiting factor on survival of many newly-independent animals, such as cheetahs Ancinonyx jubatus (Laurenson 1994) and passerine birds (Sullivan 1989), and predation events on weaned grey seal pups have been recorded in the UK (Brownlow et al 2016), there is little evidence that predation has a population-level impact on firstyear survival in UK grey seals

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Summary

Introduction

The period immediately following nutritional independence is one of the most vulnerable times in the life of an animal (Lindström 1999). Starvation is a major cause of mortality for many species during this early life stage, and individuals have a finite window of opportunity in which to develop effective foraging strategies before energy reserves are depleted (Sullivan 1989, Orgeret et al 2016). This critical learning period is most pertinent for species that have no experience of foraging at the point of nutritional independence. Early-life exploratory movements have been reported in longlived species, such as seabirds (de Grissac et al 2017) and phocid seals (McConnell et al 2002)

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