Scottish Marine and Freshwater Science Volume 6 Number 10: At-Sea Turnover of Breeding Seabirds - Final Report to Marine Scotland Science

The aim of this project was to review the potential issue of "turnover‟ of individual seabirds at sea during the breeding season and to assess how this may lead abundance estimates derived from boat or aerial surveys to underestimate the total number of b


2. Literature Review

2.1. Methodology

We conducted a review of the published literature relevant to estimating turnover in species included in the project but for which modelling was not carried out (red-throated diver, black guillemot, European shag, common eider, northern gannet and northern fulmar). We focused on studies (both UK and beyond) that provide estimates of parameters considered relevant to estimating turnover during the breeding season, such as time activity budgets, foraging trip characteristics and foraging site fidelity. The literature search was carried out in the Web of Science (all databases, 1950-2015) using the following search terms in combination with the species name: 'foraging range', 'foraging trip', 'foraging trip duration', 'site fidelity', 'foraging site fidelity' and 'foraging area fidelity', 'foraging and consistency', 'nest attendance', 'time activity budget' and 'activity budget'. Due to the limited time available and majority of the literature being focussed on the chick rearing period, the search was restricted to this stage of the breeding season.

Mean and standard deviation ( SD) were extracted for each parameter (where available), with the exception of foraging site fidelity where either various metrics or only qualitative information are reported in the literature. This parameter was, therefore, presented as a categorical variable, with two levels ('high'/'low') that directly reflect the interpretation of the authors of the original papers as to what constitutes a high or low measure of fidelity; details of the different fidelity metrics, however, are available within the cited references. In most cases parameter values were provided directly by the cited studies; in a few cases other relevant information was available which allowed us to derive the values of our parameters of interest. Derived values are presented in square brackets. Where data were available for more than one colony, estimates for each colony are presented separately; the only exceptions are two review studies (Langston 2010 and Thaxter et al. 2012) where estimates are averaged across multiple colonies and years. Where data were available for the same colony in multiple years, averaged values are presented; annual estimates, however, can be found within the cited references. Sample size of birds in each study is provided as an indication of reliability of the data; note also that estimates from recent studies are likely to be more accurate compared to those from older studies, as a result of the use of advanced bio-logging technologies in particular.

2.2. Results

Estimates of foraging trip characteristics (range, duration and frequency) were available for most species, and in many cases data for these parameters were available from multiple colonies (Table 1.2). In contrast, there was lack of information or only qualitative information available on foraging site fidelity for most species (Table 1.2). Data on daily activity budgets were of variable quality and often incomplete (Table 1.2). To fill these knowledge gaps, targeted field data collection involving the deployment of bio-logging devices and/or analysis of existing tracking datasets should be considered.

The most data-rich species was the Northern gannet, followed by the European shag. For these species parameter estimates from multiple colonies and years were typically available (Table 1.2). For the northern fulmar a reasonable amount of information was available regarding foraging trip characteristics but we found very little information on foraging site fidelity and daily activity budgets. Note also that in this species foraging behaviour of the adults changes substantially between the early (when chicks are brooded) and later (when chicks are not brooded) stages of offspring rearing which is why foraging trip characteristics are provided separately for these stages (Table 1.2). For the red-throated diver and black guillemot no recent data and very little older data were available for any of the parameters of interest (Table 1.2). Clearly, if turnover of individuals at sea is to be investigated in these species, new data collection would need to be prioritised as a first step. Due to the biology of the common eider, where females take their chicks to water soon after they hatch and do not return to the nest thereafter, the concept of a 'foraging trip' by a central-place foraging individual during chick rearing is not applicable. Linked to this, the cited estimates of foraging range indicate the general area used by the females and young once they have moved away from the nest site. Furthermore, male eiders take no part in offspring care and in the summer use different, generally more distant areas compared to females, where they initiate moult (Diéval et al. 2011). Therefore, to estimate turnover in this species, the approach to data collection would need to be sex-specific and may require (at least in females) relatively long-term deployments of the latest tracking technology (such as GPS- GSM tags, accelerometers) to obtain information of the birds' use of areas at sea at appropriate temporal and spatial scales.

Table 1.2

Parameters relevant to estimating turnover of individuals at sea during the breeding season (data from chick rearing period only; ' NA' = not available; '-' = not applicable; values in square brackets are derived based on information within the cited references - see Methods for details).

Parameter N birds Mean SD Reference
a) Red-throated diver ( Gavia stellata)
Foraging range (km) 9 11.1 NA Langston 2010
NA 4.5 NA Thaxter et al. 2012
Maximum foraging range (km) 9 12.2 NA Langston 2010
NA 9.0 NA Thaxter et al. 2012
Foraging trip duration (h) 6 1.0 NA Reimchen and Douglas 1984
16 0.9 0.6 Eriksson et al. 1990
Foraging trip frequency/day 6 5.5 NA Reimchen and Douglas 1984
16 7.0 NA Eriksson et al. 1990
Foraging site fidelity 16 high - Eriksson et al. 1990
Daily time at nest (h) NA NA NA NA
Daily resting time (h) NA NA NA NA
Daily foraging time (h) 16 [3.8] NA Eriksson et al. 1990
Daily commuting (flight) time (h) 16 [1.4] NA Eriksson et al. 1990
b) Black guillemot ( Cepphus grylle)
Foraging range (km) 56 0.7 0.5 Cairns 1987
38 5.0 NA Langston 2010
Maximum foraging range (km) 56 2.0 NA Cairns 1987
38 12.0 NA Langston 2010
Foraging trip duration (h) NA [~4.0] NA Cairns 1987; Gaston 1985
Foraging trip frequency/day NA 4.4 NA Cairns 1987
NA 5.0 NA Gaston 1985
Foraging site fidelity NA NA NA NA
Daily time at nest (h) NA 0* 0 Gaston 1985
Daily resting time (h) NA 7.0 NA Gaston 1985
Daily foraging time (h) NA 15.5 NA Gaston 1985
Daily commuting (flight) time (h) NA 1.5 NA Gaston 1985
c) European shag (Phalacrocorax aristotelis)
Foraging range (km) 29 7.0 1.9 Wanless et al. 1991
29 6.5 NA Langston 2010
29+ 5.9 4.7 Thaxter et al. 2012
Maximum foraging range (km) 29 16.4 NA Langston 2010
29+ 14.5 3.5 Thaxter et al. 2012
320 9.0 3.8 Bogdanova et al. 2014
57 4.0 3.7 Soanes et al. 2014
Foraging trip duration (h) 10 1.8 0.5 Wanless and Harris 1992
5 2.0 1.3 Gremillet et al. 1996
57 1.5 1.0 Soanes et al. 2014
Foraging trip frequency/day 10 2.8 0.4 Wanless and Harris 1992
Foraging site fidelity NA NA NA NA
Daily time at nest (h) 10 18.8 NA Wanless and Harris 1992
Daily resting time (h) 10 1.4 NA Wanless and Harris 1992
Daily foraging time (h) 10 2.5 NA Wanless and Harris 1992
Daily commuting (flight) time (h) 10 1.3 NA Wanless and Harris 1992
d) Common eider ( Somateria mollissima): females only
Foraging range (km) 10 9.3 NA Langston 2010
NA 2.4 NA Thaxter et al. 2012
Maximum foraging range (km) 55 72.0 NA Bustness and Erikstad 1993
10 38.3 NA Langston 2010
NA 80.0 NA Thaxter et al. 2012
Foraging trip duration (h) - - - -
Foraging trip frequency/day - - - -
Foraging site fidelity
- repeatability in foraging area in successive years (proportion of birds in area)
12 High - Bustness and Erikstad 1993
Daily time at nest (h) NA 0 0 Waltho and Coulson 2015
Daily resting time (h) 20 [20.6] NA Pelletier et al. 2008; Guillemette and Butler 2012
Daily foraging time (h) 20 [3.2] NA Pelletier et al. 2008; Guillemette and Butler 2012
Daily commuting (flight) time (h) 20 0.2 0.2 Pelletier et al. 2008
e) Northern gannet ( Morus bassanus)
Foraging range (km) 17 164.0 101.0 Hamer et al. 2000
62 140.1 NA Langston 2010
169+ 92.5 59.9 Thaxter et al. 2012
Maximum foraging range (km) 5 89.0 49.0 Hamer et al. 2001
20 100.0 35.0 Grémillet et al. 2006
53 238.2 108.0 Hamer et al. 2007
62 308.4 NA Langston 2010
169+ 229.4 124.3 Thaxter et al. 2012
17 106.0 43.0 Soanes et al. 2013
Foraging trip duration (h) 3 13.0 NA Garthe et al. 1999
5 11.9 6.7 Hamer et al. 2001
20 17.7 8.5 Grémillet et al. 2006
75 28.2 12.8 Lewis et al. 2005; Hamer et al. 2007
23 25.1 17.0 Votier et al. 2010
17 17.6 6.5 Soanes et al. 2013
Foraging trip frequency/day 5 [2.0] NA Hamer et al. 2001
20 [1.4] NA Grémillet et al. 2006
75 [0.8] NA Lewis et al. 2005; Hamer et al. 2007
23 [1.0] NA Votier et al. 2010
17 [1.4] NA Soanes et al. 2013
Foraging site fidelity
- repeatability in destination among successive trips (bearing and max distance)
5 Low - Hamer et al. 2001
53 High - Hamer et al. 2007
15 High - Soanes et al. 2013
18 High - Patrick et al. 2014
13 High - Patrick et al. 2014
15 Low - Soanes et al. 2013
- repeatability in duration of successive trips 18 Low - Patrick et al. 2014
13 Low - Patrick et al. 2014
Daily time at nest (h) 3 ~10.7 NA Garthe et al. 1999
22;53;12 [13.6] NA Lewis et al. 2004; Hamer et al. 2007; Ropert-Coudert et al. 2009
Daily resting time (h) 3 [~6.3] NA Garthe et al. 1999
22;53;12 [5.4] NA Lewis et al. 2004; Hamer et al. 2007; Ropert-Coudert et al. 2009
Daily foraging time (h) 3;22 [~0.6] NA Garthe et al. 1999; Lewis et al. 2004; Ropert-Coudert et al. 2009
22;53;12 [0.5] NA Lewis et al. 2004; Hamer et al. 2007; Ropert-Coudert et al. 2009
Daily commuting (flight) time (h) 3 ~6.4 NA Garthe et al. 1999
22;53;12 [4.5] NA Lewis et al. 2004; Hamer et al. 2007; Ropert-Coudert et al. 2009
f) Northern fulmar ( Fulmarus glacialis)
Foraging range (km) 51 69.4 NA Langston 2010
14+ 47.5 5.9 Thaxter et al. 2012
Maximum foraging range (km) 51 311.4 NA Langston 2010
14+ 400.0 245.8 Thaxter et al. 2012
Foraging trip duration (h)
- early chick-rearing (brooding)
14 <10 NA Furness and Todd 1984
4 10.2 4.0 Weimerskirch et al. 2001
50;48 11.2 NA Ojowski et al. 2001
- mid-/late chick-rearing (post-brooding) 23 28 NA Furness and Todd 1984
NA 24 NA Hamer et al. 1997
14 31.0 NA Phillips and Hamer2000
50;48 20.4 NA Ojowski et al. 2001
Foraging trip frequency/day
- early chick-rearing (brooding)
NA NA NA NA
- mid-/late chick-rearing (post-brooding) 23 0.8 NA Furness and Todd 1984
NA [0.9] [0.2] Hamer et al. 1997
28 0.8 NA Phillips and Hamer 2000
Foraging site fidelity 4 Low NA Weimerskirch et al. 2001
Daily time at nest (h) 28 ~1.0 NA Phillips and Hamer2000
Daily resting time (h) NA NA NA NA
Daily foraging time (h) NA NA NA NA
Daily commuting (flight) time (h) NA NA NA NA

* from 5 days after hatching no brooding occurs (Gaston 1985)

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