Ibis (2025), 167, 468–480 doi: 10.1111/ibi.

13371

Spatial consistency affects foraging effort in a

benthic diving seabird, the Black-faced Cormorant
THOMAS CANSSE,*1,2 LUC LENS,2 GRACE J. SUTTON,†,1 JONATHAN A. BOTHA‡,1 &
JOHN P.Y. ARNOULD1
1
School of Life and Environmental Sciences, Deakin University, Burwood, Australia
2
Terrestrial Ecology Unit, Ghent University, Ghent, Belgium

For animals which exploit predictable food resources, greater foraging site fidelity has
been hypothesized to lead to increased foraging efficiency. However, evidence for this in
free-ranging species is limited. In the present study, foraging site fidelity, the factors
influencing it and its effect on foraging effort were investigated in the Black-faced Cor-
morant Phalacrocorax fuscescens, a predominantly benthic forager occurring in southeast-
ern Australia. While the study population was found to have a relatively large foraging
range, individuals were highly consistent in the smaller foraging areas which they used.
Foraging effort was lower in individuals with a higher foraging site fidelity, which is likely
to indicate increased foraging efficiency. Foraging site fidelity differed greatly between
individuals but none of the assessed factors (sex, mass, size or year of study) were found
to have a significant influence. This might indicate that intrinsic factors which could not
be assessed, such as personality or experience, may influence the degree to which indi-
viduals exploit known habitats. The findings suggest that changes in prey availability or
distribution could result in reduced foraging efficiency and, consequently, negative
impacts for the species. Further investigation is required to assess within-individual forag-
ing plasticity to predict potential responses to changing prey availability and distribution.
Keywords: biologging, cormorant, foraging consistency, foraging effort, seabird.

Optimal foraging theory predicts that animals will Stephens et al. 2008, Baert et al. 2021). Species
aim to optimize their foraging efficiency, both by with a large niche width are considered to be gen-
minimizing time and energy invested in foraging, eralists while those with a narrow niche width are
as well as by maximizing collected resources over classified as specialists (Bolnick et al. 2003). Some
time (Schoener 1971, Ydenberg et al. 1994). To species have a large niche width at the species
do so, individuals can employ different strategies, level but have populations comprising individuals
such as targeting prey with a high energetic con- with narrow niche widths that each exploit differ-
tent, targeting patches of high prey abundance, ent resources (Ara
ujo et al. 2011).
and specialization towards certain foraging habitats When the abundance and distribution of
or prey types, to minimize the time needed to find resources is highly variable, generalist individuals
or capture prey (MacArthur & Pianka 1966, are believed to have an advantage over specialists
as their capacity to switch between available
†
resources can lead to more consistent reproductive
Present address: Department of Environment & Genetics, and
outputs (Partridge & Green 1987, Terraube
Research Centre for Future Landscapes, La Trobe University,
Bundoora, Australia et al. 2011, Korpim€aki et al. 2020). Conversely,
‡
Present address: Biosis Pty Ltd, Port Melbourne, Australia when resources are limited in abundance but have
a high spatial and temporal predictability, it can
*Corresponding author. be advantageous for individuals to exhibit high
Email: [email protected]
levels of foraging site fidelity (i.e. revisit the same
[Correction added on 21 January 2025, after first online publi- location repeatedly for foraging) and resource spe-
cation: The copyright line was changed.] cialization (Switzer 1993, Tinker et al. 2008).

© 2024 The Author(s). IBIS published by John Wiley & Sons Ltd on behalf of British Ornithologists' Union.
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and
distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made.
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Foraging consistency in Black-faced Cormorants 469

Such specialization can reduce the time and through experience, benthic foragers may gain
energy spent searching for prey, for example knowledge of profitable foraging patches which, in
through increased site familiarity or through an turn, can lead to the development of foraging spe-
increase in prey capture success, thereby increasing cializations which enhance efficiency (Werner
foraging efficiency (Switzer 1993, Terraube et al. 1981, Hines et al. 1997, River
on et al. 2021).
et al. 2014, Araujo & Moura 2022). However, due to oceanographic impacts of climate
As a consequence of anthropogenic impacts on change, the distribution, diversity and abundance of
the environment, individual foraging specialization prey in the marine environment are expected to be
and the associated site fidelity potentially could altered (Perry et al. 2005, Johnson et al. 2011).
have deleterious effects, both at the individual and These changes could especially impact benthic for-
population levels (Merkle et al. 2022). For exam- agers, which can exhibit foraging specializations at
ple, Northern Elephant Seal Mirounga angustiros- smaller spatial scales than pelagic foragers (Wei-
tris individuals with a high and low site fidelity merskirch 2007, Ceia & Ramos 2015, Merkle
performed similarly over the long term. However, et al. 2022).
having a high site fidelity became maladaptive Cormorants are a group of diving birds living in
under anomalous climate conditions, whereas low marine and freshwater environments across the
site fidelity became advantageous (Abrahms world (Nelson 2005). The majority of species
et al. 2018). Consequently, site fidelity can act as exclusively exploit marine environments where
an ecological trap in some species, for example as they feed predominantly on benthic fish species
a result of a decline in habitat quality or food (Nelson 2005). As benthic foragers feeding on
availability at formerly good-quality sites (Ekroos prey with relatively high levels of predictability,
et al. 2012, Kloskowski 2021). several species of cormorants have been shown to
Foraging site fidelity between individuals can dif- have high levels of individual foraging specializa-
fer by underlying factors such as individual experi- tions at several temporal scales (Bearhop
ence (Votier et al. 2017), personality (Harris et al. 2006, Camprasse et al. 2017, Morgan
et al. 2020) and sex (Phillips et al. 2017). However, et al. 2019, Fijn et al. 2022). While this foraging
these individual differences are often closely tied to specialization and consistency is often assumed to
the foraging ecology of the species as well as the have positive impacts on foraging efficiency, its
habitat type they use (Phillips et al. 2017). Conse- effects have only rarely been studied in cormorants
quently, the level of specialization, the factors (Morgan et al. 2019).
influencing it or its consequences on their foraging The Black-faced Cormorant Phalacrocorax fus-
behaviour are not well understood for most species cescens, Australia’s only endemic cormorant spe-
(Ceia & Ramos 2015). The degree of foraging site cies, has a breeding distribution stretching across
fidelity can be used as a metric to quantify individ- the continent’s southeastern coast (Nelson 2005).
ual foraging specialization (Carneiro et al. 2017). This region is experiencing some of the fastest oce-
This can subsequently be linked to intrinsic (such as anic warming in the world, which is predicted to
sex, size or mass) or extrinsic (such as year of study) impact the distribution, diversity and abundance
factors to understand the drivers of individual differ- of marine prey (Perry et al. 2005). The species
ences, as well as to foraging effort to understand the forms colonies on offshore islands ranging in size
consequences of foraging specialization on foraging from a few birds to more than a thousand breed-
efficiency (Toscano et al. 2016, Carneiro et al. 2017, ing pairs (Marchant et al. 1990), and forages exclu-
Ramellini et al. 2022). sively in the marine environment (Nelson 2005).
In the marine environment, prey resources are Previous studies have revealed that the Black-faced
highly variable in their distribution and abundance. Cormorant is primarily a benthic forager, with
This is especially the case for pelagic prey which maximum foraging depths constrained by its body
can reach high abundances but are highly variable, mass-influenced aerobic dive limit (Cansse
both spatially and temporally (Gende & Sig- et al. 2024). The species is sexually dimorphic
ler 2006, D€anhardt & Becker 2011). In contrast, (Riordan & Johnston 2013) and, consequently, the
benthic prey have lower abundances but their dis- larger males can forage in deeper regions which
tribution is predictable at smaller spatial scales results in sexual segregation of foraging areas dur-
than pelagic prey (Litzow et al. 2004, River
on ing the breeding season. Preliminary reports sug-
et al. 2021, Filatova et al. 2022). Consequently, gest that the Black-faced Cormorant displays

© 2024 The Author(s). IBIS published by John Wiley & Sons Ltd on behalf of British Ornithologists' Union.
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470 T. Cansse et al.

individual consistency in its spatial habitat use and (2020, 2022 and 2023) or 10 min (2021) for the
dive depths during the breeding season (Cansse combined GPS-pressure loggers, or every 10 min
et al. 2024), yet the degree of inter-individual vari- (2020 and 2022) or 20 min (2021) for the
ability in specialization, or its consequences, are GPS-GSM loggers. In order to reduce energy con-
not known. Such information is crucial for under- sumption, loggers were programmed to stop col-
standing foraging efficiency and predicting poten- lecting GPS data at night between 8:00 PM and
tial responses of the Black-faced Cormorant to the 4:00 AM local time. Pressure data were collected at
anticipated climate change-induced alterations in 4-s intervals as a consequence of limitations in bat-
the southeastern Australia oceanic region (Cham- tery and memory size. Similarly, differences in
bers et al. 2011, Hobday & Lough 2011, Merkle GPS collection intervals between years and loggers
et al. 2022). were a consequence of limitations in battery
The aims of the present study, therefore, were capacity. Data were remotely downloaded with a
to: (1) determine the level of individual foraging base station at the colony (Pathtrack nanoFixâ
site fidelity; (2) investigate potential factors GEO + RF loggers), or over the GSM network
influencing such behavioural consistency; and (3) (Technosmart Gipsy and Pathtrack GEO + GSM
assess its consequences for foraging effort and effi- loggers) until the logger battery expired, the
ciency in Black-faced Cormorants. device was shed or the bird was out of reception
range.
All GPS loggers used in the present study
METHODS
weighed substantially less than the recommended
3% of the body mass, a limit which has been
Fieldwork data collection
shown to have negligible effects (Wilson
The study was conducted at Notch Island et al. 2002, Vandenabeele et al. 2012). However,
(38°560 2500 S, 146°400 3300 E), northern Bass Strait, owing to logistical constraints, no comparisons
southeastern Australia. The island hosts a could be made between tagged and untagged
Black-faced Cormorant colony of approximately birds. Therefore, tag effects on the diving and for-
950 nests (Taylor et al. 2013). Data collection aging effort of individuals cannot be excluded
occurred during chick-rearing (September to Octo- (Vandenabeele et al. 2015). However, as such tag
ber) between 2020 and 2023. Adults rearing chicks effects are likely to have been negligible and simi-
aged approximately 20–40 days were captured with lar for all tagged individuals, they should not affect
a noose-pole when they returned to the breeding the interpretation of the reported results.
colony after a foraging trip. Individuals were
weighed in a cloth bag using a spring scale (25 g,
Data processing
Salter, Bristol, UK) and morphometric measure-
ments (culmen length, bill length, tarsus length and The collected GPS tracks were split into individual
bill depth) were taken using vernier callipers foraging trips using the track2kba package (Beal
(0.1 mm). Two body contour feathers were col- et al. 2021) in the R statistical environment (R
lected for molecular sexing and a uniquely num- Core Team 2022). A buffer of 0.5 km around the
bered metal leg band was applied. A combined colony was used as a threshold to start and end
remote download pressure-GPS logger (Pathtrack trips. This buffer was chosen to avoid classifying
nanoFixâ GEO + RF (20 g, 1.1  0.1% of body cormorants roosting on Notch Island, which
mass; 2020, 2021, 2022, 2023)) or remote down- extends roughly 350 m north-west and 300 m
load GPS-GSM logger (Technosmart Gipsy (25 g, south of the colony, as being on a foraging trip.
1.4  0.1%; 2020) or Pathtrack GEO + GSM Both visual inspection of the tracks and observa-
(20 g, 1.1  0.1% of body mass; 2021, 2022)) was tions during fieldwork indicated no foraging in the
then attached to two central tail feathers with immediate vicinity of the breeding colony. If no
waterproof tape (TESAâ 4651, Beiersdorf AG, data were collected for more than 12 h, or if the
Germany (Wilson et al. 1997)) before individuals return to the colony was not recorded, the trip
were released near the edge of the colony to resume was considered as incomplete. Although data gaps
normal behaviour. longer than 1 h were observed in the tracking data
Pressure data were collected at 4-s intervals, of 66% of the individuals in the present study,
and GPS location data were collected every 5 min they were not observed frequently (on average

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Foraging consistency in Black-faced Cormorants 471

0.41  0.07 gaps > 1 h day1). Missing GPS complete overlap. For each individual, the mean
points were linearly interpolated using the ‘redis- overlap of all trips for that individual was calcu-
ltraj’ function in the adehabitatLT package lated to derive a foraging site fidelity index
(Calenge 2011a) so as to match the programmed (FSFI ). In addition, for each trip, the mean over-
schedule. For individuals where pressure data were lap with all other trips for the same individual
available, these were processed using the diveMove was calculated (trip FSFI ). Subsequently, for each
package (Luque 2007), with zero-offset correction trip, the individual FSFI was subtracted from the
using the running quantile method, to obtain dive trip FSFI, which resulted in the DFSFI (i.e.
behaviour summaries. Dive locations were linearly within-individual centring; see van de Pol and
interpolated from the tracking data. Wright (2009) and Table S1). In this method, a
As only a subset of individuals had dive data positive DFSFI indicates that a trip had a higher-
available, a Hidden Markov Model (HMM) was than-average overlap with other trips of that indi-
used to identify the at-sea activities throughout vidual, while a negative DFSFI indicates a lower-
foraging trips in all individuals. This HMM used than-average overlap. By following the framework
step length and turning angle derived from the of van de Pol and Wright (2009), linear mixed
GPS tracks to identify the behavioural states of models containing FSFI and DFSFI could be
roosting, foraging (including resting on water) and applied to discern, respectively, the
commuting flight while on a foraging trip. Resting inter-individual and intra-individual effects of
on water was not classified as a separate behaviour increasing FSFI on foraging effort. Using both
as the logger sampling interval did not allow this FSFI and DFSFI in the model allows determina-
to be discerned from foraging behaviour. The tion of whether inter- and intra-individual
HMM was first fitted to the subset of individuals increases in foraging site fidelity affect foraging
for which dive behaviour was available using the effort in the same direction.
moveHMM package (Michelot et al. 2016) to Two variables indicative of foraging effort were
select the parameter values which resulted in the used: first, the proportion of time spent foraging,
best-fitting model. After fitting the HMM, tracks which was calculated as the amount of time forag-
were inspected visually, and the collected diving ing in a trip divided by the active trip duration
data were used to verify model accuracy. Subse- (i.e. trip duration minus time classified as roosting
quently, the best-fitting model was fitted to the away from the colony). Active trip duration was
tracks without dive data. As sampling intervals var- used instead of trip duration to decrease the vari-
ied between logger types and years, GPS data were ability in foraging proportion between trips, as trip
resampled to the largest time interval (20 min) duration was often increased due to birds roosting
before fitting the HMM. away from the colony. Secondly, for individuals
Data collection periods varied between the var- where dive data were available, the dive rate
ious devices used. For birds equipped with (m h1; vertical distance travelled per active trip
GPS-pressure loggers, on average 8.4  0.7 trips hour) was also investigated.
were collected (range: 1–28), while for birds with
GPS-GSM loggers, on average 15.5  2.1 trips
Statistical analyses
were collected (range: 1–53). Hence, to standard-
ize the amount of data analysed for each bird, as In a first step, the effects of sex, year,
well as to avoid using data collected potentially tarsus-length and mass on individual-level foraging
after chicks had fledged, only the first five forag- site fidelity were tested in a linear model with a
ing trips were analysed for each bird. Birds with beta distribution. Mass and sex were added as
fewer than five trips were excluded from further earlier research indicated that mass, which is
analyses. Subsequently, the 95% kernel utilization higher for males in this sexually dimorphic spe-
distribution (UD) for all locations classified as for- cies, affects diving behaviour leading to differ-
aging or commuting was calculated for each trip ences in habitat use in sexes (Cansse et al. 2024).
using the adehabitatHR package (Calenge 2011b), Thereby, these were added to the model to
and Bhattacharyya’s Affinity Index was calculated determine if these differences affected foraging
for each combination of trips (Bhattacharyya 1943, site fidelity. Tarsus-length was added as a mea-
Fieberg & Kochanny 2005). An index of 0 indi- sure of individual size to investigate whether this
cates no overlap, while an index of 1 indicates affected site fidelity. Lastly, year of study was

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472 T. Cansse et al.

added to investigate potential interannual differ-

RESULTS
ences in site fidelity.
In a second step, linear mixed models were Data were obtained from 72 birds (43 males, 29
used to assess the factors influencing foraging females) across the 4 years of study (Table 1).
effort. Two dependent variables indicative of for- Dive behaviour and GPS data were available for
aging effort, the proportion of an active trip spent 41 individuals (24 males, 17 females), and the
foraging and the dive rate, were used in two sepa- remaining birds only had GPS data. Loggers which
rate models as dependent variables. Both models collected dive behaviour collected data for
contained FSFI and DFSFI as well as sex, year of 8.3  0.5 days while the GPS-only loggers col-
study, tarsus-length (as a measure of individual lected data for 20.0  1.9 days. Comparison of
size) and overnight absence from the colony as the HMM-derived behaviour categories with
fixed factors. Overnight absence from the colony obtained dive behaviour indicated that the HMM
was added to the models as some foraging trips had similar levels of accuracy in each year of the
involved overnight roosting away from the colony, study (Table S2). Although a relatively high pro-
and it was hypothesized that this prolonged roost- portion of the commuting sections still contained
ing while on a trip could be a factor affecting for- dives (i.e. false negatives for foraging behaviour)
aging effort independently from site fidelity. Thus, (range: 54.8–66.6%, Table S2), the majority of the
adding overnight absence to the model allowed dives were included in sections classified as forag-
this potential effect to be controlled for. Sex was ing (68.7–74.2%, Table S2). This indicated that in
added as earlier research indicated differences in the derived HMM, sections classified as foraging
dive depth and habitat use between males and were largely representative for the spatial and tem-
females (Cansse et al. 2024), which potentially poral occurrence of foraging behaviour. The high
affect foraging effort. Tarsus-length was added as a proportion of commuting sections containing dives
measure of individual size to investigate whether is likely to be an effect of the large sampling inter-
size affected foraging effort. Year of study was val, thereby causing dives to be included at the
added to control for potential inter-annual varia- start or end of commuting intervals. To assess
tion in foraging effort. The individual was added as whether model outcomes were affected by the
a random factor in both models. The model con- sampling rates, the HMM was used to classify the
taining the proportion of time foraging used a beta data collected at 10-min intervals which slightly
distribution. increased accuracy (Table S3). However, model
Models which used the beta distribution were outcomes were similar, indicating the models were
fitted using the glmmTMB package (Brooks robust to some misclassifications (Table S4).
et al. 2017). All other linear mixed models were At the population level, Black-faced Cormo-
fitted using the lme4 package (Bates et al. 2015). rants used a large area for foraging reaching maxi-
For model selection throughout the study, the full mum distances of 21.9  0.7 km from the colony
model was specified, and the ‘dredge’ function in (Fig. 1). However, at the individual level, the areas
the MuMIn package (Barton 2018) was used to used were much smaller as individuals repeatedly
compare all possible models. For the models inves- visited the same area for foraging. While there
tigating inter- and intra-individual effects of FSFI, were large inter-individual differences in foraging
the FSFI and DFSFI could not be included sepa- site fidelity (FSFI; range: 0.19–0.96), the median
rately (van de Pol & Wright 2009) and, therefore, FSFI was high (0.73). This indicated that the
only models containing both or none of these vari- majority of the Black-faced Cormorants in the pre-
ables were compared in the dredge function. Can- sent study displayed high foraging site fidelity
didate models were compared using Akaike’s (Fig. S1). None of the considered variables in the
information criterion corrected for small sample model (sex, year, mass and tarsus-length) had a
sizes (AICc). Subsequently, the difference in AICc significant effect on the degree of foraging site
(DAICc) was used for model averaging on all fidelity.
models with DAICc < 4 to determine the factors On average, Black-faced Cormorants in the pre-
influencing the dependent variables for all models sent study were found to forage for 65.1  1.1%
(Burnham & Anderson 2002, Symonds & of their active trip duration. For birds where dive
Moussalli 2011). data were available, the average dive rate was

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Foraging consistency in Black-faced Cormorants 473

Table 1. Summary information for

Black-faced Cormorants Phalacrocorax Sex
fuscescens instrumented with GPS and
dive behaviour data loggers at Notch Male Female
Island, northern Bass Strait, southeastern
Australia. Year Mass (g) GPS (N ) Dive data (N ) Mass (g) GPS (N ) Dive data (N )

2020 2082  102 15 6 1699  32 14 8

2021 1892  121 10 6 1658  35 6 2
2022 2094  32 13 7 1625  57 6 4
2023 2050  41 5 5 1716  22 3 3

Figure 1. Representative foraging tracks from Black-faced Cormorants breeding at Notch Island displaying the high level of foraging
site fidelity. Each colour represents a different individual (N = 11). The black dot on the map indicates the location of Notch Island.

370.3  20.5 m h1. In the models assessing and dive rate), FSFI and DFSFI were consistently
which factors (sex, mass, tarsus-length, year of included in all of the top candidate models
study, FSFI, DFSFI and overnight absence from the (DAICc < 4), indicating that these factors affected
colony) influenced these considered measures of foraging effort (Table S5). Year of study and over-
foraging effort (i.e. proportion of time foraging night absence were included in all of the top

© 2024 The Author(s). IBIS published by John Wiley & Sons Ltd on behalf of British Ornithologists' Union.
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474 T. Cansse et al.

candidate models for dive rate. The presence or (305.4 to 18.1)), potentially indicating
absence of other predictor variables in candidate inter-annual differences in prey availability.
models was more variable (Table S5). As multiple
candidate models with DAICc < 4 were found,
DISCUSSION
model averaging was conducted for all dependent
variables. Foraging site fidelity can lead to increases in forag-
The proportion of time spent foraging in a trip ing efficiency, but evidence for this in free-ranging
decreased both with increasing FSFI (95% confi- species is limited. In the present study, a high con-
dence interval (CI) (1.84 to 0.67)) and with sistency in spatial habitat-use and foraging site
increasing DFSFI (95% CI (2.97 to 0.90)) fidelity was quantified in the Black-faced Cormo-
(Table 2). This indicates that increasing the forag- rant, a predominantly benthic forager occurring in
ing site fidelity leads to a decrease in the propor- southeastern Australia. This behavioural consis-
tion of a trip which is spent on foraging both tency was associated with reduced foraging effort,
within and between individuals (Fig. 2a). No other suggesting that individuals foraging in familiar
factors were found to have a consistent effect on locations may experience increased foraging
the proportion of time spent foraging (Table 2). efficiency.
For the subset of individuals where diving data
were available, the dive rate decreased as FSFI
Individual variation in site fidelity
increased (95% CI (565.0 to 33.1)) indicating
an inter-individual effect, where individuals with a At the population level, Black-faced Cormorants
higher FSFI had a lower dive rate (Fig. 2b). As in the present study were observed to use a rela-
DFSFI increased, the dive rate decreased (95% CI tively large area for foraging. However, individuals
(740.5 to 258.4)). This indicates that, as indi- foraged repeatedly in the same restricted location,
viduals increased their foraging site fidelity, they indicating a high level of individual specialization
reduced diving effort (intra-individual effect) in foraging sites. Similar specialization in foraging
(Fig. 2b). Dive rate was lower when an individual locations has been observed in other cormorant
was absent from the colony overnight (95% CI and shag species (Potier et al. 2015, Camprasse
(275.2 to 170.8)), indicating that overnight et al. 2017, Morgan et al. 2019), as well as in ben-
roosting may decrease effort, potentially as a con- thic foragers in general (River
on et al. 2021, Fila-
sequence of roosting in areas with good prey avail- tova et al. 2022).
ability. Dive rate was observed to be higher than The factors year of study, morphometrics (i.e.
in other years of study in 2021 (95% CI mass or tarsus-length) or sex were not found to
(73.7–402.1)) and lower in 2022 (95% CI have a significant influence on foraging site fidelity.
However, large inter-individual differences were

Table 2. Results from the models examining the factors influencing foraging proportion and dive rate.

Proportion foraging (%) Dive rate (m h1)

Predictors Est sd 95% CI Est sd 95% CI

Intercept 1.21 1.02 0.78 to 3.22 600.9 333.5 56.5 to 1258.4

FSFI 1.26 0.29 1.84 to 0.67 299.5 135.0 565.0 to 33.1
DFSFI 1.92 0.52 2.97 to 0.90 499.4 122.2 740.5 to 258.4
Overnight 0.04 0.08 0.12 to 0.21 223.0 26.5 275.2 to 170.8
Year (2021) 0.07 0.15 0.22 to 0.36 237.9 83.3 73.7 to 402.1
Year (2022) 0.05 0.12 0.18 to 0.29 161.8 72.8 305.4 to 18.1
Year (2023) 0.03 0.13 0.29 to 0.21 51.1 87.0 222.8 to 120.4
Sex (M) 0.12 0.15 0.42 to 0.18 83.8 73.0 59.7 to 227.3
Tarsus-length (mm) 0.00 0.01 0.03 to 0.04 0.6 5.5 10.1 to 11.5

The reported values were obtained after model averaging of all candidate models (DAICc < 4). Reported are the parameter esti-
mates (est), their standard deviation (sd) and 95% confidence interval (95% CI). When the CI did not cross 0 (i.e. there was an effect
of the predictor), this is indicated in bold.

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Foraging consistency in Black-faced Cormorants 475

Figure 2. Model-predicted effects of foraging site fidelity on (a) proportion of time spent foraging in a trip (%) and (b) dive rate
(m h1). Within-individual effects are indicated in dark grey (full line) and between-individual effects are indicated in light grey
(dashed line).

observed in foraging site fidelity. This could indi- important for Black-faced Cormorants to achieve
cate that other factors which are related to the high foraging efficiency.
individual, but which could not be assessed in the
present study, may influence site fidelity. For
Influence of foraging site fidelity on
example, in Black-legged Kittiwakes Rissa tridac-
foraging efficiency
tyla, bolder individuals were found to have a
higher foraging site fidelity (Harris et al. 2020). In When interpreting the effects on foraging effort
Northern Gannets Morus bassanus, foraging site in the present study, it was assumed that a lower
fidelity was higher for adults than for immatures, proportion of time at sea spent foraging was
indicating that age, potentially through individual indicative of a higher foraging efficiency. This was
familiarity with the environment, could influence assumed because benthic foraging cormorants do
this behavioural consistency (Votier et al. 2017). not locate prey while commuting (Harris
Changing environmental conditions could et al. 2012) and, as such, the commuting time to
potentially influence the distribution and abun- and from the foraging spot will have little varia-
dance of prey species for marine predators (Perry tion. As a result, if more time was spent on for-
et al. 2005, Johnson et al. 2011). Consequently, aging, as a consequence of low capture success or
the level of foraging site fidelity, and its adaptive more time needed to locate prey, the proportion
value, might differ between years if prey condi- of time spent foraging would increase and, there-
tions change (Switzer 1993, Foo et al. 2020, fore, foraging efficiency can be assumed to be
Michelot et al. 2021). While in the present study lower. Similarly, a lower dive rate was assumed
no significant differences in foraging site fidelity to be indicative of a higher foraging efficiency.
were found between years, a greater range in fidel- This was assumed as individuals with a lower
ity was observed, and lower fidelity was more dive rate are likely to need fewer dives to search
common in 2022 and 2023 than in other years. In and capture sufficient prey.
addition, dive rate (m h1) was lower in 2022, No data are available for Black-faced Cormorants
suggesting greater prey availability (Womble linking the proportion of time spent foraging or dive
et al. 2014). Furthermore, a concurrent study effort to prey capture success, so it cannot be
(Cansse et al. 2024), observed a higher proportion excluded that more time foraging indicates a higher
of pelagic dives in 2022, potentially reflecting a efficiency, as found for Scopoli’s Shearwaters Calo-
change in prey distribution and/or type. These nectris diomedea (Padget et al. 2017). However,
results could indicate that where prey is present in available data for the closely related Double-crested
high abundance but under low spatial predictabil- Cormorants Phalacrocorax auratus indicate an
ity, such as pelagic prey types (River
on increase in catch per unit effort and a decrease in
et al. 2021), foraging site fidelity becomes less search time when prey density increases (Enstipp

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 1474919x, 2025, 2, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/ibi.13371 by Capes, Wiley Online Library on [12/05/2025]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
476 T. Cansse et al.

et al. 2007). Therefore, in the present study on The temporal extent of foraging site fidelity in
Black-faced Cormorants, it was assumed that a Black-faced Cormorants is not yet known. The pre-
lower dive rate and a lower proportion of time spent sent study occurred during late chick-rearing when
foraging indicate higher prey capture rates and, provisioning of offspring is assumed to be maximal.
therefore, a higher foraging efficiency. By collecting data in different parts of the breeding
Foraging site fidelity was found to affect the season or in different years of study for the same
proportion of time spent foraging in a trip and individual, it would be possible to determine the
dive rate, both decreasing as foraging site fidelity intra- and inter-annual foraging plasticity in
increased. These results indicate that more spa- Black-faced Cormorants. Data on within-breeding
tially consistent individuals are likely to be more season plasticity could help to unravel the potential
efficient foragers. This is consistent with previous effects of sudden events, such as marine heatwaves,
research linking foraging site fidelity and foraging on foraging efficiency and reproductive success
behaviour in marine predators. For example, in (Woehler & Hobday 2023). Inter-annual informa-
Australian Fur Seals Arctocephalus pusillus doriferus tion on foraging plasticity could help us to under-
individuals with a higher foraging site fidelity were stand the extent to which individuals can adapt
found to have a higher proportion of successful to changing environmental conditions, such as cli-
dives, indicating a higher foraging efficiency mate change, over longer timespans (Chambers
(Speakman et al. 2021). et al. 2011).
Additionally, in the present study, a
within-individual effect of increasing foraging site
CONCLUSION
fidelity was found with individuals increasing their
foraging efficiency (i.e. lower proportion of the In summary, as a population, Black-faced Cormo-
trip spent foraging and lower dive rate) when they rants were found to use a relatively large area for
revisited previously exploited sites compared to foraging, but individuals were highly consistent in
when they visited new or less familiar sites. This the areas they exploited. While variation in forag-
suggests that, within individuals, increased famil- ing site fidelity was observed, no measured factors
iarity of foraging sites results in beneficial foraging were found which could explain these differences.
outcomes or that individuals actively revisit loca- Importantly, modelling results indicated that
tions where they have previously been successful. increasing foraging site fidelity, both within and
Similar findings have been reported in Little Pen- between individuals, increased foraging efficiency.
guins Eudyptula minor, where foraging site fidelity This indicates that disturbances to foraging habitat
was found to be higher after trips with higher prey could have negative impacts on individuals owing
capture success (Carroll et al. 2018). to a reduction in foraging efficiency. Further inves-
Combined, the between- and within-individual tigation is required to determine plasticity in forag-
effects of foraging site fidelity suggest that a higher ing behaviour within individuals over greater
foraging efficiency is obtained by foraging in more temporal scales. Such information would enable
familiar locations. Such increased efficiency could better predictions about the effects of habitat dis-
result in higher reproductive success, as has been turbance on foraging efficiency of Black-faced
shown in European Shags Gulosus aristotelis (Mor- Cormorants.
gan et al. 2019), African Penguins Speniscus demer-
sus (Traisnel & Pichegru 2019) and Lesser
Black-backed Gulls Larus fuscus (van den Bosch We thank Natalie Petrovski, Nicolas Laroque, Saia
et al. 2019). However, a consequence of such for- Bartes, Maddie Ledwidge and Aurore Counilh for their
aging site fidelity is that negative influences on for- assistance in the field. We also thank Best Helicopters
aging site prey availability, either by natural or (Sean Best and Cameron Lang) for their logistical sup-
anthropogenic factors, will decrease foraging effi- port. We furthermore thank the two anonymous
reviewers and the editorial team at Ibis for their com-
ciency and, consequently, reproductive success ments which significantly improved the quality of the
may decline (Merkle et al. 2022). Thereby, if prey manuscript. Open access publishing facilitated by Dea-
distributions shift, and no alternative prey is avail- kin University, as part of the Wiley - Deakin University
able, foraging sites might become unsuitable for agreement via the Council of Australian University
foraging. Librarians. [Correction added on 21 January 2025, after

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Foraging consistency in Black-faced Cormorants 477

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480 T. Cansse et al.

Figure S1. Violin plot for the mean FSFI for all Table S4. Results from the models examining
Black-Faced Cormorants in this study. the factors influencing foraging proportion and
Table S1. Example of calculation of FSFI and dive rate (m h1). These models used data which
DFSFI from trip overlaps obtained with the Bhatta- were classified with the Hidden Markov Model at
charyya Affinity Index. a 1-min interval.
Table S2. Accuracy of the Hidden Markov Table S5. Candidate models (DAICc < 4) for
Models for the different years of study for data which model averaging was applied for proportion
resampled at a 20-min interval. of time foraging and dive rate.
Table S3. Accuracy of the Hidden Markov
Models for the different years of study for data
resampled at a 10-min interval.

© 2024 The Author(s). IBIS published by John Wiley & Sons Ltd on behalf of British Ornithologists' Union.