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Received: 21 February 2017    Accepted: 5 November 2017

DOI: 10.1111/jai.13553

ORIGINAL ARTICLE

Site fidelity of intertidal fish to rockpools

J. Roma1  | M. Dias1  | C. Vinagre1  | A. C. F. Silva2

1
MARE – Marine and Environmental
Sciences Centre, Faculdade de Ciências da
Universidade de Lisboa, Lisboa, Portugal
2
CERIS, Instituto Superior
Técnico, Universidade de Lisboa, Lisboa,
Portugal
Correspondence
Joana Roma, MARE – Marine and
Environmental Sciences Centre, Faculdade de
Ciências da Universidade de Lisboa, Lisboa,
Portugal.
Email: joana.roma.isa@gmail.com
Funding information
Fundação para a Ciência e Tecnologia, Grant/
Award Number: UID/MAR/04292/2013
Summary
Gobius paganellus, Lipophrys pholis and Coryphoblennius galerita are wide-­spread inter-
tidal fish that spend their earlier life stages in rock pools, and yet very little is known
about their site fidelity behaviour. For these species, fidelity to rockpools may result in
increased fitness costs in a predicted scenario of warmer sea water, due to the low
thermal inertia of these water bodies. In this context, it is relevant to investigate these
species’ site fidelity. We made a mark-­recapture study to assess the mentioned spe-
cies’ movements within and between rockpools. We tagged a total of 530 individuals
of the aforementioned species with the Visible Implant Elastomer and tracked their
movement for a 7-­month period. We found that site fidelity and specific rockpools
conditions are important factors in distribution of intertidal blennies and gobies. We
also examined the relations between rockpool volume, depth and site fidelity. We
found that G. paganellus tends to remain in its original marking pool, with an average
recapture rate of 20.5%, but showed no evidence of inter-­pool movement. Rockpool
depth, however, proved to be important in the blennies’ movements. Our findings are
among the first to prove that a mark-­recapture method can be successfully used to
track intertidal fish movements. In particular, we showed that G. paganellus presents
site fidelity in intertidal rockpools during its early ontogeny for a period of two to three
months.

1 |  INTRODUCTION
One of the greatest challenges that researchers currently face is under-
standing the impact of global warming over biodiversity (Schwenk,
Padilla, Bakken, & Full, 2009). It is predicted that extreme climatic
events such as heat waves will increasingly become more frequent in
Europe (IPCC, 2014). The coastal habitat and in particular the inter-
tidal zone have been identified as some of the most vulnerable habi-
tats to this type of climatic pressure (Helmuth, Mieszkowska, Moore,
& Hawkins, 2006). Temperature increases have already caused shifts
in the current distribution of intertidal species (Helmuth et al., 2002).
The rocky intertidal zone is often an extremely productive habitat
(White, Hose, & Brown, 2015), subject daily to severe changes on its
physical, biological and chemical features due to its water/land inter-
face. Within this shifting zone rockpools’ temperatures vary widely
due to their relative little thermal inertia, in turn, related to the small
water volume and prolonged exposure to low-­tide conditions (Morris
& Taylor, 1983; Vinagre et al., 2013, 2016). Despite their ubiquity,
little is currently known about how the rockpools’ inhabitants inter-
act with their physical setting, both in ordinary and extreme thermal
conditions, such as in the event of heat waves due to climate change.
To correctly predict changes in the patterns of rockpool usage by its
inhabitants when faced with thermal variations, it is first necessary
to determine the still largely unknown standard behavioural patterns.
Hence, we aimed to investigate the habitat distribution behaviour of
the common intertidal gobies and blennies in the Portuguese intertidal
rockpools.
Our targeted species—Gobius paganellus, Lipophrys pholis and
Coryphoblennius galerita—are all commonly found at the intertidal
rockpools in the Eastern Atlantic coasts (Bath, 1990; Miller, 1990;
Zander, 1986), and are important prey for commercially import-
ant predator fish species (Patzner, Van Tassell, Kovačić, & Kapoor,
2011). However, much is unknown about these species’ habitat
distribution including among others, whether or not they pres-
ent any form of site fidelity, choose to move around the habitat
and/or keep coming back to it with a certain frequency (homing

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2       ROMA et al.
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F I G U R E   1   Representation of the sampled rockpools’ (rectangles of varied sizes numbered P1 through P12) location in relation to each other.
Maximum length and width are represented within the rockpools in the sampling stations (a) Cabo Raso 1, (b) Cabo Raso 2 and (c) Raio Verde 1.
behaviour) or, if they abandon their current habitat altogether at
each high tide. Available studies on the matter focused on the
distribution behaviour of other species and habitats, particularly
in coastal (Shima, McNaughtan, Geange, & Wilkinson, 2012) and
estuarine areas (Jud & Layman, 2012). In those cases, the distri-
bution behaviour was influenced by water depth, water body vol-
ume, sediment type, light penetration and the presence of aquatic
macrophytes (Connolly, 1994; Griffiths, 2001; Whitfield, 1999).
Supported by this information, and because rockpool volume and
depth are also known to strongly influence the rockpool commu-
nity structure (Martins, Hawkins, Thompson, & Jenkins, 2007), we
focused on these two factors as potentially important environmen-
tal conditions for fish distribution.
We tested the following hypothesis for each of the species sampled
in this study: (i) the species does not present rockpool fidelity and this is
independent of time; (ii) if present, the scale of the species’ movement
range between rockpools is small (<5 m) and; (iii) rockpool depth and vol-
ume contribute to explain the observed movement patterns.
2 | MATERIALS AND METHODS
2.1 | Sampling sites and techniques
Sampling was performed at three rocky beaches in the Portuguese
West coast (38º42′N, 9º28′W and 38°42′N, 9°29′W). Twelve rock-
pools (P1–P12) were sampled monthly and bimonthly (depending on
rockpool access due to tidal variation), during a 7-­month (July 2012 to
January 2013) period (Figure 1).
The rockpool average area was 4.66 ± 1.28 m2 (range: 0.79–
16.90 m2) and medium depth was 0.26 ± 0.2 m (range: 0.18–0.40 m).
The pools were categorized accordingly to: A. volume: volume 1—
below 0.8 m3 (7 pools: P5, P6, P8, P9, P10, P11 and P12) and volume
2—above 1.0 m3 (5 pools: P1, P2, P3, P4 and P7) and; B. depth: depth
1—below 0.3 m (5 pools: P7, P9, P10, P11 and P12) and depth 2—
above 0.3 m (7 pools: P1, P2, P3, P4, P5, P6 and P8).
We chose the mark-­recapture technique for this study as it is one
of the most widely accepted sampling methodologies for the distribu-
tion studies of small fishes (Imbert, Beaulaton, Rigaud, & Elie, 2007).
The visible implant elastomer (VIE) was chosen as the internal tag as it
causes little to no damage or disturbance to the living organism, espe-
cially to such small sample subjects (some were as small as 2 cm or less).
This implant tag has been successfully used for rockpool fishes in Knope,
Tice, and Rypkema (2017). The individuals were caught in rockpools with
hand-­held nets, tagged with VIE by injecting a hypodermic syringe just
above the left lateral line and their length, location and species registered,
followed by immediate release into their original rockpool. The target ani-
mals were tagged for each pool, and subsequently the marked individuals
were examined for recaptures on a monthly or bi-­monthly basis. Each
pool was assigned a different colour.
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2.2 | Statistical analysis
The following independent variables and experimental design were
used to test for significant differences in the response variables: rock-
pool volume (fixed, two levels: below 0.8 m3 and above 1.0 m3) and
rockpool depth (fixed, two levels: below 0.3 m and above 0.3 m). Three
response variables were assessed: (i) average number of recaptures, (ii)
average size of all marked individuals and, (iii) average size of all recap-
tured ­individuals. These were chosen to examine the influence of rock-
pool characteristics on the movement patterns and population structure
associated with the movements. A PERMANOVA analysis was used to
assess the hypotheses stated in the introduction. Prior to analysis, the
data were fourth root transformed in order to obtain a more balanced
weight distribution between variables. The number of used permutations
was 9,999 and the similarity measure used was the Euclidean distance.
Depending on the PERMANOVA results, we performed a
SIMPER similarity analysis to identify which response variable had
the most contribution for the significant differences found in the
PERMANOVA analysis. We also made a multi-­dimensional scal-
ing (MDS) analysis to obtain a visual ordination of the differences
between samples.
3 | RESULTS
In total, 530 fish were tagged in this study, of which 83.2% were G.
paganellus (441), 9.4% were L. pholis (50) and 7.4% were C. galerita
(39).
The total number of tagged animals for volume 1 pools (<0.8 m3)
was 153, and for volume 2 (>1.0 m3) was 377. The total number of
tagged animals for depth 1 pools (<0.3 m) was 196, and for depth 2
(>0.3 m) was 334.
3.1 | Gobius paganellus
The highest number of tagged individuals in a single rockpool was 107
and the lowest was 12. With the exception of a single rockpool, G.
paganellus was recaptured at least once in all 12 sampled rockpools.
The average recapture rate across all rockpools was 20.5%. The highest
recapture number in a single event was 44 individuals in P2, although
the highest average percentage (of all recapture events of previously
marked individuals in a single pool) was 35.8% in P4 (Figure 2). No
marked G. paganellus were recaptured after the 4th recapture event,
4 months after the tagging time. The average recapture rate in pool
volume <0.8 m3 was 20.8% and 21.1% for pools >1.0 m3. The aver-
age recapture rate of pools with depth <0.3 m was 23.1% and 19.3%
for those with depth >0.3 m. Concerning the recaptures rates within
the same pool where individuals were initially tagged (henceforth,
“home pool”), the average size of marked individuals (3.6 ± 1.0 cm,
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4       ROMA et al.
F I G U R E   2   Overview of Gobius paganellus recaptured tagged individuals, represented as the average percentage of recaptured individuals in
all recapture occasions, per sampled rockpool (P1 through P12).
range 2–9.5 cm) was smaller than that of individuals later recaptured
(4.5 ± 1.4 cm, range 2.7–9.9 cm).
As for external recaptures, i.e., the number of marked individuals
found outside their home pool, only one rockpool (P5) registered 8.9%
of its tagged individuals in other pools.
3.2 | Lipophrys pholis
In total, 50 L. pholis individuals were tagged in all 12 pools, but only
five individuals were recaptured afterwards in four of those pools. No
tagged L. pholis were further recaptured after the 1st recapture event
(within the same month of tagging). The average size of individuals
when they were first captured (5.1 ± 1.5 cm, range 1.9–11.3 cm)
was larger than that of individuals recaptured in later occasions
(4.4 ± 2.2 cm, range 3.9–10.5 cm). Concerning external recaptures,
only one rockpool registered one of its individuals in other pools, rep-
resenting 20% of its total marked individuals.
3.3 | Coryphoblennius galerita
A total of 39 C. galerita individuals were tagged in all 12 pools, but only
six individuals were recaptured afterwards in five of those pools. No
marked individuals were further recaptured after the 1st recapture
event (within the same month of tagging). The average size of marked
individuals (5.0 ± 1.6 cm, range 3–11 cm) was smaller than individuals
later recaptured (6.8 ± 3.0 cm, range 4.2–10.5 cm). No external recap-
tures were registered.
There were no significant differences between rockpool volumes
(Table 1a) for the tested parameters (average number of recaptured
individuals, average size of marked individuals and average size of
recaptured individuals). However, the same parameters varied signifi-
cantly with rockpool depth (Table 1b and Figure 3). The SIMPER anal-
ysis showed that all three analysed parameters were greater at deeper
rockpools, i.e., rockpools ≥0.3 m.
4 | DISCUSSION
Our study was the first of its kind to use this mark-­recapture meth-
ods applied to the distribution of intertidal fish species in rockpools.
The main hypotheses of this study were successfully addressed by
the methodology. However, it was not possible to validate our sec-
ond hypothesis (the scale of its movement range between rockpools
is smaller than 5 m) for all studied species since there were very few
recaptured individuals that were caught outside their home pools, and
hence no accurate conclusion could be derived from our data.
Next, the specific results obtained for each species will be dis-
cussed, addressing the remaining hypothesis: (i) it does not have rock-
pool fidelity and this is independent of time; and (ii) rockpool depth
and volume contribute to explain the movement patterns.
4.1 | Gobius paganellus
Gobius paganellus generally remained in the same rockpool, suggesting
a site fidelity behaviour and avoidance to cross into nearby pools. This
fidelity behaviour is potentially associated with: (i) increased predator
refuge conditions, such as comparatively higher algae canopy cover
or increased crevice availability (Baeza & Thiel, 2007; Baeza, Farías,
Luppi, & Spivak, 2010); (ii) territorial and food defence behaviours
(Baeza & Thiel, 2003, 2007; Emlen & Oring, 1977) and; (iii) preference
ROMA et al. |
      5

T A B L E   1   Results of PERMANOVA
Source df SS MS Pseudo-­F P(perm) Unique perms
analysis on Coryphoblennius galerita
according to the following experimental (a)
design: (a) rockpool volume (fixed, two Volume 1 3.8786 3.8786 1.3103 0.3404 512
levels: below 0.8 m3 and above 1.0 m3) and
Residuals 22 65.121 2.9601
(b) rockpool depth (fixed, two levels: below
0.3 m and above 0.3 m) Total 23 69
(b)
Depth 1 13.138 13.138 5.1741 0.0101 512
Residuals 22 55.862 2.5392
Total 23 69

F I G U R E   3   MDS analysis of the

analysed Coryphoblennius galerita variables
(average number of recaptured individuals,
average size of marked individuals and
average size of recaptured individuals) at
each rockpool and analysed by respective
depth.

for a habitat that individuals were previously familiar with (Carlson,
2005).
Mostly, only juveniles of G. paganellus (i.e., fish <7 cm, the average
size at first maturity) were collected. During the early winter sampling
period almost no individuals were found in the sampling areas. This
absence may be explained by high mortality rates in earlier ontogenic
stages, or by simply migrating outside the sampling area due to a bio-
logical driver, such as reproductive behaviour. The reproductive sea-
son occurs between January and June (Azevedo & Simas, 2000), hence
potentially explaining the absence of tagged individuals. Gibson (1972,
1999) also proposes that this species favours rockpools in the lower
intertidal zone. However, no movements between different areas of
the intertidal zone or even to the subtidal have yet been documented.
Site fidelity behaviour can lead to the prediction that when faced
with global warming effects, G. paganellus living in rockpools may
face challenging if not adverse conditions, potentially affecting their
physiology and ecology. While during the sampling days of the study
rockpool water temperatures never surpassed 28°C in the shallowest
rockpools. If these same temperatures rise as high as 33.1°C (previ-
ously verified as this species’ thermal tolerance limit) (Vinagre et al.,
2013), this can lead to thermal stress or even desiccation, compro-
mising the survival of this species (Brierley & Kingsford, 2009). In
southern Europe where our study was located, days with tempera-
tures above 35°C are expected to increase to more than 40 days per
year (Miranda, Coelho, Tomé, & Valente, 2002). Within this context
and among the IPCC temperature variation prediction models, one
already predicts a temperature increment as high as 4.8°C within this
century alone (IPCC, 2014). Considering that this species is known to
exclusively inhabit the intertidal habitat, and if this becomes inhospi-
table due to high temperatures, the fish may migrate into habitats with
lower temperatures. This would ensure future survival in more resilient
habitats, such as deeper rockpools lower in the intertidal zone or even
channels found in this type of habitat (Faria & Almada, 2001a, 2001b;
Vinagre et al., 2013).
4.2 | Lipophrys pholis and Coryphoblennius galerita
We did not acquire enough data (i.e., sampled individuals) of these
species to be able to ascertain if they display any site fidelity or inter-­
rockpool behaviours. Few individuals were caught and tagged and
even less were later recaptured. While L. pholis individuals were all
juveniles (adult size ~16 cm), we found adult C. galerita measuring up
to 9.5 cm.
Around October and at 1 year of age, L. pholis is known to aban-
don intertidal rockpools in favour of near-­subtidal rock fissures and
corridors (Faria & Almada, 2001a, 2001b; Faria, Almada, & Gonçalves,
1996; Gibson, 1972), potentially explaining its disappearance from
our study area around the same time. We could find no evidence
in literature that attested the same for C. galerita, however Nieder
(1993) found that C. galerita apparently exhibits both site fidelity
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6       ROMA et al.
and movement around the same rockpool for only 2 months, which
might explain its apparently premature disappearance from the
study site. Almada, Dores, Pinheiro, Pinheiro, and Santos (1983) ver-
ified that both blennies possibly show site fidelity not to a single
rockpool, but rather to a group of rockpools. Later, Jorge, Almada,
Gonçalves, Duarte-­Coelho, and Almada (2012) in a laboratory set-
ting, verified a homing behaviour in adult L. pholis individuals which
is a behaviour often associated with site fidelity, albeit not a con-
firmation. Altogether, these studies indicate that some type of site
fidelity could be shown by C. galerita.
However, we successfully proved that C. galerita seems to prefer
deeper rockpools, a choice also verified by Nieder (1993) who observed
more blennies in deeper pools and a theory previously confirmed by
Martins et al. (2007). Deeper pools may offer different prey availability
and additional predator refuge opportunity due to larger algae canopy
cover, which are characteristics that fishes displaying homing and site
fidelity behaviours often seem to favour (e.g., Abecasis, Bentes, & Erzini,
2009; Bellquist, Lowe, & Caselle, 2008; Harasti, Lee, Gallen, Hughes, &
Stewart, 2015; Mason & Lowe, 2010; March, Alós, Grau, & Palmer, 2011).
In conclusion, the study successfully showed that intertidal fish
species can display site fidelity behaviours to rockpools, and that pool
characteristics may strongly influence their movement choices. Future
studies using individual tagging procedures would enable a more com-
prehensive assessment of intertidal fish movements.
ACKNOWLE DG E MEN TS
We would like to thank all those involved in the fieldwork. This work
was funded by the “Fundação para a Ciência e Tecnologia”, Portugal,
through the researcher position awarded to C. Vinagre and the strate-
gic project UID/MAR/04292/2013 granted to MARE.
O RCI D
J. Roma  http://orcid.org/0000-0002-5884-4623
M. Dias  http://orcid.org/0000-0003-0447-6009
C. Vinagre  http://orcid.org/0000-0003-2146-7948
A. C. F. Silva  http://orcid.org/0000-0003-4734-6538
REFERENCES
Abecasis, D., Bentes, L., & Erzini, K. (2009). Home range, residency and
movements of Diplodus sargus and Diplodus vulgaris in a coastal lagoon:
Connectivity between nursery and adult habitats. Estuarine, Coastal and
Shelf Science, 85, 525–529.
Almada, V. C., Dores, J., Pinheiro, A., Pinheiro, M., & Santos, R. S. (1983).
Contribuição para o estudo do comportamento de Coryphoblennius
galerita (L.) (Pisces, Blenniidae). Memórias do Museu do Mar: Série
Zoológica, 2, 166.
Azevedo, J. M. N., & Simas, A. M. V. (2000). Age and growth, reproduction
and diet of sublittoral population of the rock goby Gobius paganellus
(Teleostei: Gobiidae). Hydrobiologia, 440, 129–135.
Baeza, J. A., Farías, N. E., Luppi, T. A., & Spivak, E. D. (2010). Refuge size, group
living and symbiosis: Testing the “resource economic monopolization”
hypothesis with the prawn Betaeus lilianae and description of its part-
nership with the crab Platyxanthus crenulatus. Journal of Experimental
Marine Biology and Ecology, 389, 85–92.
Baeza, J. A., & Thiel, M. (2003). Predicting territorial behavior in symbiotic
crabs using host characteristics: A comparative study and proposal of a
model. Marine Biology, 142, 93–100.
Baeza, J. A., & Thiel, M. (2007). The mating system of symbiotic crusta-
ceans. A conceptual model based on optimality and ecological con-
straints. In J. E. Duffy, & M. Thiel (Eds.), Reproductive and social behavior:
Crustaceans as model systems (pp. 245–255). Oxford: Oxford University
Press.
Bath, H. (1990). Blenniidae. In J. C. Quero, J. C. Hureau, C. Karrer, A.
Post, & L. Saldanha (Eds.), Check-list of the fishes of the eastern tropi-
cal Atlantic (CLOFETA) (Vol. 2, pp. 905–915). Lisbon, Paris: JNICT/SEI/
UNESCO.
Bellquist, L. F., Lowe, C. G., & Caselle, J. E. (2008). Fine-­scale movement pat-
terns, site fidelity, and habitat selection of ocean whitefish (Caulolatilus
princeps). Fisheries Research, 91, 325–335.
Brierley, A. S., & Kingsford, M. J. (2009). Impacts of climate change on ma-
rine organisms and ecosystems. Current Biology, 19, R602–R614.
Carlson, P. (2005). Homing, habitat fidelity and kin recognition in Eurasian
perch (Perca fluviatilis) (Master thesis dissertation), Uppsala University,
Sweden.
Connolly, R. M. (1994). Removal of seagrass canopy: Effects on small fish
and their prey. Journal of Experimental Marine Biology and Ecology, 184,
99–110.
Emlen, S. T., & Oring, L. W. (1977). Ecology, sexual selection and the evolu-
tion of mating systems. Science, 197, 215–223.
Faria, C., & Almada, V. (2001a). Agonistic behaviour and control of ac-
cess to hiding places in two intertidal blennies, Lipophrys pholis and
Coryphoblennius galerita (Pisces: Blenniidae). Acta Ethologica, 4, 51–58.
Faria, C., & Almada, V. (2001b). Microhabitat segregation in three rocky
intertidal fish species in Portugal: Does it reflect interspecific competi-
tion? Journal of Fish Biology, 58, 145–159.
Faria, C., Almada, V. C., & Gonçalves, E. J. (1996). Juvenile recruitment,
growth and maturation of Lipophrys pholis (Pisces: Blenniidae), from the
west coast of Portugal. Journal of Fish Biology, 49, 727–730.
Gibson, R. N. (1972). The vertical distribution and feeding relationships of
intertidal fish on the Atlantic cost of France. Journal of Animal Ecology,
41, 189–207.
Gibson, R. N. (1999). Movement and homing in intertidal fishes. In M. H.
Horn, K. L. M. Martin, & M. A. Chotkowski (Eds.), Intertidal fishes: Life in
two worlds (pp. 97–125). San Diego, CA: Academic Press.
Griffiths, S. P. (2001). Factors influencing fish composition in an Australian
intermittently open estuary. Is stability salinity-­dependent? Estuarine,
Coastal and Shelf Science, 52, 739–751.
Harasti, D., Lee, K. A., Gallen, C., Hughes, J. M., & Stewart, J. (2015).
Movements, home range and site fidelity of snapper (Chrysophrys
auratus) within a temperate marine protected area. PLoS ONE, 10,
e0142454.
Helmuth, B., Harley, C. D. G., Halpin, P. M., O’Donnell, M., Hofmann, G. E.,
& Blanchette, C. A. (2002). Climate change and latitudinal patterns of
intertidal thermal stress. Science, 298, 1015–1017.
Helmuth, B., Mieszkowska, N., Moore, P., & Hawkins, J. (2006). Living on
the edge of two changing worlds: Forecasting the responses of rocky
intertidal ecosystems to climate change. Annual Review of Ecology,
Evolution, and Systematics, 37, 373–404.
Imbert, H., Beaulaton, L., Rigaud, C., & Elie, P. (2007). Evaluation of visible
implant elastomer as a method for tagging small European eels. Journal
of Fish Biology, 71, 1546–1554.
IPCC (2014). The regional impacts of climate change: An assessment of vulner-
ability. UK: Cambridge University Press.
Jorge, P. E., Almada, F., Gonçalves, A. R., Duarte-Coelho, P., & Almada, V. C.
(2012). Homing in rocky intertidal fish. Are Lipophrys pholis L. able to
perform true navigation? Animal Cognition, 15, 1173–1181.
ROMA et al.
Jud, Z., & Layman, C. A. (2012). Site fidelity and movement patterns of in-
vasive lionfish, Pterois spp., in a Florida estuary. Journal of Experimental
Marine Biology and Ecology, 414–415, 69–74.
Knope, M. L., Tice, K. A., & Rypkema, D. C. (2017). Site fidelity and hom-
ing behaviour of intertidal sculpins revisited. Journal of Fish Biology, 90,
341–355.
March, D., Alós, J., Grau, A., & Palmer, M. (2011). Short-­term residence
and movement patterns of the annular seabream Diplodus annularis
in a temperate marine reserve. Estuarine, Coastal and Shelf Science, 92,
581–587.
Martins, G. M., Hawkins, S. J., Thompson, R. C., & Jenkins, S. R. (2007).
Community structure and functioning in intertidal rock pools: Effects
of pool size and shore height at different successional stages. Marine
Ecology Progress Series, 329, 43–55.
Mason, T. J., & Lowe, C. G. (2010). Home range, habitat use, and site fidelity
of barred sand bass within a southern California marine protected area.
Fisheries Research, 106, 93–101.
Miller, P. J. (1990). Gobiidae. In J. C. Quero, J. C. Hureau, C. Karrer, A.
Post, & L. Saldanha (Eds.), Check-list of the fishes of the eastern tropical
Atlantic (CLOFETA) (Vol. 2, pp. 925–951). Lisbon, Paris: JNICT/SEI/
UNESCO.
Miranda, P. M. A., Coelho, F. E. S., Tomé, A. R., & Valente, M. A. (2002). 20th
century Portuguese climate and climate scenarios. Scenarios, impacts
and adaptation measures – SIAM project. In: F.D. Santos, K. Forbes &
Moita R (Eds.), Climate change in Portugal: Scenarios, impacts and adapta-
tion measures (SIAM Project). Lisbon, Portugal: Gradiva, 454 pp.
Morris, S., & Taylor, A. C. (1983). Diurnal and seasonal variation in physico-­
chemical conditions within intertidal rock pools. Estuarine, Coastal and
Shelf Science, 17, 339–355.
Nieder, J. (1993). Distribution of juvenile blennies (Coryphoblennius galer-
ita and Lipophrys pholis, Pisces, Blenniidae) in small tide-­pools on the
Algarve coast of southern Portugal: Result of low-­tide lottery or strate-
gic habitat selection? Bonner Zoologische Beiträge, 44, 133–140.
|
      7
Patzner, R. A., Van Tassell, J. L., Kovačić, M., & Kapoor, B. G. (Eds.) (2011).
The biology of gobies (p. 685). Enfield, NH: Science Publishers.
Schwenk, K., Padilla, D. K., Bakken, G. S., & Full, R. J. (2009). Grand chal-
lenges in organismal biology. Integrative and Comparative Biology, 49,
7–14.
Shima, J. S., McNaughtan, D., Geange, S. W., & Wilkinson, S. (2012).
Ontogenetic variation in site fidelity and homing behaviour of a tem-
perate reef fish. Journal of Experimental Marine Biology and Ecology,
416–417, 162–167.
Vinagre, C., Dias, M., Roma, J., Silva, A., Madeira, D., & Diniz, M. S. (2013).
Critical thermal maxima of common rocky intertidal fish and shrimps –
A preliminary assessment. Journal of Sea Research, 81, 10–12.
Vinagre, C., Leal, I., Mendonça, V., Madeira, D., Narciso, L., Diniz, M. S., &
Flores, A. A. V. (2016). Vulnerability to climate warming and acclima-
tion capacity of tropical and temperate coastal organisms. Ecological
Indicators, 62, 317–327.
White, G. E., Hose, G. C., & Brown, C. (2015). Influence of rock-­pool charac-
teristics on the distribution and abundance of inter-­tidal fishes. Marine
Ecology, 36, 1332–1344.
Whitfield, A. K. (1999). Ichthyofaunal assemblages in estuaries: A South
African case study. Reviews in Fish Biology and Fisheries, 9, 151–186.
Zander, C. D. (1986). Blenniidae. In P. J. P. Whitehead, M.-L. Bauchot, J.-
C. Hureau, J. Nielsen, & E. Tortonese (Eds.), Fishes of the north-eastern
Atlantic and the Mediterranean (pp. 1096–1112). Paris: UNESCO.
How to cite this article: Roma J, Dias M, Vinagre C, Silva ACF.
Site fidelity of intertidal fish to rockpools. J Appl Ichthyol.
2017;00:1–7. https://doi.org/10.1111/jai.13553