Marine Biodiversity Records, page 1 of 7.

# Marine Biological Association of the United Kingdom, 2014

doi:10.1017/S175526721400089X; Vol. 7; e97; 2014 Published online

Potential hybridism between free-ranging

Risso’s dolphins (Grampus griseus)
and bottlenose dolphins (Tursiops truncatus)
off north-east Lewis (Hebrides, UK)
nicola k. hodgins1, sarah j. dolman1 and caroline r. weir2,3
1
Whale and Dolphin Conservation, 38 St Paul Street, Chippenham, Wiltshire SN15 1LJ, UK, 2Ketos Ecology, 4 Compton Road,
Kingsbridge, Devon TQ7 2BP, UK, 3School of Biological Sciences (Zoology), University of Aberdeen, Tillydrone Avenue, Aberdeen,
AB24 2TZ, UK

Hybrid cetaceans have been documented to occur both in the wild and in captivity. Identifying wild hybrid individuals can be
problematic in the absence of genetic techniques, but published accounts indicate that intermediate morphological character-
istics are often present. Between 2010 and 2013, a land-based and boat-based study of the Risso’s dolphin (Grampus griseus)
was carried out in nearshore waters around the Eye Peninsula located on north-east Lewis, Scotland. Three atypical indivi-
duals were photographed which exhibited morphological features intermediate between Grampus and the common bottlenose
dolphin (Tursiops truncatus). These individuals were typically larger in body size than Tursiops, and had a dorsal fin shape
and size consistent with Grampus. Two individuals had coloration most similar to Tursiops and the third exhibited extensive
white linear scarring consistent with Grampus. The intermediate morphology was most apparent in the head shape, with all
three individuals exhibiting a defined (in contrast to Grampus) but very short (compared with Tursiops) rostrum and two
having an unusually steep (compared with Tursiops) forehead. On one occasion, one of the atypical individuals was observed
within a mixed-species school of Grampus and Tursiops. There were four further sightings of atypical dolphins associated
with Tursiops-only schools. Atypical dolphins were not recorded within Grampus-only schools. These observations are con-
sistent with hybridization between free-ranging Risso’s and bottlenose dolphins, the first such occurrence to be documented for
these species in UK waters. The context and significance of these hybridization events are unknown.

Keywords: cetacean, odontocete, hybrid, morphology, species association, interspecific mating, Atlantic Ocean, photo-identification,
Grampus griseus, Tursiops truncatus

Submitted 11 February 2014; accepted 27 July 2014

INTRODUCTION recently since 2010 (this study). Worldwide the Risso’s

dolphin preferentially occurs over topography with steep-
The north-east Atlantic waters to the north-west of Scotland sloped bottoms near the outer edge of the continental shelf
are inhabited by a diversity of baleen and toothed whale or upper slope (Jefferson et al., 2014), and its occurrence in
species including the Risso’s dolphin (Grampus griseus, nearshore shallow waters around Lewis is therefore less
Cuvier, 1812) and the common bottlenose dolphin (Tursiops typical.
truncatus, Gervais, 1855) (hereafter referred to as ‘bottlenose Risso’s and bottlenose dolphins are seen throughout
dolphin’). Both of these species occur through a range of habi- the Hebrides, but bottlenose dolphins are more commonly
tats in this geographical region, including deep waters along recorded from The Little Minch southwards (particularly
and seaward of the shelf break and nearshore shallow waters around Mull, Coll, Tiree, and Barra) while the Risso’s
(Weir et al., 2001; Reid et al., 2003). dolphin is more frequently observed in The Minch and
The regular occurrence of Risso’s dolphins in the shallow along the Atlantic seaboard (HWDT, 2014). Studies in the
coastal waters of the Minch (located between Lewis and main- area in the 1990s never observed bottlenose dolphins
land Scotland: Figure 1) has been well-documented (Evans around the Eye Peninsula (Alison Gill, personal communica-
et al., 1993; Atkinson et al., 1998; Weir et al., 2001; Reid tion). However, recent survey effort (between 2010 and 2013)
et al., 2003; Dolman & Hodgins, 2013). This species was the in the same area has recorded bottlenose dolphins in 2011 and
focus of dedicated studies around the Eye Peninsula on 2012, including some mixed-species associations with the
Lewis during the 1990s (Atkinson et al., 1998) and more Risso’s dolphin (WDC, unpublished data).
The occurrence of mixed-species associations in sympatric
cetacean species provides opportunities for interspecific sexual
Corresponding author:
interaction and the potential for hybridization (Bérubé, 2009).
N.K. Hodgins Although infrequently observed, hybrids between various
Email: [email protected] odontocete species have been recorded, both in the wild and

1
2 nicola k. et al.

Fig. 1. Location of the Eye Peninsula study area (north-east Lewis, UK) and places mentioned in the text. The inset shows the location of the shore-based study site
from which four sightings were recorded (black triangle) and the locations of three boat-based sightings (grey squares) (see Table 1).

in captivity (Sylvestre & Tasaka, 1985; Reyes, 1996; Herzing & 2011 and 2012 during cetacean fieldwork off Lewis, north-
Johnson, 1997; Karczmarski, 1997; Herzing et al., 2003; west Scotland. We provide evidence to suggest that these indi-
Bérubé, 2009). Suspected wild cetacean hybrids are typically viduals may be the result of hybridism between wild Risso’s
observed without prior knowledge of parental interactions. and bottlenose dolphins.
Therefore their detection and identification in the field is
problematic and the number of documented incidences in
the wild to date is limited (for a review, see Bérubé, 2009).
Collecting genetic evidence to verify the occurrence of live MATERIALS AND METHODS
wild hybrid dolphins is logistically difficult, and has associated
welfare considerations (biopsying wild cetaceans is intrusive This fieldwork utilized a combination of land- and boat-based
and potentially risky; e.g. Bearzi, 2000). However, morpho- methods as briefly described below.
logical analyses of captive-born hybrid dolphins indicate Land-based surveys were conducted from a fixed land-
that the offspring consistently exhibit intermediate character- based site (approximately 50 m above sea level) at Tiumpan
istics of both parent species (Sylvestre & Tasaka, 1985; Head, on the Isle of Lewis (Figure 1). Data were collected
Zornetzer & Duffield, 2003). Consequently, the occurrence using a standardized scan sampling protocol (Pierpoint
of individuals exhibiting intermediate morphological et al., 1998), during which a single observer slowly scanned
characteristics in the field may represent a valid method of across a predefined sector of sea using 7 × 50 reticulated bin-
identifying wild-born hybrids. oculars with built-in compass. Whenever cetaceans were
This paper describes three ‘atypical’ dolphins with mixed sighted, standardized data were logged including species,
morphological characteristics that were photographed in group size, behaviour, distance from shore and direction of
 potential hybridism between two species of dolphin 3

Table 1. Survey effort (in sea state ≤3) and sightings of Risso’s (Grampus RESULTS
griseus) and bottlenose (Tursiops truncatus) dolphins during field survey
work at north-east Lewis between 2010 and 2013. A total of 26 groups of Risso’s dolphins, six groups of bottle-
Year Survey effort (h) Number of sightings
nose dolphins and three mixed-species groups were recorded
off north-east Lewis between 2010 and 2013 (Table 1). Three
individual ‘atypical’ dolphins (ATD1 –3) were photo-
Shore- Boat- Grampus Tursiops Mixed Grampus identified during 2011 and 2012 (Table 2), the only years in
based based griseus truncatus and Tursiops which bottlenose dolphins were recorded. All of the sightings
2010 48.6 26.6 14 0 0
of atypical animals occurred in close proximity to Tiumpan
2011∗ 16.5 39.1 7 1 1 Head (Figure 1).
2012 15.4 30.8 2 5 2 Two of the atypical dolphins were photographed on single
2013 0 14.6 3 0 0 occasions: ATD1 (Figure 2A) within a mixed-species school of
Risso’s and bottlenose dolphins in 2011 and ATD3
∗
, excludes one sighting comprising an atypical dolphin (ATD2) seen trav- (Figure 2C) with five bottlenose dolphins during 2012.
elling with a dolphin of uncertain identity (IND4). ATD2 (Figure 2B) was photographed in two sightings in
2011 and two sightings in 2012, once together with an
travel. A total of 80.4 h of land-based survey effort (sea animal of uncertain species (IND4 (Figure 2D): see below)
state ≤3) was carried out between 2010 and 2012 (Table 1). and three times in association with schools of bottlenose dol-
Vessel-based surveys were focused primarily on photo- phins (including one further occasion where IND4 was
identification methods, carried out under licence and there- present) (Table 2).
fore designed to maximize encounters with dolphins. One atypical dolphin was observed within a mixed-species
Different vessels were used in each year. In 2010, the survey school of Risso’s and bottlenose dolphins. With the exception
vessel was ‘MV Puffin’, approximately 5 m motor boat. In of sighting number two (uncertain group composition), the
2011, the survey vessel was ‘MV Fish n’ Trips’, a 6 m motor remaining sightings all involved atypical animals travelling
boat and in 2012 a 6.5 m rigid inflatable boat (RIB) also within bottlenose-only schools (Table 2). Atypical dolphins
named ‘MV Fish n’ Trips’ was used. were never recorded within a Risso’s-only school.
The boat survey route depended on prevailing weather The morphological characteristics of the three atypical dol-
conditions and on whether sightings had been reported phins are described below.
from land-based sites. Two observers searched continuously
from the port and starboard sides respectively and the vessel
position was continually recorded at 1 min intervals using a
handheld Garmin GMAP 76CSx GPS. Whenever cetaceans
ATD1
were seen standardized data (including position, species, This was immediately noted to be a particularly large animal,
group size and behaviour) were logged and the boat was man- with an estimated body size approximately 50% larger than
oeuvred carefully towards the animals to attempt photo- the accompanying bottlenose dolphins. The dorsal fin was
identification. A total of 111.1 h of boat-based survey effort prominent and very broad-based, being double the size of
(sea state ≤3) was carried out between 2010 and 2013 the bottlenose dolphins and slightly larger than the fins of
(Table 1). the accompanying Risso’s dolphins. The head shape was inter-
Photo-identification was attempted from both the shore mediate between Risso’s and bottlenose dolphins, with a
site (when dolphins passed sufficiently close to the cliff) and sloped forehead and a very short (cf. bottlenose) but defined
from the boat. Photographs of dorsal fins and other body rostrum (Figure 2A). Overall body coloration was a uniform
marks were taken using a Canon 7D digital SLR camera dark grey with little scarring or white lesions/patches visible;
with a 100 – 400 mm lens. The photography and subsequent however, the weather was overcast and details of body color-
cataloguing of individual dolphins was carried out by a ation were obscured. This animal was observed within a
single person (the lead author), according to standard proto- mixed-species school of bottlenose and Risso’s dolphins
cols (Würsig & Jefferson, 1990). (Table 2).

Table 2. Details of seven sightings off north-east Lewis in which three atypical dolphins (ATD), consistent with the potential hybridization between free-
ranging Risso’s (Grampus griseus) and bottlenose (Tursiops truncatus) dolphins, were photographed at north-east Lewis between 2010 and 2013.

Sighting Date Sighting Platform Group type No. of animals ATD

Number time Reference
Number
Start End Total Atypical Grampus Tursiops Unknown∗
dolphins griseus truncatus

1 24 August 2011 14:53 15:04 Land Mixed species 7 1 4 2 0 ATD1

2 25 August 2011 09:39 09:52 Land Uncertain∗ 2 1 0 0 1 ATD2
3 25 August 2011 10:13 10:17 Land Single species 9 1 0 7 1 ATD2
4 21 August 2012 17:08 18:00 Land Single species 8 1 0 7 0 ATD2
5 22 August 2012 10:21 11:27 Boat Single species 9 1 0 8 0 ATD2
6 19 September 2012 10:25 10:59 Boat Mixed species 9 0 1 7 1 N/A∗
7 4 October 2012 09:20 09:43 Boat Single species 6 1 0 5 ATD3
∗
, these records relate to a further individual (IND4) which was also suspected to be atypical.
4 nicola k. et al.

presence of three mother/calf pairs, and IND4 on one occa-

sion (Table 2), but was not observed with Risso’s dolphins.

ATD3
This animal resembles a bottlenose dolphin in overall body
size and appearance (coloration and dorsal fin shape).
However, the head shape is atypical, with a more sloped and
less-rounded melon than a bottlenose dolphin and a less dis-
tinct crease than usual (Figure 2C). The rostrum is particularly
short when compared with a typical bottlenose dolphin.

IND4
In addition, a further individual (IND4) was photographed
that was also suspected to be atypical (Figure 2D). This
animal was observed twice in association with ATD2 and
once in a mixed-species school of Risso’s and bottlenose dol-
phins (Table 2). It had similar overall appearance to a bottle-
nose dolphin, with a uniform dark body colour and areas of
lesioning. One poor-quality image also suggested the presence
of a rounded melon and a short well-defined beak. However, it
was noted to be 25% larger in body size than accompanying
adult bottlenose dolphins and with a prominent large dorsal
fin (Figure 2D). The species affiliation of this animal was
uncertain.

DISCUSSION

Opportunities for hybridism

As noted by Bérubé (2009), cetacean hybrids are most
common within genera where the different species have
similar life histories and habitat requirements. Clearly, two
species must overlap in habitat in order to provide opportun-
ities for social interaction and mating. Additionally, oppor-
tunities for interspecific mating would be greatest amongst
species that regularly form mixed-species associations.
Fig. 2. (A) Photograph of atypical dolphin 1 (ATD1) photographed off Risso’s dolphins and bottlenose dolphins are two species
north-east Lewis. ATD1 exhibits morphological characteristics consistent
which overlap in habitat in many geographical regions and
with hybridization between Risso’s and bottlenose dolphins; (B) photograph
of atypical dolphin 2 (ATD2) photographed off north-east Lewis. ATD2 are known to form mixed-species schools. The occurrence
exhibits morphological characteristics consistent with hybridization between of mixed-species schools comprising these two species have
Risso’s and bottlenose dolphins; (C) photograph of atypical dolphin 3 been reported from areas as varied as California (Bearzi,
(ATD3) photographed off north-east Lewis. ATD3 exhibits morphological 2005), the Gulf of Mexico (Maze-Foley & Mullin, 2006), the
characteristics consistent with hybridization between Risso’s and bottlenose
dolphins; (D) photograph of one suspected atypical dolphin (IND4)
west coast of Africa (Weir, 2011) and the Indian Ocean
photographed off north-east Lewis. (Ballance & Pitman, 1998). However, most of those associa-
tions have been observed in offshore, deep waters rather
than from the coastal neritic habitat documented in this
paper. This is presumably because the Risso’s dolphin is pre-
ATD2 dominantly found in habitat seaward of the shelf edge in most
This animal had similar (possibly slightly larger) body and of its geographical range (Jefferson et al., 2014). Nevertheless
dorsal fin size to a Risso’s dolphin and was estimated to be the species does occur in a wide range of habitats from the
approximately 25% larger than accompanying bottlenose dol- coast to deep oceanic waters, and the shelf waters along the
phins. It was also Risso’s-like in coloration, exhibiting a Atlantic seaboard of Britain and Ireland are one notable geo-
uniform grey base colour but with extensive white linear scar- graphical region where the species regularly inhabits relatively
ring and some white lesioning on the flanks (Figure 2B). shallow, coastal waters (Reid et al., 2003). A mixed-species
Although body size, coloration and the broad dorsal fin association between bottlenose and Risso’s dolphins has
were similar to a Risso’s dolphin, the head comprised a been recorded in very shallow, nearshore habitat on at least
rounded melon, distinct crease and well-defined short beak one previous occasion (Jefferson et al., 2014). In this paper
resembling the morphology of a bottlenose dolphin. we report three mixed-species associations between the two
Moreover, this animal was observed accompanying bottlenose species in the coastal waters off Lewis, including at least one
dolphins on three occasions, two of which involved the in which an atypical individual was present. Collectively,
 potential hybridism between two species of dolphin 5

these accounts indicate that the associations between these the colour of a bottlenose dolphin with a small and distinct
two species regularly occur throughout the world and in a rostrum, a dorsal fin shape that was the same as a Risso’s
wide range of habitats, and clearly provide opportunity for dolphin, and a melon a little bit more predominant than in
hybridism between the species. bottlenose dolphins (Sylvestre & Tasaka, 1985). None
reached physical maturity. However, a hybrid dolphin born
to a female bottlenose dolphin and a male Risso’s dolphin at
hybridism between risso’s and bottlenose Minami Chita Beachland in Japan in 1993 (Cetabase, 2013)
dolphins is still alive, indicating that these hybrids can live long
The atypical individuals described here are consistent with enough to reach physical maturity. The morphological fea-
hybridism between free-ranging Risso’s dolphins and bottle- tures described in these captive hybrids are very similar to
nose dolphins for three reasons: (1) the morphological those we describe here for wild atypical animals photographed
characteristics of the three individuals appeared intermediate off Lewis.
between those two species; (2) their occurrence within schools The functional explanations for the formation of mixed-
of one (or both) of those two species; and (3) the three obser- species cetacean associations primarily comprise foraging
vations of mixed-species schools of those two species within advantages and predator avoidance, although there could be
the study area. In particular, the pronounced, broad-based additional social and reproductive advantages (Stensland
falcate dorsal fins seen on all three of the individuals were con- et al., 2003). While interspecific mating could happen
sistent with Risso’s dolphin, while their short, defined beaks during relatively brief interactions between species, our obser-
were consistent with (though shorter than) the bottlenose vations of associations between bottlenose and Risso’s dol-
dolphin (however, the forehead in ATD1 was far more phins around Lewis are suggestive of a more stable
sloped than in a bottlenose dolphin). Given the absence of affiliation between the species. Predation (from sharks or
molecular data to confirm the genetic affiliation of these killer whales, Orcinus orca, (Linnaeus, 1758)) is unlikely to
animals, we cannot be certain that these animals are hybrids be a primary driver for multi-species associations in Scottish
between Risso’s dolphins and bottlenose dolphins and waters, although killer whales are occasionally sighted in the
cannot rule out that these are hybrids involving other sympat- study area (Nicola Hodgins, personal observation).
ric species such as Lagenorhynchus or Globicephalus. Strandings evidence from stomach contents indicates that
However, other instances where hybrids have been initially Risso’s and bottlenose dolphins exhibit different prey prefer-
described by morphological traits and then later confirmed ences in Scottish waters. The octopus Eledone cirrhosa
by molecular analysis (e.g. Spilliaert et al., 1991; Bérubé & (Lamarck, 1798) comprised almost 90% of the total recon-
Aguilar, 1997) suggest that the identification of hybrids structed prey weight of stranded Risso’s dolphins (mostly
based on observed morphological features is reasonable. from the north and west coasts) (Macleod et al., 2013),
This is not the first evidence for hybridism between wild while sampled bottlenose dolphins (mostly from the east
Risso’s dolphins and bottlenose dolphins in north-west coast) fed upon a variety of benthic and mid-water fish and
Europe. On 31 May 1933 three dolphins stranded in some cephalopods (Santos et al., 2001). Consequently, the
Blacksod Bay, County Mayo in Ireland, and all showed inter- functional explanation for this association remains unclear,
mediate morphological and skeletal characteristics relating to and they may rather have some type of social basis as has
both the bottlenose dolphin and the Risso’s dolphin (Fraser, been suggested for other odontocete species associations
1940). Two had short well-defined beaks (6.4 and 8.9 cm (e.g. Melillo et al., 2009). Whatever the driver may be, the
respectively) and the third animal lacked a beak and had a existence of these associations provides opportunity for
strongly-sloped forehead (Fraser, 1940). This seems compar- hybridism.
able in description to the short beaks photographed in Hybrids originating in captive situations are most likely to
ATD2 and ATD3, and the sloped forehead photographed in be the result of lack of mate choice due to their artificial con-
ATD1. Generally, the stranded animals described by Fraser finement. However, this is not always the case; a captive
(1940) had rostrums that were shorter and wider, tooth hybrid in Japan born to a female bottlenose dolphin and a
counts lower, and craniums wider than found in the bottle- male Risso’s dolphin was apparently conceived even though
nose dolphins, and he considered hybridism between Risso’s a mature male bottlenose dolphin was present in the same
and bottlenose dolphins to represent the most likely explan- pool at the time of the matings (Sylvestre & Tasaka, 1985).
ation. Despite the observations reported in this paper, no The driver for interspecific matings amongst wild cetacean
wild hybrids between the two species have been reported species is even less clear, although such matings have been
stranded to date in UK waters (Paul Jepson & Andrew documented for a number of sympatric cetacean species (see
Brownlow, personal communications). Bérubé, 2009). The context of these matings may be varied
Several confirmed incidences of hybridism between Risso’s and not necessarily reproductive, with alternative explana-
and bottlenose dolphins have been recorded in captivity tions including ‘practice’ matings by immature animals to
(Sylvestre & Tasaka, 1985; Shimura et al., 1986). While we improve adult reproductive success and to diffuse aggressive
acknowledge that the circumstances of captivity produce tension between species (Melillo et al., 2009). Not all
behaviours that would not occur naturally in the wild, interspecific matings seem to lead to conception and hybrid-
these records at least provide evidence that hybridism ization. For example, matings between wild bottlenose and
between the two species is possible and that such hybrids Atlantic spotted (Stenella frontalis, Cuvier, 1829) dolphins in
may reach maturity. Sylvestre & Tasaka (1985) compiled 13 the Bahamas are very frequent, and yet only a single possible
accounts of hybrids born to captive Risso’s and bottlenose dol- hybrid has been reported (Herzing et al., 2003). Nevertheless,
phins, all of which occurred at Enoshima Marineland in the number of reported hybrids is likely to increase as
Japan. The offspring bore intermediate morphological traits advances in molecular analysis techniques and biopsy sam-
between their two parental species, one individual having pling are made. Indeed, DNA analysis is now providing
6 nicola k. et al.

evidence of hybridization events that occurred in the past. For K., Simmonds M., Wittich A. and Wright A.J. (eds) Grampus griseus
example, recent evidence suggests that past hybridization 200th anniversary: Risso’s dolphins in the contemporary world. Report
between spinner (Stenella longirostris, Gray, 1828) and from the European Cetacean Society Conference Workshop, 26th
striped (S. coeruleoalba, Meyen, 1833) dolphins was of suffi- European Cetacean Society Conference, Galway, Ireland, 23– 29
March 2012. European Cetacean Society Special Publication Series
cient frequency to produce a new species, the Clymene
No. 54, pp 44–53.
dolphin (S. clymene, Gray, 1846) (Amarel et al, 2013).
The observations of three atypical dolphins off north-east Evans P.G.H., Lewis E.J., Parsons E. and Swann C. (1993) A survey of
Lewis are consistent with hybridization between free-ranging whales and dolphins in Hebridean waters. Oxford: SeaWatch
Risso’s and bottlenose dolphins, the first such occurrence to Foundation, 11 pp.
be documented for these species in UK waters. The context Fraser F.C. (1940) Three anomalous dolphins from Blacksod Bay, Ireland.
and significance of these hybridization events is unknown. Proceedings of the Royal Irish Academy; Section B: Biological,
Since wild cetacean hybrids are rare, the occurrence of four Geological and Chemical Science 45 (1938–1940), 413–455.
atypical dolphins consistent with hybridism in one small geo- Herzing D.L. and Johnson C.M. (1997) Interspecific interactions between
graphical region is highly unusual. However, the three hybrid Atlantic spotted dolphins (Stenella frontalis) and bottlenose dolphins
animals reported by Fraser (1940) in Ireland indicate that this (Tursiops truncatus) in the Bahamas, 1985–1995. Aquatic Mammals
is not an altogether unprecedented scenario. If these indivi- 23.2, 85–99.
duals do indeed represent hybrids, it raises interesting ques- Herzing D.L., Moewe K. and Brunnick B.J. (2003) Interspecies interac-
tions regarding their fertility, lifespan and relatedness. tions between Atlantic spotted dolphins, (Stenella frontalis) and bottle-
nose dolphins, (Tursiops truncatus), on Great Bahama Bank, Bahamas.
Aquatic Mammals 29.3, 335 –341.
ACKNOWLEDGEMENTS HWDT (2014) Hebridean Whale and Dolphin Trust unpublished data
and distribution maps. Available at: http://www.whaledolphintrust.
The financial support of WDC, DEFRA, BBC Wildlife Fund, co.uk (accessed 27 January 2014).
Elite Couriers and David Henriques enabled this field Jefferson T.A., Weir C.R., Anderson R.C., Ballance L.T., Kenney R.D.
project. The authors would like to thank Scottish Natural and Kiszka J.J. (2014) Global distribution of Risso’s dolphin
Heritage for granting WDC photo-id licences (Licence Grampus griseus: a review and critical evaluation. Mammal Review
Numbers 10991 and 13371) and everyone who made the 44, 56–68.
surveys possible, especially Lewis Mackenzie at Hebrides Karczmarski L., Thornton M. and Cockcroft V.G. (1997) Description of
Fish n’Trips, Tim Atkinson and the SNH Stornoway office. selected behaviours of humpback dolphins, Sousa chinensis. Aquatic
The authors are also grateful for comments from three Mammals 23.3, 127–133.
anonymous referees that helped improve this manuscript.
Macleod C.D., Santos M.B., Burns F., Brownlow A. and Pierce G.J.
(2013) Can habitat modelling for the octopus Eledone cirrhosa help
identify key areas for Risso’s dolphin in Scottish waters?
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