General characteristics

The common bottlenose dolphin (Tursiops truncatus) is perhaps one of the best-known cetaceans and likely what most people picture when they think of a dolphin. It is a widespread and abundant species, occurring in tropical and temperate coastal and pelagic marine waters worldwide. Bottlenose dolphins are familiar from appearances in films and TV shows, as well as from being exhibited in numerous zoos and aquariums.

Their name comes from their distinct rostrum (often referred to as a beak), which is typically 7–8 cm long. Their mouths are gently curved, dipping slightly downward from the tip of the beak and upwards at the base, giving them the appearance of smiling. Their beak contains 76–98 conically-shaped teeth.

The bottlenose dolphin is a light grey to black colour dorsally, with white or cream colour underneath. Some individuals may also have lighter colour on the sides.

Size and age

The bottlenose is a relatively large, robust dolphin. The body is streamlined with a tall, falcate (curved) dorsal fin located in the middle of their back. There are considerable regional variations in body size, to the extent that different “morphotypes” or “ecotypes” are recognised throughout its range (Oudejans et al. 2015). In some areas, distinct inshore and offshore populations can be found, with the inshore dolphins generally being smaller in size than the offshore dolphins. The differences between the two types are large enough that they may eventually be classified as separate species (Hammond et al. 2012).

Average body length for the coastal ecotype of this dolphin is around 2.6 m, with weights between 190–260 kg. Offshore dolphins can reach 3.8 m in length for males and 3.7 for females. An average body weight offshore is around 450 kg, although weights up to 650 kg have been recorded. Males of both ecotypes are typically larger than females, although considerable variation in size-at-age for both sexes at all ages has been noted (Read et al. 1993). Calves are 84–140 cm long at birth and weigh approximately 20 kg (Wells & Scott 1999).

Bottlenose dolphins are a long-lived species. Dolphins found stranded along the Mississippi coast in the Gulf of Mexico were found to be up to 30 years old (Mattson et al. 2006). In a long-term study of dolphins in Florida, males were observed to live for more than 40 yrs and females for more than 50 (Connor et al. 2000).

Taxonomy

All bottlenose dolphins around the world were previously recognised as Tursiops truncatus, but recently the genus has been split into two species: T. truncatus and T. aduncus. The latter species is called the Indo-Pacific bottlenose dolphin. There is also a separate subspecies of bottlenose dolphin recognised in the Black Sea, T. t. ponticus (Hammond et al. 2012).

Life History and Ecology

Behaviour

Surface behaviour and diving

Bottlenose dolphins are very active at the surface and can often be observed breaching or spy hopping. They may ride the bow waves of ships and in some locations, are well known for following fishing vessels (Byrd & Hohn 2017, Jones & Sayigh 2002, Wells & Scott 1999). They are also often observed associating with other species of dolphins and whales.

Although active at the surface, bottlenose dolphins spend most of their time underwater. One tagged female in Florida was found to spend about 87% of her time submerged (Mate et al. 1995). Bottlenose dolphins tend to make numerous short dives from the surface, rather than long deep dives. Dolphins in the Moray Firth, Scotland, were observed to spend most of their time within the surface layers of the water column. Relatively little time was spent below 10 m, although dolphins were occasionally seen diving to depths of around 50 m (Hastie et al. 2006).

Inter- and intraspecific aggression

Bottlenose dolphins demonstrate aggressive behaviour. Physical violence (body slamming, tail hitting, charging, jawing and biting), hostile head-to-head posturing, and acoustic threats have all been observed (Robinson 2014), and many dolphins carry scars and rake marks from such encounters. Adult male bottlenose dolphins have also been seen attempting to kill dolphin calves. Robinson (2014) describes one event in which an adult male chased, repeatedly rammed, tossed out of the water and attempted to asphyxiate a newborn calf. A potential driver of this behaviour is that if the mother loses her calf in this way, she may become available for mating.

Bottlenose dolphins can also act aggressively towards other species, especially harbour porpoises. Violent and sometimes fatal attacks on harbour porpoises by bottlenose dolphins have been reported in several locations (Santos & Peirce 2003, Spitz et al. 2006). Around the Moray Firth, Scotland, 63% of the harbour porpoises found stranded appeared to have been attacked by bottlenose dolphins and died from trauma, specifically bone fractures and internal injuries (Ross & Wilson 1996).

Sounds

Bottlenose dolphins produce a variety of sounds that are typically separated into three categories: whistles, burst-pulse sounds, and echolocation clicks (Jones & Sayigh 2002).

Whistles are used in communication, in social interactions, and in maintaining contact with each other. Individual dolphins produce a “signature whistle”, which is a unique frequency modulation pattern that identifies the individual. This is likely used to maintain social bonds. The burst-pulse vocalisations may also be used to communicate (Jones & Sayigh 2002). Dolphin communication is complex and dolphins have been observed to modify their sounds through learning. One study observed dolphins using their whistles to “address” each other, as a whistle produced by one dolphin would be matched by a dolphin emitting the same whistle type (Janik 2000a).

Echolocation clicks are high-frequency sounds produced in order to locate prey and to navigate (Nuuttila et al. 2017). They are often emitted in “click trains”, a long sequence of sound that usually increases in speed as the dolphin gets closer to its prey. Another type of sound that has been observed associated with feeding is termed a “bray”. Dolphins in the Moray Firth were observed to emit these low-frequency sounds when feeding on salmonid fish (Janik 2000b). It is thought that the sounds are used to help the dolphins capture this particular prey.

Social interactions

Bottlenose dolphins are a highly social species. They are commonly found in small groups of 2–15 individuals (Wells & Scott 1999), although larger groups are sometimes observed in offshore areas. The social structure is often described as “fission-fusion”, in which individuals associate in small groups that may change on a daily or even hourly basis (Connor et al. 2000).

In some locations, associations between animals of a similar age and reproductive state are commonly observed, with adult dolphins associating with dolphins that they grew up with or spent time with as juveniles (Robinson et al. 2017). Such long-term alliances may be shaped by kin selection, mating strategies, ecological factors such as feeding strategies, or perhaps a combination of these factors (Louis et al. 2017). Bottlenose dolphins in the Shannon estuary, however, do not appear to associate in such groups, but rather have strong and persistent bonds between individuals (Baker et al. 2017).

Cooperative foraging behaviour has been observed in bottlenose dolphins, in which groups work together to herd schools of prey fish, sometimes encircling them in open water and at other times driving them up onto a sloping shore (Wells & Scott 1999).

Strandings

Bottlenose dolphins are found stranded on coastlines throughout their range and at various times of the year. Along the eastern coast of the US, from Florida to Maine, an average of 150 dolphins stranded each year in the period 1972–1997 (McLellan et al. 2002).

In some locations, strandings are more frequent at certain times of the year. Along the coast of North Carolina in the US, adult dolphins stranded most frequently during the spring and fall, and less frequently from June–August (Thayer et al. 2003). Neonates in this area stranded more often in April and May, with lower levels in the fall and winter. Along the north-central Gulf of Mexico coast, strandings of both adult and neonate dolphins peaked in March and April (Mattson et al. 2006), while along the coast of South Carolina, about half of  annual dolphin strandings occurred between April and July (McFee & Hopkins-Murphy 2002).

Why do bottlenose dolphins strand?

There are various reasons proposed for why dolphins may strand. Strandings may occur when dolphins follow a high tide in to coastal waters and become trapped when the tide recedes. Dolphins have also been seen to sometimes intentionally beach themselves in order to catch fish that they have chased towards shore. This foraging strategy has been observed throughout their range (Silber & Fertl 1995).

Other stranded animals show signs of interactions with fishing gear, which may have contributed to their death prior to being found on shore. Between 1997 and 2008, 17% of all dolphins found stranded  along the coast of North Carolina showed evidence of contact with some type of fishing gear (Byrd & Hohn 2017). In some cases, gear is still attached to the stranded dolphin, while in others, the dolphin shows lesions that have been caused by gear. On the coast of France, 21% of stranded bottlenose dolphins were found to have lesions caused by fishing gear (ICES 2018).

One study of strandings of common dolphins, bottlenose dolphins and harbour porpoises in north-western Spain looked at various meteorological, oceanographic, prey abundance and fishing-related variables to see if any pattern was apparent. They found that the local ocean meteorology (particularly the strength and direction of the winds) along with the winter North Atlantic Oscillation Index best explained the stranding pattern, and that there was no association with fishing effort or landings occurring in the same area (Saavedra et al. 2017).

Sexual maturity and breeding

The age at sexual maturity for bottlenose dolphins varies by both sex and location.

In Florida, females were found to be sexually mature at 5–12 years old, and males from 10–13 years. Females along the central US Atlantic coast were found to mature between 7–13 years old (Wells & Scott 1999). In the Moray Firth, Scotland, females were observed to be mature from 6–13 years old, and produced their first calf at a mean age of 8 (Robinson et al. 2017).

Calves

In the Western North Atlantic, calves are primarily born in the late spring and early summer. In Sarasota Bay Florida, the calving peak occurs between May and July (Wells & Scott 1990), with similar timing observed off the coast of North Carolina (Thayer et al. 2003). In the Moray Firth, calves are produced throughout the summer, from May to October, but with a seasonal pulse that corresponds to the peak in summer water temperatures (Robinson et al. 2017).

Gestation lasts for about 12 months in bottlenose dolphins and a single calf is produced. At birth, calf lengths range from 84–104 cm (Wells & Scott 1999). The calf usually remains with its mother for several years. Milk is the primary food for the first year, but nursing has been observed to continue for up to 5 years (Connor et al. 2000). Females generally produce a calf every 2 or 3 years, but the interval may  be longer in some cases. Dolphins in the Moray Firth were observed to have a mean interbirth interval closer to 4 years (Robinson et al. 2017). Factors such as maternal age and size, breeding experience, dominance, individual associations, group size and other social factors are all likely to influence reproductive success.

Food and feeding

Bottlenose dolphins have a varied diet, which changes with habitat and season. Studies in various parts of their range have shown that they take a wide variety of locally abundant prey items, adapting both their foraging tactics and prey selection to local conditions.

Spatial differences

In North Carolina, for example, at least 31 different species of prey items from 21 different families were found in the stomachs of coastal bottlenose dolphins (Gannon & Waples 2004). Pelagic and demersal fish are the main components of the diet, but they also take cephalopods and crustaceans.

Dolphins in Cardigan Bay, Wales, were found to prey on sandeels (Ammodytidae sp.), cod (Gadus morhua), whiting (Merlangius merlangius), sprat (Sprattus sprattus), Atlantic herring (Clupea harengus), mackerel (Scomber sp.) and a variety of flatfish (Nuuttila et al. 2017). Dolphins found stranded on the Scottish coast were found to have fed mainly on cod, saithe (Pollachius virens), whiting, and to a lesser extent, salmon (Salmo salar), haddock (Melanogrammus aeglefinus), and cephalopods (Santos et al. 2001). Crustacean and polychaete remains were also found in a couple of stomachs in this study. Dolphins feeding offshore in the northeastern Atlantic appear to concentrate on shoaling fish such as Atlantic herring and whiting (Nuuttila et al. 2017). Whiting and hake (Merluccius merluccius) were also found to be important prey items for dolphins feeding off the northwest coast of Spain (Santos et al. 2007).

Differences in inshore and offshore feeding have also been noted in the US. Dolphins found stranded along the North Carolina coast had fed primarily on weakfish (Cynoscion regalis), Atlantic croaker (Micropogonias undulatus), and spot (Leiostomus xanthurus), as well as on inshore squid (Loligo sp.) and striped anchovy (Anchoa hepsetus) (Gannon & Waples 2004). Dolphins that were found stranded inside estuaries had a diet dominated by Atlantic croaker, while dolphins found stranded elsewhere had diets dominated by weakfish. Inshore squid was commonly eaten by dolphins in the ocean, but not by those in the estuaries. This study also found that there was considerable variation in the diet by season.

Differences between sexes

Variation in the diet between the sexes has also been noted in some locations. Dolphins in Sarasota Bay, Florida, take small fish largely associated with seagrass habitat such as pigfish (Orthopristis chrysoptera), pinfish (Lagodon rhomboids), mojarra (Gerreid sp.) but also larger species such as striped mullet (Mugil cephalus) (Rossman et al. 2015). This study found that while the males appeared to specialise on the seagrass-associated prey, the females in the area had broader foraging habits and took prey from a variety of habitats.

Predators

Sharks appear to be the main predators of bottlenose dolphins, although they may also be taken by killer whales (Orcinus orca). Dolphins in many locations show scarring from shark bites and dolphin remains have been found in shark stomachs. Off the southeastern coast of the United States, bull (Carcharhinus leucas), tiger (Galeocerdo cuvier) and dusky sharks (Carcharhinus obscures) have all been observed to prey on bottlenose dolphins (Wells & Scott 1999).

Health – diseases and parasites

Like all other cetaceans, bottlenose dolphins can be affected by various diseases and may be infected by parasites. They are commonly found to have external lesions or scars on their bodies, either from shark attacks or attacks by other dolphins, or sometimes from interactions with fishing gear.

Diseases

One disease that has been apparent in bottlenose dolphins is morbillivirus, which can rapidly affect many animals in an area. This virus has been responsible for several “Unusual Mortality Events” along the east coast of the United States. In a 10 month period from June 1987 to March 1988, a ten-fold increase in stranding rates was seen along the coast, which was later determined to be caused by morbillivirus (McLellan et al. 2002). Another outbreak of morbillivirus occurred along the coast between July 2013 and March 2015, killing some 1650 dolphins (Hayes et al. 2016).

A more complete discussion of some of the other diseases and conditions that affect bottlenose dolphins may be found in Eissa & Abu-Seida (2015).

Parasites

Parasites may be found in or on various parts of the bottlenose dolphin’s body. Whale “lice” (a type of Cyamid amphipod) have been found on the skin, and in one case were associated with lesions which penetrated the skin into the underlying adipose tissue (Woodard et al. 1969). These amphipods do not appear to harm the dolphin.

Internally, dolphins have been found to host various types of helminth parasites. The nematode Anisakis sp. is a common parasite in the guts of various cetaceans and is present in bottlenose dolphins as well. In one case, this parasite was also found infecting the skin of a dolphin (Van Beurden et al. 2015). Other nematodes are present in the lungs. The lungworm Halocercus lagenorhynchi is commonly found in dolphins and was present in 77% of stranded dolphins examined in Florida (Fauquier et al. 2009). Infection with these lungworms leads to mucopurulent bronchiolitis and pneumonia (Woodard et al. 1969). Parts of the lungs containing the nematodes eventually become walled off, sclerotic, and calcified. In most of the dolphins examined in Florida, these calcified lesions were mild, chronic, and did not cause the death of the dolphins.

The digestive tracts of dolphins may harbour various species of nematodes, cestodes and digeneans or trematodes. Quiñones et al. (2013) found 7 different helminth species in 15 bottlenose dolphins from the western Mediterranean: 3 digeneans: Pholeter gastrophilis, Synthesium tursionis, and Brachycladium atlanticum; 1 nematode Anisakis sp.; and 3 cestodes: Tetrabothrius forsteri, Diphyllobothrium sp., and Strobilocephalus triangularis. Synthesium tursionis was the most prevalent of these parasites, found in over 70% of the dolphins examined (Quiñones et al. 2013). Another trematode that has been found in dolphins is Campula palliata, which may infect the liver (Woodard et al. 1969).

Distribution and habitat

Bottlenose dolphins have a worldwide distribution in temperate and tropical waters. They are found in waters across the Atlantic Ocean, in both the northwest and northeast. Off the coast of North America, they are generally found in waters with surface temperatures ranging from about 10–32°C (Wells & Scott 1999).

In the northwest Atlantic during the summer months, bottlenose dolphins are seen inshore off New England and along the US coast south to Florida. Offshore they range as far north as Nova Scotia (Wells & Scott 1999).

In the northeast Atlantic, they are regularly seen off the coasts of Portugal, Spain, France, the UK and Ireland (Hammond et al. 2013). Around the UK and Ireland, they have a patchy distribution with resident populations in Cardigan Bay (Wales) the Moray Firth (north-east Scotland) and the Shannon estuary (Ireland), but are also often sighted in waters to the north and west of Scotland, up to around the Faroe Islands. They are also found in the North Sea and have been recorded along the Norwegian coast as far north as the Lofoten Islands.

Bottlenose dolphins inhabit a wide range of habitats throughout their worldwide distribution. Some groups of coastal dolphins may be found in estuaries, bays, lagoons and other shallow inshore regions, occasionally ranging far up into rivers. Offshore, some dolphins are resident around islands or archipelagos, e.g. around the Azores (Silva et al. 2008) or the Madeira Islands (Dinis et al. 2016). Other offshore bottlenose dolphins exhibit low site fidelity, ranging instead over large areas.

Migrations and movements

Some coastal dolphins maintain long-term multi-generational home ranges in certain bays or estuaries. In such locations, females generally range less than males and have more site-fidelity (Robinson et al. 2017, Wells & Scott 1990). Other coastal dolphins have more complex movements and may alternate between residing in one location and making occasional long-range travels (Wells & Scott 1999). Dolphins from around northern Scotland, for example, were seen to travel distances up to 1,277 km (Robinson et al. 2012), while a dolphin tagged off the Florida coast in the Gulf of Mexico covered a distance of 2,050 km in 43 days (Wells et al. 1999). Coastal dolphins around Ireland have also been documented making large-scale movements around Ireland and between Atlantic coastal waters and the North Sea (Oudejans et al. 2015).

Offshore dolphins generally show low site fidelity and range over much greater areas in larger groups (Oudejans et al. 2015). At the northern edge of their range in the western north Atlantic, they seasonally migrate, moving south in the winter months and north in the summer (Hammond et al. 2012). Seasonal patterns of movement have also been observed in bottlenose dolphins off Wales, with dolphins moving inshore and becoming more commonly seen in late summer (Nuuttila et al. 2017). This may correspond to seasonal concentrations of migratory fish that the dolphins are preying on. During the winter, dolphins in this area were observed moving further north and dispersing over a wider area. In the waters off Ireland, surveys carried out in 2015 and 2016 found an eightfold increase in bottlenose dolphin abundance during winter compared to summer (ICES 2018).

North Atlantic stocks

Bottlenose dolphins are a wide-ranging species and for the most part, do not have clearly defined stocks in the North Atlantic. There is some evidence of population structure in certain estuaries or around island groupings where there may be a resident population of dolphins, however even in those areas, there are transient dolphins and much gene flow.

Northwest Atlantic

A number of stocks are recognised in the Northwest Atlantic along the east coast of the United States. Here there are 5 coastal stocks of bottlenose dolphins (Hayes et al. 2016):

• Northern Migratory and Southern Migratory Coastal,
• South Carolina/Georgia Coastal,
• Northern Florida Coastal,
• Central Florida Coastal,
• Western North Atlantic Offshore.

These stocks are differentiated by their abundance, migratory patterns and stable isotope ratios, and show varying degrees of genetic differentiation (Toth et al. 2012). Along the southern US coast there are also a number of smaller populations recognised that inhabit the estuarine systems of some of the larger rivers.

Northeast Atlantic

In the Northeast Atlantic, bottlenose dolphin stock structure is not well understood. As in the west, differences between inshore and offshore dolphins have been noted. In the waters around Ireland, three genetically distinct populations have been identified: a resident population in the Shannon estuary, a coastal population in the waters off western Ireland, and a likely offshore population (Oudejans et al. 2015). These appear to be separate breeding populations. Other resident populations around the UK are found in Cardigan Bay (Wales) and the Moray Firth (north-east Scotland).

On the northwest coast of Spain, stable isotope analysis of skin and muscle tissue from stranded dolphins was used to assess dietary variation, habitat segregation and population substructure. It revealed the presence of two clear groups of Galician bottlenose dolphins: a northern and southern group (Fernández et al. 2011).

Further investigation throughout the Northeast Atlantic is needed in order to more fully understand bottlenose dolphin population structure in this region.

Current abundance and trends

Determining population numbers for bottlenose dolphins is difficult as they are a highly mobile species that can sometimes occur in very large groups. This makes them hard to survey, although there has been some work done in various parts of their range in order to produce population estimates. It is estimated that there is a minimum of 600,000 bottlenose dolphins world-wide (Hammond et al. 2012).

Northeast Atlantic

In the Northeast Atlantic, several surveys have been carried out for different small cetacean species, including bottlenose dolphins. One example is the multinational Small Cetacean Abundance in the North Sea and Adjacent Waters (SCANS) surveys, a series of surveys that took place in 1994, 2005 and 2016.

The 1994 survey was not able to generate a population estimate for bottlenose dolphins. The 2005 survey gave an estimated abundance of dolphins in the survey area of 16,641 (CV = 0.42, 95% CI: 7,618–36,351) (Hammond et al. 2017). The SCANS III survey in 2016 gave a population estimate for the same area of 27,697 (CV = 0.23, 95% CI: 17,662–43,432) (Hammond et al. 2017).

Another survey, the Cetacean Offshore Distribution and Abundance in the European Atlantic (CODA) survey, investigated European Atlantic waters beyond the continental shelf in 2007 and estimated a population of 19,295 (CV = 0.25) bottlenose dolphins for the survey area (CODA 2009).

In the North Atlantic, a series of surveys called the North Atlantic Sighting Surveys (NASS) have been carried out since the late 1980s for various species of cetaceans. Between 1987 and 2015, a total of 91 dolphins were seen on these surveys. The number seen was too low to be able to make a population estimate for bottlenose dolphins in the North Atlantic.

Some of the inshore populations around Ireland and the UK have been surveyed. In Cardigan Bay, Wales, the most recent population estimate was 258 (95% CI: 161–508) for the bay area (ICES 2018). Around 400–500 are found in Scottish and Irish coastal waters particularly the Shannon River estuary and the Moray Firth (Berrow et al. 2010, Thompson et al. 2011 in Robinson et al. 2012).

Northwest Atlantic

In the Northwest Atlantic, surveys have been done primarily along the east coast of the United States. In the summers of 2010 and 2011, aerial surveys were flown off the coast covering waters from Florida to the lower Bay of Fundy. The best population estimate for the offshore stock of bottlenose dolphins in the Northwest Atlantic is 77,532 (CV=0.40) (Hayes et al. 2016).

The same surveys also provided population estimates for the inshore stocks along the US east coast. The inshore northern migratory coastal stock is estimated at 11,548 (CV=0.36), the southern migratory coastal stock is estimated at 9,173 (CV=0.46), the South Carolina/Georgia coastal stock at 4,377 (CV=0.43), the northern Florida coastal stock at 1,219 (CV=0.67), and the central Florida coastal stock at 4,895 (CV=0.71) (Waring et al. 2015). This makes the total inshore population around 31,000 dolphins. In addition, there are various smaller populations of several hundred dolphins living in estuarine systems along the southern part of the US east coast. Depending on the size of the estuary, there may be 100–400 dolphins in each (Waring et al. 2015).

Stock status

It is difficult to establish population trends for bottlenose dolphin populations, as most parts of their range have not been systematically surveyed. Even in areas where repeated surveys have taken place, the methods used are not always comparable and the time-series is short relative to the lifecycle of the dolphins. Another problem with detecting trends is that the precision of the population estimates that are obtained is generally low, making it difficult to infer with any confidence whether populations are decreasing, stable or increasing.

Northeast Atlantic

In the Northeast Atlantic, the multinational Small Cetacean Abundance in the North Sea and Adjacent Waters (SCANS) surveys, are one of the few systematic surveys that have included estimates for bottlenose dolphins. These surveys took place in 1994, 2005 and 2016, although an estimate could not be made for 1994. The 2016 survey found a larger population of dolphins in the survey area than the 2005 survey did (Hammond et al. 2107). It is not clear whether this reflects an increase in the population or a change in distribution.

In many locations around the Northeast Atlantic Ocean, coastal bottlenose dolphin populations declined or disappeared during the late 1800s or early 1900s. Currently, however, most of the populations in this region appear to be stable (NAMMCO 2016).

On a smaller scale, the population in Cardigan Bay, Wales, has been monitored for the past 17 years. Over that time, the number of dolphins has fluctuated but the population estimate obtained in 2017 is not significantly different from the estimate made in 2001 (ICES 2018).

Stranding rate can also be used as an indirect measure of population, although other factors also influence strandings. In northwestern Spain, the stranding rate did not appear to reflect any trend in dolphin abundance over the period 2000–2013 (Saavedra et al. 2017).

Northwest Atlantic

In the Northwest Atlantic, three different abundance estimates are available for the offshore stock. The summer 1998 surveys estimated 29,774 (CV=0.25), summer 2002/2004 surveys estimated 81,588 (CV=0.17) and the summer 2011 surveys 77,532 (CV=0.40) (Hayes et al. 2016). A trend analysis, however, has not been done using these estimates due to methodological differences between the surveys.

Management

The common bottlenose dolphin is a widespread and abundant species. It is listed on the IUCN Red List in the category of “Least Concern” (Hammond et al. 2012). While there may be threats acting on local populations, none of these are considered to be causing a global population decline.

Although not considered endangered, bottlenose dolphins are listed in Appendix II of the Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES), as well as in Appendix II of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). These conventions provide for international cooperation in the management of species like the bottlenose dolphin that are widely distributed.

The bottlenose dolphin is also recorded in the European Union (EU) Habitats Directive as a Species of Special Interest. As such, it requires the designation of Special Areas of Conservation (SAC) by EU Member States for its protection. One example of an SAC is Cardigan Bay, Wales, where there is a resident population of around 200 dolphins.

Hunting and utilisation

Recent catches

Bottlenose dolphins are hunted in certain parts of the world for human consumption and/or for use as bait in fisheries. These catches currently occur in Peru, Sri Lanka, Taiwan and Japan (Hammond et al. 2012). In the Faroe Islands, a small number of bottlenose dolphins have been hunted occasionally in combination with the pilot whale drives. Since 2000, a total of 45 have been hunted there, with none taken since 2009 (https://www.whaling.fo/).

Catches in NAMMCO member countries since 1992

Country	Species (common name)	Species (scientific name)	Year or Season	Area or Stock	Catch Total	Quota (if applicable)
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2023	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2022	Faroe Islands	98
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2021	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2020	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2019	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2018	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2010-2017	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2009	Faroe Islands	1
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2007-2008	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2006	Faroe Islands	17
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2004-2005	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2003	Faroe Islands	3
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2002	Faroe Islands	18
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	2001	Faroe Islands	6
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	1997-2000	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	1996	Faroe Islands	21
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	1995	Faroe Islands	0
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	1994	Faroe Islands	8
Faroe Islands	Common bottlenose dolphin	Tursiops truncatus	1992-1993	Faroe Islands	0

Other human impacts

By-catch

Bottlenose dolphins are vulnerable to by-catch in a variety of fishing gear throughout their range, including in gillnets, driftnets, purse seines, trawls, long-lines, and on hook-and-line gear. The levels of by-catch are, however, relatively low compared to other cetaceans such as common dolphins or harbour porpoises. Dolphins may get caught by not seeing nets, by chasing fish into them, or by taking fish from nets (depredation). In some areas, they are known to follow fishing vessels out to where they set or retrieve gear (Byrd and Hohn 2017), perhaps increasing the likelihood that they will become caught or entangled in gear.

Levels of by-catch are not well known in most parts of the bottlenose dolphin’s range due to a lack of reporting. One estimate of some fishery-related mortality has been made for the western North Atlantic offshore stock. A mean annual death of 39.4 (CV=0.29) dolphins due to interactions with the northeast sink gillnet, northeast bottom trawl, mid-Atlantic bottom trawl, and pelagic longline fisheries was determined (Hayes et al. 2016). However, the total mortality for this stock is unknown as there are other fisheries occurring in this region with unreported by-catch.

Age-class and season have appeared to play an important role in levels of dolphin by-catch off North Carolina. A study found that the by-catch risk for older calves and subadults was 1.5 (in the summer) to 3.5 (in the spring) times greater than that for an adult or young-of-year dolphins (Byrd & Hohn 2017).

No by-catch of bottlenose dolphins has recently been reported for any of the fisheries taking place in Norway, Iceland, Greenland or the Faroe Islands (NAMMCO 2017).

Noise/disturbance

Coastal populations of bottlenose dolphins may be exposed to various types of human activity occurring in their habitat. This includes vessel traffic, construction, oil and gas exploration and other industrial activities. Dolphins may also be directly or indirectly disturbed or harassed by boat traffic, including tourist ships on “dolphin watching” excursions.

Noise is of particular concern for bottlenose dolphins since they use sound extensively for both foraging and communication. Dolphins in the Sado estuary, Portugal, changed both their call rate and whistle characteristics in the presence of certain vessels (Luis et al. 2014). The overall call rate decreased in the presence of operating vessels, especially ferry boats. In Florida, dolphins were observed to increase their rate of whistle production at the onset of a vessel approach (Buckstaff 2004). These changes in vocalisations could potentially disturb important social interactions, such as mating or mother-calf communication.

Dolphins also react to the presence of vessel traffic. Dolphins in Sarasota Bay, Florida, swam closer together, changed heading, and increased swimming speed significantly in response to an approaching vessel (Nowacek et al. 2001).

Behavioural changes such as these could have long-term effects on the species if animals that are feeding, resting, or breeding are disturbed by noise or vessel traffic. In Shark Bay, Australia, the presence of tourist vessels (which often approach quite close to the dolphins), was found to contribute to declining dolphin numbers in the area (Bejder et al. 2006). In this and other cases, the effect of noise and disturbance could be that dolphins get displaced and are forced into areas of poorer habitat. This is of particular concern for small coastal populations inhabiting a single bay or estuary.

 

Contaminants

Like other marine mammals, bottlenose dolphins ingest persistent organic pollutants (POPs) in their diet by consuming fish that have been exposed to these compounds from coastal runoff and human industrial activities. Coastal bottlenose dolphins in particular may be exposed to contaminants as their habitats may be adjacent to areas of high human population and industrial activity. Since they are a relatively long-lived species, contaminant levels may build up over time to quite high levels.

PCBs

Polychlorinated biphenyls (PCBs) are of particular concern for bottlenose dolphins. In a large Europe-wide analysis of stranded (n = 929) or biopsied (n = 152) cetaceans, it was found that bottlenose dolphins in the Northeast Atlantic and Mediterranean had mean PCB levels in their blubber that were among the highest recorded in cetaceans globally (Jepson et al. 2016). These concentrations are higher than the level at which severe toxic effects are known to occur. High levels of PCB contamination can, for example, cause immunosuppression, which could have been a significant contributing factor in the death of many of the stranded individuals in the European study that showed fatal infectious diseases on necropsy (Jepson et al. 2016).

High PCB levels in female dolphins may also be transferred to their young. A disproportionately high maternal transfer of accumulated PCBs was noted in primiparous females in the Moray Firth, Scotland (Robinson et al. 2017), which could result in brain cell damage, thyroid inhibition and immunosuppression in the foetus. This may be a contributing factor to the low survival rate of first-born calves in this area (Robinson et al. 2017).

Elevated PCB levels are also seen in coastal bottlenose dolphin populations in the US. Male dolphins in some parts of Florida have been found to have PCB levels more than an order of magnitude greater than the threshold for adverse health effects (Hammond et al. 2012). High levels of PCBs in the blubber have also been noted in North and South Carolina (Waring et al. 2015). As in the Moray Firth, these high levels of PCBs are correlated with higher levels of mortality in first-born offspring, and so could be affecting the reproductive success of these populations.

Heavy metals

Bottlenose dolphins in Florida were also found to have high concentrations of total mercury in their blood and skin, among the highest levels reported worldwide (Titcomb et al. 2017). Such high levels of mercury may have adverse health effects for the dolphins, particularly in hepatic, renal, endocrine, hematological, and immune parameters (Titcomb et al. 2017).

The exposure of bottlenose dolphins to these environmental pollutants and the subsequent effects on their population health is an area of concern and ongoing research.

Climate change

Climate change could have a number of impacts on bottlenose dolphins either directly or indirectly. For example, if water temperatures increase, this could have a direct impact on the habitat available for dolphins and consequently their distribution. Changes in water temperature could also affect the distribution of the dolphins’ prey species and so indirectly affect dolphin distribution.

Research in NAMMCO member countries

The common bottlenose dolphin is included in the NAMMCO-coordinated Trans North Atlantic Sighting Surveys (T-NASS). While bottlenose dolphins are not the primary target species, sightings of them are recorded during the surveys. So far, however, there have not been enough sightings made to be able to produce a population estimate.

Baker, I., O’Brien, J., McHugh, K., Simon, N., Ingram, S. N. and Berrow, S. (2017). Bottlenose dolphin (Tursiops truncatus) social structure in the Shannon Estuary, Ireland, is distinguished by age- and area-related associations. Marine Mammal Science, 34(2), 458–487. https://doi.org/10.1111/mms.12462

---

Bejder, L., Samuels, A., Whitehead, H., … and Krützen, M. (2006). Decline in relative abundance of bottlenose dolphins exposed to long‐term disturbance. Conservation Biology, 20, 1791–1798. https://doi.org/10.1111/j.1523-1739.2006.00540.x

---

Buckstaff, K. C. (2004). Effects of watercraft noise on the acoustic behavior of bottlenose dolphins, Tursiops truncatus, in Sarasota Bay, Florida. Marine Mammal Science, 20, 709–725. https://doi.org/10.1111/j.1748-7692.2004.tb01189.x

---

Byrd, B. L. and Hohn, A. A. (2017). Differential risk of bottlenose dolphin (Tursiops truncatus) bycatch in North Carolina, USA. Aquatic Mammals, 43(5), 558–569. https://doi.org/10.1578/AM.43.5.2017.558.

---

Cetacean Offshore Distribution and Abundance in the European Atlantic (CODA). (2009). SMRU, Gatty Marine Labratory, University of St. Andrews, UK. Available at http://biology.st-andrews.ac.uk/coda/

---

Connor, R. C., Wells, R. S., Mann, J. and Read, A. J. (2000). The Bottlenose Dolphin: Social Relationships in a Fission-Fusion Society. In J. Mann, R. C. Connor, P. L. Tyack and H. Whitehead (eds.), Cetacean Societies: Field Studies of Dolphins and Whales, 91–126. Chicago: The University of Chicago Press.

---

Dinis, A., Alves, F., Nicolau, C., Ribeiro, C., Kaufmann, M., Cañadas, A. and Freitas, L. (2016). Bottlenose dolphin Tursiops truncatus group dynamics, site fidelity, residency and movement patterns in the Madeira archipelago (North-East Atlantic). African Journal of Marine Science, 38(2), 151–160. https://doi.org/10.2989/1814232X.2016.1167780.

---

Eissa, A. E. and Abu-Seida, A. M. (2015). Synopsis on the most common pathologies of dolphins. Journal of Fisheries and Aquatic Science, 10, 307–322. https://doi.org/10.3923/jfas.2015.307.322

---

Fauquier, D. A., Kinsel, M. J., Dailey, M. D., Sutton, G. E., Stolen, M. K., Wells, R. S. and Gulland, F. M. D. (2009). Prevalence and pathology of lungworm infection in bottlenose dolphins Tursiops truncatus from southwest Florida. Diseases of Aquatic Organisms, 88(1), 85–90. https://doi.org/10.3354/dao02095

---

Fernández, R., García-Tiscar, S., Santos, M. B., López, A. Martínez-Cedeira, J. A., Newton, J. and Pierce, G. J. (2011). Stable isotope analysis in two sympatric populations of bottlenose dolphins Tursiops truncatus: evidence of resource partitioning? Marine Biology, 158, 1043–1055. https://doi.org/10.1007/s00227-011-1629-3

---

Gannon, D. P. and Waples, D. M. (2004). Diets of coastal bottlenose dolphins from the U. S. mid-Atlantic coast differ by habitat. Marine Mammal Science, 20(3), 527–545. https://doi.org/10.1111/j.1748-7692.2004.tb01177.x.

---

Genov, T. 2023. Tursiops truncatus (Europe assessment). The IUCN Red List of Threatened Species 2023: e.T22563A219008383. Accessed on 21 December 2023.

---

Hammond, P. S., Lacey, C., Gilles, A. et al. (2017). Estimates of cetacean abundance in European Atlantic waters in summer 2016 from the SCANS-III aerial and shipboard surveys. SCANS III final report. 40 pp.

---

Hammond, P. S., Macleod, K., Berggren, P., … and Vázquez, J. A. (2013). Cetacean abundance and distribution in European Atlantic shelf waters to inform conservation and management. Biological Conservation, 164, 107–122. https://doi.org/10.1016/j.biocon.2013.04.010

---

Hammond, P. S., Bearzi, G., Bjørge, A. et al. (2012). Tursiops truncatus. The IUCN Red List of Threatened Species 2012: e.T22563A17347397. http://dx.doi.org/10.2305/IUCN.UK.2012.RLTS.T22563A17347397.en.

---

Hastie, G. D., Wilson, B. and Thompson, P. M. (2006). Diving deep in a foraging hotspot: acoustic insights into bottlenose dolphin dive depths and feeding behaviour. Marine Biology, 148, 1181–1188. https://doi.org/10.1007/s00227-005-0143-x

---

Hayes, S. A., Josephson, E., Maze-Foley, K. and Rosel, P. E. (eds.) (2016). US Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 2016. NOAA Tech Memo NMFS NE 241. 274 pp. Available at https://www.fisheries.noaa.gov/tags/marine-mammal-stock-assessments

---

International Council for the Exploration of the Sea (ICES). (2018). Report of the Working Group on Marine Mammal Ecology (WGMME), 19-22 February 2018, La Rochelle, France. ICES CM 2018/ACOM:28. 120 pp

---

Janik, V. M. (2000a). Whistle matching in wild bottlenose dolphins (Tursiops truncatus). Science, 289, 1355–1357. https://doi.org/10.1126/science.289.5483.1355

---

Janik, V. M. (2000b). Food-related bray calls in wild bottlenose dolphins (Tursiops truncatus). Proceedings of the Royal Society B, 267(1446), 923–927. https://doi.org/10.1098/rspb.2000.1091

---

Jepson, P., Deaville, R., Barber, J., … and Law, R. J. (2016). PCB pollution continues to impact populations of orcas and other dolphins in European waters. Scientific Reports, 6, 18573. https://doi.org/10.1038/srep18573

---

Jones, G. J. and Sayigh, L. S. (2002). Geographic variation in rates of vocal production of free-ranging bottlenose dolphins. Marine Mammal Science, 18(2), 374–393. https://doi.org/10.1111/j.1748-7692.2002.tb01044.x.

---

Louis, M., Buanic, M., Lefeuvre, C., Le Nilliot, P., Ridoux, V. and Spitz, J. (2017). Strong bonds and small home range in a resident bottlenose dolphin community in a marine protected area (Brittany, France, Northeast Atlantic). Marine Mammal Science, 33(4), 1194–1203. https://doi.org/10.1111/mms.12419.

---

Luis, A. R., Couchinho, M. N. and dos Santos, M. E. (2014). Changes in the acoustic behavior of resident bottlenose dolphins near operating vessels. Marine Mammal Science, 30(4), 1417–1426. https://doi.org/10.1111/mms.12125

---

Mate, B., Rossbach, K. A., Nieukirk, S. L., Wells, R. S., Irvine, A. B., Scott, M. D. and Read, A. J. (1995). Satellite-monitored movements and dive behavior of a bottlenose dolphin (Tursiops truncatus) in Tampa Bay, Florida. Marine Mammal Science, 11(4), 452–463. https://doi.org/10.1111/j.1748-7692.1995.tb00669.x

---

Mattson, M. C., Mullin, K. D., Ingram, G. W. and Hoggard, W. (2006). Age structure and growth of the bottlenose dolphin (Tursiops truncatus) from strandings in the Mississippi Sound region of the north‐central Gulf of Mexico from 1986 to 2003. Marine Mammal Science, 22(3), 654–666. https://doi.org/10.1111/j.1748-7692.2006.00057.x

---

McFee, W. E. and Hopkins-Murphy, S. R. (2002). Bottlenose dolphin (Tursiops truncatus) strandings in South Carolina, 1992–1996. Fisheries Bulletin, 100, 258–265.

---

McLellan, W. A., Friedlaender, A. S., Mead, J. G. et al. (2002). Analysing 25 years of bottlenose dolphin (Tursiops truncatus) strandings along the Atlantic coast of the USA: do historic records support the coastal migratory stock hypothesis? Journal of Cetacean Research and Management, 4(3), 297–304. Available at https://archive.iwc.int/pages/search.php?search=%21collection15&k=

---

North Atlantic Marine Mammal Commission (NAMMCO). (2017). Report of the By-Catch Working Group, May 2017, Copenhagen, Denmark. Available at https://nammco.no/topics/sc-working-group-reports/

---

North Atlantic Marine Mammal Commission (NAMMCO). (2016). NAMMCO Annual Report 2016. NAMMCO, Tromsø, Norway, 363 pp. Available at https://nammco.no/topics/annual-reports/

---

Nowacek, S. M., Wells, R. S. and Solow, A. R. (2001). Short‐term effects of boat traffic on bottlenose dolphins, Tursiops truncatus, in Sarasota Bay, Florida. Marine Mammal Science, 17(4), 673–688. https://doi.org/10.1111/j.1748-7692.2001.tb01292.x

---

Nuuttila, H., Courtene-Jones W, Baulch S., Simon, M. and Evans, P. G. H. (2017). Don’t forget the porpoise: acoustic monitoring reveals fine scale temporal variation between bottlenose dolphin and harbour porpoise in Cardigan Bay SAC. Marine Biology, 164(3), 1–16. https://doi.org/10.1007/s00227-017-3081-5.

---

Oudejans, M. G., Visser, F., Englund, A., Rogan, E. and Ingram, S. N. (2015). Evidence for distinct coastal and offshore communities of bottlenose dolphins in the North East Atlantic. PLoS ONE, 10(4), e0122668. https://doi.org/10.1371/journal.pone.0122668

---

Quiñones, R., Giovannini, A., Raga, J. A. and Fernández, M. (2013). Intestinal helminth fauna of bottlenose dolphin Tursiops truncatus and common dolphin Delphinus delphis from the western Mediterranean. Journal of Parasitology, 99(3), 576–579. https://doi.org/10.1645/GE-3165.1

---

Read, A. J., Wells, R. S., Hohn, A. A., Scott, M. D. (1993). Patterns of growth in wild bottlenose dolphins, Tursiops truncatus. Journal of Zoology, 231, 107–123. https://doi.org/10.1111/j.1469-7998.1993.tb05356.x

---

Robinson, K. P. (2014). Agonistic intraspecific behavior in free-ranging bottlenose dolphins: calf-directed aggression and infanticidal tendencies by adult males. Marine Mammal Science, 30(1), 381–388. https://doi.org/10.1111/mms.12023.

---

Robinson, K. P., Sim, T. M. C., Culloch, R. M.., … and Pierce, G. C. (2017). Female reproductive success and calf survival in a North Sea coastal bottlenose dolphin (Tursiops truncatus) population. PLoS ONE, 12(9), 1–16. https://doi.org/10.1371/journal.pone.0185000.

---

Robinson, K. P., O’Brien, J. M., Berrow, S. D., … and Whooley, P. (2012) Discrete or not so discrete: Long distance movements by coastal bottlenose dolphins in UK and Irish waters. Journal of Cetacean Research and Management, 12(3), 365–371. https://archive.iwc.int/pages/search.php?search=%21collection15&k=

---

Ross, H. M. and Wilson, B. (1996). Violent interactions between bottlenose dolphins and harbour porpoises. Proceedings of the Royal Society B, 263, 283–286. https://doi.org/10.1098/rspb.1996.0043

---

Rossman, S., McCabe, E. B., Barros, N. B., … Wells, R. S. (2015). Foraging habits in a generalist predator: Sex and age influence habitat selection and resource use among bottlenose dolphins (Tursiops truncatus). Marine Mammal Science, 31(1), 155–168. https://doi.org/10.1111/mms.12143

---

Saavedra, C., Pierce, G. J., Gago, J., … and Santos, M. B. (2017). Factors driving patterns and trends in strandings of small cetaceans. Marine Biology, 164, 165. https://doi.org/10.1007/s00227-017-3200-3

---

Santos, M. B., Fernández, R., López, A., Martínez , J. A. and Pierce, G. J. (2007). Variability in the diet of bottlenose dolphin, Tursiops truncatus, in Galician waters, north-western Spain, 1990–2005. Journal of the Marine Biological Association of the United Kingdom, 87, 231–241. https://doi.org/10.1017/S0025315407055233

---

Santos, M. B. and Pierce, G. J. (2003). The diet of harbour porpoise (Phocoena phocoena) in the northeast Atlantic. Oceanography and Marine Biology, 41, 355–390. Available at https://www.researchgate.net/publication/228580356

---

Santos, M. B., Pierce, G. J., Reid, R. J.,Patterson, I. A. P., Ross, H. M. and Mente, E. (2001). Stomach contents of bottlenose dolphins in Scottish waters. Journal of the Marine Biological Association of the United Kingdom, 81(5), 873–878. https://doi.org/10.1017/S0025315401004714

---

Silber, G. K. and Fertl, D. (1995). Intentional beaching by bottlenose dolphins (Tursiops truncatus) in the Colorado River delta, Mexico. Aquatic Mammals, 21, 183–186. Available at https://www.aquaticmammalsjournal.org

---

Silva, M. A., Prieto, R., Magalhães, S., Seabra, M. I., Santos, R. S. and Hammond, P. S. (2008). Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure. Marine Biology, 156, 179–192. https://doi.org/10.1007/s00227-008-1075-z

---

Spitz, J., Rousseau, Y. and Ridoux, V. (2006). Diet overlap between harbour porpoise and bottlenose dolphin: An argument in favour of interference competition for food? Estuarine Coastal and Shelf Science, 70, 259–270. https://doi.org/10.1016/j.ecss.2006.04.020

---

Thayer, A. G., Read, A. J., Friedlaender, A. S., … and Rittmaster, K. A. (2003). Reproductive seasonality of western Atlantic bottlenose dolphins off North Carolina, U.S.A. Marine Mammal Science, 19(4), 617–629. https://doi.org/10.1111/j.1748-7692.2003.tb01120.x

---

Titcomb, E. M., Reif, J. S., Fair, P. A., Stavros, H.-C. W., Mazzoil, M., Bossart, G. D. and Schaefer, A. M. (2017). Blood mercury concentrations in common bottlenose dolphins from the Indian River Lagoon, Florida: Patterns of social distribution. Marine Mammal Science, 33(3), 771–784. https://doi.org/10.1111/mms.12390

---

Toth, J. L., Hohn, A. A., Able, K. W. and Gorgone, A. M. (2012). Defining bottlenose dolphin (Tursiops truncatus) stocks based on environmental, physical, and behavioral characteristics. Marine Mammal Science, 28, 461–478.  https://doi.org/10.1111/j.1748-7692.2011.00497.x

---

Van Beurden, S. J., IJsseldijk, L. L., Cremers, H. J.,Gröne, A., Verheije1, M. H. and Begeman, L. (2015). Anisakis spp. induced granulomatous dermatitis in a harbour porpoise Phocoena phocoena and a bottlenose dolphin Tursiops truncatus. Diseases of Aquatic Organisms, 112, 257–263. https://doi.org/10.3354/dao02818

---

Waring, G. T., Josephson, E., Maze-Foley, K. and Rosel, P. E. (eds.) (2015). US Atlantic and Gulf of Mexico Marine Mammal Stock Assessments – 2015. NOAA Tech Memo NMFS NE 238; 501 pp. Available at https://www.fisheries.noaa.gov/resource/document/us-atlantic-and-gulf-mexico-marine-mammal-stock-assessments-2005

---

Wells, R. S. and Scott, M. D. (1999). Bottlenose dolphin Tursiops truncatus (Montagu, 1821). In S. H. Ridgway and R. Harrison (eds.), Handbook of marine mammals, Vol. 6: The second book of dolphins and the porpoises, 137-182. San Diego, CA, USA: Academic Press.

---

Wells, R. S. and Scott, M. D. (1990). Estimating bottlenose dolphin population parameters from individual identification and capture-release techniques. Report of the International Whaling Commission, Special Issue 12, 407–415.

---

Wells, R. S., Rhinehart, H. L., Cunningham, P., Whaley, J., Baran, M., Koberna, M. and Costa, D. P. (1999). Long distance offshore movements of bottlenose dolphins. Marine Mammal Science, 15(4), 1098–1114. https://doi.org/10.1111/j.1748-7692.1999.tb00879.x

---

Woodard, J. C., Zam, S. G., Caldwell, D. K. and Caldwell, M. C.  (1969). Some parasitic diseases of dolphins. Veterinary Pathology, 6, 257–272. https://doi.org/10.1177/030098586900600307

International Union for the Conservation of Nature (IUCN) Red List

Whale and Dolphin Conservation Society – Common bottlenose dolphin

National Geographic – Common bottlenose dolphin

Seaworld – Bottlenose dolphin

Animal Diversity Web – Bottlenose dolphin

Videos:

BBC Earth – Dolphins beach themselves to feed

PBS – Dolphins beach themselves to catch fish

Bottlenose dolphin “bray” calls

Dolphin Research Center: Echolocation

Dolphin Research Center: Blindfolded imitation study

Dolphin Research Center: Followup blindfolded imitation study