FIELD GUIDE TO

Whales, Dolphins
and
Porpoises

Mark Carwardine

Illustrated by
Martin Camm
With additional illustrations by
Rebecca Robinson
Toni Llobet

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 For my Dad

BLOOMSBURY WILDLIFE
Bloomsbury Publishing Plc
50 Bedford Square, London, WC1B 3DP, UK
29 Earlsfort Terrace, Dublin 2, Ireland

BLOOMSBURY, BLOOMSBURY WILDLIFE and the Diana logo are trademarks of Bloomsbury Publishing Plc

First published in the United Kingdom 2022

This electronic edition published in 2022 by Bloomsbury Publishing Plc

Copyright © Mark Carwardine, 2022

Copyright © 2022 illustrations as credited on page 283

Mark Carwardine has asserted his right under the Copyright, Designs and Patents Act, 1988,
to be identified as Author of this work.

All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means,
electronic or mechanical, including photocopying, recording, or any information storage or retrieval system,
without prior permission in writing from the publishers.

Bloomsbury Publishing Plc does not have any control over, or responsibility for, any third-party websites
referred to in this book. All internet addresses given in this book were correct at the time of going to press.
The authors and publisher regret any inconvenience caused if addresses have changed or sites have ceased
to exist, but can accept no responsibility for any such changes.

A catalogue record for this book is available from the British Library.

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ISBN: PB: 978-1-4729-6997-2

ePub: 978-1-4729-6996-5
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Maps and scale drawings by Julie Dando, Fluke Art

Design by Julie Dando, Fluke Art

To find out more about our authors and books, visit www.bloomsbury.com and sign up for our newsletters

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 CONTENTS
How to use this book 4
The challenges of identification 6
Cetacean topography 10
Quick ID guides
Bow-riding dolphins and porpoises 12
Identifying whales by their flukes 14
Identifying whales by their blows 16
Identifying male beaked whales by their lower jaws 18
Cetaceans of the North Atlantic Ocean 20
Cetaceans of the South Atlantic Ocean 22
Cetaceans of the North Pacific Ocean 24
Cetaceans of the South Pacific Ocean 26
Cetaceans of the Indian Ocean 28
Cetaceans of the Arctic Ocean 30
Cetaceans of the Southern Ocean 31

Right and bowhead whales (family Balaenidae) 32

Pygmy right whale (family Neobalaenidae) 48
Grey whale (family Eschrichtiidae) 50
Rorquals (family Balaenopteridae) 54
Sperm whales (families Physeteridae, Kogiidae) 90
Narwhal and beluga (family Monodontidae) 98
Beaked whales (family Ziphiidae) 104
Blackfish (family Delphinidae) 154
Shorter-beaked oceanic dolphins (family Delphinidae) 180
Longer-beaked oceanic dolphins (family Delphinidae) 214
River dolphins (families Platanistidae, Iniidae, Pontoporiidae, Lipotidae) 252
Porpoises (family Phocoenidae) 262

Caring for whales, dolphins and porpoises 276

Glossary 277
Species checklist 280
Sources and resources 282
Artists’ biographies 283
Image credits 283
Acknowledgements 284
Index 286

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 HOW TO USE THIS BOOK
SPECIES NAMES AND AUTHORITY
The currently accepted common name of the species is
given in English. Many have alternative names – some
in common usage, many rare or regional and others
historical – and these are all provided as well. Names
in other languages are not given, purely for reasons
of space (each species has lots of different names in
almost every language, so the number of possibilities
is huge). Common names can vary but each species
has only one scientific name, which is given in italics.
The authority (the person credited with first publishing
the name of the species) is also provided. If the
original name has changed – e.g. the species has been
transferred to a new genus (not the one in which it was
originally described) – the original author’s name and
date are put in parentheses, for example: (Linnaeus,
1758). If it is not in parentheses, it is still in the original
genus.
IUCN STATUS
This is given for each species (together with the
year of assessment). This is the official status on the
International Union for Conservation of Nature Red List
of Threatened Species, which is the most authoritative,
objective and comprehensive list of species that have
been rigorously evaluated for their risk of extinction.
There are eight categories: Extinct (no reasonable doubt
that the last individual has died); Extinct in the Wild
(known only to survive in captivity or as a naturalised
population well outside its historic range); Critically
Endangered (extremely high risk of extinction in the
wild); Endangered (very high risk of extinction in the
wild); Vulnerable (high risk of extinction in the wild);
Near Threatened (likely to qualify for a threatened
category in the future); Least Concern (does not qualify
for a more at-risk category); and Data Deficient (not
enough data to make an assessment). A ninth category
– Not Evaluated – is for species not yet assessed.
POPULATION
Counting cetaceans is notoriously difficult and, inevitably,
estimates are of variable accuracy (and, of course, some
are more recent than others). However, while most
population figures should be viewed cautiously, this
section provides what is known and gives a guide to the
estimated abundance of the species.
CLASSIFICATION AND TAXONOMY
The guide follows the taxonomic arrangement, and
the scientific names of species and subspecies,
recommended by the Society for Marine Mammalogy’s
Committee on Taxonomy. This authoritative list is
4 INTRODUCTION
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updated annually. It is in a constant state of flux, with
new species being discovered and named and other
species being combined or split as more information
comes to light. Notes about the taxonomy of each
species are given as background information and to
point out any possible future changes.
SIZE
The species silhouette shows the typical size of an adult
to scale against a human diver (all human silhouettes
represent 1.8m). L = length, WT = weight. Size ranges
for adult males, adult females and calves are given,
together with the maximum lengths and weights
recorded to date. All measurements are in metric.
AT A GLANCE
An abbreviated list of key features is provided for quick
reference – outlining the main features to concentrate
on to achieve a correct identification – including
distribution, size and key physical and behavioural
features. Size is based on a simple scale: small (up to
3m), medium (4–10m), large (11–15m), and extra large
(more than 15m).
ILLUSTRATIONS
The main illustration for each species shows a typical
side view (if males and females look different they
are illustrated separately). Other images show the
opposite side view (if it is different), the upperside
and underside, any subspecies, regional variations or
age variations, and any relevant close-ups of dorsal
fins, beaks, flippers or other features (sometimes
including comparisons with similar species). Each
illustration has its own annotations, pointing out the
main distinguishing features useful for identification
and any other interesting characteristics. See Cetacean
Topography on p. 10 for a breakdown of the names of
the various parts of a cetacean. There are additional
illustrations showing the shape and size of the blow or
spout from behind (for the larger species) and a typical
dive sequence.
DISTRIBUTION
The distribution maps show the known and/or
presumed range of each species. For many, there is very
little information and putting together a distribution
map is like putting together a jigsaw puzzle with a few
or many parts missing. The accompanying text explains
in more detail what is known about the distribution
and where there are gaps in our knowledge, as well
as providing more information on habitat and depth
preferences (within the mapped range, each species
29/11/2021 12:09
 occurs only in appropriate habitats and depths), FOOD AND FEEDING
migrations and other movements, and extralimital This section includes short notes on the main prey
records. species and foraging behaviour.
BEHAVIOUR DIVING AND BLOW
Behaviour such as breaching, spyhopping, lobtailing and The way a cetacean surfaces – the dive sequence –
reaction to boats can provide valuable clues in cetacean can be very distinctive in many species and is often
identification, and details are given in this section. Bear a useful identification feature. This is illustrated,
in mind, though, that behaviour can vary enormously together with a short description of the main features
between individuals, from one region to another, with to look for. This section also includes information on
the seasons and according to many other factors. dive depth and dive times (with the maximum recorded
to date) and blow characteristics (if a distinctive blow
TEETH/BALEEN PLATES
is visible).
The number of teeth is normally given as a count for
each row (i.e. two rows in the upper jaw and two in the GROUP SIZE AND STRUCTURE
lower jaw). However, for simplicity, the figures in this There is a great deal of variation in cetacean
guide show the range in total number of teeth in the groupings, between individuals, regions and seasons,
upper and the same for the lower jaw. Baleen plates are but the most commonly observed group sizes and
present only in the upper jaw and the figure gives the structures are given, together with further information
total range for each side. on variability.

Species names Authority

NORTHERN BOTTLENOSE WHALE Huge, rounded melon (often overhanging

rostrum) with very steep (almost square-
Uniform medium to dark grey,
chocolate- to olive-brown or tan
ADULT MALE

IUCN status Hyperoodon ampullatus (Forster, 1770)

Two conical teeth
shaped) forehead as it matures coloration (brownish or yellowish
tinge may be enhanced by film or
at tip of lower Much of melon and face (back to Robust patches of diatoms)
The northern bottlenose whale is the largest beaked whale in the North Atlantic, one of the best-known members of jaw (rarely visible eyes) white or cream-coloured body
the family and one of the few to have been targeted by whalers on a large scale. The heads of males and females look outside closed
Population so different that early anatomists believed them to be two separate species. References to ‘bottlenose whales’ in the
northern North Pacific refer to Baird’s beaked whales.
mouth and often
wear down or
Distinct junction
between beak and
Tip usually
pointed
melon (but no crease) Prominent, erect but
fall out)
IUCN status Near Threatened (2020). relatively small, falcate

Classification Population Probably in the order of 20,000+. Possibly c. 100,000 before intense whaling began in the 1880s, but this dorsal fin (up to 30cm)
was reduced to a few tens of thousands by the time whaling ceased in the 1970s. Trend unknown. two-thirds of the way
along back
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Taxonomy Other names Northern bottle-nosed whale, bottlehead, flathead, steephead, common bottlenose whale.
DISTRIBUTION Cold temperate to Arctic waters in the North Atlantic. Ranges from the ice edge to at least c. 37°N.
However, rarely seen south of c. 55°N – with one notable exception: a well-studied population in a submarine canyon
Other names called The Gully, 200km south-east of the Atlantic coast of Halifax, Nova Scotia, Canada. Usually frequents waters
deeper than 500m (with a preference for 800–1,800m) over the continental slope. Rarely strays over the continental Medium-length, thick, well- Two shallow,
Small, blunt
flippers (fit Little or no May be mottled with
defined beak (highly variable V-shaped throat into ‘flipper linear scarring round or oval white
shelf, except in submarine canyons, and prefers areas with complex seabed topography, such as the continental shelf
appearance – beak looks grooves pockets’ on scarring (healed bites from
edge, oceanic islands and seamounts. Generally occurs in open water.
Distribution BEHAVIOUR Males may use their large, bulbous heads to headbutt one other. When resting at the surface, both sexes
smaller relative to melon
growing larger)
sides) cookiecutter sharks)

and all ages may hang in the water at a 45° angle, with the entire melon and beak above the surface. Breaching and
ADULT MALE GREY FORM
lobtailing are not uncommon. Can be very curious towards boats and even large ships, and will often approach closely.
FOOD AND FEEDING Mainly deepwater squid; sometimes fish; rarely prawns, sea cucumbers and starfish. Most

Behaviour feeding appears to be on or near seabed in deep water; probably suction-feeder.

TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only (up to 5cm tall).
GROUP SIZE AND STRUCTURE Typically 1–10, rarely more than 20. Group size varies according to region. There is
some segregation by age and sex. Females appear to form loose, fluid associations, but pairs of males form long-term
relationships that can last anything from days to 1–2 years.

Food and Melon bulbous but Much of melon and

ADULT FEMALE
Huge melon with very steep,
OLD ADULT MALE

feeding relatively smaller and less

square than adult male
face slightly paler

No linear scarring
flattened forehead (giving
distinctly squarish profile)

SIZE
L:  7.5–9m,  7–8.5m; Much of melon and
Teeth/baleen WT: c. 5–8t; MAX: 9.8m, 10t
Calf – L: 3–3.5m; WT: c. 300kg
face very pale (white
or cream-coloured)

No erupted Beak may be thicker

AT A GLANCE Cold, deep waters of North
teeth than on male 80°

Atlantic • Medium size (larger than other

Group size and 70°
beaked whales in region) • Grey, tan or
brownish coloration • Huge, squared-off,
structure 60°

50°
bulbous white or cream-coloured melon
(especially in male) • Medium-length, thick,
well-defined beak • Prominent falcate dorsal
fin two-thirds of the way along back • Little
40°

DIVE Sequence Rolls forward, sometimes with head, back and dorsal fin visible simultaneously; rarely flukes. 30°
or no linear scarring • Male’s teeth not clearly
• Depth Routinely dives to 800+m (average in one study 1,065m); maximum recorded 2,339m. • Duration Routinely 20°
visible • Often inquisitive and may approach
30–40 minutes; maximum recorded 94 minutes; possibly capable of 2 hours. 10°
stationary vessels
BLOW Low, puffy blow (1–2m), often clearly visible and canted forward.

114 BEAKED WHALES NORTHERN BOTTLENOSE WHALE 115

Diving and blow Size Distribution map At a glance

HOW TO USE THIS BOOK 5

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 THE CHALLENGES OF IDENTIFICATION
Identifying whales, dolphins and porpoises at sea can Remember that only a small portion of the animal (the
be enormously satisfying, but also quite challenging. In top of the head and back, for example) may be visible
fact, it can be so difficult that even the world’s experts at any one time. Larger species don’t necessarily show
are unable to identify every species they encounter – on more of themselves than smaller species, so size can be
most official surveys, at least some sightings have to be quite deceptive. It is therefore better to use four simple
logged as ‘unidentified’. categories: small (up to 3m), medium (4–10m), large
The trick is to use a relatively simple process of (11–15m), and extra large (more than 15m).
elimination, running through a mental checklist of 14
key features every time a new animal is encountered at
sea. It is not often possible to use all of these features
together and one alone is rarely enough for a positive Narwhal
identification. The best approach is to gather information
on as many as possible before drawing any firm
conclusions.
1. Geographical location There is not a single place
in the world where every cetacean species has been Killer whale
Sperm whale
recorded. In fact, there are not many places with records
of more than a few dozen, so this immediately helps to 4. Unusual features Some cetaceans have very
cut down on the number of possibilities. unusual features, which can be used for a quick
2. Habitat Just as cheetahs live on open plains rather identification. These include the extraordinary long tusk
than in jungles, and snow leopards prefer mountains of the male narwhal, the enormous dorsal fin of the male
to wetlands, most whales, dolphins and porpoises are killer whale and the wrinkly skin of the sperm whale.
adapted to specific marine or freshwater habitats. In 5. Dorsal fin The size, shape and position of the dorsal
this respect, marine charts can be surprisingly useful fin varies greatly between species and is a particularly
identification aids. Knowing the underwater topography useful aid to identification. Don’t forget to look for any
could help to tell the difference between a minke distinctive colours or markings on the fin.
whale (normally found over the continental shelf) and
a superficially similar northern bottlenose whale (more
likely to be seen over submarine canyons or in deep
waters offshore).
3. Size It is difficult to estimate size accurately at sea,
unless a direct comparison can be made with the length Pacific white-sided dolphin
of the boat, a passing bird or an object in the water.

Spectacled porpoise
SMALL (up to 3m) – Vaquita

MEDIUM (4–10m) – Cuvier’s beaked whale Pygmy beaked whale

LARGE (11–15m) – Bryde’s whale

EXTRA LARGE (more than 15m) – Blue whale

6 INTRODUCTION

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 6. Flippers The length, colour and shape of the
flippers, as well as their position on the animal’s body,
vary greatly from one species to another. It is not always
possible to see them, but flippers can be useful for
identification in some species – in the humpback whale,
for example, they are unmistakable.
Bowhead whale
Humpback whale
Long-finned pilot whale
7. Body shape Much of the time, whales, dolphins
and porpoises do not show enough of themselves to
provide an overall impression of their shape. Sometimes,
however, this can be a useful feature. Is the animal
stocky or slim, for example? The shape of the melon
(forehead) can also be distinctive.
Omura’s whale
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Franciscana
Rough-toothed dolphin
Atlantic spotted dolphin
8. Beak The presence or absence of a prominent beak
is a particularly useful identification feature in toothed
whales. Broadly speaking, river dolphins, beaked whales
and half the oceanic dolphins have prominent beaks,
while porpoises, belugas and narwhals, killer whales
and their allies, sperm whales and the remaining
oceanic dolphins do not. There is also great variation
in the beak length from one species to another. And try
to see if there is a smooth transition from the top of
the head to the end of the snout (as in rough-toothed
dolphins, for example) or a distinct crease (as in Atlantic
spotted dolphins).
Deraniyagala’s beaked whale
Dwarf sperm whale
THE CHALLENGES OF IDENTIFICATION 7
29/11/2021 12:09
 9. Colour and markings Many cetaceans are
surprisingly colourful and have distinctive markings
such as body stripes or eye patches. Bear in mind that Heaviside’s dolphin
colours at sea vary according to water clarity and light
conditions, and the animal can appear much darker
than normal if viewed against the sun.

Beluga

Pygmy right whale

10. Flukes The flukes can be important in identifying

larger whales. Some species lift their flukes high
into the air before they dive, while others do not, and
that alone can help to tell one from another. It is also
worth checking the shape of the flukes, looking for any
distinctive markings and noticing whether or not there is
a deep notch between the trailing edges.
Antarctic minke whale

Grey whale Sei whale

North Pacific right whale

Southern right whale

11. Blow or spout The blow is particularly distinctive

in larger whales. It varies in height, shape, density and
visibility between species and can be extremely useful for
identification, especially on calmer days. But identifying
a blow is not easy – if it is raining or windy, the blow can
be bent out of shape, and there are variations between
individuals – and the first blow after a deep dive tends to
Sperm whale be stronger than the rest. However, experienced observers
can often tell one species from another just by the blow,
even from a considerable distance.

8 INTRODUCTION

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12. Dive sequence The dive sequence can be
surprisingly distinctive in many species. Variations
include: the angle at which the head breaks the surface;
how much of the head and beak (if present) are visible;
whether or not the dorsal fin and blowhole are visible
at the same time; whether the animal arches its back to
dive (and how much it arches) or whether it merely sinks
below the surface; the time interval between breaths;
and the number of breaths before a deep dive.
13. Behaviour Some species are more active at the
surface than others, so any unusual behaviour can
sometimes be useful for identification purposes. Did it
leap out of the water, for example, or was its behaviour
quite cryptic? The reaction to boats can also be helpful:
common bottlenose dolphins may race over and bow-
ride, while the similar-looking Atlantic humpback
dolphins tend to be more shy and will not bow-ride.
Spinner dolphin
14. Group size Since some species are highly
gregarious, while others tend to live alone or in small
groups, it is worth noting the number of animals seen
together. Estimating group size is notoriously difficult,
because the animals are mobile and frequently change
direction and, at the time of counting, any number of
000 intro WDP FG.indd 9
Fin whale
Strap-toothed beaked whale
Dwarf sperm whale
them can be hidden beneath the surface. Estimating the
size of a large school of active dolphins is especially
challenging – the tendency is usually to underestimate.
It is often tempting to guess the identification of an
unusual whale, dolphin or porpoise that you have
not seen very clearly. However, working hard at
identification – and then enjoying the satisfaction of
knowing that an animal has been identified correctly –
is what makes a real expert in the long term. It is
perfectly acceptable to record simply ‘unidentified
dolphin’, ‘unidentified whale’ or ‘unidentified beaked
whale’ if a more accurate identification is not possible.
If you write detailed notes at the time, and then see the
same species again in the future, it may be possible to
turn a sighting previously recorded as ‘unidentified’ into
a positive identification, days, weeks, months or even
years later.
It does get easier with practice. After a while, you
look at a whale and it triggers a switch in your brain.
You know what it is likely to be – a single species or,
perhaps, several possibilities – from its ‘jizz’ (the
overall impression). At a glance, you get an
overall impression that is more instinctive
than something that can easily be put
into words.
Part of the fun is that cetaceans
are unpredictable. Never say ‘never’
on a whale-watching trip: just
because the distribution maps
suggest you’re unlikely to see a
particular species in a particular area, it doesn’t mean
to say it couldn’t pop up anywhere at any time; and just
because the guidebooks say that one species of whale
doesn’t fluke, it doesn’t mean to say the particular
individual next to your boat won’t prove everyone wrong
by lifting its tail high into the air.
THE CHALLENGES OF IDENTIFICATION 9
29/11/2021 12:09
 CETACEAN TOPOGRAPHY
two blowholes
rostrum or eye PARTS OF A BALEEN WHALE
ear
upper jaw

baleen plates tailstock or

(suspended from caudal peduncle
upper jaw –
visible only when
mouth open)
mouthline
flukes

mandible or anus
lower jaw axilla
keel
(‘armpit’) genital slit
flipper or
ventral pleats or throat pectoral fin navel or umbilicus
grooves (present only on
rorquals) (underside)
chevron
PARTS OF A BALEEN WHALE
two blowholes
splashguard dorsal fin
fluke

median notch

dorsal ridge (visible

rostral ridge (present or present in only a
only on rorquals) few species)

PARTS OF A TOOTHED WHALE

blowhole (one)
dorsal fin
melon
eye tailstock or
caudal peduncle
rostrum ear
or upper
jaw
flukes

median
notch

beak or
snout
anus keel

mandible or axilla (‘armpit’) genital slit

mouthline
lower jaw navel or umbilicus
flipper or pectoral fin

10 INTRODUCTION

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 PARTS OF A TOOTHED WHALE DORSAL FIN

single blowhole leading edge trailing

edge

anterior insertion nicks and

point notches
posterior insertion
point

crease

lower boundary of
dorsal fin
HOW TO SEX A CETACEAN

female mammary slits

umbilicus genital aperture anus

male
dian notch

PARTS OF A CETACEAN SKELETON

thoracic vertebrae no skeletal support for
dorsal fin
cervical vertebrae (fused
in some species)
lumbar vertebrae
cranium
rostrum caudal vertebrae

ribcage no skeletal support

mandible for flukes
scapula
(shoulder blade) phalanges
(finger bones)

CETACEAN TOPOGRAPHY 11

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 QUICK ID GUIDES
BOW-RIDING DOLPHINS AND PORPOISES

False killer whale

Short-finned
pilot whale
Risso’s dolphin

Pygmy killer
whale

Long-finned pilot whale

Killer whale

Melon-headed whale Northern right

whale dolphin Southern right
whale dolphin

12 QUICK ID GUIDES

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 Clymene dolphin

Rough-toothed
dolphin
Indo-Pacific Atlantic
bottlenose spotted dolphin
Striped dolphin
dolphin

Common Pantropical
bottlenose spotted dolphin
dolphin Common dolphin
Spinner dolphin

Dusky dolphin Fraser’s dolphin

White-beaked Hourglass Dall’s porpoise

dolphin dolphin

Harbour porpoise
Chilean dolphin
Atlantic white-
sided dolphin

Hector’s dolphin

Peale’s dolphin
Pacific white-sided dolphin Heaviside’s dolphin
Commerson’s dolphin

BOW-RIDING DOLPHINS AND PORPOISES 13

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 IDENTIFYING WHALES BY THEIR FLUKES

North Atlantic right whale

North Pacific right whale

Southern right whale

Bowhead whale

14 QUICK ID GUIDES

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 Blue whale

Grey whale

Humpback whale

Killer whale

Sperm whale

IDENTIFYING WHALES BY THEIR FLUKES 15

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 IDENTIFYING WHALES BY THEIR BLOWS
The height and intensity of a whale’s blow, or spout, depends on many factors, including behaviour, the size of the
individual, when it occurs during the surfacing sequence, air temperature, the quality of the light and wind conditions.
It is therefore important to bear in mind that a single whale’s blow can vary from virtually invisible to tall and dramatic.
Indeed, blow heights have been seriously underestimated in the past, not least because they largely disappear against
a pale sea or sky background.
These illustrations show picture-perfect blows (from behind the whales) on the first surfacing after a long dive – in
ideal conditions – and represent the maximum heights. Not all cetaceans have clearly visible blows, but these are the
ones that are most distinctive and most useful for identification purposes.

North Atlantic right whale North Pacific right whale

up to 7m up to 7m

Southern right whale Bowhead whale

up to 7m up to 7m

Blue whale Fin whale

up to 12m up to 10m

16 QUICK ID GUIDES

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 Sei whale Bryde’s whale Omura’s whale
up to 10m up to 10m up to 7m

Common minke whale Antarctic minke whale

up to 3m up to 4m

Humpback whale
up to 10m

Grey whale Sperm whale

up to 5m up to 6m

Killer whale
up to 5m

Long-finned pilot whale Short-finned pilot whale

up to 1m up to 1m

IDENTIFYING WHALES BY THEIR BLOWS 17

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 IDENTIFYING MALE BEAKED WHALES BY THEIR LOWER JAWS
With a few exceptions, most adult male beaked Flattened, triangular, Posterior teeth hidden
forward-pointing teeth when mouth closed
whales have dramatically reduced dentition, with
only one or two pairs of erupted teeth in the lower
jaw and none in the upper jaw (most females have
no erupted teeth at all). The number, position, size
and shape of the males’ teeth are usually the best
clues to their identification. Anterior teeth up to 10cm high and
10cm at longest point (twice as large
as peg-like posterior teeth)
Flattened, Posterior teeth Baird’s beaked whale
triangular, forward- hidden when
pointing teeth mouth closed Two pairs of teeth near tip
of lower jaw (anterior tooth
much larger than posterior)

Anterior teeth twice as large

as peg-like posterior teeth
Arnoux’s beaked whale Sato’s beaked whale

Two forward-pointing, conical Two teeth erupt at tip of

teeth c. 8cm long (including lower jaw (lean forward)
portion buried in jawbone)

Rarely second pair Northern bottlenose whale

erupts behind main pair
Cuvier’s beaked whale Pair of apical teeth at tip
of lower jaw (present in Only beaked whale with
Two teeth erupt at male and female – but full set of functional
tip of lower jaw unerupted in female) teeth in both jaws
(lean forward)

Southern bottlenose whale Shepherd’s beaked whale

Pair of pear-shaped Two relatively large, laterally
teeth at tip of compressed triangular teeth
lower jaw c. 1–2cm behind tip of lower jaw

Longman’s beaked whale Perrin’s beaked whale

Sharply Posterior Teeth very large and

Two small, pointed margin of laterally compressed (taller
conical teeth crown tooth convex than they are wide – teeth
at apex of arch in ginkgo-toothed usually
Anterior wider than they are tall)
margin of
tooth nearly
straight

Pygmy beaked whale Deraniyagala’s beaked whale

18 QUICK ID GUIDES

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 Laterally flattened Teeth remarkably similar Tooth Tooth as wide or wider than
Tooth shape
Teeth tilt teeth can be quite (in size and shape) to compressed it is tall (6.5cm by 11.5cm) cf.
resembles leaf
slightly wide (c. 6–10cm) those of ginkgo-toothed laterally Deraniyagala’s beaked whale
of ginkgo tree
forward and up to 10cm tall beaked whale (taller than it is wide)
(variable)

Gray’s beaked whale Ginkgo-toothed beaked whale

Two relatively large, laterally Laterally flattened teeth
compressed triangular teeth less splay slightly inward (cf. Tooth 16–17cm high (including
than 1cm behind tip of lower jaw Andrews’ beaked whale) portion buried in jawbone),
9cm at widest point

Hector’s beaked whale Hubbs’ beaked whale

Out of socket, tooth measures c. 15–18cm
Flattened tooth tall, 8–9cm wide, 4.5cm deep (but typically
less than 2cm extends above gums) Flattened, Teeth can be heavily
erupts from top triangular
of bony arch worn in older animals
teeth
(angled forward
at c. 45°)

Blainville’s beaked whale Sowerby’s beaked whale

Acorn-shaped Teeth relatively small Tooth 30cm high,

teeth angled (c. 5cm) and may become 10cm wide, 2.5cm thick
slightly forward heavily worn with age

True’s beaked whale Stejneger’s beaked whale

Older animals may Laterally flattened teeth Tooth up to 14cm

Tooth c. 7cm high, have deeply worn splay slightly outward high (including
Triangular, flattened 4cm long, 1cm deep ‘groove’ or ‘hollow’ at up to 20° (cf. Hubbs’ portion buried in
tooth (may wear
on anterior margin beaked whale) jaw), 8cm at
down with age)
widest point

Gervais’ beaked whale Andrews’ beaked whale

Long, curved,
Spade-shaped teeth lean
flattened white teeth
backwards at c. 45° angle
(up to 33cm)

Strap-toothed beaked whale Spade-toothed whale

IDENTIFYING MALE BEAKED WHALES BY THEIR LOWER JAWS 19

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 CETACEANS OF THE NORTH ATLANTIC OCEAN
(including the Caribbean Sea, Gulf of Mexico, Mediterranean Sea, Black Sea, Baltic Sea)
Please note: many species on this spread have very restricted ranges within this broad region, and it is not impossible
to see species from other parts of the world outside their normal range. Relative sizes (for average-length males) are
correct for the region.

Blue whale

Common minke
whale

Fin whale

Narwhal

Sei whale

Beluga

Bryde’s whale

Rice’s whale

Omura’s whale
Harbour porpoise

Pygmy sperm
Dwarf sperm whale
whale
Humpback whale

Sperm whale

20 QUICK ID GUIDES

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 North Atlantic right
whale

Killer whale

Risso’s dolphin

Long-finned pilot
whale

Bowhead whale

Short-finned pilot
whale

Blainville’s beaked
whale Sowerby’s beaked
whale

False killer
whale

Cuvier’s beaked
whale Gervais’ beaked
Melon-headed
whale whale

True’s beaked
Pygmy killer whale
whale
Northern bottlenose
whale

Rough-toothed
Clymene dolphin dolphin White-beaked
dolphin
Atlantic spotted
dolphin

Pantropical
spotted dolphin Spinner dolphin

Guiana dolphin Atlantic white-sided

dolphin Common bottlenose
dolphin

Fraser’s dolphin
Striped dolphin
Common dolphin Atlantic humpback
dolphin

CETACEANS OF THE NORTH ATLANTIC OCEAN 21

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 CETACEANS OF THE SOUTH ATLANTIC OCEAN
Please note: many species on this spread have very restricted ranges within this broad region, and it is not impossible
to see species from other parts of the world outside their normal range. Relative sizes (for average-length males) are
correct for the region.

Blue whale

Dwarf minke whale

Fin whale

Franciscana

Sei whale

Antarctic minke
whale
Burmeister’s
porpoise

Pygmy right whale

Spectacled
porpoise
Bryde’s whale

Omura’s whale
Southern right whale

Pygmy sperm whale

Humpback
whale

Dwarf sperm
whale

Sperm whale

22 QUICK ID GUIDES

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 Southern bottlenose
whale
Arnoux’s beaked whale

Killer whale
Shepherd’s beaked whale
Gray’s beaked whale

Hector’s beaked
whale Andrews’ beaked
Long-finned pilot whale
whale

Spade-toothed whale
Strap-toothed beaked
whale
Short-finned pilot whale

True’s beaked whale

Blainville’s beaked whale

False killer whale

Spinner dolphin Cuvier’s beaked whale Gervais’ beaked whale

Melon-headed
whale

Hourglass Southern right

dolphin whale dolphin
Risso’s dolphin
Pygmy killer whale
Commerson’s
Peale’s dolphin dolphin
Rough-toothed
Clymene dolphin
Heaviside’s
dolphin
dolphin
Dusky dolphin
Chilean dolphin
Pantropical
spotted dolphin
Guiana dolphin
Atlantic spotted Common bottlenose
dolphin dolphin

Common dolphin Striped dolphin

Fraser’s dolphin Atlantic humpback dolphin

CETACEANS OF THE SOUTH ATLANTIC OCEAN 23

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 CETACEANS OF THE NORTH PACIFIC OCEAN
(including the Gulf of California, Gulf of Alaska, Bering Sea, Sea of Okhotsk, Sea of Japan, Philippine Sea,
Yellow Sea, East China Sea, South China Sea)
Please note: many species on this spread have very restricted ranges within this broad region, and it is not impossible
to see species from other parts of the world outside their normal range. Relative sizes (for average-length males) are
correct for the region.

Blue whale

Common minke
whale

Fin whale

Dall’s porpoise Harbour

porpoise
Sei whale
Vaquita

Indo-Pacific Narrow-ridged
Bryde’s finless porpoise finless porpoise
whale

Omura’s whale
Humpback whale

Dwarf sperm
whale Pygmy sperm
Grey whale whale

Beluga

Sperm whale

24 QUICK ID GUIDES

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 North Pacific right whale
Cuvier’s beaked whale

Bowhead whale
Killer whale

Melon-headed whale

Baird’s beaked whale

Short-finned pilot whale Pygmy killer whale

False killer whale Blainville’s beaked whale Sato's beaked whale

Perrin’s beaked
whale Hubbs’ beaked whale Ginkgo-toothed beaked
whale

Deraniyagala’s
Peruvian beaked beaked whale
Stejneger’s beaked whale whale

Longman’s beaked
Risso’s dolphin Indo-Pacific whale
humpback dolphin
Rough-toothed
dolphin Irrawaddy dolphin
Common
bottlenose dolphin

Indo-Pacific Pantropical
spotted dolphin Spinner dolphin
bottlenose dolphin
Striped dolphin
Pacific white-sided
dolphin
Common dolphin Fraser’s dolphin
Northern right
whale dolphin

CETACEANS OF THE NORTH PACIFIC OCEAN 25

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 CETACEANS OF THE SOUTH PACIFIC OCEAN
Please note: many species on this spread have very restricted ranges within this broad region, and it is not impossible
to see species from other parts of the world outside their normal range. Relative sizes (for average-length males) are
correct for the region.

Omura’s whale

Blue whale

Dwarf minke whale

Fin whale

Sei whale
Antarctic minke
whale
Burmeister’s
Indo-Pacific
porpoise
finless porpoise

Pygmy right whale

Spectacled
porpoise
Bryde’s whale

Southern right whale

Humpback whale Pygmy sperm

whale

Dwarf sperm
whale

Sperm whale

26 QUICK ID GUIDES

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 Peruvian beaked whale
Arnoux’s beaked whale

Killer whale Shepherd’s beaked

whale
Gray’s beaked whale

Hector’s beaked
whale
Long-finned pilot whale Andrews’ beaked
whale

Strap-toothed Spade-toothed
beaked whale whale

True’s beaked whale

Short-finned pilot whale

Blainville’s beaked Deraniyagala’s beaked

whale whale

False killer whale Cuvier’s beaked

whale

Ginkgo-toothed beaked
whale Longman’s beaked
whale
Pygmy killer whale

Melon-headed Southern bottlenose

whale whale
Australian snubfin Hourglass dolphin
dolphin
Irrawaddy Peale’s dolphin
dolphin Dusky dolphin
Pantropical
Chilean dolphin
spotted dolphin
Rough-toothed
dolphin Commerson’s
dolphin
Indo-Pacific
Spinner
bottlenose dolphin Hector’s dolphin
dolphin
Australian
Common
humpback dolphin
bottlenose dolphin
Southern right whale Indo-Pacific
dolphin humpback dolphin
Striped dolphin

Fraser’s dolphin Risso’s dolphin

Common dolphin

CETACEANS OF THE SOUTH PACIFIC OCEAN 27

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 CETACEANS OF THE INDIAN OCEAN
(including the Mozambique Channel, Red Sea, Persian Gulf, Arabian Sea, Bay of Bengal)
Please note: many species on this spread have very restricted ranges within this broad region, and it is not impossible
to see species from other parts of the world outside their normal range. Relative sizes (for average-length males) are
correct for the region.

Blue whale

Irrawaddy
dolphin
Dwarf minke
whale

Fin whale

Australian snubfin
dolphin

Sei whale
Antarctic minke
whale

Omura’s whale

Bryde’s whale

Pygmy right whale

Southern right whale

Pygmy sperm whale

Humpback whale

Dwarf sperm whale

Sperm whale

28 QUICK ID GUIDES

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 Ginkgo-toothed beaked whale
Arnoux’s beaked whale

Killer whale Shepherd’s beaked

whale
Gray’s beaked whale

Hector’s beaked whale

Long-finned pilot
whale Andrews’ beaked whale

Strap-toothed Spade-toothed whale

beaked whale

Short-finned pilot True’s beaked whale

whale

Deraniyagala’s beaked
Blainville’s beaked
whale
whale

False killer whale Cuvier’s beaked whale

Indo-Pacific
finless porpoise Longman’s beaked whale
Pygmy killer
whale Spectacled
porpoise
Melon-headed
whale Southern bottlenose
Southern right whale
whale dolphin Rough-toothed
dolphin
Commerson’s Hourglass dolphin
dolphin

Pantropical
Indo-Pacific
spotted dolphin
humpback dolphin Indian Ocean
humpback Dusky dolphin
dolphin
Spinner dolphin
Australian
humpback dolphin Indo-Pacific
bottlenose dolphin Common bottlenose
dolphin
Striped dolphin

Common dolphin Fraser’s dolphin

Risso’s dolphin

CETACEANS OF THE INDIAN OCEAN 29

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 CETACEANS OF THE ARCTIC OCEAN
(including the Greenland Sea, Barents Sea, White Sea, Kara Sea, Laptev Sea, East Siberian Sea, Chukchi Sea,
Beaufort Sea, Davis Strait, Baffin Bay, Hudson Bay)
Please note: many species on this spread have very restricted ranges within this broad region, and it is not impossible
to see species from other parts of the world outside their normal range. Relative sizes (for average-length males) are
correct for the region.

Blue whale

Harbour
porpoise Common minke whale

Fin whale Beluga

Humpback whale
Sei whale

Narwhal
Sowerby’s beaked
whale

Grey whale

Northern bottlenose whale

Common bottlenose
dolphin
Baird’s beaked whale
Killer whale

Long-finned pilot whale

Sperm whale

White-beaked
dolphin Atlantic white-sided
Bowhead whale
dolphin

30 QUICK ID GUIDES

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 CETACEANS OF THE SOUTHERN OCEAN
(including the Weddell Sea and the Ross Sea)
Please note: many species on this spread have very restricted ranges within this broad region (south of 60°S), and it
is not impossible to see species from other parts of the world outside their normal range. Relative sizes (for average-
length males) are correct for the region.

Arnoux’s beaked whale

Sperm whale

Cuvier’s beaked whale

Southern right whale

Southern bottlenose whale

Spectacled porpoise
Pygmy right whale

Humpback whale
Antarctic minke whale

Gray’s beaked whale

Dwarf minke whale

Sei whale Strap-toothed

beaked whale

Long-finned pilot
whale

Fin whale
Southern right
whale dolphin
Commerson’s
dolphin

Killer whale

Blue whale

Hourglass dolphin Peale’s dolphin

CETACEANS OF THE ARCTIC OCEAN AND SOUTHERN OCEAN 31

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 NORTH ATLANTIC RIGHT WHALE
Eubalaena glacialis (Müller, 1776)

The North Atlantic right whale is one of the most closely studied – and most endangered – large whales in the world.
The few animals around today are the survivors of nearly 1,000 years of commercial exploitation and, although hunting
has stopped, they face new human-induced threats and are widely considered to be in very real danger of extinction.
IUCN status Critically Endangered (2020).
Population c. 356 (early 2021). Pre-whaling population unknown (one estimate of 9,000–21,000 based on ecological
carrying capacity). Reduced to fewer than 100 by 1935. Decreasing (declined dramatically since a post-whaling peak of
483 in 2011). In recent years, more animals have died than have been born.
Classification Mysticeti, family Balaenidae.
Taxonomy No recognised forms or subspecies; split from the North Pacific right whale in 2000 due to genetic
differences between both the whales and their lice (the two species were previously lumped together as ‘northern
right whale’, Eubalaena glacialis).
Other names Atlantic right whale, northern right whale.
DISTRIBUTION Historically, there were two largely isolated populations on either side of the North Atlantic. However,
the eastern population is considered functionally extinct. Western North Atlantic animals are highly mobile, but only
pregnant females and a few other animals undertake predictable seasonal migrations. Occurs mainly in temperate and
sub-polar coastal waters, including shallow basins, and in relatively deeper areas over the continental shelf.
Eastern population Historically, North Atlantic right whales probably ranged from the only known breeding ground,
in Cintra Bay, off Western Sahara, to feeding grounds in the Bay of Biscay, off western Britain, around Iceland, across
the Norwegian Sea to the North Cape (northern Norway) and possibly in the Mediterranean. Rarely seen in the north-
east Atlantic nowadays, all identified individuals since 1960 have been migrants from the west, so a remnant eastern
population seems unlikely (although a calf with foetal folds – suggesting that it had been born nearby – was observed
in the Canary Islands, Spain, in December 2020).
Western population There has been a broad-scale distribution shift since 2011. Due to warming waters, sightings have
decreased dramatically in the Gulf of Maine (with the exception of narrow bands across the Scotian Shelf) and the
Bay of Fundy, which used to be critical foraging grounds. There has been a general northeastward shift, with sightings
increasing along Roseway Basin and in the Gulf of St Lawrence. Acoustic data also reveal an increase in occurrence
further into the Atlantic, from north of Cape Hatteras to south of Cape Cod, and along the edge of the continental shelf.
In November and December, pregnant females (sometimes accompanied by a small number of juveniles and non-
calving females) migrate south along the eastern seaboard of North America to the only known calving ground, in the
relatively sheltered, shallow coastal waters of northern Florida and southern Georgia (mainly between Savannah and
St Augustine); for some individuals, the calving grounds may extend as far north as Cape Fear, North Carolina, and
occasionally further west into the Gulf of Mexico. They return to northern feeding grounds in March and April. Most
animals (including most juveniles and adult males) do not migrate to these calving grounds; their wintering range is
unknown.

DIVE Sequence May lift head almost clear of water on surfacing after long dive;
flukes often raised quite high before deep dive. • Depth Often at or near surface;
can easily reach near-bottom depths (200+m) on continental shelf feeding grounds.
• Duration Typical deep-feeding dives 10–20 minutes; maximum 40 minutes.
BLOW Bushy, V-shaped (when viewed from in front or behind) or oval and bushy
(from side). • Up to 7m high (height highly variable). • Jets often asymmetric
in height. • V more widely spaced at base than in humpback (which sometimes
produces V-shaped blows).

32 RIGHT AND BOWHEAD WHALES

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 Top of head sprinkled
with callosities (naturally
light to dark grey, but
appear white, cream or
yellowish due to presence
of whale lice)
Upper
margin of
lower ‘lip’
serrated
Very strongly Extremely stocky body ADULT
arched jawline (bordering on rotund)
Individuals sometimes appear
Massive head Maximum girth can
mottled (caused by uneven
(up to one-third exceed c. 60 per cent
sloughing of patches of skin)
of body length) of total length
Smooth, broad back with no ADULT
dorsal fin or ridge
Predominantly black

Largest
callosity (the
‘bonnet’) on
tip of rostrum
Many individuals have irregular white patch
around navel on underside (highly variable and
Pattern of callosities may extend laterally onto sides and towards chin
No pleats or
varies between – but not as extensive as in southern right whale)
grooves on
individuals but
throat
distributed in generally White scarring (from entanglement,
consistent locations Large, broad vessel strikes and killer whale attacks)
Eyes just above flippers up to mainly on tailstock and flukes, but
corners of mouth 1.7m long sometimes elsewhere on body

Callosities on rostrum tend

to be more continuous than
in southern right whale

SIZE
L: ♂ 13–16m, ♀ 15–17m; Distinctive gap
WT: 36–55t; MAX: 18m, 90t between two
Calf – L: 4–5m; WT: 1,000kg rows of baleen
Female typically c. 1m longer than plates at front
male. Average length decreased by 1m
(7 per cent) in past 40 years, due to
dragging around fishing nets and lines
(less energy for growth). Rim of lower ‘lip’ has several crenulations
or scallop-shaped indentations, but varies
from many to a few or no callosities

70°

60° AT A GLANCE North Atlantic • Extra-large

size • Extremely stocky body • Predominantly
50°
black • Smooth back with no dorsal fin or
ridge • Low body profile at surface • Massive
??? head covered in light-coloured callosities
40°

30° • Very strongly arched jawline • No pleats

or grooves on throat • V-shaped blow •
Rectangular, broad, paddle-shaped flippers
20°

primary feeding range migratory corridor calving area

secondary range eastern historical range

NORTH ATLANTIC RIGHT WHALE 33

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 BEHAVIOUR Generally slow moving and may rest at the surface for long periods. Frequently engages in active surface
behaviour, however, and will breach, spyhop, lobtail and flipper-slap repeatedly. Often shows little or no avoidance
behaviour in the presence of boats, and can be inquisitive and approachable.
FOOD AND FEEDING Mostly calanoid copepod crustaceans, but also other small invertebrates, including smaller
copepods, amphipods, krill, pteropods and larval barnacles. Normally skim-feeds; no feeding on winter breeding
grounds.
BALEEN 205–270 plates (each side of the upper jaw). Long, thin plates averaging 2–2.8m long; grey-brown to black.
GROUP SIZE AND STRUCTURE Normally one to two, up to 12 in occasional loose aggregations; much larger
aggregations may form on temporarily rich feeding grounds or in breeding groups.

ADULT

Rostrum very narrow Pointed tips

(widens slightly towards tip) Flukes black on
upperside and
underside

Deep
median
notch

Widely separated blowholes Wide flukes (up to 6m

angled slightly to sides across – more than 30
per cent of body length) Smooth trailing edge
(cf. humpback whale)

Pale grey areas thicken and Smooth, pale grey areas on head and
roughen during first few rostrum may be apparent on newborns
months after birth (callosities CALF
(where callosities will develop)
not fully developed and
colonised by whale lice until
7–10 months old)

FLUKES BREACH

34 RIGHT AND BOWHEAD WHALES

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 CALLOSITIES Scientists use the unique callosity
patterns on the heads of right whales (as well as
scars and other distinguishing features) to tell one
individual whale from another.

Chin callosity

Bonnet
Mandibular
island
(‘dribble’)
Peninsula

Rostral
‘Lip’ patch
island

Post-blowhole
callosity

Coaming
Eyebrow callosity

NORTH ATLANTIC RIGHT WHALE 35

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 NORTH PACIFIC RIGHT WHALE
Eubalaena japonica (Lacépède, 1818)

In 1874, whaling captain and naturalist Charles Scammon remarked that North Pacific right whales were ‘scattered
over the surface of the water as far as the eye can discern’. But commercial whaling was so intense that they suffered
the most dramatic and rapid depletion of all the great whales.
IUCN status Endangered (2017). Eastern North Pacific population Critically Endangered (2017).
Population Uncertain, but possibly c. 400 in the west and c. 30 in the east. An estimated 26,500–37,000 taken by
commercial whalers 1839–1909. Pre-whaling population believed to have been 30,000+, though never documented in
detail. Trend unknown.
Classification Mysticeti, family Balaenidae.
Taxonomy No recognised forms or subspecies, though two recognised populations occur, one on either side of the
North Pacific. Split from the North Atlantic right whale in 2000 due to genetic differences between both the whales
and their lice (the two were previously lumped as the ‘northern right whale’, Eubalaena glacialis).
Other names Pacific right whale, northern right whale.
DISTRIBUTION Formerly abundant in cold temperate waters across much of the North Pacific, but currently occupies
only a fraction of this former range. There appear to be two distinct populations: one of several hundred individuals
(western North Pacific, centred around the Sea of Okhotsk); and one of just tens of individuals (eastern North Pacific,
primarily in the Bering Sea and Gulf of Alaska). There appear to be seasonal migrations from higher-latitude summer
feeding grounds to potential lower-latitude winter breeding grounds. No calving grounds have been located. Tends
to be more pelagic than the North Atlantic right whale. Most sightings in the past 20 years have occurred in the
southeastern Bering Sea (where there has been greatest research effort). North Pacific, North Atlantic and southern
right whales are separated by Arctic ice and warm equatorial waters, and it is estimated that there has been no
interchange between the three populations for millions of years.
Western population Historical whaling records indicate that the principal summer feeding grounds were in the Sea of
Okhotsk (between Sakhalin Island and Kamchatka), around the Kuril and Commander islands, along the east coast of
Kamchatka and in the central Bering Sea – north of 40°N. These regions are still considered to be important summer
habitats. In the autumn, there was a southward shift in distribution to at least 30°N and possibly 25°N. The lack of
evidence for coastal winter breeding grounds suggests that they may breed in open-ocean waters offshore. There is
some historical evidence of two distinct stocks of right whales in the western North Pacific, kept apart by the Japanese
islands. The ‘Sea of Japan stock’ migrated along the western coast of Japan, between summering grounds in the Sea
of Okhotsk and unidentified wintering grounds south of Japan; and the ‘Pacific stock’ travelled along the eastern coast
of Japan, between summering grounds around the Kuril Islands and in the western Bering Sea and unidentified (but
possibly the same) wintering grounds.
Eastern population Historical whaling records indicate that the principal summer feeding grounds were in the eastern
Bering Sea and the Gulf of Alaska – north of 40°N. In the autumn, there was a southward shift in distribution to
unknown wintering grounds. Since the 1990s, most sightings of the eastern population have been concentrated in

DIVE Sequence May lift head almost clear of water on surfacing after long dive;
at surface, water washes over back creating distinctive white water; flukes
often raised quite high before deep dive. • Depth Often at or near surface.
• Duration Typical deep-feeding dives 10–20 minutes.
BLOW Bushy V-shaped blow (when seen from in front or behind) or oval
and bushy (from side). • Up to 7m high (highly variable height). • Jets often
asymmetric in height. • V more widely spaced at base than in grey or humpback
(which sometimes produces V-shaped blows). • If wind dissipates blow, it can
be difficult to see (as body so low in water).

36 RIGHT AND BOWHEAD WHALES

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 Top of head Extremely stocky body White blotches sometimes ADULT
sprinkled with Very strongly (bordering on rotund) occur elsewhere on body
callosities (naturally arched jawline
light to dark grey,
but appear white, Massive head Individuals sometimes
(up to one-third Maximum girth can appear mottled (caused
cream or yellowish
Upper of body length) exceed c. 60 per cent by uneven sloughing of
due to presence of
margin of of total length patches of skin)
whale lice)
lower ‘lip’
serrated Smooth, broad back
with no dorsal fin
or ridge

Predominantly
black

Largest
callosity (the
‘bonnet’) on Many individuals have irregular white
tip of rostrum No pleats or patch around navel on underside
grooves on (highly variable and may extend
throat laterally onto sides and towards chin)
Pattern of callosities Large, broad flippers
varies between up to 1.7m long White scarring (from entanglement,
individuals but vessel strikes and killer whale attacks)
distributed in generally mainly on tailstock and flukes, but
consistent locations Eyes just above sometimes elsewhere on body
corners of mouth

Male may have

proportionately greater
areas covered by
callosities than female

SIZE Distinctive gap

L: ♂ 14–17m, ♀ 15–18m;
WT: 50–80t; MAX: 19.8m, 100t
Calf – L: 4–5.5m; WT: 700–1,000kg
Female typically c. 1m longer than
male.
between two rows of
baleen plates at front
Rim of lower jaw has several crenulations or
scallop-shaped indentations, but varies from
many to a few or no callosities

75°

AT A GLANCE Northern North Pacific •

60°
Extra-large size • Extremely stocky body •
Predominantly black • Smooth back with
no dorsal fin or ridge • Low body profile at
surface • Massive head covered in light-
45°

30°
coloured callosities • Very strongly arched
jawline • No pleats or grooves on throat •
15°
V-shaped blow • Rectangular, broad, paddle-
0° shaped flippers
15°

primary range sightings hotspots

NORTH PACIFIC RIGHT WHALE 37

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 two areas during summer: particularly in the southeastern Bering Sea, west of Bristol Bay, Alaska (c. 57–59°N), where
the whales appear to select relatively shallow waters c. 70m deep over the mid-continental shelf; but also over the
continental shelf and slope south of Kodiak Island in the Gulf of Alaska. Possible wintering grounds for the eastern
population are obscure (calving likely takes place offshore).
BEHAVIOUR There have been relatively few direct observations of living North Pacific right whales in recent decades.
They are generally slow moving and may rest at the surface for long periods, but have been observed breaching,
spyhopping, lobtailing and flipper-slapping. They show little or no avoidance behaviour in the presence of boats and
can be inquisitive and approachable, although a legacy of whaling means some individuals may still be very sensitive
to vessels.
FOOD AND FEEDING Mostly calanoid copepods, but will take other small invertebrates, including smaller copepods,
amphipods, krill, pteropods and larval barnacles. Normally skim-feeds (swimming slowly with mouth open through
patches of concentrated prey at or near the surface); will also filter-feed at depth (up to 300m); at least one observation
of lunge-feeding; no feeding on winter breeding grounds.
BALEEN 205–270 plates (each side of the upper jaw). Long, thin plates averaging 2–2.8m long; grey-brown to black.
GROUP SIZE AND STRUCTURE Normally one to two, though larger aggregations of 30+ may form on temporarily rich
feeding grounds or breeding grounds.
ADULT
Rostrum very Pointed tips
narrow (widens
slightly towards tip) Flukes black on
upperside and
underside

Deep
median
notch

Wide flukes (up to 6m

Widely separated blowholes across – more than 30
angled slightly to sides per cent of body length) Smooth trailing edge
(cf. humpback whale)

Smooth, pale grey areas on

Pale grey areas thicken and head and rostrum may be
roughen during first few apparent on newborns (where
months after birth (callosities callosities will develop)
CALF
not fully developed and not
colonised by whale lice until
7–10 months old)

38 RIGHT AND BOWHEAD WHALES

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 FLUKES
CALLOSITIES AND LICE
CALLOSITIES Callosities are areas of thick, irregular,
calloused tissue – found only on right whales –
that develop around the sparse hairs scattered
about the whale’s head. They are named after their
resemblance to the ‘callus’ or thickened skin that
occurs naturally in many animal species. Pockmarked
with ridges and depressions, they feel like hard
rubber to touch and, from a distance, look a little
like barnacles. The callosity tissue is naturally light
to dark grey, but it is home to colonies of thousands
of creamy-white or yellowish cyamid crustaceans
or ‘whale lice’, which obscure the underlying
colour. Callosities occur in approximately the same
places that humans have facial hair: above the
eyes (eyebrows), along the rostrum, between the
blowholes and the tip of the snout (moustache), and
along the margins of the lower ‘lips’ and jaw (beard).
Their height can change throughout a whale’s life
(growing upwards and breaking off repeatedly), but
their overall size and placement on the head remains
the same; consequently, their shape and size serve
as ‘fingerprints’ or ‘distinguishable faces’ that enable
researchers to tell one individual right whale from
another. The function of the callosities is unknown.
One theory is that they are designed specifically
to attract dense populations of whale lice, which
stand on their hind legs to catch copepods – perhaps
alerting the whale and helping to steer it towards
denser concentrations of its tiny prey.
000 WDP FG.indd 39
BREACH
LICE Right whales carry large populations of three
species of cyamid crustaceans or ‘whale lice’. Two
of these are host-specific (i.e. found on no other
whale species) and one (Cyamus ovalis) is also
found on sperm whales. Recent genetic evidence
suggests that these three species should be split
into nine species, because they are sufficiently
different on their North Atlantic, North Pacific and
southern hemisphere hosts. Whale lice occur among
the callosities and are common in creases and folds
elsewhere on the body.
Cyamus gracilis
6mm long
Predominantly yellow
Typically c. 500 per whale
Mainly in pits and grooves between
elevated patches of callosity tissue
Cyamus erraticus
12–15mm long
Predominantly orange
Typically c. 2,000 per whale
Mainly on smooth skin in genital
and mammary slits, and in large
concentrations in wounds
Also found in large patches on heads
(where there is no callosity tissue) of
young calves (disappearing when calf is
c. 2 months old)
Cyamus ovalis
12–15mm long
Predominantly white
Typically c. 5,000 per whale
Coat callosities at average density
of one adult per cm2 (main reason
for pale colour of callosities)
NORTH PACIFIC RIGHT WHALE 39
29/11/2021 12:11
 SOUTHERN RIGHT WHALE
Eubalaena australis (Desmoulins, 1822)

The southern right whale is one of the world’s best-known large whales: research at Peninsula Valdes (Argentina) has
been ongoing since 1971 and is one of the longest-running studies following the lives of known individual large whales.
While a long way from recovering from the ravages of whaling, the southern right whale is not as seriously endangered
as its two northern relatives.
IUCN status Least Concern (2017). Chile–Peru sub-population Critically Endangered (2017).
Population Minimum 15,000–20,000. Pre-whaling population (1770) estimated at 70,000+. Minimum 150,000 killed
by whalers (1770–1900) plus 3,300+ taken by the Soviet Union after protection in the 1960s. The population dropped
to c. 300 by the 1920s. Increasing (though sharp declines observed in South Africa and Namibia since 2015).
Classification Mysticeti, family Balaenidae.
Taxonomy No recognised subspecies; although barely differs morphologically, is genetically distinct from North
Atlantic and North Pacific right whales. There are five main colour morphs.
DISTRIBUTION Circumglobal distribution in the southern hemisphere, approximately 20–60°S (occasionally to 16°S
on both coasts of South America and to at least 65°S along the Antarctic Peninsula). Migrates between low-latitude
coastal winter breeding grounds (typically May–December – precise timing varies with region) and high-latitude,
predominantly offshore feeding grounds. Feeds mostly in the mid Southern Ocean south of 40°S, with some individuals
reaching the pack ice. Specific feeding areas include the Falkland Islands, South Georgia and Shag Rocks, the Antarctic
Peninsula and south of Australia. Prefers to calve in sheltered, nearshore, shallow waters and bays with sandy bottoms
(probably for protection from predatory killer whales and large sharks). There appears to be some interchange between
breeding areas off the same continent but little interchange between continents. Major mating and calving areas are:
Southern Africa Mainly South Africa, from Saint Helena Bay in the west to Port Elizabeth in the east, but also
Namibia (occasionally into southern Angola) and Mozambique, with small numbers off eastern Madagascar; there is a
small, separate population around Tristan da Cunha.
Southern South America Mainly Argentina (especially Peninsula Valdes) but also an expanding range in southern
Brazil, with very small numbers (probably separate populations) in Uruguay, and Peru and Chile.
Australia Mainly along the southern coasts of Australia (including Tasmania) as far as Perth in the west and Sydney in
the east, with occasional occurrences further north. They probably form two distinct populations (southwestern/south
central and southeastern/eastern). In recent years, they have been reoccupying historical calving grounds along the
southern Australian coast.
Sub-Antarctic islands of New Zealand Mainly off the Auckland and Campbell islands. Historically, there were
winter breeding grounds around the North and South Islands of mainland New Zealand but, after extensive whaling
in the nineteenth and twentieth centuries, no southern right whales were seen for nearly four decades (1928–63);
however, they have been sighted every year since 1988.
BEHAVIOUR Generally slow moving and may rest or log at the surface for long periods. It can move surprisingly quickly,
however, and frequently engages in active surface behaviour, including repeated breaching, spyhopping, lobtailing

DIVE Sequence May lift head almost clear of water on surfacing after long
dive; flukes often raised quite high before deep dive. • Depth Often at or near
surface; sometimes reaches greater depths of 200+m (possibly 300+m).
• Duration Typical feeding dive 10–20 minutes; maximum 50 minutes.
BLOW Bushy V-shaped blow (viewed from in front or behind) or oval and
bushy (from side). • Up to 7m high (highly variable height). • Jets often
asymmetric in height. • V more widely spaced at base than in humpback
whales (which sometimes produce V-shaped blows). • Can be difficult to see
if wind dissipates blow (as body so low in water).

40 RIGHT AND BOWHEAD WHALES

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 Pattern of callosities
varies between
individuals but
is distributed in
consistent locations
Top of head has raised
patches of roughened skin
(callosities) – naturally light
to dark grey, but appear
creamy or yellowish due to
presence of whale lice
ADULT
Very strongly Some individuals
Around 3–6 per cent of individuals
arched mouthline have varying
have white or light grey blazes
amounts of white
on back (uncommon in northern
on upperside
Extremely stocky body hemisphere right whales)
(bordering on rotund)
May appear mottled pale
grey (caused by uneven
Massive head Maximum girth can sloughing of patches of skin)
(c. 25–33 per exceed c. 60 per
cent of body cent of total length Smooth, broad back with
length) no dorsal fin or ridge

Predominantly
black

Many individuals have irregular

Largest callosity white patch on underside around
(the ‘bonnet’) on navel (highly variable and may extend
tip of rostrum Eyes just above laterally onto sides and towards chin)
corners of
mouth (below Large, broad
flippers up to Extent and shape of
large callosity)
No pleats or 1.7m long white patch does not
Callosities often on upper grooves on throat change with time
margin of lower ‘lips’ (cf.
northern relatives)
Male tends to
have more robust
callosities than
female
Distinctive gap
between two
SIZE
L: ♂ 13.5–16m, ♀ 14–16.5m; rows of baleen
WT: 36–55t; MAX: 17m, 80t plates at front
Calf – L: 4–5.5m; WT: 700–1,000kg
Female typically c. 1m longer than
male.
Rim of lower ‘lip’ has several crenulations or
scallop-shaped indentations and can have long
callosity patches (cf. northern relatives)

80°

70° AT A GLANCE Cold temperate southern

60°
hemisphere • Extra-large size • Extremely
stocky body • Predominantly black with
50°

40°

variable white on underside • Smooth back

30°

20°
10°
0° with no dorsal fin or ridge • No pleats or
10°
20°
grooves on throat • Low body profile at
30°

40°
surface • Massive head covered in light-
50°
coloured callosities • Very strongly arched
jawline • V-shaped blow • Frequently
60°

70°

80°
engages in active surface behaviour

SOUTHERN RIGHT WHALE 41

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 and flipper-slapping. Unlike northern hemisphere counterparts, it will also ‘sail’ by tipping its head down, with its
tail sticking out of the water (perpendicular to the wind) and being blown along for a short distance (then righting
itself, swimming back upwind, and doing it again); or may simply hang upside down, flukes suspended in the air, for
several minutes. Tactile behaviour (touching and rubbing conspecifics) is fairly common. It frequently shows little or no
avoidance behaviour in the presence of boats, and it can be inquisitive and approachable.
FOOD AND FEEDING Exclusively zooplankton (mostly copepods north of 40°S and krill – especially Antarctic krill –
south of 50°S); will also take squat lobster, juvenile pelagic crabs, mysid shrimps, pteropods, larval barnacles, other
small invertebrates; prey must be concentrated in exceptionally high densities to elicit feeding. Normally skim-feeds
(swimming slowly with mouth open through dense patches of prey, near surface or at moderate depth); little feeding on
winter breeding grounds.
BALEEN 200–270 plates (each side of the upper jaw). Long, thin plates averaging 2–2.8m long; dark grey to black.
GROUP SIZE AND STRUCTURE Normally one to two (away from aggregation sites on breeding grounds or on rich
feeding grounds), up to 12 in loose associations; loose aggregations of up to 100 may form on temporarily rich feeding
grounds.
May be scarring on back of ADULT
Rostrum very narrow (widens Peninsula Valdes adults and Pointed tips
slightly towards tip) calves caused by kelp gulls
feeding on skin and blubber

Deep
median
notch

Widely separated
blowholes angled
slightly to sides
Smooth, pale grey circular areas
on head and rostrum may be
apparent on newborns (where
callosities will develop)
Flukes black on
both sides
Wide flukes (up to 6m
across – more than 30 Smooth trailing edge
per cent of body length) (cf. humpback whale)
Large patches of orange whale lice disappear CALF
when calf is c. 2 months old (replaced by white
whale lice as calloused tissue begins to appear)

Pale grey areas begin

to thicken and roughen
into calloused tissue
when 3 months
old (callosities not
fully developed and
colonised by whale lice
until 7–10 months old)
Very young calf paler than adult
(before it sheds natal skin)

SAILING
FLUKES

42 RIGHT AND BOWHEAD WHALES

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 DORSAL SKIN PATTERNS
(five main colour morphs recognised)

‘black’ or ‘wild type’

'white-blaze’

‘grey-morph’

‘partial-grey-morph’

‘partial-grey-morph with white-blaze’

SOUTHERN RIGHT WHALE 43

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 BOWHEAD WHALE
Balaena mysticetus Linnaeus, 1758

The only large whale found exclusively in the Arctic, the bowhead is well adapted to life in its freezing home. With a
layer of blubber up to 28cm thick and the ability to create its own breathing holes by breaking through ice up to 60cm
thick, it can live at higher latitudes than any other baleen whale.
IUCN status Least Concern (2018). Sea of Okhotsk sub-population Endangered (2018); East Greenland–Svalbard–
Barents Sea sub-population Endangered (2018).
Population Minimum 30,000. Pre-commercial whaling population 71,000–113,000. Increasing.
Classification Mysticeti, family Balaenidae.
Taxonomy No recognised forms or subspecies (though four separate stocks).
Other names Greenland/Arctic whale, Greenland/Arctic right whale.
DISTRIBUTION Circumpolar in the Arctic and sub-Arctic, mainly 54–85°N (it is the only baleen whale living exclusively
in this region). Closely associated with the pack ice and its seasonal movements, migrating to the High Arctic in
summer and retreating southward in winter with the advancing ice edge (the winter range is poorly known, but it is
believed to live in areas near the ice edge, in polynyas and unconsolidated pack ice). May travel long distances (up to
200km per day) between high-productivity feeding areas. Mainly pelagic, but does occur in coastal waters. There is
some geographic segregation by sex.
Four ‘stocks’ (sub-populations) are currently recognised (mainly based on geographical separation): Bering–
Chukchi–Beaufort (Alaska, Canada and Russia); Sea of Okhotsk (Russia); Eastern Canada–Western Greenland (formerly
considered two stocks: Hudson Bay–Foxe Basin, Canada, and Baffin Bay–Davis Strait, Canada and Greenland); and East
Greenland–Svalbard–Barents Sea (Greenland, Norway and Russia). There is some recent evidence of limited mixing in
the extreme summer ranges between the first and third stocks (due to diminishing ice in the Northwest Passage).
BEHAVIOUR Generally a slow, deliberate swimmer, but capable of bursts of speed up to 21km/h. Frequently breaches,
flipper-slaps, lobtails and spyhops, and may inspect or play with objects in the water. During breaches, up to 60 per
cent of the body leaves the water and the whale usually falls back into the water on its back or side. Often quite
approachable by boat and may closely investigate people standing on the floe edge. It can swim beneath ice, making
breathing holes by breaking through ice up to 60cm thick with the raised part of its massive head. Often seen in
association with belugas and narwhal.
FOOD AND FEEDING Catholic diet (more than 100 prey species known) but prefers small to medium-sized crustaceans
(mostly 3–30mm-long), especially copepods and euphausiid krill; also feeds on mysids and gammarid amphipods. Feeds
throughout water column, anywhere from surface to seabed, under ice as well as in open water (where may ‘skim’
through concentrated prey at surface, swimming slowly with mouth open).

DIVE Sequence Frequently flukes before deep dive (tail may tip to right).
• Depth Frequently less than 30m when foraging in summer; dives deeper during
winter (often 250+m) and when travelling; maximum recorded 582m (West Greenland,
2011). • Duration Typically 1–20 minutes (depending on behaviour – foraging dives in
Beaufort Sea average 3.4–12.1 minutes); tends to dive for longer under heavy pack
ice than in open water; maximum documented 61 minutes under natural conditions
(80 minutes when harpooned and being chased by whalers).
BLOW Tall, bushy, V-shaped blow up to 7m (typically 5m, but height highly variable).
• Jets usually different heights. • May appear as a single blow if seen from side or
in wind.

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 Prominent triangular hump (known as Very rotund body (max. ADULT
‘crown’ or ‘stack’) in front of blowholes girth can exceed 70 per
cent of total length)
Enormous head (can be
35–40 per cent of body length) Distinctive indentation No dorsal fin,
Broad, rounded,
between head and smooth back hump or ridge
Resembles right back (‘neck’)
whale (without Predominantly
pale callosities black (some lighter-
on head) coloured animals
occasionally seen)
Small lump
on tailstock

Variable white Additional white marks may

chin (usually with be caused by scarring from
‘necklace’ of black banging against sea ice
spotting)
Broad-based paddle-
Strongly arched shaped flippers Diffuse white or whitish
rostrum and mouthline Blubber 5.5–28cm markings on tailstock and
No longitudinal thick (depending centre of flukes (become
grooves on where on body) whiter with age – older
throat or belly animals may have all-
white tails and flukes)

SIZE
L: ♂ 14–17m, ♀ 16–18m;
WT: 60–90t; MAX: 19.8m, 107t
Calf – L: 4–4.5m (max. 5.2m);
WT: 900kg

40° 20° 0° 20° 40°

Bering–Chukchi–
ATLANTIC
60°
OCEAN Beaufort stock
Arctic Circle

70°
Eastern Canada–Western
60° 60° Greenland stock
80°
East Greenland–Svalbard– AT A GLANCE Arctic and sub-Arctic • Extra-
80° 80° Barents Sea stock large size • Predominantly black • Smooth
Sea of Okhotsk stock back with no dorsal fin or ridge • Enormous
100°
ARCTIC
100°
head • V-shaped bushy blow • Two distinct
humps in profile • No callosities or barnacles
OCEAN

120° 120°

PACIFIC
140° 140°
OCEAN
160° 180° 160°

BOWHEAD WHALE 45

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 BALEEN 230–360 plates (each side of the upper jaw). Baleen plates are up to 4m (maximum 5.2m) long – the longest of
any whale; dark grey to brownish black, usually with lighter fringes.
GROUP SIZE AND STRUCTURE Bowheads are usually solitary, but are sometimes seen in small groups of two to
three (up to 14). There are occasional loose aggregations of as many as 60 at productive feeding grounds and during
migration. In summer, groups are often segregated by sex and age.
ADULT

Pointed tips
Narrow rostrum

Deeply
notched

Broad, triangular
flukes up to 7m across
Blowholes widely Shallow, concave trailing
separate edge with smooth margins
Lighter grey colour (often
with dark blotching)

MOULTING ADULT

Head hump less developed CALF

(double-humped profile less Slimmer body shape
obvious or absent) Paler and greyer

Usually lacks
distinct white
markings

Characteristic double-
humped surface profile
distinguishes bowhead
from right whales

46 RIGHT AND BOWHEAD WHALES

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 BOWHEAD MIGRATIONS: BERING–CHUKCHI–BEAUFORT STOCK
60° 40° 20° 0° 20° 40° 60° 80°
80°

Bowhead 100°
whale baleen 100°
plate ARCTIC
OCEAN

80°
120°

BEAUFORT
SEA

CANADA
120°
CHUKCHI SEA
70°

140° ALASKA
RUSSIA Arctic Circle

BERING SEA
60°

160° 180° 160° 140°

Apr–May: migrate north (most to eastern Beaufort Sea, some

Chukotka coast)
May–Jul: most in Cape Bathurst region in Amundsen Gulf (some as far
west as Chaunskaya Bay, Chukotka)
Mid-Jul–Oct: Beaufort sea whales migrate west to Chukotka coast,
then all slowly move south into Bering Sea as winter approaches
Nov–Mar: over the continental shelf and north of southern boundary
of sea ice
Primary feeding areas

FLUKES

BREACHING

CHIN VARIATIONS

Variable amounts 'Necklace' of black

of white spotting variable

BOWHEAD WHALE 47

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 PYGMY RIGHT WHALE
Caperea marginata (Gray, 1846)

The smallest and most poorly known baleen whale, the pygmy right whale is rarely observed at sea but can be
positively identified when seen well. Despite its name, it is placed in a separate family and not regarded as one of the
‘true’ right whales.
IUCN status Least Concern (2018).
Population No estimates of abundance, but likely to be reasonably common (at least in some parts of its range).
Trend unknown.
Classification Mysticeti, family Neobalaenidae (though see Taxonomy).
Taxonomy No recognised forms or subspecies. New fossil evidence suggests that it should be moved to the family
Cetotheriidae.
DISTRIBUTION Circumpolar in both coastal and oceanic mid-latitude temperate waters of the southern hemisphere.
Occurs mainly from 30–55°S, preferring water temperatures of 5–20°C; however, if cold currents are present (such
as the Benguela Current off south-west Africa), it has been known to reach 19°S. May be resident year-round in some
parts of the range, but elsewhere there is limited evidence of inshore movements during spring and summer. The
presence of numerous oval cookiecutter shark scars on some individuals suggests that at least some time is spent in
tropical or warm temperate waters. Calving and nursery grounds are not known, though there is one possible location
off the coast of Namibia; most very young animals have been observed north of 41°S. There have been no confirmed
sightings south of the Antarctic Convergence.
BEHAVIOUR Relatively few confirmed sightings at sea. No records of breaching, spyhopping or lobtailing. Apparently
swims slowly at the surface, but it is capable of very rapid acceleration and speed (leaving a conspicuous wake).
Smaller vessels have been approached quite closely, but larger vessels are more often avoided.
FOOD AND FEEDING Calanoid and cyclopoid copepods, amphipods, small krill and possibly other plankton. Believed to
skim-feed (rather than gulp); presence of bird feathers in stomachs may indicate surface-feeding.
BALEEN 213–230 plates (each side of upper jaw).
GROUP SIZE AND STRUCTURE Sightings close to shore tend to be of 1–2 animals (often mother–calf pairs), but groups
of up to 14 have been observed. Aggregations of 80 (590km south of Western Australia, in 1992) and more than 100
(40km south-west of Victoria, Australia, in 2007) have been reported in the open ocean.
ADULT
Head broadest at eyes
(narrows sharply into
long, narrow rostrum) Broad flukes with
pointed tips

Deep
median
notch

Single median ridge

from blowholes to Underside of flukes pale grey
tip of rostrum or white with dark trailing edge

DIVE Sequence Surfaces inconspicuously and fairly quickly (rarely for more than a few seconds); ‘throws’ head out
of water at an angle (often possible to see arched mouthline); does not raise flukes clear of water before diving. •
Depth Probably not deep diver (dives short, and heart and lungs small). • Duration From 40 seconds to 4 minutes.
BLOW Blow often inconspicuous. • When visible, ranges from narrow and columnar to small and oval.

48 PYGMY RIGHT WHALE

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 Head large but One (sometimes two) white ADULT
not as large to pale grey chevrons Medium to dark grey
No callosities (relative to body across back (roughly or blue-grey upperside Falcate dorsal fin two-
size) as in right above or behind flippers) (some individuals may thirds (up to three-quarters)
whales have brownish tinge) of the way along back
Blunt rostrum May be
(relative to numerous
minke whale) cookiecutter
shark scars

Moderately arched Pale grey to

jawline (similar to but white underside
Flippers small, Flippers medium to dark
not as high as in right
narrow and slightly grey above (no white
whales) Body shape slimmer than
rounded at tip band, cf. minke whale)
and paler below in right whales (more
No throat grooves (some individuals streamlined, like rorquals)
have two shallow throat creases –
reminiscent of those in grey whales)
Long, narrow creamy-white baleen
Some individuals plates (up to 68cm long) with outer ADULT
Few or no white
have blue-grey margin of brown, dark grey or black
to pale grey
chevrons upperside
ADULT VARIATION
(not to scale)

Distinctive white or cream-coloured

Moderately arched gumline above baleen (visible even
jawline (less pronounced when mouth only slightly open)
than in adult)
CALF

SIZE
L: ♂ 5.9–6.1m, ♀ 6.2–6.3m;
WT: 2.9–3.4t; MAX: 6.5m, 3.9t
Young animals may be slimmer
Calf – L: 1.6–2.2m; WT: unknown
and lighter than adults

80°

70°

60°

50°

40°
AT A GLANCE Temperate waters of southern
30°

20°
hemisphere • Medium size • Proportionately
10°
0°
0°
large head with arched jawline • Falcate
10°
20°
dorsal fin two-thirds of the way along back •
30°
Light-coloured chevrons on back • Indistinct
blow • Often ‘throws’ head out of water at an
40°

50°

60°
angle on surfacing
70°

primary range secondary range

single extralimital record from The Gambia, West Africa, at 13°N

PYGMY RIGHT WHALE 49

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 GREY WHALE
Eschrichtius robustus (Lilljeborg, 1861)

The grey whale is an inveterate traveller: the round-trip distance between its winter breeding grounds and summer
feeding grounds can exceed 20,000km. Among the world’s most-watched whales, it is instantly recognisable thanks to
its mottled grey colouring and the small hump instead of a dorsal fin.
IUCN status Least Concern (2017). Western North Pacific sub-population Endangered (2018).
Population 20,580 (2020) following an ‘unusual mortality event’ (cf. 26,960 in 2016). Western North Pacific population
c. 175 (2020). Pre-whaling levels uncertain, but the most widely accepted estimate is 15,000–24,000 (although one
DNA study estimated 76,000–118,000). Population stable (but fluctuates extensively).
Classification Mysticeti, family Eschrichtiidae.
Taxonomy No recognised forms or subspecies; two sub-populations (Eastern North Pacific or ENP and Western North
Pacific or WNP), though there are no anatomical differences and there is evidence of mixing on the Mexican breeding
grounds (more than 50 are known to have migrated from Russian summer feeding grounds to Baja winter breeding
grounds).
Other names Gray whale (American spelling), grayback, California gray whale, Pacific gray whale; historically –
mussel-digger, mud-digger, scrag whale, hardhead, devilfish.
DISTRIBUTION Mainly over shallow continental shelf waters of the North Pacific and adjacent seas. Primarily coastal,
but does feed far from shore on the shallow flats of its feeding grounds and can navigate deep oceans on migration. The
ENP population migrates between winter breeding grounds in Baja California, Mexico, and summer feeding grounds
predominantly in the Bering, Chukchi and Beaufort seas; it appears to be expanding north-west as Arctic ice opens
up. The WNP population migrates between winter breeding grounds (probably in the South China Sea), and summer
feeding grounds in the Sea of Okhotsk and off southern and southeastern Kamchatka, Russia. There is evidence of
some mixing between stocks during the winter breeding and summer feeding seasons. Historically, grey whales also
occurred in the North Atlantic (see p. 53). There is one record from the South Atlantic – an individual spent nearly five
weeks in Walvis Bay, Namibia, in 2013.
Migration Breeding and feeding grounds are widely separated, demanding long coastal migrations (spanning up to 50°
of latitude). The ENP population makes an exceptionally long migration, hugging the length of the North America coast,
usually within 10km of shore. The shortest return journey – between San Ignacio Lagoon, Mexico, and Unimak Pass,
Alaska – is c. 12,000km, but many individuals swim considerably further; the longest documented migration of any
mammal (excluding lost individuals) was a female grey whale that completed a 22,511km round trip between Sakhalin
Island, Russia, and Baja California, Mexico. Since 2011, some individuals have remained in the Arctic for longer –
exceptionally even year-round – to obtain sufficient food. This is presumed to result from reduced ice cover (due to
global warming) affecting prey distribution and availability. With the retreat of sea ice, the summer distribution has also

DIVE Sequence On initial surfacing, head appears to slope downward from blowholes (giving appearance of
shallow triangle); ‘knuckles’ clearly visible as back arches slightly to dive; flukes raised high into air before
deep dive. • Depth Mainly seafloor feeder (prefers 30–60m; range of 3–120m, maximum known 170m) but
feeds opportunistically in mid-water and at surface.
• Duration On migration, 3–7 minutes; feeding dives
typically 5–8 minutes; in breeding lagoons, 50 per cent
of dives less than 1 minute; when resting, up to c. 26
minutes.
BLOW Bushy blow up to 5m (highly variable height). •
May be V-shaped, tall and bushy, tree-shaped or heart-
shaped (when spray falls inward) when seen from front
or rear. • May be single bushy column.

50 GREY WHALE

000 WDP FG.indd 50 29/11/2021 12:11

 ADULT
Large white skin lesions
Highly variable appearance common (unknown cause –
caused by scarring and possibly from exposure to
abrasions from injuries or damaging UV light or even
Head arched downward previous barnacle and lice
between blowhole and Arctic ‘frostbite’)
infestations 8–14 fleshy bumps
snout (less arched than (‘dorsal crenulations’
right whales, more or ‘knuckles’) on
arched than rorquals) Robust body upperside of tailstock
(stockier than most Hump two-thirds (between fin and flukes)
rorquals, slimmer of the way along
than right whales) back
Head slender and
small in relation Small, low hump
Heavily to body size instead of dorsal fin
encrusted with (variable size and
barnacles and shape)
whale lice
(mainly on head)

Long, slightly Light to dark grey or grey-

arched mouthline brown with white mottling
(more mottled with time)
Relatively short, broad, paddle-
shaped flippers (usually with
pointed tips – more rounded from
Some parallel linear
abrasion in older individuals) Unique cyst-like structure
scarring, especially on
flippers and flukes (rake (10–25cm diameter) on
More vibrissae than any other whale
marks – tooth scars – from ventral surface of caudal
(widely spaced bristles emerge from
killer whale attacks) peduncle (unknown
small dimples mainly on upper and lower
function)
jaw); many obliterated by barnacles and
scarring in older animals

75°
75°

AT A GLANCE Coastal or shallow waters of

60° North Pacific and adjacent seas • Light to
dark grey or grey-brown with white mottling
45°
• Large size • Low hump (instead of dorsal
fin) • Head (and other parts of body) encrusted
30° with barnacles and lice • ‘Knuckles’ on upper
??
side of tailstock (between fin and flukes)
• Low V-shaped or heart-shaped bushy
15°

WNP range ENP winter breeding range

Pacific coast feeding secondary range
blow • Frequently flukes upon deep dive
group summer/autumn range
?? possible WNP winter range
possible overlap during summer
ENP migration
ENP summer range

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 expanded in recent decades, especially to more northerly feeding areas (to at least 71°N); observations as far east as
the Canadian Beaufort Sea and as far west as Wrangel Island in the Chukchi Sea (and even into the East Siberian Sea)
are now common. The southernmost record in the North Pacific is a stranded adult in El Salvador (c. 14°N) in 2010.
PACIFIC COAST FEEDING GROUP Some 200–240 grey whales – known as the Pacific Coast Feeding Group (Canada) or
the Pacific Coast Feeding Aggregation (USA) – do not migrate all the way to the Arctic, and instead spend the summer
and autumn feeding in a well-defined coastal region between northern California and south-east Alaska.
BEHAVIOUR One of the most active large whales at the surface, frequently breaching (often several times in a row,
exceptionally 40–50 times), spyhopping, and waving tail or flippers in the air. May appear to ‘play’ in the surf and
rub against pebble beaches, rocks, piers and boats, possibly to ease skin irritations caused by ectoparasites. Often
inquisitive and may approach boats. ‘Friendly’ or ‘curious’ behaviour is most common in Mexican breeding lagoons.
FOOD AND FEEDING Variety of benthic and planktonic prey; in northern seas, benthic amphipods usually account for
90 per cent of diet; south of Aleutian Islands, main prey often planktonic mysids but also benthic amphipods and other
species. Will opportunistically take pelagic species such as red crabs and crab larvae, mysids, fish eggs and larvae,
baitfish and squid. Most feeding during summer, fasting thereafter (except opportunistically on migration). Swims
slowly along seabed, sucking up sediment, then filters out prey. May also skim-feed like right whales or gulp-feed like
rorquals, to exploit free-swimming prey.
BALEEN 130–180 plates (each side of the upper jaw). Among the shortest and coarsest baleen plates of any whale,
just 5–50cm long.
GROUP SIZE AND STRUCTURE Grey whales usually migrate alone or in twos or threes, but up to 16 can be seen
in unstable groups. Mother–calf pairs tend to migrate alone. More than 1,000 can congregate in a single winter
aggregation area and breeding lagoon. On summer feeding grounds, usually alone or in pairs, but several hundred may
be scattered across food-rich areas. Towards the end of the feeding season, recently weaned young animals may form
groups of up to 12 or more.
Pointed tips (rounded ADULT
due to abrasion in older
Head elongate and individuals)
triangular from above
Broad grey flukes
3–3.6m wide

Distinct
median
notch

Flukes frequently
Longitudinal marked or scarred
double blowholes
Convex, scalloped
trailing edges
No barnacles or lice
(although obtained from CALF
mother shortly after birth)

100–170 vibrissae (more prominent than At birth, darker than adult

on adult – particularly on rostrum and (uniform dark charcoal grey –
along margins of upper and lower jaws) may look almost black)
Less mottled (may have a few
swirls of grey, black and white)

SIZE
L: ♂ 11–14.6m, ♀ 11.6–15m;
WT: 16–30t; late-term pregnant
females can weigh an additional 5t;
MAX: 15.6m, 40t
Calf – L: 4.2–4.9m; WT: c. 1.1t

52 GREY WHALE

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 GREY WHALE HYPOTHETICAL
HISTORICAL RANGE IN NORTH ATLANTIC
70°

Grey whales once occurred on both sides of the North

? Atlantic (indeed, Lilljeborg’s original description of the
?
60°
?
? species in 1861 was from a subfossil skeleton found
50° in Sweden). There is very little information on their
?
range or migratory routes, but there were possibly
40°
? two discrete sub-populations (partly overlapping in
30° ? Iceland). The population was predominantly extinct
by the late 17th or early 18th centuries, probably
due to (or at least hastened by) hunting by Basque,
20°

10° Icelandic and Yankee whalers. There have been three

exceptional sightings of lone individuals in recent
Possible summer feeding areas Possible migratory routes years: Israel/Spain in 2010; Namibia in 2013; and
Possible winter breeding areas Morocco/Italy/France/Spain/Gibraltar in 2021.

ADULT HEAD ADULT HEAD UNDERSIDE

right side

Baleen plates
creamy to pale
yellow (and
shorter than in Throat has 2–7
any other whale) (usually 2–3) short, deep
Most grey whales are right-dominant feeders and grooves (c. 1.5m long)
right side of head often differs from left side due
stinct to abrasion during seafloor feeding: more heavily
dian scarred, fewer barnacles and whale lice, shorter
tch and more worn baleen plates
ADULT FLUKES
FLUKES (showing shape
change with age)
ped

Younger Older

BARNACLES AND LICE

Cryptolepas rhachianecti Cyamus scammoni Cyamus kessleri Cyamus ceti Cyamus eschrichtii
(whale barnacle) (found only on greys) (found only on greys) (also found on bowheads) (found only on greys)
Grey whale calves are born free of external parasites, but rapidly acquire them as they grow. Adults have more than
any other cetacean: consisting of one species of barnacle (up to 5.5cm in diameter and host-specific to grey whales)
and four species of whale louse (1–2.7cm long).

GREY WHALE 53

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 BLUE WHALE
Balaenoptera musculus (Linnaeus, 1758)

The largest animal known to have existed on Earth, the blue whale can be remarkably inconspicuous and difficult to
see. But a close encounter with this true gargantuan is unforgettable. It was hunted relentlessly worldwide, until every
population was severely depleted, and came dangerously close to extinction.
IUCN status Endangered (2018). Antarctic blue whale Critically Endangered (2018).
Population c. 10,000–25,000. Pre-whaling population c. 300,000, including 239,000 Antarctic blue whales. During the
period 1868–1978, 382,595 were killed by commercial whalers. Increasing.
Classification Mysticeti, family Balaenopteridae.
Taxonomy Five subspecies are currently recognised: northern blue whale (B. m. musculus), Antarctic or ‘true’ blue
whale (B. m. intermedia), northern Indian Ocean blue whale (B. m. indica), pygmy blue whale (B. m. brevicauda) and
Chilean blue whale (B. m. chilensis). The first two share a similar external form (though they differ genetically and
acoustically).
Other names Sulphur-bottomed whale or sulphur-bottom (after the diatom film that can form on its body), Sibbald’s
rorqual.
DISTRIBUTION Occurs from the tropics to the edge of the pack ice in both hemispheres, though distribution is patchy
and it is rare in most equatorial waters and in the centre portions of major ocean basins. Most populations are
migratory – moving between productive, higher-latitude summer and early autumn feeding areas, and lower-latitude
winter breeding and feeding areas – but at least one population (in the northern Indian Ocean) is largely resident year-
round. Unlike most other baleen whales, blue whales feed year-round so food availability probably dictates distribution
for the majority of the year; they will forage in productive areas anywhere. Seasonal movements can be extensive, but
they are complex and poorly understood.
No specific breeding grounds have been discovered conclusively in any ocean (they do not appear to be as well
defined as for humpback, grey and right whales) but they are believed to be in tropical and sub-tropical waters. One
probable breeding ground is the Costa Rica Dome (or Papagayo upwelling) in the eastern tropical Pacific; another in the
Galapagos Islands (for the Chilean subspecies). The Gulf of California, Mexico, is a definite nursing area and possible
calving area.
The species is mainly oceanic and associated with waters deeper than the continental shelf, roaming widely across
ocean basins, but also inhabits some shelf and coastal waters (such as in Mexico’s Gulf of California, the southern
California Bight in the US, Canada’s Gulf of St Lawrence, and Iceland’s Skjalfandi Bay). It prefers habitats marked by
steep submarine topographic features that enhance upwelling. There are still many gaps in our knowledge of any
overlap in distribution between different subspecies, especially in the southern hemisphere.

DIVE (‘true’ blues) Sequence Distinctive blowhole ‘splashguard’ appears as rounded

hump; massive elongated expanse of back rolls into view; many individuals raise
flukes before a sounding dive. • Depth Foraging dives typically to maximum 250m but
capable of 300+m (deepest recorded 370m off Isla Monserrat, Baja California); dives
deeper during middle of the day, following diel vertical movements of prey to feed
at shallower depths up to surface later in the day and at night; non-foraging dives
typically shallower than c. 70m. • Duration Feeding dives typically 8–15 minutes,
but 20 minutes not uncommon (considerably shorter – average 3 minutes – in New
Zealand); occasionally up to 30 minutes (maximum recorded 36 minutes).
BLOW Slender, columnar blow can be at least 12m high (highly variable height).
• Denser and broader than that of fin or sei whale.

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 ADULT
Appears turquoise
Body has light to extensive when swimming
mottling (dark spots on pale just below surface Xenobalanus
Strikingly large
background, pale spots on barnacles may dangle
fleshy ‘splash guard’
dark background, or both) from tip of dorsal fin
immediately in front Relatively small dorsal
of blowholes fin (up to 35cm tall (and from flukes in
Some individuals – smallest fin of all warmer waters)
have chevrons rorquals in proportion to
Relatively flat curving back from body size) three-quarters Extremely deep
head (22–27 behind blowholes of the way along back tailstock from side
per cent of (especially large
body length) adult females)
No mottling
on head or
flukes

Both lower jaws

uniformly bluish grey Turquoise below surface
Slender, (predominantly bluish-grey
60–88 ventral pleats or throat streamlined above but varies from slate
grooves extend to or near navel body grey-blue on cloudy days to
Long, bluntly pointed flippers c. 3–4m long silvery on bright, sunny days)
(c. 15 per cent of total body length)
In cold waters, all or parts of body (especially
underside) may be covered in yellow to greenish diatom
Flippers with bluish-grey upperside films (when diatoms fresh) or opaque rust-coloured
(usually with thin white border diatom films (when diatoms mature and dying)
or tip and occasionally mottled),
whitish underside
ADULT ANTARCTIC/NORTHERN

Jaws capable of
opening to nearly 90°

80°

70°

60°
AT A GLANCE Worldwide (though patchy
distribution) • Extra-large size • Streamlined
50°

40°

30°

20°
body shape • Mottled bluish-grey colour
10°
0°
• Turquoise underwater (when viewed from
10°
20°
surface) • Small to large dorsal fin three-
30°

40°
quarters of the way along back • Prominent
50°
blowhole ‘splash guard’ • Extremely deep
tailstock • Often raises flukes on diving
60°

70°

80°

BLUE WHALE 55

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 BEHAVIOUR Some blue whales raise their flukes when diving (c. 18 per cent in the north-west Atlantic and north-east
Pacific, 25 per cent in Mexico’s Gulf of California and 55 per cent off Sri Lanka). Capable of burst speeds of up to
35km/hr if being chased by boats or killer whales (when swimming very fast, it sometimes almost porpoises above
the surface, throwing up a large rooster-tail and pushing a mass of water in front as it flees). There have been a few
observations of breaching blue whales – usually youngsters – leaping out of the water at a c. 45° angle, but this is very
rare. Behaviour around boats varies from avoidance through indifference to inquisitiveness.
FOOD AND FEEDING Mainly euphausiid crustaceans (i.e. krill); some other crustaceans (including copepods, mysids
and amphipods); occasionally small schooling fish and cephalopods. Unlike most baleen whales, probably does not
fast during winter (continues to feed on breeding grounds); dives below dense layer of krill, turns upwards and lunges,
rolling sideways or doing full barrel-roll and opening mouth, then drifts slowly forward as closes mouth; typically up to
6–7 lunge-feeds per dive (maximum 15); when feeding near surface, often surfaces slowly on one side or upside down
(with one flipper and part of flukes above water).
BALEEN 260–400 plates (each side of the upper jaw). Baleen plates are black, broad-based, and each c. 1m long
(slightly longer in ‘true’ blues, cf. pygmy blue).
GROUP SIZE AND STRUCTURE Usually alone or in pairs, though groups of 3–6 are known in some areas during
summer. May be scattered in loose aggregations of 50+ on good feeding grounds.
Relatively larger, ADULT PYGMY
More silvery
wider head
Rostrum blue or grey-blue Proportionately more
shorter May be less compact than ‘true’ Shorter tailstock
mottled Body more blue whales ( brevicauda means
Tip of rostrum tadpole-shaped ‘short-tailed’) – hence
more rounded proportionately longer
(from above) body in front of dorsal fin

Shorter baleen
plates Slightly fewer ventral
pleats or throat grooves

SIZE – NORTHERN
L: ♂ 23–26m, ♀ 24–27m; WT: 70–135t; MAX: 28.1m, 150t
Calf – L: 6–7m; WT: 2–3t
Female longer than male (in all subspecies).

SIZE – ANTARCTIC
L: ♂ 24–27m, ♀ 24–29m; WT: 75–150t; MAX: 33.58m, 190t
(though these max. sizes were estimated by non-standard
measurements at whaling stations)
Calf – L: 7–8m; WT: 2.7–3.6kg

SIZE – NORTHERN INDIAN OCEAN

L: ♂ 20–22m, ♀ 21–23m; WT: 70–95t; MAX: 24m, 130t

SIZE – PYGMY
L: ♂ 20–22m, ♀ 21–23m; WT: 70–95t; MAX: 24m, 130t
Tends to be shorter than ‘true’ blues, but with a relatively
heavier body weight.

SIZE – CHILEAN
L: ♂ 22–24m, ♀ 22–24m; WT: unknown; MAX: 25.6m
Intermediate in size between pygmy and Antarctic blue
whales.

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 ADULT ANTARCTIC/NORTHERN

Head broad and U-shaped
(more blunt than in fin or
sei whales)
ADULT ANTARCTIC/NORTHERN
Flippers have anteriorly
convex leading edge
Relatively narrow
tailstock from above

Prominent
median
notch

Single prominent ridge

Broad flukes up to 7m
along centre of rostrum from
across (c. 24 per cent
blowholes to tip of rostrum
of body length)

ADULT ANTARCTIC/NORTHERN

Looks like giant tadpole

when feeding

CALF

Similar in appearance
and shape to adult

BLUE WHALE 57

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 30°
Antarctic blue whale
20°
distribution
10°
0°
10°
? ?
20° ? ?
? ? ? summer range
?
30°
? ?
40°
? ? ? possible/probable
50° ? ? winter range (present
60° ? ? year-round in southern
Indian Ocean, with
70° peaks during winter
and spring)
80°

80°
Known and possible
key ranges of the four
70° recognised populations
of northern blue
60°
whales (distinguishable
50° acoustically but not
40° morphologically)
30°

20°
10°
0°
10°
20°

30°

Known and possible key

20° ranges of the five recognised
populatons of pygmy blue
10° whales
0°
Madagascan pygmy
10° blue whale
Possible Chagos
20°
20° pygmy blue whale
30° Sri Lankan
10°
pygmy blue whale
40°
0° Australian
pygmy blue whale
50° 10°
New Zealand
pygmy blue whale
20°

30°

20° 20°

40°
10°
10°

0° 50°
0°

10°
10° 60°

20°

20°
summer feeding range
30°

30° possible winter range

40°

40°
50°

50°
60°

Northern Indian Ocean blue whale distribution summer

Chilean feedingdistribution
blue whale range
possible winter range

58 RORQUALS

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 WHY ARE BLUE WHALES SO BIG?
Blue whales feed on krill – small animals that
swarm in mind-boggling numbers (up to 1,500
individuals per square metre) but live in patches
that might be hundreds or thousands of kilometres
apart. Anything that feeds on krill needs to be able
to swim great distances quickly and efficiently
(the relative energy cost of travelling declines
as body size increases); able to store energy for
days, weeks or even months at a time, to fill the
gaps between krill patches (about a quarter of a
blue whale’s body mass consists of blubbery fat
reserves); and able to ingest huge numbers of
the tiny prey when it is available (feeding on a
few krill at a time would be far too inefficient).
Therefore, the ultimate krill-eating predator is a
large-bodied, big-gulping, filter-feeding whale.
And there are no size constraints thanks to the
buoyancy of water.

FLUKES
Many individuals have Relatively smooth and
radial pattern of light straight (or slightly sinuous)
streaks on underside trailing edge

Tips distinctly Broad, triangular

pointed flukes (some
variation in shape)

DORSAL FIN VARIATIONS Highly variable dorsal fin (from small

nubbin to triangular, hooked or falcate)

PSEUDO-STALKED BARNACLE Xenobalanus globicipitis

This curious dark, worm-like animal (up to 5cm
long) hangs from the trailing edges of the tails,
dorsal fins, and flippers of at least 34 cetacean
species – especially baleen whales such as
blues – in tropical, sub-tropical and temperate
waters worldwide. It is sometimes on the
rostrum, and even on baleen plates and teeth, as
well. There can be just one or as many as 100 in
a cluster. They burrow into the skin (and blubber)
to various depths and, once attached with the
shell base embedded in the host, do not move.

BLUE WHALE 59

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 FIN WHALE
Balaenoptera physalus (Linnaeus, 1758)

The second-longest whale, after the blue, the fin whale is also one of the fastest (it’s been dubbed the ‘greyhound of
the sea’). The distinctive asymmetrical pigmentation on its lower jaw – largely dark on the left and white on the right
– has never been satisfactorily explained (it is also a feature of Omura’s whale, and some sei and dwarf minke whales,
but is more marked on the fin whale).
IUCN status Vulnerable (2018); Mediterranean sub-population Vulnerable (2011).
Population Minimum 150,000. At least 915,000 killed by commercial whalers. Increasing.
Classification Mysticeti, family Balaenopteridae.
Taxonomy Four subspecies are recognised: North Atlantic fin whale (B. p. physalus) in the North Atlantic, North
Pacific fin whale (B. p. velifera) in the North Pacific, southern fin whale (B. p. quoyi) in most of the southern hemisphere
and pygmy fin whale (B. p. patachonica) off the west coast of South America (south to c. 55°S).
Other names Finback, finner, razorback, common rorqual, herring whale, finfish.
DISTRIBUTION In summer, found in cool temperate to polar waters worldwide, in all major oceans in both hemispheres.
It is rarely found in the tropics (except in certain cool-water areas such as off Peru) or in high latitudes near the ice
edge. Movements are complex: some populations appear to be migratory (especially in the southern hemisphere), with
a general shift to higher latitudes for feeding in summer and lower latitudes for breeding (and less feeding) in winter,
but they do not follow a simple pattern and breeding grounds remain uncertain (assuming such areas exist). Resident
or semi-resident populations occur in the Gulf of California (Mexico), the Gulf of Alaska (USA), the East China Sea (off
Japan), and possibly southern California and the central and western Mediterranean Sea (where there appears to be
some mixing with seasonal visitors from the North Atlantic).
Density tends to be higher near or seaward of the continental shelf edge, but it is frequently seen over the shelf and
close to shore where the water is deep enough. Typically, it frequents water deeper than 200m (100m in some regions)
wherever topographic and oceanographic conditions concentrate prey.
BEHAVIOUR Capable of swimming exceptionally fast, reaching 37km/h for short bursts. Rarely breaches (more
when feeling harassed). Often forms mixed schools with blue whales and sometimes associates with pilot whales
and dolphins; often seen in large feeding aggregations with humpback whales, minke whales, Atlantic white-sided
dolphins and other species. Typically, neither avoids boats nor approaches them, but it can be quite approachable and
is sometimes curious.
FOOD AND FEEDING Opportunistic, depending on locality, season and availability. Northern hemisphere: mainly
krill, also copepods, schooling fish (including herring, mackerel, cod, pollock, capelin, sardines, sand lance and blue
whiting), some small squid. Southern hemisphere: almost exclusively krill, but also other planktonic crustaceans. Feeds

DIVE Sequence Body remains low in water; dorsal fin normally appears after
blow has dissipated (though blowholes and fin sometimes visible at same
time, especially in younger animals); rarely shows flukes. • Depth Frequently
dives to 100m (in Ligurian Sea, frequently to 180m); maximum recorded
474m; some feeding at surface. • Duration Typically 3–10 minutes;
maximum 25 minutes.
BLOW Very tall, columnar blow up to 10m high (highly variable height).
• Typically more dense than sei whale blow. • Only blue whale blow is
regularly taller (humpback and sei whale blows can occasionally be as tall).

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 ADULT – LEFT SIDE
Chevrons more
visible in good light Dark grey or brownish-grey Rear half especially may be
(sometimes faint or upperside and sides (often heavily pockmarked with light oval
obscured by diatoms) appears dark with silvery sides) cookiecutter shark and lamprey
bites (varies with locality)
No blaze on
Usually one or more light
left side Sleek, streamlined body
grey V-shaped ‘chevrons’
Raised on back behind head (cf. (slimmer than blue whale)
‘splashguard’ U-shaped on sei whale) Tall, falcate dorsal fin (up
around blowholes with apex pointing forward to 60cm) seven-tenths of
Relatively No dark eye the way along back
flattened or ear stripes No mottling
head on left side (cf. blue whale) Leading edge of dorsal
fin highly variable but
averages 33° angle from
horizontal (cf. sei whale)

May be distinct flipper

Left lower ‘lip’ largely dark
grey or dark brownish-grey
(highly variable extent
towards throat)
Long, tapered flippers with
pointed tips (shorter and
broader in northern fin whales)
‘shadows’ on lower sides
Upperside of flippers can be
dark grey, brownish-grey or
creamy white
Underside of flippers
light grey to white
Fades to white underside
(sometimes yellowish or
brownish in colder waters
due to film of diatoms – can
become blotchy as diatom layer
sloughed off)

SIZE (NORTHERN)
L: ♂ 18–22m, ♀ 20–23m;
WT: 40–50t; MAX: 24m, 90t
Calf – L: 6–6.5m; WT: 1–1.7t
SIZE (SOUTHERN)
L: ♂ 23–25m, ♀ 24–26m;
WT: 60–80t; MAX: 27m, 120t
Calf – L: 6–7m; WT: 1–1.9t
Females are 5–10 per cent longer than
males; northern hemisphere animals
are smaller.

80°

70°

60°

50°

40°
AT A GLANCE Worldwide • Dark grey or
30°

20°
brownish-grey upperside • Extra large size
10°
0°
• Light grey V-shaped chevrons on back
10°
20°
• Asymmetrical lower ‘lip’ coloration • Single
30° prominent ridge on rostrum • Backward-
sloping dorsal fin • Rarely raises flukes on
40°

50°

60°
diving • Alone or in pairs or small groups
70°

80°

primary range secondary range

FIN WHALE 61

001 WDP FG.indd 61 29/11/2021 12:14

 intensively in summer, consumes much less in winter; lunge-feeder (often rolling on side – typically to the right); mouth
opens to almost 90° angle; no evidence of cooperative feeding.
BALEEN 260–480 (average c. 350–390) plates (each side of the upper jaw). Northern fin whales have slightly more
plates on average. Longest plates c. 80cm.
GROUP SIZE AND STRUCTURE Frequently seen alone, but often in small groups of 2–7; large, loose aggregations of
several dozen (up to 100 in exceptional cases) may occur in highly productive areas. Group composition tends to be
dynamic (with individuals frequently moving between groups).

ADULT – RIGHT SIDE

May be other streaks or swirls

of light grey extending from
underside upward (especially
over right flipper) and from eye
Typically has dark eye and
ear stripes on right side
White coloration can
extend onto right upper
‘lip’ and head as 'blaze'
(highly variable)

Right lower ‘lip’ largely creamy

white (in good light visible even
several metres below surface)
Darker
coloration
ADULT PYGMY

May have nearly

black baleen plates
Noticeably smaller than
southern fin whale (c. 18–24m)

ADULT

Baleen plates predominantly dark

blue-grey to nearly black (except
front 20–30 per cent on right side,
which are all whitish or yellowish)

Often striated with grey bands

and fringed with horizontal lines
of yellowish-white, brownish-
grey or olive-green 50–100 longitudinal throat pleats
(extend slightly beyond umbilicus)

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 Sharply pointed rostrum Pale chevron ADULT
Head slender
(more pointed than blue very clear Tailstock
and V-shaped Long, narrow
whale but less so than from above compressed
body shape
sei or Bryde's whales) laterally

Prominent
median
notch

Prominent ridge
Single prominent along tailstock
Chevron tends to (between dorsal Broad flukes (up to
longitudinal ridge on Smooth
be more V-shaped fin and flukes)
rostrum (not as sharply 5m from tip to tip trailing edge
(cf. U-shaped in
defined as in sei whale) in 20m animal)
sei whale)

CALF

DORSAL FIN

Leading edge usually rises at Tip not hooked

FLUKES
shallower angle (average c. 33°) (cf. Omura's
than in other balaenopterids whale)

Less erect and lower Taller, more falcate

than on sei whale or and set further
Bryde’s whale forward than on
blue whale

Shape varies from falcate and rounded

to triangular and pointed (tip always
strongly directed backward)
Underside of flukes light grey to white with
dark grey border (rarely raised above surface)
FIN/BLUE HYBRIDISATION
Hybrids between fin whales and blue whales
have been reported since the 19th century,
most originating from the successful mating
COOKIECUTTER SHARK BITES
of male fin whales with female blue whales.
The offspring share characteristics from both
species. Fin/blue hybridisation is believed to be
more common than the limited genetic evidence
suggests, likely due to reduced population
sizes causing a significant disruption to the
whales’ reproductive dynamics. There is limited
evidence that, in certain circumstances, some
first-generation hybrids might be fertile and Raw bite wound Healed bite wound
able to breed with one of the parental species.

FIN WHALE 63

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 SEI WHALE
Balaenoptera borealis Lesson, 1828

The enigmatic sei whale is the third-longest whale, yet it is surprisingly poorly known. This is partly because in the past
it was often confused in whaling records and scientific accounts with Bryde’s (and possibly Omura’s) whales.
IUCN status Endangered (2018).
Population c. 80,000. Pre-whaling population c. 230,000. At least 325,000 killed by whalers. Increasing.
Classification Mysticeti, family Balaenopteridae.
Taxonomy Two subspecies are recognised: northern sei whale (B. b. borealis) and southern sei whale
(B. b. schlegelii).
Other names Coalfish whale, sardine whale, lesser fin whale, pollack whale, northern rorqual.
DISTRIBUTION Ranges from the tropics to the poles in both hemispheres, but most abundant in mid-latitude temperate
zones. Distribution is poorly documented and most information comes from whaling catches. Migrates between higher-
latitude (cold temperate to sub-polar) summer and autumn feeding grounds and lower-latitude (warm temperate to
sub-tropical) winter breeding grounds. Compared with some other rorquals, migrations are less extensive, feeding and
breeding grounds are less distinct, and it generally does not range as far north or south. It tends to be less predictable
than other rorquals. It may abruptly disappear from areas where it had occurred regularly for years, and suddenly
appear in other areas where it had been absent for years (or even decades); irruptions of sei whales are known as ‘sei
whale years’ or ‘invasion years’.
Generally considered a pelagic species with an offshore distribution along and beyond the continental shelf edge,
especially in areas characterised by complex submarine topography such as seamounts and ridges. However, in some
areas (e.g. Chile and the Falkland Islands) it regularly enters shelf waters and can be found at relatively shallow depths
(less than 40m) and close to shore, including inside inlets and channels. Prefers sea surface temperatures of 8–18°C
(occasionally up to 25°C).
BEHAVIOUR One of the swiftest of the rorquals, capable of swimming at 25km/h (even 55km/h in short bursts,
according to some whaling records). Breaching is rare but usually at a low angle and ends in a belly-flop. The Sei
whale has been seen in association with Peale’s dolphins in the Falkland Islands. Most individuals avoid boats, or are
indifferent, but some can be curious, repeatedly approaching and swimming alongside.
FOOD AND FEEDING Diverse diet varies regionally; mainly dense concentrations of minuscule copepods and krill, but
also amphipods, squid, schooling fish (including sand lance, lumpfish, capelin, anchovy, herring, saury, lanternfish);
in the North Atlantic it prefers pelagic copepods; around the Falkland Islands, mainly lobster krill. In some areas,
especially when feeding near the surface, often associated with large flocks of feeding seabirds. Unusually among
baleen whales, has two modes of feeding: normally ‘skims’ like right whales, but sometimes ‘lunges and gulps’ like

DIVE Sequence Surfaces at shallow angle; tip of rostrum usually just breaks
the surface; blowhole and dorsal fin often (but not always) visible at same
time; tends to sink below surface (back relatively flat, though sometimes
arches before deep dive); fin disappears last; very rarely, if ever, raises flukes;
sometimes dives and surfaces in predictable line (remaining visible just below
surface), frequently leaving long series of flukeprints on surface – but often
impossible to predict and can be erratic in surfacing behaviour. • Depth Varies
according to vertical migrations of prey: limited evidence from Japan suggests
averages c. 10–12m at night, c. 16–19m in day. • Duration On feeding grounds
typically 1–3 breaths at the surface over 20–30 seconds, then longer dives of up
to 13 minutes.
BLOW Blow up to 9–10m high and columnar to bushy (highly variable height –
typically 3–5m). • Generally more diffuse than fin whale blow.

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 ADULT
Often has fin whale-like
(NORTHERN)
lighter-coloured chevron (less Relatively tall, erect
extensive than on fin whale and dorsal fin (average 55cm)
U-shaped – highly variable in slightly less than two-
brightness and extent – more thirds of the way along
visible in good light) back (further forward
than on other rorquals)
Some individuals
have white streak Dark brown-grey
behind eye Lighter ‘brush stroke’ upperside (can Leading edge of fin rises at
between eye and appear bluish-grey steep angle from back (steeper
dorsal fin (highly to steely black in than in fin whale – typically
Head 21–25 variable between poor light) c. 46° from horizontal)
per cent of individuals and more
body length visible in good light) Fin often has distinct
backward bend
Mid brown- halfway to two-thirds
grey sides up (cf. Omura’s whale
Somewhat – though some overlap
arched head with Bryde’s whale)
with slightly
downturned
tip

Deep tailstock

Lower ‘lips’ dark to light grey

Lighter brown-grey Sleek,
on both sides (varies according
(sometimes creamy streamlined
to light conditions and some
white) underside body
individuals may show mild
form of fin whale asymmetry)
May be heavily pockmarked
Relatively small, slender, (especially rear half) with
32–65 (average 50) relatively short light oval cookiecutter shark
pointed flippers (c. 9 per
longitudinal throat pleats on underside and lamprey bites
cent of body length)
(short for rorqual – terminate midway
between flippers and umbilicus)

80°

AT A GLANCE Sub-tropical to sub-polar

70°

60°

50° offshore waters worldwide • Large size

40°

30°
• Sleek body • Dark upperside, lighter
20°
10°
underside • Pale ‘brush strokes’ on sides
0°
10°
• May have fin-whale-like chevron (U-shaped)
20°

30°
• Single prominent ridge on rostrum
40°
• Rostrum has downturned tip • Tall and erect
dorsal fin (highly variable) • Symmetrical head
50°

60°

70° colouring • Dorsal fin and blowholes may be

80° visible simultaneously

primary range secondary range

SEI WHALE 65

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 other rorquals; most feeding occurs during summer (winter consumption is low); individuals may remain in a specific
feeding area for several weeks if prey densities are sufficient; there are no reports of cooperative feeding.
BALEEN 219–402 (average c. 350) plates (each side of the upper jaw). Longest plates c. 80cm; tend to be narrower than
in other rorquals.
GROUP SIZE AND STRUCTURE Varies according to location and season – often seen alone or in small, fluid groups of
2–5. Larger groups may travel together, and loose aggregations numbering tens of individuals can form in productive
feeding areas. Sightings of apparent social groups have been recorded, perhaps engaged in courtship behaviour,
involving high-speed chases and swimming on their sides with tail flukes emerging from the water.

ADULT
Relatively narrow, fairly pointed Chevron tends to be
rostrum (intermediate between more U-shaped (cf.
broadly U-shaped blue whale V-shaped on fin whale) Relatively small flukes
rostrum and more sharply (width c. 25 per cent
pointed fin whale rostrum) of body length)

Median
notch

Single prominent
longitudinal ridge
Relatively straight
on rostrum
trailing edge

Often has ‘jointed axis’ on

dorsal fin (backward bend
May have fin-whale-like lighter- halfway to two-thirds up)
ADULT coloured chevron (less extensive than
(SOUTHERN) on fin whale and U-shaped – highly
variable in lightness and extent –
more visible in good light)

Slightly larger
than northern
subspecies
Lower ‘lips’ typically dark
to light grey on both sides
(varies according to light
conditions) but some
individuals may show SIZE (NORTHERN)
mild form of fin whale L: ♂ 12–15m, ♀ 13–16m;
asymmetry WT: 16–27t; MAX: 19.5m, 30t
Calf – L: c. 4.4–4.6m; WT: 600–700kg
North Pacific slightly larger than North
Atlantic; female slightly larger than male.
SIZE (SOUTHERN)
L: ♂ 13–16m, ♀ 14–17m;
WT: 22–38t; MAX: 20m, 46t
Calf – L: c. 4.5–4.8m; WT: 650–780kg
Female slightly larger than male.

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 CALF

ADULT

Some anterior plates may

be nearly white (may also be
paler markings on upper ‘lip’)

Baleen generally dark grey or black (may

have diffuse, longitudinal yellowish-brown
streaks) with very fine bristles

Median
notch
FIN VARIATIONS
Tip can be
pointed or
rounded

Shape and curve can vary from triangular

to falcate to backswept

FLUKES

Dark on upperside (may Lighter on underside

have lighter leading edge)

WHAT’S IN A NAME?
‘Sei whale’ comes from the Norwegian ‘seihval’ – off northern Norway at the same time (presumably
‘seje’ for a fish known in English as pollock, saithe, feeding on the same prey). It is normally pronounced
coley or coalfish (a close relative of codfish) and ‘say’ or ‘sigh’ (the Norwegian pronunciation is
‘hval’ for ‘whale’. The two species often appeared halfway between the two).

SEI WHALE 67

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 BRYDE’S WHALE
Balaenoptera edeni Anderson, 1879

One of the least known and more elusive of the large baleen whales, ‘Bryde’s whale’ is actually a complex of subspecies
and possible species with taxonomic issues that are yet to be resolved. They all have one particular characteristic in
common: three parallel longitudinal ridges on the rostrum (all other rorquals, except Rice’s whale and some Omura’s
whales, have a single ridge). The name is pronounced ‘bree-duss’ (the correct Norwegian pronunciation) or, frequently,
‘broo-duss’.
IUCN status Least Concern (2017).
Population No overall global estimate, though broad guesstimate might be 90,000–100,000. Whalers killed at least
30,000 during 1911–87. Trend unknown.
Classification Mysticeti, family Balaenopteridae.
Taxonomy Two subspecies are recognised: the larger, more pelagic and globally distributed Bryde’s whale
(B. e. brydei), otherwise known as the ‘large-form Bryde’s whale’, ‘offshore Bryde’s whale’ or ‘ordinary Bryde’s whale’;
and the smaller, predominantly coastal Eden’s whale (B. e. edeni), of the western Pacific and Indian Oceans. Given
strong genetic and morphological differences, and habitat partitioning, it is highly likely that these should be given full
species status (as Bryde’s whale and Eden’s whale, respectively). The term ‘pygmy Bryde’s whale’ was erroneously
used for whales now known to be Omura’s whale, described as a new species in 2003 (but originally considered part
of the Bryde’s whale complex)
Other names Eden’s whale, tropical whale, sittang.
DISTRIBUTION Circumglobal distribution in tropical, sub-tropical and some warm temperate waters in the Atlantic,
Pacific and Indian oceans, primarily between 40°N and 40°S. Tends to concentrate in water warmer than 16°C, in
areas with exceptionally high productivity. It occurs in some semi-enclosed seas, such as the Red Sea and Persian
Gulf, but is not found in the Mediterranean. Primarily pelagic or coastal, depending on the subspecies. No extensive
north–south migrations are known, although at least some offshore animals make shorter, general movements towards
lower latitudes in winter and mid-latitudes in summer. Other populations – especially inshore in mid-latitudes – remain
year-round in highly productive waters (e.g. Mexico’s Gulf of California, New Zealand’s Hauraki Gulf and the Gulf of
Thailand).
BEHAVIOUR Occasionally breaches (typically coming out of the water vertically), sometimes multiple times in a row (70
times on one exceptional occasion off Ogata, Japan). When feeding, it typically makes sudden changes in direction,
both underwater and at the surface. Behaviour around vessels ranges from taking flight to unconcerned, or even
sometimes curious.
FOOD AND FEEDING Mainly small schooling fish (including pilchard, anchovy, mackerel, herring, sardine, lanternfish);

DIVE Sequence Surfaces at shallow angle; tip of rostrum usually breaks surface
first; dorsal fin usually seen after blowholes submerged (sometimes simultaneously,
especially in younger individuals); tends to arch back before deep dive (cf. sei
whale); does not raise flukes; often impossible to predict and can be erratic in
surfacing behaviour; frequently does not leave telltale flukeprints behind on surface
(cf. sei whale). • Depth Often feeds at or close to surface; maximum 300m. •
Duration 5–15 minutes; maximum 20 minutes.
BLOW Blow up to 9–10m high and columnar to bushy (highly variable height – often
only 3–4m and even shorter in Eden’s); tends to be lower in coastal edeni than
pelagic brydei. • Often exhales underwater, then surfaces with little or no visible
blow (especially if frightened by killer whales or vessel approaching too close).

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 ADULT
Relatively uniform dark grey Tall, strongly falcate dorsal
Three prominent, parallel or bluish-black upperside (may fin (up to 46cm) two-thirds
longitudinal ridges on extend down to include throat to three-quarters of the
rostrum (poorly developed grooves and flippers) can way along back (variable
in some individuals) appear brownish or golden size and shape)
in certain lights
Head 24–26
per cent of Dorsal fin rises at steep
body length angle from back (usually
Relatively Sleek, stream­
less erect than in sei
flat rostrum lined body
whale, less backswept
than in fin whale)

Lower ‘lips’ typically Yellowish or creamy-

uniform dark grey
or bluish-black (not
asymmetrical, cf. fin
whale and Omura’s
whale)
40–70 longitudinal throat
pleats on underside (unusually
long – reach to or past
umbilicus – cf. sei whale)
Throat can white underside May be heavily
be pinkish to (may have pinkish to pockmarked (especially
bright pink strongly pink tinge) rear half) with light
Relatively slender, oval cookiecutter shark
pointed flippers bites (more common in
Flippers dark (c. 8–10 per cent offshore animals – rare
grey or bluish- body length) in coastal animals)
black (both sides) Diffuse boundary fusing
between dark upperside
and light underside

SIZE (LARGER, PELAGIC FORM – BRYDE’S)

L: ♂ 12–14.5m, ♀ 12.5–15m;
WT: c. 15–25t; MAX: 15.6m, 25t
Calf – L: 3.8–4m; WT: 600–750kg
SIZE (SMALLER, COASTAL FORM – EDEN’S)
L: ♂ 10–11.5m, ♀ 11–13m;
WT: 12–17t; MAX: 11.7m, 17t
Calf – L: 3.4–4m; WT: 600–700kg
Female slightly larger than male in both forms.

80°

70° AT A GLANCE Tropical to warm temperate

60° waters worldwide • Large size • Sleek,
50°

40°
streamlined body • Uniform dark grey
30°

20°
upperside, lighter underside • Throat
10°
0°
sometimes pinkish • Three parallel
10°
20°
longitudinal ridges on rostrum • Tall, strongly
30°
falcate dorsal fin two-thirds to three-
quarters of the way along back • Dorsal fin
40°

50°

60°
usually visible after blowholes submerged •
Symmetrical lower ‘lip’ coloration • Typically
70°

arches back and tailstock on diving

80°

possible (minimum) range of Eden’s whale (B. e. edeni)

BRYDE’S WHALE 69

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 also squid, krill, pelagic red crabs, other zooplankton; mostly an opportunistic feeder, switching prey preference
according to availability, geographical location, season and year. Wide variety of foraging techniques; active lunge-
feeder (often attracting seabirds and other pelagic predators); may skim-feed at surface like right whale; sometimes
uses bubble nets to corral prey; in Gulf of Thailand, uses passive feeding technique among schooling fish called ‘trap-
feeding’ or ‘tread-water feeding’ (there are several subtle variations in this technique but, basically, the whale hangs
nearly vertically for several seconds, with mouth wide open at surface, allowing fish to swim or wash inside, then lifts
head up and closes mouth); in New Zealand, uses ‘chin-slaps’ to aggregate zooplankton prey, then side-lunges through
concentrated patch.
BALEEN 250–280 plates (each side of upper jaw) but up to 365 (including many rudimentary plates). Longest plates
c. 50cm; may be more slender in Eden’s whale.
GROUP SIZE AND STRUCTURE Generally seen alone, but sometimes in small groups of 2–3, and occasionally loose
aggregations of 10–20 on prime feeding grounds.

Prominent median ridge

Three ridges not always (running from blowholes
easy to see (especially to tip of rostrum) Shorter auxiliary ridge on each
with water washing side of median ridge (poorly
over rostrum) developed in some individuals)

Slight
median
notch

Relatively broad flukes

(width c. 23–24 per
Relatively narrow, fairly pointed V-shaped cent of body length)
rostrum (intermediate between broadly
U-shaped blue whale rostrum and more
sharply pointed fin whale rostrum)
ADULTS

Plates tend to be yellowish or creamy white in

the anterior quarter to one-third of mouth, often
darkening to slate-grey or dark grey (particularly
on outside) in posterior three-quarters to two-
thirds of mouth

Some individuals have asymmetrical

coloration of throat and baleen plates

CALF

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 FIN VARIATIONS
Height and
Tip usually strength of sickle
pointed shape variable
Fin of some individuals may have slight backward bend halfway to
two-thirds up (cf. Omura’s whale – not as obvious as in sei whale)
FLUKES
light
median
otch
THE BRYDE’S WHALE COMPLEX
The so-called ‘Bryde’s whale complex’ originally
comprised a single species, Bryde’s whale. But
research has revealed that it consists of several
different species: the original Bryde’s whale,
Omura’s whale (formally separated in 2003) and
Rice’s whale (formally separated in 2021). Bryde’s
whale itself currently consists of two subspecies –
or species – and the precise taxonomic status of
these is still in dispute.
Eden’s whale
Appears to be restricted to the northern Indian
Ocean and western Pacific Ocean, roughly between
the equator and 40°N. Genetic studies confirm its
presence in Oman, India, Bangladesh, Myanmar,
Singapore, Indonesia, the Philippines, China, Hong
Kong and as far as south and southwestern Japan.
It is uncertain if its range extends to Australia.
There are no verified records from the Atlantic
Ocean. Primarily in coastal waters (with some
records very close inshore) and over the continental
shelf; it has not been recorded offshore. It seems to
be resident year-round and there is no evidence of
long-distance migrations.
001 WDP FG.indd 71
Often
hooked
at tip
Underside of
flukes typically
creamy white
Bryde’s whale
Circumglobal distribution in tropical and sub-
tropical waters of the Pacific, Atlantic, and Indian
Oceans, and the Caribbean Sea. Genetic studies
confirm its presence in many countries outside
the range of Eden’s whale. The subspecies in
Australia is still uncertain, but in New Zealand
it has been confirmed as brydei. All Bryde’s
whales in the Atlantic Ocean are believed to be
this form. Primarily offshore, but its distribution
is more cosmopolitan than previously thought
and appears to include some coastal habitat
(the resident population in the Hauraki Gulf,
New Zealand, for example, is primarily coastal).
Some offshore populations are known to migrate,
but these migrations appear to be relatively
short for baleen whales (typically over 20°–30°
latitude). Inshore populations tend to be resident
year-round. Recent genetic studies demonstrate
that broadly sympatric populations of migratory
offshore Bryde’s whales and resident inshore
Bryde’s whales off South Africa – which also
differ in size and prey preference – are this large-
form Bryde’s.
BRYDE’S WHALE 71
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 OMURA’S WHALE
Balaenoptera omurai Wada, Oishi and Yamada, 2003

Omura’s whale is one of the most recently described living species of baleen whale. Scientifically named in 2003, it
was initially known from only a handful of specimens and probable sightings at sea. But in the past few years our
knowledge of this slender, tropical whale has grown substantially.
IUCN status Data Deficient (2017).
Population Unknown. Hunted commercially – misidentified as Bryde’s whale – but numbers lacking. Trend unknown.
Classification Mysticeti, family Balaenopteridae.
Taxonomy No recognised forms or subspecies. Previously considered part of the Bryde’s whale complex.
Other names Small-form Bryde’s whale, pygmy Bryde’s whale, dwarf Bryde’s whale or dwarf fin whale.
DISTRIBUTION Knowledge of this whale’s distribution remains rather limited, but there are now many detailed and
verified records from 21 range states. Most are from the Indo-Pacific, on both sides of the equator, but it is also known
from several geographically distant records in the Atlantic (suggesting that the normal distribution could extend into
that ocean). There are no records from the central and eastern Pacific – despite extensive survey effort – and this
could represent a genuine gap in distribution. Records of ‘small Bryde’s whales’ elsewhere may yet prove to be this
species. No long-distance migrations are known. Most sightings are in coastal and shallow waters. Studies in north-
west Madagascar show the species has a preference for shallow water – primarily 10–25m but ranging from 4m to
202m – with sea surface temperatures of 27.4–30.2°C. In this region, it is distributed exclusively over the continental
shelf, within c. 10–15km of the shelf break, and avoids both deep water off the shelf and very shallow coastal water and
bays. There is some evidence of occurrence in deeper water in the Solomon Sea and off the Cocos Islands, and on the
Mid-Atlantic Ridge, and this may be the case in other parts of the range (or at certain times of year).
BEHAVIOUR Breaching has been observed.
FOOD AND FEEDING Mostly krill, minuscule zooplankton and/or fish eggs. Frequently rolls at the surface when lunge-
feeding; observed feeding in areas with high densities of whale sharks (perhaps feeding on the same prey).
BALEEN 180–210 plates (each side of the upper jaw). Fewer plates than any other Balaenoptera species; longest plates
23–28cm.
GROUP SIZE AND STRUCTURE Generally seen alone, in mother–calf pairs, in temporary pairings of two adults (lasting no
more than 10 minutes), or in loose aggregations of up to six animals within a few to several hundred metres of each other.

Broad, flat head Relatively straight ADULT

Single prominent longitudinal
ridge from blowhole to tip of trailing edge
rostrum (most Bryde’s whales
have three) Broad flukes

Some populations show heavy

Front of rostrum scarring from cookiecutter
fairly U-shaped May have weak lateral ridge on either side of central ridge shark bites (others show no
(not unlike blue (detectable only at some angles in certain light conditions evidence of these bites)
whale) and more pronounced in young animals and calves)

DIVE Sequence Head breaks surface at relatively shallow angle; dorsal fin typically emerges
just as head and blowholes submerge; does not fluke on diving. • Depth Unknown, but
probably typically less than 100m. • Duration Unknown.
BLOW Diffuse, bushy blow, rarely conspicuous (height highly variable).

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 ADULT
Asymmetrical coloration Light-coloured, single- Blue-grey Relatively small, LEFT SIDE
banded chevron just upperside Slender, low, strongly falcate
of inner lower ‘lips’ streamlined
opposite to outer lower behind flipper (less dorsal fin (variable)
pronounced on left side) body
‘lips’ (lighter on left, Dorsal fin two-
darker on right) thirds of the way
No dark stripes along back

80–90 longitudinal Cream-coloured to

Darkly pigmented, blue-grey left
lower ‘lip’ (‘lip’ area always dark
but may be light underneath
towards throat), cf. right side
Dorsal fin may
ADULT
RIGHT SIDE be hooked
throat pleats (extend light grey underside
Moderately long, slender as far as umbilicus)
flippers (light upperside
with lighter leading edge) Diagnostic light-
coloured blaze
Overall, white pigmentation 4–5 dark stripes originating in front
appears more extensive on (including eye and of eye that sweeps
right side than left ear stripes) across back (absent
from left side)

Light-coloured chevron in front of

Lightly pigmented,
dorsal fin (more pronounced and White leading edge of flipper more cream-coloured
often double-banded on right side) extensive on right side (gradually to light grey right
fades into darker pigmentation) lower ‘lip’
FLUKES

SIZE
L: 8–12m;
WT: unknown (probably less than 20t) Underside of
Calf – L: c. 3–4m; WT: unknown flukes off-white Irregular dark
Females presumed to be slightly larger margins
than males (as in other rorquals).

80°

70°

60°
AT A GLANCE Mainly Indo-Pacific (also
Atlantic) • Shallow, nearshore tropical and
50°

40°

30°

20°
sub-tropical waters • Large size • Distinctly
10°
0°
countershaded • Asymmetrical lower-‘lip’
10°
20°
coloration • Relatively small, low, strongly
30°

40°
falcate dorsal fin • Body pigmentation
50° similar to fin whale • Single prominent
60°
ridge on rostrum • Alone or in small, loose
aggregations
70°

80°

confirmed range visual, acoustic and other records

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 RICE’S WHALE
Balaenoptera ricei Rosel, Wilcox, Yamada and Mullin, 2021

A small population of whales belonging to the ‘Bryde’s whale complex’ has been known in the Gulf of Mexico since an
individual stranded there in 1965. But recent research reveals that these whales are genetically and morphologically
distinct (and geographically separated from other Bryde’s whale populations). They were identified as belonging to a
new species in 2021.
IUCN status Not yet evaluated as a separate species, but Critically Endangered (2017) as a subpopulation of Bryde’s
whale. With such a small population and such a limited range, arguably the most endangered whale species in the
world.
Population Current best estimate 33 (certainly fewer than 100). In 2010, an estimated 17 per cent of the population
died in the wake of an oil spill.
Classification Mysticeti, family Balaenopteridae.
Taxonomy The genetic differences between Rice’s whale and Bryde’s whale are two to three times greater than the
differences between the three recognised species of right whale. No recognised forms or subspecies.
Other names Gulf of Mexico Bryde’s whale, Gulf of Mexico whale.
DISTRIBUTION Based on 181 sightings from 1989–2019, the distribution appears to be restricted predominantly to
the continental shelf break near the De Soto Canyon, c. 100km south of Mobile, Alabama (US), in the northeastern
Gulf of Mexico. There have also been two possible sightings (in the early 1990s) and one confirmed sighting (in 2017)
in the western Gulf of Mexico and unique vocalisations have been heard there, south of Louisiana. These records
suggest three possibilities: some individuals venture out of their core range into the western side; there is another, as
yet unidentified, population; or, most likely, these are the remnants of a formerly more broadly distributed population.
Whaling data does suggest a wider past distribution (records of ‘finback’ whales from the north-central Gulf, south of
the Mississippi, and in the southern Gulf on the Campeche Banks are almost certainly misidentified Rice’s whales).
There have been no confirmed sightings outside the Gulf of Mexico, and it is believed to be non-migratory. Single
strandings in South Carolina and North Carolina are believed to have been extralimital strays. It is the only year-round
resident baleen whale in the Gulf of Mexico and there is no genetic evidence for any other Bryde’s-like whale species
or subspecies in the Gulf. The nearest confirmed populations of other members of the Bryde’s whale complex are in the
southern Caribbean south to Venezuela and Brazil, the eastern North Atlantic and the eastern South Atlantic off South
Africa. Sightings and strandings of all other baleen whale species in the Gulf are rare and considered extralimital.
During the 181 sightings, the whales were observed almost entirely in water 151–352m deep (with two exceptions, in
117m and 408m respectively). A satellite-tagged individual remained in waters 100–400m deep for a month.
BEHAVIOUR Unknown, but likely similar to other members of the Bryde’s whale complex.
FOOD AND FEEDING Prey unknown. Pelagic schooling fish such as herring, anchovies and sardines are the preferred
prey of most whales in the Bryde’s whale complex. Some populations also target squid, krill, pelagic red crabs and
other species. Lanternfish and hatchetfish are abundant in the region, and may be taken. Feeding techniques unknown.
BALEEN c. 264 plates (each side of the upper jaw).
GROUP SIZE AND STRUCTURE Generally seen alone, or in pairs. May occasionally form larger groups in productive
feeding areas.

DIVE Depth Unlike Bryde’s whales, which tend to feed near the surface, limited
evidence suggests foraging close to the seafloor (deepest recorded 271m), but rarely
in water deeper than 350m; some evidence indicates that they spend a considerable
amount of time shallower than 15m at night. • Duration Unknown.

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 Relatively uniform dark charcoal grey Sleek, stream-
Tall, strongly falcate
Relatively upperside (may appear brownish in lined body
dorsal fin two-thirds of
flat rostrum some lighting conditions)
the way along back

Lower ‘lips’ typically

uniform dark charcoal c. 54 longitudinal throat Relatively slender, Lighter (sometimes
grey on both sides (not pleats on underside pointed flippers uniformly pinkish) underside
asymmetrical cf. fin whale (unusually long – some dark grey (both sides)
and Omura’s whale) reach to or past umbilicus) ADULTS

Three prominent, parallel longitudinal May be diffuse white washes around

ridges from blowhole to tip of rostrum base of dorsal fin and/or along sides
(not always easy to see – especially (some individuals only) but no white
with water washing over rostrum) chevrons or blazes on back or sides

Central ridge
Relatively narrow, larger, lateral
fairly pointed ridges smaller
V-shaped rostrum
Cream-coloured
anterior baleen
plates on both
sides, posterior
plates black with
cream-coloured
SIZE fringe
L: ♂ 11.26m (largest adult measured; FLUKES
an immature was 11.05m); ♀ 12.65m
(largest adult measured);
WT: possibly c. 12–17t
Calf – L: 4.7m (one calf of unknown Underside of
age that stranded alive); WT: unknown flukes lighter
Females presumed to be slightly larger
than males (as in other rorquals).

40°

AT A GLANCE Core range in the northeastern

Gulf of Mexico • Larger than Omura’s whales,
smaller than Bryde’s whales (roughly the same
30°

size as Eden’s whales) • Sleek, streamlined

20° body • Uniform dark charcoal grey upperside,
lighter to pinkish underside • Three parallel
longitudinal ridges on rostrum • Tall, strongly
falcate dorsal fin two-thirds of the way along
10°

back • Symmetrical lower ‘lip’ coloration

0°

core range
core range

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 COMMON MINKE WHALE
Balaenoptera acutorostrata Lacépède, 1804

The common minke whale is the smallest rorqual and the second smallest of all the baleen whales (after the pygmy
right whale). It has three disjunct populations: in the North Atlantic, the North Pacific and the southern hemisphere.
IUCN status Least Concern (2018).
Population Minimum c. 200,000 mature individuals. At least 170,000 killed by whalers. Trend unknown.
Classification Mysticeti, family Balaenopteridae.
Taxonomy Two subspecies are currently recognised: North Atlantic minke whale (B. a. acutorostrata); and North
Pacific minke whale (B. a. scammoni). The ‘dwarf minke whale’ may be a third valid subspecies.
Other names Northern minke whale, dwarf minke whale; formerly lesser/least rorqual, little piked whale, pikehead,
lesser finback, sharp-headed finner, little finner.
DISTRIBUTION In the northern hemisphere, the North Atlantic minke whale ranges to at least c. 80°N during summer.
The wintering grounds – probably in the southern North Atlantic – are poorly known but extend at least to the Caribbean
and possibly West Africa. The North Pacific minke whale ranges to at least 70°N during summer. The wintering grounds
– probably in the southern North Pacific – are also poorly known but extend to at least 15°N. Northern minke migrations
are not as well defined as in some other baleen whales. There is a tendency for the distribution to shift from high-
latitude summer feeding areas to lower-latitude winter breeding areas (although some individuals are resident in cold
temperate regions year-round). During summer, they appear to be most abundant in cold temperate to polar waters
(where they are known to penetrate areas with extensive ice floes and polynyas). At this time of year, they occur in
inshore coastal waters more frequently than any other rorqual, and will enter bays, inlets, fjords and even some large
rivers (such as the St Lawrence River, Canada). Winter sightings are uncommon, suggesting that when in lower latitudes
they are mainly offshore. Some populations – such as around the Isle of Mull (Scotland, UK) and the San Juan Islands
(Washington, USA) – have high site fidelity, with certain individuals returning each year to feed in particular locations.
The dwarf minke occurs only in the southern hemisphere and may or may not be circumglobal (relatively little is
known about its distribution). Occurs in both coastal and offshore waters off South Africa, southern Mozambique,
Australia, New Zealand (North and South Islands), New Caledonia, eastern South America (from northern Brazil to
northern Argentina), and Chilean Patagonia. Records cover most of the year (March–December), but there are strong
indications that at least some populations are migratory. The only known predictable aggregation of dwarf minke
whales is off the northern Great Barrier Reef in Australia, predominantly in June–July. The most northerly confirmed
records are from 2°S off the northern coast of Brazil and 11°S in the western Pacific off Australia. It partly overlaps with
the Antarctic minke whale during summer in the sub-Antarctic but is not as polar. Most sightings in the sub-Antarctic
have been in December–March south of Australia and New Zealand – between 55°S and 60°S, with one record as far
as 65°S – probably because this is where there has been most research effort. However, it is also likely to occur in sub-
Antarctic waters south of South America and South Africa. It is not known from the northern Indian Ocean.
BEHAVIOUR Breaches fairly frequently, sometimes completely clearing the water, and performs other aerial behaviours
such as head rises and spyhops (particularly in icy areas). Rarely lobtails or fipper-slaps. Can be quite curious towards

DIVE Sequence Surfaces at a distinct angle c. 20–40°; tip of rostrum pointed up and
distinctively breaks surface first; blowhole and dorsal fin generally visible at same
time; typically arches back and tailstock quite high before sounding dive; does not
raise flukes. • Depth Feeds mostly at or near surface; dwarf minkes observed at
20–40m. • Duration Northern typically 3–10 minutes, maximum 20 minutes; dwarf
minke up to 12.5 minutes.
BLOW Diffuse, upright blow up to 3m high (height highly variable), more bushy than
columnar. • Generally less distinct than any other large whale blow (often invisible).

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 ADULT NORTHERN HEMISPHERE
Variable pale shoulder streak behind Brush strokes and
flippers (typically well-defined intermediate shades of
leading edge) may extend to form grey on sides and back
chevron pattern over back (roughly (highly variable)
symmetrical with other side) Relatively tall, falcate
dorsal fin slightly less
Dark grey, than two-thirds of
Head 22–23 per the way along back
brownish-grey
cent of body length (variable shape)
or blackish
upperside Dorsal fin may
Sharply pointed, No white shoulder be hooked at tip
flattened rostrum patch (cf. dwarf
minke) May have distinct
dorsal and ventral
keels

May be
50–70 moderately pockmarked
short longitudinal Relatively sleek,
with light
throat pleats on Pointed tip streamlined body
round or oval
underside (terminate cookiecutter
Snowy-white to creamy-
between flippers Sharply demarcated shark bites
white underside (may
and umbilicus) brilliant white band flush pink when active)
in middle of flipper Slender, pointed
(often clearly visible flippers (c. 12 per cent
when animal just of body length)
below surface)

SIZE (NORTH PACIFIC AND NORTH ATLANTIC)

L: ♂ 7.5–8.5m, ♀ 8.5–9.5m;
WT: 6–8t; MAX: 9.8m, 9.2t
Calf – L: 2.2–2.8m; WT: 350–450kg
The female is longer than the male in all subspecies.
SIZE (DWARF)
L: ♂ 6–7m, ♀ 6.5–7.2m;
WT: 4–5t; MAX: 7.8m, 6.4t
Calf – L: 2–2.3m; WT: 250–350kg

80°

70°

60°

50° AT A GLANCE Tropics to poles worldwide

• Medium size • Dark grey, brownish-grey
40°

30°

20°
10°
or blackish upperside, white underside •
0°
10° Variable swathes of lighter grey on sides and
20°

30° back • Sharply pointed rostrum breaks surface

40°

50°
first • Single longitudinal ridge on rostrum
60°
??
• Relatively tall, falcate dorsal fin two-thirds
70°
of the way along back • Unique, bright white
80°
flipper bands •Indistinct or invisible blow
primary range known minimum distribution of
possible range dwarf minke whale (based on reported
sightings – largely reflecting most
secondary range
sighting and hunting effort)

COMMON MINKE WHALE 77

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 boats in some parts of the world – and will swim around stationary vessels or alongside moving vessels for minutes or
even hours at a time. Elsewhere, it can be difficult to approach.
FOOD AND FEEDING Northern hemisphere minkes feed on wide variety of small schooling fish (including sand
lance, salmon, capelin, cod, mackerel, sprat, pollack, whiting, herring, haddock, anchovy and lanternfish) and small
invertebrates (including euphausiids and copepods). Dwarf minkes prefer lanternfish, but opportunistically feed on other
fish and possibly krill. Feeding technique varies significantly according to prey and location; entrapment manoeuvres
include circles, gyres, ellipses, figures-of-eight, hyperbolas, head-slaps and underwater blows; engulfing manoeuvres
include oblique, lateral, vertical and ventral lunges.
BALEEN 231–290 plates in North Pacific, 270–325 in North Atlantic, 200–300 in dwarf (each side of the upper jaw).
Longest plates c. 21cm; in northern animals, usually white, creamy or yellowish; in dwarf minke, about half of the
plates posteriorly appear dark grey or brown (due to a narrow, dark fringe); all have symmetrical plate coloration.
GROUP SIZE AND STRUCTURE Typically solitary, sometimes in twos or threes, but there can be larger, temporary
aggregations in good feeding areas. Social structure appears to be complex, with evidence for some segregation by
age, sex and/or reproductive class.

ADULT NORTHERN HEMISPHERE

Single prominent Roughly symmetrical
longitudinal ridge chevrons on back (highly
on rostrum variable – often not visible)

Median
notch

Relatively small flukes

Narrow, sharply (width c. 23–28 per
pointed V-shaped Diagnostic white cent of body length)
Smooth
rostrum patch on each flipper trailing edge
(variable in extent)

CALF NORTHERN HEMISPHERE

Coloration highly variable
between individuals

ADULT NORTHERN HEMISPHERE FLIPPERS FLUKES Borders and trailing

edge dark grey

Size and shape of white

flipper band varies
between individuals Predominantly ivory-
white underside

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 ADULT DWARF
Complex lateral
Dark grey coloration with Coloration
upperside three dark grey highly variable
Most complex
fields descending between
coloration of
Dark grey band on from back individuals
any baleen Relatively larger
neck continues as whale dorsal fin than
dark throat patch
common minke

Tailstock
slightly
longer

Lower jaw
dark grey or
brownish- Ivory-white Side swirl in shape
grey (variable underside of Salvador Dali
extent) Roughly moustache
Tip of flipper Flipper tends to have
triangular,
usually much longer band of Ivory-white side streaks and
pale grey
remains dark ivory-white extending to blazes typically extend from
throat patch
base and onto body as underside onto back
highly distinctive broad
Median white shoulder patch
otch

55–75 throat pleats Underside of flukes

mostly ivory-white
(dark grey trim at tips
and onto trailing edges)

Underside of flippers
Symmetrical large, dark grey similar to upperside
throat patches in front of flippers
(do not meet in middle)

ADULT DWARF FLIPPERS ADULT DWARF

May have light grey ‘blowhole

streaks’ visible from above (left
streak consistently curves to left)

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 ANTARCTIC MINKE WHALE
Balaenoptera bonaerensis Burmeister, 1867

The Antarctic minke whale was declared a new species in 1998, when it was formally split from the slightly smaller
common minke whale (although the two species are believed to have diverged 4.7–7.5 million years ago). It is actually
more closely related to sei and Bryde’s whales than to its namesake.
IUCN status Near Threatened (2018).
Population Minimum 515,000. At least 140,000 killed by whalers. Trend unknown.
Classification Mysticeti, family Balaenopteridae.
Taxonomy No recognised forms or subspecies.
Other names Southern minke whale.
DISTRIBUTION Endemic to the southern hemisphere. It is believed to be circumpolar, and occurs both nearshore and
offshore. There have been four extralimital records in the North Atlantic: Suriname on the Atlantic coast of South
America; Togo in the Gulf of Guinea; Louisiana in the northern Gulf of Mexico; and north of the Arctic Circle off Jan
Mayen. Two individuals identified as hybrids with common minke whales have been recorded in Svalbard.
Most abundant south of 60°S during the southern summer – south of the Antarctic Circumpolar Current – and known
to reach at least 78°S in the Ross Sea. It occurs in greatest densities near the ice edge and is probably the most ice-
affiliated rorqual (it is least abundant in ice-free waters). Found from the ice edge to several hundred kilometres inside
the ice in anything from brash to floes – even with nearly 100 per cent ice cover – following leads and utilising polynyas
for breathing; it may even use its pointed, stout rostrum to break through newly formed or tightly packed brash ice to
create breathing holes. It partly overlaps with the dwarf minke whale (B. acutorostrata) during summer in the sub-
Antarctic, but tends to be much more polar.
Migrations are not as well defined as in some other baleen whales. Many Antarctic minkes appear to overwinter
in the Antarctic, but others may migrate from their high-latitude feeding grounds to dispersed lower-latitude winter
breeding areas. Most calving seems to take place north of the Antarctic Convergence. There may be poorly defined
breeding grounds roughly from 10°S to 30°S in the Pacific (between 170°E and 100°W), as well as west of Easter Island
in Chile, off northeastern and eastern Australia, western South Africa and Brazil’s northeastern coast. Intriguingly,
Antarctic minke whale vocalisations have been heard simultaneously during winter and spring in the eastern Weddell
Sea and off Western Australia, indicating a very widespread distribution – or a seasonal migration by one segment of
the population and a year-round presence in Antarctic waters by another.
BEHAVIOUR Breaches fairly frequently, sometimes completely clearing the water, and performs other aerial behaviours
such as head rises and spyhops. In dense pack ice, frequently lifts its head right out of the water to breathe. Behaviour
around vessels ranges from taking flight to unconcerned or even curious; tends to be more approachable when feeding.
There appears to be a growing tendency towards curious and ‘friendly’ behaviour among single smaller whales around
the Antarctic Peninsula, with increasing close encounters with rigid-hulled inflatable boats and kayaks.

DIVE Sequence Surfaces at distinct angle c. 20–40°; tip of rostrum pointed up and
distinctively breaks surface first; blowhole and dorsal fin generally visible at same
time; typically arches back and tailstock quite high before sounding dive; does not
raise flukes. • Depth Generally feeds in upper 100m of water column (up to 150m);
dive depths may change from day to night with vertical migration of prey. • Duration
Typically 1–5 minutes; maximum c. 15 minutes; typically surfaces 2–15 times between
deep dives.
BLOW Diffuse, upright blow up to 3.5–4m high (height highly variable), more bushy than
columnar. • Generally more distinct than common minke whale blow but highly variable
(conspicuous in colder Antarctic waters, but can be invisible at lower latitudes).

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Variable pale grey shoulder
streak behind flippers (may
extend to form chevron
pattern over back)
Sharply pointed,
flattened rostrum
ADULT
Brush strokes and lobes of
Dark grey,
pale to medium grey along
brownish-grey or
sides (greatly undulating zone
blackish upperside
of transition between light
underside and dark upperside)
Prominent, tall, falcate
dorsal fin slightly less
Relatively sleek,
No white shoulder than two-thirds of
streamlined body
patch (cf. dwarf the way along back
minke whale) (variable shape)
May have
distinct dorsal
and ventral keels

May have dark

60–70 longitudinal throat
pleats on underside
(terminate up to 30cm
anterior to umbilicus)
Slender, pointed
flippers (c. 13 per
cent of body length)
'flipper shadow'
mark
No transverse white Flippers uniform pale
band on flippers grey (sometimes with
white leading edge)
(cf. common minke
whale)
May be pockmarked
with light round or oval
cookiecutter shark bites
Snowy-white to creamy-
white underside (may
flush pink when active)

ADULT FLIPPER VARIATIONS

Extent of pale grey and
white bordering variable

SIZE
L: ♂ 8–9m, ♀ 8.5–9.5m;
WT: 7–9t; MAX: 10.7m, 11t
Calf – L: 2.6–2.8m; WT: 350–500kg
Female is longer than the male.

80° AT A GLANCE Tropics to poles in southern

70° hemisphere • Medium size • Dark grey,
60°
brownish-grey or blackish upperside, pale
underside • Variable undulating swathes
50°

40°

of lighter grey on sides and back • May

30°

20°
10°
0° be blotchy ochre with diatoms • Sharply
10°
20°
pointed rostrum breaks surface first • Single
30°

40°
longitudinal ridge on rostrum • Tall, falcate
50°
dorsal fin two-thirds of the way along back
• Light grey chevron across back (at level of
60°

70°

80°
flippers) • Often has light grey pigmentation
trailing from blowholes • Pale grey flippers (no
primary range extralimital records hybrids with common minke whales white band) • Distinct blow in high latitudes

ANTARCTIC MINKE WHALE 81

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 FOOD AND FEEDING Primarily Antarctic krill on summer feeding grounds in offshore waters (e.g. Weddell Sea);
sometimes smaller krill in coastal shelf waters (e.g. Ross Sea); known to feed on krill under the ice; occasionally
amphipods and Antarctic silver fish. Lunges into large prey aggregations (often rolling to one side); most feeding may
be during southern summer, but little information; no accounts of cooperative feeding, though groups of feeding whales
observed around Antarctic Peninsula; lunge-feeds up to 22–24 times per dive.
BALEEN 261–359 plates (each side of the upper jaw). Longest plates c. 31cm; asymmetrical coloration: most plates are
black, except for the first few on the left side and the first third on the right side (which are yellowish-white).
GROUP SIZE AND STRUCTURE On summer feeding grounds, typically solitary or in groups of 2–6; larger aggregations
of up to 50 are uncommon, but are seen occasionally around the Antarctic Peninsula (socialising as well as feeding).
Groups of 2–5 are common in warmer waters during winter. Groups of one male and two females, or two males and
three or more females, have been reported in the western South Atlantic. There may be some segregation by age, sex
and/or reproductive status.

Single prominent
longitudinal ridge
on rostrum
ADULT
Pale swirling pigmentation
behind blowholes (reminiscent
of smoke billowing out) Relatively small flukes
(width c. 25 per cent of
body length)

Median
notch
(variable)

Narrow, sharply Underside of flukes snowy-

pointed V-shaped white to creamy-white
rostrum Usually no white May have pale grey with dark grey trim Smooth trailing
patch on flipper chevron on back
edge
between flippers

May be tinged yellowish-orange ADULT WITH DIATOMS

or ochre with film of diatoms
(or diatoms appear as distinct
patches)

ADULT

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 CALF
ADULT DORSAL FIN VARIATIONS
Median
notch
variable)
DIATOMS
Many cetacean species often have a thin yellowish,
brownish, greenish or orangish film – either
continuous or in irregular patches – over their
skin. This is formed by microscopic single-celled
algae, called diatoms. There are countless tens
of thousands of species of diatoms – they are the
key primary producers in the ocean – and most are
free-living. Only four genera and a small number
of species have been found on cetacean skin. The
cold-water Bennettella (formerly Cocconeis) ceticola
is the most common species on baleen whales
and killer whales, in particular, and can cover their
bodies after an extended stay in polar waters; it has
never been found free-living.
Diatom films are most common on cetaceans
living in cold waters. Normally, whales slough and
regenerate their skin continually (to repair scars,
sunburn etc.) but the build-up of diatoms strongly
indicates that this is not happening in cold waters. It
is believed that, to conserve body heat in subfreezing
temperatures, the whales reduce blood flow to their
outer skin, which precludes normal skin growth and
sloughing (allowing the diatoms to accumulate).
Therefore the extent of the diatom film is a rough
001 WDP FG.indd 83
Most dorsal fins
have pointed tips
indicator of how long a whale has spent in cold
waters. In the Antarctic, for example, it takes about
a month to develop, so its extent can be used to
judge the length of time an animal has been in the
region. There is some evidence of thicker layers of
diatoms on fatter individuals, also suggesting that
they have been feeding in productive colder waters
for longer.
The diatom layer can get so thick that it may
cause significant drag, slowing the animals down
(diatoms are a big problem for ships – reducing
speed by up to 5 per cent – hence the use of anti-
fouling paints). Indeed, Antarctic killer whales make
rapid migrations into tropical waters where they
incidentally shed the diatoms when they regenerate
their skin tissue; round trips typically last 5–7 weeks,
and they return to the cold waters of the Antarctic
looking much ‘cleaner’.
The dominant diatom species on Antarctic minke
whales is Bennettella ceticola, but three other
species are known to occur in smaller numbers. As
the southern summer progresses, both the extent of
diatom films and the number of individuals exhibiting
them increases.
ANTARCTIC MINKE WHALE 83
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 HUMPBACK WHALE
Megaptera novaeangliae (Borowski, 1781)

Renowned for its spectacular breaching, lobtailing and flipper-slapping, its complex and melodious song, and its remarkably
long flippers, the humpback whale is one of the most familiar and best known of all the large whales. Black-and-white
markings on the undertail are distinctive and readily identifiable, enabling researchers to tell one individual from another.
IUCN status Least Concern (2018). Arabian Sea sub-population Endangered (2008). Oceania sub-population
Endangered (2008).
Population Minimum 140,000 worldwide (compared to an all-time low of 5,000–10,000 due to whaling). Pre-whaling
population c. 240,000. At least 300,000 killed by whalers. Increasing.
Classification Mysticeti, family Balaenopteridae.
Taxonomy Three subspecies are recognised: North Atlantic (M. n. novaeangliae); North Pacific (M. n. kuzira); and
southern (M. n. australis). A fourth subspecies, the Arabian Sea humpback whale (M. n. indica), has been proposed.
Other names Hump-backed whale.
DISTRIBUTION Worldwide. Migrates between mid- to high-latitude summer feeding grounds and low-latitude winter
breeding grounds (most populations migrate through deep oceanic waters). During summer, frequents coastal,
continental shelf and offshore waters. During winter, breeds around oceanic islands, offshore seamounts and reef
systems. Significant numbers also occur at mid- to high latitudes during winter – in British Columbia, Norway, Iceland
and other locations – but it is unclear whether these are overwintering or simply very late-leaving migrants. Some
populations are responding to climate change by expanding their range into higher polar latitudes (significant declines
in sea ice are also resulting in longer feeding seasons). The breeding area in Central America used by southern
hemisphere humpbacks during the austral winter is also used by North Pacific humpbacks during the boreal winter (it’s
unclear whether they ever overlap at the same time).
Winter breeding distribution There are 15 known and two suspected winter breeding grounds: two (plus one
suspected) in the North Atlantic, five (plus one suspected) in the North Pacific, seven in the southern hemisphere and
one in the Arabian Sea; the isolated population of c. 90 individuals in the Arabian Sea is unique in being resident. All
are between 30°N and 40°S, mostly centred around c. 20° latitude. They are typically in warm, relatively shallow water
(less than 200m) surrounded by much deeper water. The preferred sea temperature on the breeding grounds is c. 25°C
(in the range of 21.1–28.3°C). There is high fidelity to natal breeding grounds and little interchange between them.
Summer feeding distribution Individuals usually return to the same feeding grounds used by their mothers, with
relatively little interchange between feeding areas. Habitat preferences include the continental shelf break, submarine
channels, oceanic fronts, eastern boundary currents and ice-edge zones (areas of upwelling); in the southern
hemisphere, feeding habitat is often closely linked to regions of marginal sea ice. The preferred sea temperature on
feeding grounds is typically below 14°C.
North Atlantic Most North Atlantic humpbacks breed in the West Indies, primarily on the oceanic side of many
Caribbean islands, and in northern Venezuela (the most populous areas are off the northern Dominican Republic and

DIVE Sequence Blowholes appear first and remain in view as dorsal fin appears;
distinctive sloping back forms shallow triangle with surface of sea; body arches, forming
high triangle, making hump on back especially evident; flukes lifted high on many dives.
• Depth On summer feeding grounds often follows diel movement cycle of prey (deeper
during day); most foraging in upper 120m (upper 25m during bubble-netting), but capable
of 400+m; on breeding grounds usually shallow. • Duration Depends on season, location
and behaviour; singing dives up to 20 minutes; resting dives on breeding grounds 15–30
minutes; foraging dives typically 3–10 minutes (up to 15); maximum c. 40 minutes.
BLOW More variable than in any other large whale (bushy, columnar or, very occasionally,
V-shaped). • Usually tall and columnar (may be bushier at top), up to 10m high (height
highly variable – often only 4–5m).

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 Body often dotted ADULT MALE NORTHERN HEMISPHERE
22–64 knobs (called with barnacles Dark grey to
tubercles) on top of and whale lice black upperside
head and much of (Cyamus boopis) and sides Dorsal fin
lower jaw Low, broad-based dorsal fin
sits on raised (up to 30cm) slightly more
Acorn barnacles usually Stocky fleshy hump than two-thirds of the way
found along rostrum, body along back
‘lips’ and throat
Dorsal fin highly
variable (from small
and blunt to tall
and falcate)

Relatively
Distinctive 14–35 ventral narrow
cluster of acorn pleats or throat Less extensive tailstock
barnacles on tip grooves (extend white on underside
of chin (known as to umbilicus or compared to southern Black-and-white
the ‘cutwater’) beyond) hemisphere animals pigmentation on
underside of flukes
White circular scarring on lower Flippers up to 5m long
jaw (scars from when acorn (23–33 per cent of total body
length) and weigh up to 1t Underside varies from black to
barnacles have fallen off) white or mottled black and white
Scalloped leading edge with knobs or (variable between individuals and
tubercles (including two more prominent by population – more white in North
ones dividing margin into thirds) often Atlantic, less in North Pacific)
encrusted with acorn barnacles

SIZE
L: ♂ 11–15m, ♀ 12–16m;
WT: 25–35t; MAX: 18.6m (historically
– rarely more than 16m today), 40t
Calf – L: 4–4.6m; WT: 0.6–1t
Adult females typically 1–1.5m longer
than males.

80°

70° AT A GLANCE Worldwide distribution

60°
• Large size • Predominantly dark grey
to black upperside • Variable amount of
50°

40°

30°

20°
white on underside • Stocky body • Small
10°
0°
dorsal fin sits on (variable) hump on back •
10°
20°
Exceptionally long white (or black-and-white)
30°

40°
flippers • Distinctive knobs on head
50° • Strongly arches back when diving •
60°
Usually flukes on sounding dive • Variable
(and individually distinctive) black-and-white
70°

pigmentation on underside of flukes

80°

primary range secondary range

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 over the offshore reef systems of Silver, Navidad and Mouchoir banks). Historical whaling records reveal Cape Verde
(possibly extending to the continental shelf of Senegal and Western Sahara) as another breeding ground, with up to
4,000 whales before whaling commenced. However, there are relatively few whales today – far fewer than are known
to exist in the eastern North Atlantic (and unaccounted for in the West Indies), suggesting the existence of a third, as
yet undetermined, breeding ground.
North Pacific There are five known and one suspected breeding populations in the North Pacific: the main Hawaiian
islands, accounting for about half of all North Pacific humpbacks (about half feeding off northern British Columbia
and south-east Alaska, and half in the northern Gulf of Alaska and the Bering Sea); mainland Mexico (feeding mainly
off California); Revillagigedo Islands (feeding mostly from northern California to Alaska); Central America, along the
Pacific coast from southern Mexico to Costa Rica (feeding almost exclusively off California and Oregon); southern Japan
(mainly the Okinawa Islands) to Taiwan and the northern Philippines, and east to the Mariana and Marshall Islands
(feeding mainly off eastern Kamchatka, but also across a broad band in the western Pacific). There is also an unknown
breeding ground somewhere else in the western North Pacific – possibly in the Northwestern Hawaiian Islands and/
or the Mariana Archipelago (between the Philippines and Japan); this is inferred from sightings in the Russian Far
East and around the Aleutian Islands that cannot be linked to any known breeding population. Baja California and the
Ogasawara Islands are not believed to be primary migratory destinations but are more likely to be transiting areas.
Southern hemisphere Seven separate breeding populations migrate to feed in the Southern Ocean (in six specific
areas designated by the International Whaling Commission as I–VI): Pacific coasts of Central and South America, from
northern Peru to Costa Rica (with records as far north as Nicaragua) but particularly Colombia (feeding in Antarctic
Area I, particularly off the western Antarctic Peninsula, as well as in southern Chile); coastal waters off Brazil from
3–23°S, particularly around Abrolhos Bank (feeding in the Scotia Sea, i.e. Antarctic Area II); western Africa, centred
around the Gulf of Guinea (feeding south of 18°S, in waters off Namibia and western South Africa, and in Antarctic Area
III); southeastern Africa and Madagascar (feeding in Antarctic Area III); northwestern Australia (feeding in Antarctic
Area IV); northeastern Australia (feeding in Antarctic Area V); and Oceania or the South Pacific islands (feeding in
Antarctic Areas V, VI and I, including the western Antarctic Peninsula).
BEHAVIOUR More demonstrative at the surface than any other large whale. Breaching, flipper-slapping and lobtailing
are common. May lie on side or back, holding one or both flippers in the air. All these activities occur year-round and in

ADULT Tips of flukes often encrusted

Head shape broadly
intermediate between blue with clusters of acorn and
whale (U-shaped) and fin stalked barnacles
Single low,
longitudinal median whale (sharply pointed)
Parallel linear scars on
ridge on rostrum
tail (result of unsuccessful
(highly variable)
attacks by killer whales)

Deep
median
notch

Flukes dark
grey to black
on upperside
Median ridge
generally obscured Tips usually
by knobs or tubercles pointed Serrated trailing
edge diagnostic
to this species

CALF

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 a variety of contexts, so clearly perform various functions (probably including communication, mate attraction, parasite
removal, prey corralling, expression of excitement or annoyance, and even play). Both sexes and all ages frequently
breach (sometimes fully) on both the feeding grounds and the breeding grounds, alone or in small groups, once or many
times consecutively. Shows little fear of boats: frequently very inquisitive (especially juveniles).
FOOD AND FEEDING Krill; wide variety of schooling fish; occasionally squid; some mysids, copepods and benthic
amphipods; generalist in northern hemisphere (with regional preferences); in southern hemisphere, mainly Antarctic krill.
Adaptable gulp- or lunge-feeder using diverse techniques; uniquely among large whales, uses bubble nets (circular nets
of large bubbles) or bubble clouds (bursts of tiny bubbles) to corral schooling prey (individually or in groups); one of few
baleen whales to group-feed. Other feeding techniques include flick-feeding (upside down, repeatedly sweeping tail
forward at surface before surfacing and lunging with mouth open), lobtail-feeding (slapping surface above school of fish,
then diving to blow bubble screen) and trap-feeding (at surface, opening mouth wide, using flippers to push fish into the
‘trap’); little or no feeding on winter breeding grounds; opportunistic feeding during migration.
BALEEN 270–400 plates (each side of the upper jaw). Plates dark grey to black, often with white or brownish-white
longitudinal streaks (anterior-most plates may be lighter); max. length 85–107cm.
GROUP SIZE AND STRUCTURE On winter breeding grounds, usually occurs singly or in small groups, with seven main
groupings: singer (usually a lone male); adult female with a male escort; competitive or surface-active group (multiple
males following a single female and competing with one another for the position of female’s primary escort); lone
(non-singing) whales travelling to join others; groups of juveniles; mother–calf pair (sometimes accompanied by the
previous year’s juvenile); and a mother–calf pair plus male escort. Five main groupings on summer feeding grounds:
single individual or pair of either sex; mother–calf pair; small, usually temporary feeding association; larger (and, for
most individuals, ephemeral) feeding association of up to 24 whales, working together to corral and capture prey by
bubble-net feeding; and (much rarer) super-groups of up to 200 whales. Migrates typically in small, fluid groups with
individuals joining and leaving on a regular basis.
ADULT SOUTHERN HEMISPHERE

A few anomalous all-

white (possibly albino White on underside often
or leucistic) individuals more extensive than most
observed northern hemisphere
animals (can extend up
sides and even onto back)
Typically more
white on underside
of tail (cf. northern
hemisphere)

ADULT FEMALE NORTHERN HEMISPHERE

Grapefruit-sized hemispherical
lobe underneath tailstock behind
genital slit (absent in male)

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 80°
main breeding areas

70°
main feeding areas
60°

50° main links between

40° feeding and breeding
30° areas (not necessarily
20° migration routes)
10°
0°

Humpback whale North Atlantic

30° main breeding areas

20°
10°
main feeding areas
0°
10°
20° feeds and breeds in
30°
?? same area
40°

50° main links between

60°
feeding and breeding
areas (not necessarily
70°
migration routes)
80°

Area VI Area I Area II Area III Area IV Area V

Humpback whale southern hemisphere (and Arabian Sea) showing IWC Management Areas I–VI

75°
main breeding areas

main feeding areas

main links between

60° feeding and breeding
areas (not necessarily
migration routes)

45° occasional links

between feeding
and breeding areas
30°

15°

0°

15°

Humpback whale North Pacific

10°
key aggregation
areas
20° known to be
present
may be present
30°

40°
TO ANTARCTIC
AREA IV
TO ANTARCTIC
AREA V

Humpback whale Australia

FLUKES

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 FLUKE COMPARISONS Underside of flukes varies from virtually all
black to virtually all white with countless
combinations of black and white in between

General proportion Black-and-

of black and Coloration of underside of flukes ranked
from one (nearly all white) to five (nearly white patterns
white varies individually
geographically all black) – more than 80 per cent of
humpbacks in Australia are in category one distinctive
(cf. less than 10 per cent in North Pacific)

FLIPPER COMPARISONS

North Atlantic, North Pacific

type one (one in three
individuals) and Antarctic
Peninsula – white underside,
mostly white upperside (with
varying amounts of black)
North Pacific type two – Western Australia – white
white underside, mostly underside, mostly black
black upperside (two in upperside (with varying
three individuals) amounts of white)

TUBERCLES
Each tubercle is a hair follicle,
with a single coarse sensory
Barnacles tend to hair about 1–3cm long
attach to tubercles growing out of the centre

A tubercle is about
the size of golf ball

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 SPERM WHALE
Physeter macrocephalus Linnaeus, 1758

The largest odontocete, or toothed whale, the iconic sperm whale – well known from Moby Dick – is designed for life
in the ocean depths. This animal of extremes shows the greatest sexual size difference among cetaceans; moreover, its
brain is the world’s largest, and it dives deeper, and for longer, than almost any other whale.
IUCN status Vulnerable (2008). Mediterranean sub-population Endangered (2006).
Population c. 360,000. Pre-whaling population c. 1.1 million. 1.03 million killed by whalers. Trend unknown.
Classification Odontoceti, family Physeteridae.
Taxonomy No recognised forms or subspecies; the scientific name was highly controversial in the past (Physeter
macrocephalus vs P. catodon), but macrocephalus is now used almost universally.
Other names Cachalot.
DISTRIBUTION One of the most widely distributed marine mammals (after killer whales). Found in the deeper parts of
all oceans, from the tropics to the edge of the polar pack ice. Occurs in many deep, semi-enclosed seas, including the
Mediterranean Sea, Gulf of Mexico, Caribbean Sea, Sea of Japan and Gulf of California; mostly absent from enclosed
seas and semi-enclosed seas with shallow entrances (e.g. the Black Sea, Red Sea and Persian Gulf). Generally more
abundant where there is high productivity, usually from upwelling (in areas dubbed the ‘grounds’ by Yankee whalers).
Prefers bathymetric and oceanographic features that concentrate prey, including frontal boundaries, eddies, submarine
canyons and steep continental shelf edges. Daily movements depend on prey abundance: 10–20km when plentiful to
90–100km in poor feeding conditions.
For most of their adult lives, male and female sperm whales are widely separated. Females and young males tend to
remain in the tropics, sub-tropics and warm temperate waters year-round, usually between 40°S and 50°N (sometimes
higher in the North Pacific), corresponding roughly to sea surface temperatures above 15°C. They are usually in waters
deeper than c. 1,000m (less around oceanic islands and in Mexico’s Gulf of California). After leaving natal groups,
young males gradually move to higher latitudes: the larger and older the male, the higher the average latitude.
Large adult males usually frequent waters deeper than 300m, with sea surface temperatures down to c. 0°C;
they will approach close to shore where there is sufficiently deep water (such as in submarine canyons). They spend
much time in more productive latitudes above 40°, often near the edge of the polar pack ice. However, they do return
sporadically to warm-water breeding grounds to mate (on an unknown schedule). Over time, they range widely – often
moving across entire ocean basins and sometimes between them.
BEHAVIOUR The two main behavioural states comprise foraging (c. 75 per cent of their time) and resting/socialising.
When foraging, they make repeated deep dives. Groups of family units spread out over 1+km, diving for extended
periods between breathing intervals at the surface. Adult males typically forage alone. While resting or socialising,
often in the afternoon, females and young gather at or near the surface, close together, when they may lie still and
quiet (sometimes for hours), or they may be active, vocalising, rolling, touching one another, breaching and lobtailing.
Large males also lie quietly at the surface, or socialise if with females.

DIVE Sequence At surface, several individuals may cluster, facing

same direction; before sounding dive, lifts head higher out of water for
penultimate breath; body straightens out and back bends in a ‘stretch’;
may drop below surface temporarily; reappears and accelerates forward;
takes final breath; back arches high out of water showing rounded hump
and ‘knuckles’; throws flukes high into air (though usually not if disturbed).
• Depth Renowned deep diver; adult female typically to 200–1,200m; adult
males often less than 400m; deepest recorded 2,035m but possibly 3+km.
• Duration Typically 30–50 minutes; maximum 138 minutes (the second-
longest mammalian dive, after Cuvier’s beaked whale, Ziphius cavirostris).
BLOW Bushy or ‘puffy’ blow directed forward and to left. • Blow projects
up to 6m (height highly variable).

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 ADULT MALE
Primarily dark grey
Single blowhole (sometimes dark bluish-grey, Dorsal fin two-thirds of the
at front of head Disproportionately large, may appear dark brownish- way along back (slightly
and to left squarish head (25–36 per grey in bright sunlight) further back than in female)
cent of total length)
Dorsal fin a low,
Surface of head
thick and usually
smooth (no c. 30 per cent of young males
rounded hump
wrinkles) Margin between head have rough white or yellowish
and trunk may appear as calluses on dorsal fin (usually
Some mature distinct crease (making absent in adult male)
males have spermaceti organ appear
white patches swollen, especially in
around head larger males) Series of large bumps
(often called knuckles
or crenulations) along
dorsal ridge (behind
dorsal fin)

Upper jaw May have prominent

tends to 2–10 short, post-anal keel
overhang tip deep throat
of lower jaw grooves (less Large wrinkles
Upper ‘lips’, obvious in older or ‘corrugations’
lower jaw cover most of Flukes all dark
animals)
Narrow, underslung and interior body behind eyes on both sides
lower jaw (barely of mouth
visible from side) often white or May have white
creamy-white blotches on
underside (variable
Short, wide, spatulate extent and pattern) Skin on flippers
flippers held close to body and flukes smooth
White scratches and scars common
(possibly as protection (no wrinkles)
(especially on head) of large adults
from predators)
(made by other male sperm whales
and cephalopod prey)

80°

70°

60°

50° AT A GLANCE Deep, ice-free oceanic waters

worldwide • Large to extra-large size •
40°

30°

20°
10°
Primarily dark grey • Huge squarish head •
0°
10°
Thick, low, rounded dorsal fin • ‘Knuckles’
20°

30°
from dorsal fin to flukes • Wrinkly, prune-like
40°

50°
skin • Bushy blow directed forwards and to
60° left • Often motionless (or swims leisurely)
70° at surface • Flukes usually raised on diving
80°

females and visiting males adult males secondary range

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 Breaching is most common when groups come together or split, and between periods of foraging, and tends to
occur in bouts (with individuals breaching repeatedly). Females breach more than lone adult males. Frequently spyhops
– particularly females when there are mature males around during the breeding season, or in response to killer whale
vocalisations. A behaviour apparently unique to sperm whales is ‘driftdiving’. They hang passively and upright in the
water, with their heads up or down, just below the surface; they are probably sleeping (in one study for up to 31.5
minutes). Generally seems oblivious to boats, but will dive prematurely if a boat approaches too rapidly or too closely.
Juveniles are often curious and may come close to investigate.
FOOD AND FEEDING Mainly deepwater squid (25+ species, including giant squid and jumbo squid); 60+ species
of medium- to large-sized deep-sea fish; male typically takes larger individuals than females; occasionally takes
octopuses, crustaceans, jellyfish etc. Feeds mainly in water column but some evidence of also feeding along seabed;
details of how it catches prey unknown (probably draws it into mouth by suction; unlikely to use powerful clicks for
acoustic stunning); males take fish from some longline fisheries.
TEETH Upper jaw 0; lower jaw 36–52.
GROUP SIZE AND STRUCTURE Six main types of social group: family unit or nursery group (about 10 females and their
young); temporary cohesive group, consisting of 2+ family units sharing the same dialect (and therefore belonging to
the same vocal clan); clan, containing many family units (hundreds or thousands of females and their calves); bachelor
school (loose aggregations of young males of similar size and age, 4–21 years old); lone adult male (usually from late
20s onwards); and associations of adult males (long-term preferred companionships).
ADULT FEMALE
Less pronounced Dorsal fin slightly
head (25–31 per cent c. 75 per cent of adult
further forward
of total length) females have rough white
or yellowish calluses on
dorsal fin (secondary sexual
characteristic)

Less scarring
on head May have nicks and notches
along trailing edge
ADULT MALE

Deep,
V-shaped
median
notch

Single
S-shaped Broad, triangular flukes
Blowhole at front of (largest in proportion to
blowhole head and offset to left body size of any whale)
Relatively straight
to slightly convex
male
SIZE trailing edge
L: ♂ 14–16.5m, ♀ 10.5–12m;
WT: ♂ 35–55t, ♀ 10–20t;
MAX: ♂ 19.2m, 70t, ♀ 12.5m, 24t
female Calf – L: 3.5–4.5m; WT: c. 1t
Extreme sexual dimorphism (adult male
typically three times heavier and c. 40 per
cent longer than adult female).

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 ADULT MALE

Upper jaw has small

vestigial teeth (rarely
break through gums)

Lower jaw can be opened Lower jaw much

nearly perpendicular to body narrower than upper

Large, conical teeth on lower

jaw (fit into sockets in upper CALF
jaw when mouth closed)
Same dark grey
as adult
FLUKES

Mouthline Flukes very

relatively short long

Skin wrinkled all over at birth Little or no

(skin of head and flippers scarring
becomes smooth with age)

THE SPERMACETI ORGAN

The spermaceti organ complex – which Blowhole
Skull Spermaceti Museau
dominates the sperm whale’s head – is organ de singe
the world’s most powerful natural sonar Right nasal (monkey
Deep, Frontal air sac passage
V-shaped or echolocation system. It consists of a muzzle)
median complex array of soft structures, cradled
notch above the lower jaw and in front of the Distal
skull, in the whale’s highly modified and air
asymmetric head. Known by whalers as sac
the ‘case’, the spermaceti organ itself
is up to c. 5m long, and enclosed by a
tough muscular sheath. It consists of a
white spongy tissue soaked in a liquid
wax called spermaceti oil. The organ is
used mainly for forming, focusing and Acoustic
broadcasting extremely powerful, highly wax lenses
directional clicks (recent evidence Junk in junk
suggests that it is not used to debilitate Left nasal Lower jaw
prey with sound, as was previously Position passage
Long, flat rostrum (designed to
thought). of eye support spermaceti complex)

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 PYGMY SPERM WHALE
Kogia breviceps (Blainville, 1838)

Pygmy and dwarf sperm whales are typically seen floating on the surface, with just the top of the head and back (as far
as the dorsal fin) exposed. They are very difficult to spot in anything but calm conditions, and sightings of both species
tend to be brief.
IUCN status Least Concern (2019).
Population Unknown, although frequent strandings in some areas implies that it is more common than the lack of
sightings suggests. Trend unknown.
Classification Odontoceti, family Kogiidae.
Taxonomy No recognised forms or subspecies.
Other names Lesser cachalot, short-headed cachalot, lesser sperm whale, short-headed sperm whale.
DISTRIBUTION Tropical to warm temperate waters worldwide. Generally inhabits waters along the outer continental
shelf and beyond, particularly over and near the continental slope. In the North Atlantic, it is closely associated with
the Gulf Stream. Prefers more temperate seas and relatively deeper, more pelagic waters than the dwarf sperm whale.
There is no evidence of long-distance migrations. Most information comes from strandings.
BEHAVIOUR Difficult to spot, except in extremely calm seas. Surfacing patterns are hard to predict, and it tends to
be shy, undemonstrative and tricky to approach closely. Aerial behaviour is extremely rare, though it does sometimes
breach. Between dives, it tends to raft motionless at the surface – from a distance resembling a piece of driftwood –
with the top of the head, back and dorsal fin exposed, and the tail hanging down underwater.
FOOD AND FEEDING Mostly deepwater squid; will take some fish and shrimps; more diverse diet and averages larger
prey than dwarf sperm whale. Feeding mostly on or near seabed; anatomy suggests powerful suction feeding.
TEETH Upper jaw 0; lower jaw 20–32.
GROUP SIZE AND STRUCTURE Usually solitary, but up to six of varying age and sex composition. Strandings usually
involve single animals (maximum known was three: one male and two females).
Flippers can be pressed ADULT
Blunt, rounded tight against body
snout
Broad flukes

Slight
median
notch

More body
Blowhole lies well
scarring
back and slightly Blunter,
to left of centre OLDER ADULT
squarer (not to scale)
head

More scarring
around mouth

DIVE Sequence Inconspicuous blow; floats motionless in same position on surface (with front of melon to dorsal
fin visible); usually sinks vertically out of sight but (particularly if startled) may roll forward with little arching of
back; does not show flukes. • Depth Unknown, but believed to forage at greater depths than dwarf sperm whale.
• Duration 12–15 minutes (based on limited evidence); maximum recorded 18 minutes.

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 Blunt, squarish head
(proportionately larger,
less square and with
longer snout than on
dwarf sperm whale)
Head becomes
blunter and
squarer
with age
Dark bluish-grey Tip of dorsal fin rounded
ADULT
to brownish-black and usually below
upperside highest point (variable)
Robust body Small, falcate dorsal fin well
(not unlike small behind midpoint of back
sperm whale) (slightly further back than on
dwarf sperm whale)
Dorsal fin less than
5 per cent of total
body length

Lighter cream or
ivory underside
(sometimes with
May have
No throat Small, broad pinkish tinge)
circular scars
around mouth grooves flippers set far
(caused by forward near head Scarring on both sexes
squid bites) (attributed to fighting during
Narrow May be slightly Crescent-shaped, light- mating season or shark attacks)
underslung darker coloration coloured marks usually
lower jaw around eye present on sides of head
(dubbed ‘false gills’)

Tiny, underslung
lower jaw with long,
sharp teeth (fit into
sockets in upper jaw)

CALF

SIZE
L: ♂ 2.7–3.5m, ♀ 2.7–3.5m;
WT: 315–450kg; MAX: 3.8m, 515kg
Calf – L: c. 1–1.2m; WT: c. 50–55kg

80°

70°

60°

50°
AT A GLANCE Deep tropical to warm
40° temperate waters worldwide • Small size
• Generally appears dark grey at sea • Blunt,
30°

20°

squarish head • Small, falcate dorsal fin

10°
0°
10°
20° slightly further back than on dwarf sperm
30°

40°
whale • Floats motionless on surface
50°
between dives • Back has distinctive bulge
when logging
60°

70°

80°

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 DWARF SPERM WHALE
Kogia sima (Owen, 1866)

With their long, sharp teeth, underslung lower jaws and gill-like markings on either side of the head, dwarf and pygmy
sperm whales are often mistaken for sharks when they strand. Their appearance may be a form of mimicry, to help
avoid predation.
IUCN status Least Concern (2020).
Population Unknown, although frequent strandings in some areas implies that it is more common than the lack of
sightings suggests. Trend unknown.
Classification Odontoceti, family Kogiidae.
Taxonomy No recognised forms or subspecies; however, recent genetic studies suggest there may be two distinct
species, one in the Atlantic, the other in the Indo-Pacific.
Other names Owen’s pygmy whale, snub-nosed cachalot.
DISTRIBUTION Tropical to warm temperate waters worldwide. Generally inhabits deep waters over or near the edge of
the continental shelf. In the North Atlantic, it is closely associated with the Gulf Stream. Prefers more tropical seas and
relatively shallower, less pelagic waters than the pygmy sperm whale (sometimes frequenting more coastal areas) and
probably does not range as far into high latitudes. Most information comes from strandings.
BEHAVIOUR Difficult to spot, except in extremely calm seas. Surfacing patterns are hard to predict. Rarely allows close
approach. Aerial behaviour is extremely rare, though it does sometimes breach. Between dives, it tends to raft (or log)
motionless at the surface – where, from a distance, it looks like a piece of driftwood – with the top of the head, back
and dorsal fin exposed, and the tail hanging down underwater.
FOOD AND FEEDING Mostly mid- and deepwater squid; will take some fish and shrimps; less diverse diet and averages
smaller prey than pygmy sperm whale. Feeding mostly on or near seabed; anatomy suggests powerful suction feeding.
TEETH Upper jaw 0–6; lower jaw 14–26.
GROUP SIZE AND STRUCTURE Usually solitary, but up to 12 (maximum 16 recorded) of varying age and sex
composition. Group size varies according to location and season: e.g. in the Bahamas 1–8 in summer, 1–12 in winter;
average in Hawaii is 2.7. Individuals in a group appear to be loosely associated – when one appears, there are often
others several hundred metres away. Strandings usually involve single animals (maximum known was four immatures:
one male and three females).
Broad flukes
ADULT

Slight
median
notch

Blowhole lies
well back and Flippers can be SIZE
slightly to left L: ♂ 2.1–2.4m, ♀ 2.1–2.4m;
pressed tight
of centre WT: 135–270kg; MAX: 2.7m, 303kg
against body
Calf – L: c. 0.9–1.1m; WT: 40–50kg

DIVE Sequence Inconspicuous blow; floats motionless in same position on surface (with front of melon to dorsal fin
visible); usually sinks vertically out of sight but (particularly if startled) may roll forward with little arching of back;
does not show flukes. • Depth Unknown, but believed to forage at shallower depths than pygmy sperm whale (600–
1,200m suggested). • Duration 7–15 minutes (possibly up to 30 minutes); brief 1–3 minutes at surface in between.

96 SPERM WHALES

001 WDP FG.indd 96 29/11/2021 12:15

 Blunt, squarish head Dark bluish-grey ADULT
Height, position
(proportionately smaller to brownish-black
and shape of dorsal
and squarer than on upperside (colour
fin highly variable Relatively large,
pygmy sperm whale) quite variable) Robust body
(in extreme cases falcate dorsal fin
May be (not unlike small overlaps with pygmy just in front of
Head becomes slightly darker sperm whale) sperm whale) midpoint of back
blunter and coloration
squarer around eye Dorsal fin more than 5 per
with age cent of total body length

Tip of dorsal fin

pointed and usually at
highest point (variable)

Narrow, Small, broad

underslung flippers set far Lighter cream or
lower jaw with forward near head ivory underside
long, sharp (sometimes with May have scarring
Crescent-shaped, light-coloured pinkish tinge) from cookiecutter
teeth (fit into marks usually present on sides
sockets in shark bites (quickly
of head (dubbed ‘false gills’) re-pigment to
upper jaw) Scarring on both sexes
(attributed to fighting background colour)
May be circular Two or more short,
longitudinal throat during mating season
scars around mouth or shark attacks)
(caused by squid grooves (similar to those
bites) on beaked whales)

COMPARISON OF SILHOUETTES
• Distinctive rounded bulge on back (between blowhole and
dorsal fin) visible when logging.
• Distinct ‘neck’ behind relatively large head.
• Smaller, rounded dorsal fin, further back.
Pygmy sperm whale • Sometimes dorsal fin invisible until animal rolls out of sight.

• Flatter profile in water (usually without a prominent bulge),

reminiscent of an upside-down surfboard.
• Tends to float lower in water.
• Larger, pointed dorsal fin, further forward.
Dwarf sperm whale • Dorsal fin not unlike that of bottlenose dolphin.

80°

70°

60°

50°
AT A GLANCE Deep tropical to warm
40° temperate waters worldwide • Small size
• Generally appears dark grey at sea • Blunt,
30°

20°

squarish head • Tall, pointed, falcate dorsal

10°
0°

fin slightly further forward than on pygmy

10°
20°

30°

40°
sperm whale • Floats motionless on surface
50°
between dives • Back appears flat when
logging
60°

70°

80°

DWARF SPERM WHALE 97

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 NARWHAL
Monodon monoceros Linnaeus, 1758

The narwhal can be difficult to see: living in remote regions of the High Arctic, it spends half the year in dense pack ice
under continuous darkness. However, it does have predictable migratory patterns, and the male is unmistakable, with
its extraordinary long, spiralling tusk.
IUCN status Least Concern (2017).
Population c. 170,000, excluding north-east Greenland and the Russian Arctic (for which no estimates are available).
Trend unknown.
Classification Odontoceti, family Monodontidae.
Taxonomy No recognised forms or subspecies.
Other names Narwhale, unicorn whale, sea unicorn.
DISTRIBUTION Mainly in the Atlantic sector of the Arctic (60–85°N, most commonly 70–80°N). Occasional stragglers
in the Pacific sector. Relatively rare in Svalbard and south of the Arctic Circle. Discontinuous range, separated by
Greenland. Winter and summer ranges up to 2,000km apart. Vagrants have been recorded in Newfoundland, UK,
Germany, Belgium and the Netherlands (in the North Atlantic); and as far south as the Alaska Peninsula and the
Commander Islands (in the North Pacific).
Winter distribution Wintering areas tend to be deep, offshore, ice-covered habitats along the continental slope. Two-
thirds of all narwhals winter in deep water offshore under dense pack ice in Baffin Bay and Davis Strait (between Baffin
Island and Greenland), in two distinct ‘grounds’ (northern and southern); the east Greenland population winters offshore
in deep waters of the Greenland Sea. On the wintering grounds, they spend six months in continuous darkness, in air
temperatures as low as -40°C, where there is often less than 3 per cent open water.
Summer distribution Spends about two months each summer in ice-free bays, fjords and island passages in the
Canadian Arctic, western and eastern Greenland, Svalbard, and the northwestern Russian High Arctic. In some parts
of the range, glacial fronts are an important summer habitat. It prefers deep water but readily enters shallow water to
hide from hunting killer whales.
Migrations There are predictable migrations between the summer and winter grounds, as the ice retreats in spring and
refreezes in autumn, and in many areas narwhals pass certain promontories, bays and fjords at precisely the same time
each year. Migrations last about two months each way.
ADULT MALE
Dark brown dorsal surface
Right tusk (measuring (strongest on top and front of
c. 30cm) usually remains head, along dorsal ridge and along
embedded in skull borders of flippers and flukes)

Erupted tusk appears on

left side of upper jaw and
angles slightly to left

DIVE Sequence Visible but inconspicuous blow (clearly audible on calm days and at close range); male’s tusk
sometimes (but not always) appears above surface (usually briefly); alternatively, may see impression of tusk just
below surface; may fluke before deep dive (rarely before shallow dive). • Depth Summer 13–850m, though usually
less than 50m; winter much deeper (typically spending over three hours per day at depths of at least 800m, during
18–25 dives); over half winter dives reach 1,500m; maximum recorded 1,800m. • Duration Mostly 7–20 minutes,
maximum documented 25 minutes.

98 NARWHAL AND BELUGA

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 Variable light and dark mottling ADULT MALE
Relatively small (dark mottling especially on top
head with and front of head, dorsal ridge Distinctive serrated,
pronounced melon and appendages) fleshy dorsal ridge (up
to 5cm high) on rear
Typically darker on top Robust body half of back
of head and cape
No dorsal fin
Scarring on head
fairly common

Majority of adult males have Flippers short,

long, straight tusk like gnarled square and Lighter underside
and twisted walking stick close to head

Female typically ADULT FEMALE

smaller than male

Tusks (measuring up to
c. 25cm) normally remain in
tooth sockets within upper jaw

SIZE
L: ♂ 4.3–4.8m (excluding tusk of up to
3m), ♀ 3.7–4.2m;
WT: 700–1,650kg; MAX: 5m, 1,800kg
Calf – L: 1.5–1.7m; WT: c. 80kg

80°

70°

60°

50°

40°
AT A GLANCE High Arctic • Small to medium
30°

20°
size • Long tusk of male • Relatively small
10°
0°
bulbous head • Little or no beak • No dorsal
10°
20°
fin (but slight dorsal ridge) • Variable light and
30° dark mottling
40°

50°

60°

70°

80°

NARWHAL 99

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 BEHAVIOUR Not given to spontaneous exuberance such as breaching or speed-swimming, but occasionally spyhops
and lunges. Frequently logs (or rafts), with the top of its head and back visible, and may roll around at the surface while
socialising. Spends less time at the surface in choppy or rough seas. Male may wave its tusk in the air or rest it on the
back of another individual. Before a deep dive, a narwhal often swims directly towards the ice edge, then flukes about
5–30m away. Navigates easily under ice and can travel several kilometres between breathing opportunities, or uses
head or back to break through ice several centimetres thick; the only barrier to its movements is fast ice without cracks.
Often associates with bowhead whales; rarely forms mixed herds with belugas. Often shy and wary of boats (especially
where hunted), but in Canada, at least, can be less nervous of people standing on the floe edge or shore.
FOOD AND FEEDING Fish (especially Greenland halibut, Arctic cod; also Arctic eelpout, polar cod, roughhead grenadier,
capelin), squid (especially small deepwater Gonatus spp.), shrimp (especially deep-sea prawn); precise diet varies with
region and season. Most feeding in winter (November–March), very little or no feeding in summer (July–September),
will feed under sea ice in spring and autumn; probably sucks prey into mouth and swallows it whole; no evidence of
cooperative hunting.
TEETH Upper jaw 2 (usually only 1 erupts, typically only in the male); lower jaw 0.
GROUP SIZE AND STRUCTURE Most narwhal pods or ‘clusters’ typically contain 2–10 individuals (range 1–50). Mostly
these comprise a single sex: all males, or all females with young. Groups often combine to form large dispersed herds,
each with up to 600 clusters, containing hundreds or even thousands of individuals and a mix of sexes and ages.

OLDER ADULT MALE

Typically remains dark
on top and front of
Tusk typically to c. 2m head, along dorsal ridge
(maximum 3m in old males) and along borders of
flippers and flukes

Flippers often curl

up at the tips (more
exaggerated in Mottling becomes lighter with
older males) age (older males, especially, can
Tips often curl up
appear almost white at a distance)
(more exaggerated
in older males)
Pinkish at birth, turning
uniform mid-grey, blue-grey
CALF
or brownish-grey (develops
mottled pattern after 2 years)

No tusk

JUVENILE MALE As it matures, pale spotting begins

to appear on belly and sides

Small tusk may be

present in male (begins
to erupt at 2–3 years old)

100 NARWHAL AND BELUGA

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 ADULT FEMALE WITH TUSK 1 in 30 females has 1 tusk (though
estimates vary since tusked females are
easy to confuse with juvenile males)

Female tusks shorter (maximum

1.2m), less robust and whiter
(do not collect as much algae
on surface as male tusks)

Tusk usually ends in ADULT MALE

shiny white point Tusk typically Little or no beak
1.8–2.7m long

Tusk length, girth,

morphology, wear and
colour highly variable Short, upturned
mouthline
Right tusk usually
shorter and less robust
1 in 500
males has
two tusks

One record of female

with two tusks
ADULT MALE WITH TWO TUSKS (collected in 1684)

The tusk always spirals anticlockwise

from the whale’s-eye view

ADULT MALE FLUKES ADULT FEMALE FLUKES

UPPERSIDE UPPERSIDE
Strongly convex trailing
edges (giving ‘back-to-front’
appearance), especially in Deep median Trailing edges of flukes
older males notch less convex, even straight
Tips may
(more dolphin-like)
curl upwards
(especially in
older males)

Upperside of flukes
Concave generally darker
leading edges

NARWHAL 101

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 BELUGA
Delphinapterus leucas (Pallas, 1776)

Ancient mariners used to call the pale white beluga the ‘sea canary’ because of its great repertoire of groans, roars,
whistles, squawks, moos, buzzes and trills. When seen from above the surface, its ghost-like glow is hard to mistake
for any other species.
IUCN status Least Concern (2017). Cook Inlet sub-population Critically Endangered (2018).
Population c. 200,000 (though many parts of the range remain unsurveyed). Trend unknown.
Classification Odontoceti, family Monodontidae.
Taxonomy No recognised forms or subspecies.
Other names Beluga whale, white whale; historically – sea canary.
DISTRIBUTION Cold waters of the Arctic and sub-Arctic, ranging from 47–82°N. Wide choice of habitat, including
estuaries, coastal waters (as shallow as 1–3m), continental shelves and deep ocean basins, in open water and loose
ice (it generally avoids dense pack ice, though commonly overwinters in polynyas). Will swim up rivers. There is an
isolated population in the St Lawrence River estuary, Canada, which is resident year-round. Dense concentrations are
common in shallow coastal waters in summer.
BEHAVIOUR Rarely given to aerial displays, though it can be more demonstrative (spyhopping, tail-waving and lobtailing)
in nearshore concentrations. Able to turn its head sideways (the cervical vertebrae are unfused, making the neck more
flexible), which is unusual in cetaceans. Little fear of shallow water (if stranded, it is often able to wait and refloat on
the next tide – assuming it is not found by a polar bear first). Sometimes shows curiosity towards boats and frequently
towards snorkellers and divers. Often associates with bowhead whales, though rarely forms mixed herds with narwhals.
The annual moult, which involves rubbing along the seafloor to remove sloughed skin, is rare in cetaceans.
FOOD AND FEEDING Mainly fish such as salmon, herring, Greenland halibut, smelt, Arctic and polar cod and capelin,
but also squid, octopuses, shrimps, crabs, clams, mussels and even marine worms and large zooplankton. Sucks prey
into mouth with flexible ‘lips’; some evidence of cooperative hunting (e.g. in groups of 3–5, hunting smelt in Russia’s
Sea of Okhotsk), but usually hunts alone (even within a group).
TEETH Upper jaw 16–20; lower jaw 16–20. Teeth are often heavily worn – even down to the gums in older animals.
GROUP SIZE AND STRUCTURE Usually in groups of 5–20 (although large adults are occasionally seen alone). Many
groups together may form herds of hundreds, or more than 1,000, which can be mixed or segregated by age and sex.
Group structure tends to be fluid.
Female up to 25 per cent
smaller than male ADULT FEMALE
Less robust build
than male

Flipper tips rarely curled

DIVE Sequence Swims in slow rolling motion (look for white arc that appears, grows, shrinks, then disappears);
flukes occasionally appear at low angle above water; indistinct low, steamy blow (often not visible). • Depth
Regularly 300–600m, sometimes beyond 800m (maximum recorded 956m). • Duration 9–18 minutes (feeding
dives typically 18–20 minutes); maximum documented 25 minutes.

102 NARWHAL AND BELUGA

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 Very pale to pure Tough, serrated ADULT MALE
Relatively small, white (during moult, May have Robust dorsal ridge
bulbous head especially in early yellowish tinge (sometimes visible along May be some darker
summer, some adults caused by layer rotund) body midline of back pigmentation on dorsal
Very short, can be light grey) of diatoms (used to break ridge and borders of
wide beak (especially ice) appendages Distinct
in spring) Transverse Trailing edges notch in
nicks sometimes middle
No dorsal darker
fin

Cleft Small flukes

upper
‘lip’
May have rake marks
Visible neck Body often wrinkled and and severe scarring
region (may give Small rounded flippers ‘blubbery’, with folds of caused by killer
appearance of Unusually flexible (tips curled upwards fat along belly and sides whales or polar bears
‘shoulders’) neck (due to unfused in adult male – more (insulating blubber up to
neck vertebrae) pronounced with age) 15cm thick)

CALF

SIZE
L: ♂ 3.7–4.8m, ♀ 3.0–3.9m;
WT: 500–1,300kg; MAX: 5.5m, 1.9t
Lightens with age – lighter grey in first year, Calf – L: 1.5–1.6m; WT: 80–100kg
May have
remains grey 5–10 years, then gradually changes Body size varies considerably between
slight beak sub-populations.
to white – usually pure white by 5–12 years

FLUKES IMMATURE
ADULT FACIAL EXPRESSIONS FLUKES ADULT Trailing edge more
convex in older animals Trailing edge
usually straight

Can alter shape of ‘lips’ and melon Tail changes

(impressive array of facial expressions) shape as it grows

80°

70°

60°

AT A GLANCE Arctic and sub-Arctic • Small

50°

40°

to medium size • Very pale to pure white, pale

30°

20°
10°
0° grey or yellowish with no mottling • Robust
10°
20°
body • Small, bulbous head • No dorsal fin
30°

40°
• Surfaces often with distinctive slow rolling
50°
motion
60°

70°

80°

BELUGA 103

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 BAIRD’S BEAKED WHALE
Berardius bairdii Stejneger, 1883

The largest of all the beaked whales, Baird’s beaked whale is one of the easiest members of the family to identify in the
North Pacific. It is strikingly similar to Arnoux’s beaked whale (though the two species are widely separated geographically).
Baird’s is one of few beaked whale species to be commercially hunted; Japanese hunting began in the early 1600s, reached
a peak after the Second World War (322 were killed in 1952) and continues to this day (with an annual quota of c. 66).
IUCN status Least Concern (2020).
Population Unknown. Regional estimates: c. 7,100 in Japanese waters, c. 7,960 in the California Current. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Giant bottlenose whale, North Pacific bottlenose whale, four-toothed whale, northern fourtooth whale.
DISTRIBUTION In summer, deep, offshore, cool temperate to sub-polar waters in the northern North Pacific and the
adjacent Sea of Japan, Sea of Okhotsk and Bering Sea, at least from c. 30–62°N. Appears to be migratory in most
areas, with seasonal peaks in abundance. In winter, it is believed to move into deeper waters away from the continental
slope, with at least some time spent in the sub-tropics and possibly tropics. Prefers continental slope waters, 1,000–
3,000m deep, and areas with complex topography such as submarine canyons, seamounts and ridges.
BEHAVIOUR Moderately aerially active. Will sometimes breach repeatedly, leap in a low arc or, especially in mixed
groups, jump on top of one another. Will also spyhop, flipper-slap and lobtail. May swim belly up or sideways or roll at
the surface. Reaction to vessels varies.
FOOD AND FEEDING Pelagic and benthic fish, squid, octopuses, some crustaceans. Often (but not exclusively) down to
seafloor; pebbles in stomachs may reflect bottom-feeding; suction feeder.
TEETH Upper jaw 0; lower jaw 4.
GROUP SIZE AND STRUCTURE Typically 3–20, with some regional variation; occasionally up to 50. There may be
some sexual segregation. Lone individuals are rare. Groups can be so tightly packed that individuals may be in physical
contact with each other.
May be flattened area Tailstock ADULT MALE
Crescent-shaped blowhole Flippers fit into or depression along compressed
(ends point backward – cf. all ‘flipper pockets’ centre of back between Flukes wide
laterally
other beaked whale genera) blowhole and dorsal fin relative to
body length

Front of melon may

appear largely white Linear scars begin to appear
Most individuals have no median
due to scarring around time of sexual maturity
notch or slight depression (may
(accumulate with age)
even be slight prominence)

DIVE Sequence Typically in tight school, all surfacing in unison; body appears very long; blows continuously while
swimming slowly (easily identifiable from distance); shallow roll with little arching of back (more arching before
deep dive); rarely raises flukes before deep dive. • Depth Deep dives routinely to 1,000m or more; maximum
recorded 1,777m (Japan) but probably capable of deeper. • Duration Maximum recorded 67 minutes (Japan),
unsubstantiated reports of up to two hours.
BLOW Strong, low, puffy or rather shapeless blow (up to c. 2m), quite conspicuous in calm weather (but tends to
dissipate quickly). • Blow sometimes angled slightly forward.

104 BEAKED WHALES

001 WDP FG.indd 104 29/11/2021 12:15

 Bulbous melon Long, fairly Extensive single and Upperside of older males ADULT MALE
with moderately slender, slightly closely paired linear may appear whitish (due
steep forehead spindle-shaped scarring (tooth rake to heavy scarring)
(more bulbous body marks made mainly by
with age) conspecifics) especially Small triangular to slightly
Predominantly on upperside falcate dorsal fin (25–32cm high)
slate-grey to two-thirds of the way along back
Long, well- Relatively black
defined tube- small head Tip can be very
like beak rounded (especially
(50–60cm) on older animals)

Deep
Long, sinuous May be extensive
Small flippers tailstock
mouthline light oval scarring
set far forward More extensive
scarring in Some animals from cookiecutter
Two pairs of teeth near tip of Lighter underside older animals have brownish or shark bites (possibly
lower jaw (only larger pair at front (may be irregular greenish-brown also from Pacific
visible outside closed mouth) white area along tinge (due to lampreys)
ventral midline) diatoms)
One of the least sexually
dimorphic beaked
whales (both sexes have Far less linear ADULT FEMALE
Melon slightly scarring
erupted teeth and look May be more
less bulbous
remarkably similar) extensive scarring
from cookiecutter
shark bites

Pair of V-shaped throat grooves

(additional smaller pairs may be present)
SIZE
L: ♂ 9.1–10.7m, ♀ 9.8–11.1m;
WT: 8–12t; MAX: 13m, 12.8t
Calf – L: 4.5–4.9m; WT: unknown
OLD ADULT MALE
Lower jaw extends c. 10cm further forward
than upper jaw (exposing front teeth)
Teeth may appear white and
‘flash’ brightly in sunlight
Larger pair of teeth may be worn
down (sometimes invisible) and
heavily infested with stalked (goose)
barnacles in older individuals

75°

AT A GLANCE Cool offshore waters of

northern North Pacific • Predominantly dark
60°

with heavy scarring • Medium to large size

45°
• Bulbous melon • Long, slender beak •
Two visible teeth at tip of lower jaw • Small,
30°
rounded fin two-thirds of the way along back
15° • Tightly packed groups surface in unison
0°

15°

BAIRD’S BEAKED WHALE 105

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 ARNOUX’S BEAKED WHALE
Berardius arnuxii Duvernoy, 1851

Despite closely resembling the relatively well-studied Baird’s beaked whale, which lives far away in the North Pacific,
Arnoux’s beaked whale is very poorly known. Both species are unusual among beaked whales for several reasons: four
teeth erupt in both males and females, there is little sexual dimorphism and the males seem to live much longer than
the females.
IUCN status Least Concern (2020).
Population Unknown. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Southern beaked whale, four-toothed whale, giant bottlenose whale, New Zealand beaked whale.
DISTRIBUTION Circumpolar in southern hemisphere, favouring deep, offshore, cold temperate to polar waters, most
abundant from 40–77°S. There was a sighting of 12 right in front of Scott Base, Antarctica (77°51’S), in 2021. Often
found close to the ice edge and in areas with extensive ice cover. Normally prefers deeper waters over the continental
slope and beyond, but may be more coastal along the western side of the Antarctic Peninsula, where there are
numerous intrusions of deepwater channels and canyons.
BEHAVIOUR Breaches fairly frequently. Will also spyhop, flipper-slap and lobtail. Reaction to vessels varies.
FOOD AND FEEDING Deepwater squid, possibly some deepwater fish. Probably often (but not exclusively) down to
seafloor; suction feeder.
TEETH Upper jaw 0; lower jaw 4.
GROUP SIZE AND STRUCTURE Typically 6–15; several smaller groups may join together for periods. A group of about
80 was followed by researchers in the Antarctic for several hours; the whales eventually split into smaller groups of
8–15 and dispersed into loose pack ice.

May be flattened area or ADULT

depression along centre of back Flukes wide
between blowhole and dorsal fin relative to
body length

Crescent-shaped
blowhole (ends point
backward – cf. all other
beaked whale genera)
Flippers fit into Tailstock
compressed Most individuals have no median
‘flipper pockets’ notch or slight depression (may
laterally
even be slight prominence)

DIVE Sequence Typically in tight school, all surfacing in unison; body appears very long; shallow roll with arching of
back (more arching before deep dive); when logging or swimming slowly, may lift head and beak out of water and
sink backwards under surface; rarely raises flukes before deep dive. • Depth Probably greater than 500m, possibly to
3,000m. • Duration Typically 15–25 minutes; maximum recorded 70 minutes.
BLOW Strong, low, puffy or rather shapeless blow (up to c. 2m), quite conspicuous in calm weather (but tends to
dissipate quickly). • Blow sometimes angled slightly forward. • Blows continuously while swimming slowly (easily
identifiable from distance).

106 BEAKED WHALES

001 WDP FG.indd 106 29/11/2021 12:15


Bulbous melon with
moderately steep
forehead (not as bulbous
or steep as in southern
bottlenose whale)
Long, well-
defined tube-
like beak
Predominantly dark slate- ADULT
Long, fairly slender, slightly grey to blue-black (some
Head region individuals have brownish
generally spindle-shaped body (more
slender than southern tinge due to diatoms –
paler appear browner in sunlight)
bottlenose whale)
Relatively Small triangular to
small head slightly falcate dorsal fin
(25–30cm high) two-thirds
of the way along back

May be light oval

Two pairs of teeth Small flippers Lighter Both sexes scarring from
near tip of lower far forward underside have single and cookiecutter
jaw of both sexes Lower jaw Pair of V-shaped throat closely paired shark bites
(only larger pair at noticeably grooves (additional smaller linear scarring,
front visible outside longer than pairs may be present) made mainly by
closed mouth) upper jaw conspecifics (less
extensive on female)
Older animals (especially
Melon more males) may be lighter colour
Front of melon OLD ADULT MALE
bulbous with due to so much scarring
may be whiter age
due to scarring Tip can be very
rounded

Larger pair of teeth may be worn down

in older animals (sometimes invisible)
or heavily infested with stalked (goose)
barnacles SIZE
L: ♂ 8–9.3m, ♀ 8–9.3m;
WT: c. 6–7t; MAX: 9.8m, 10t
Calf – L: c. 4–4.6m; WT: unknown

80°

70°

60°
AT A GLANCE Cool offshore waters, mainly
in sub-Antarctic and Antarctic • Slate-grey
50°

40°

to light brown with heavy scarring • Medium

30°

20°
10°
0° size • Bulbous melon • Long, slender beak
10°
20° • Two visible teeth at tip of lower jaw •
30°

40°
Small, rounded fin two-thirds of the way
50°
along back • Tightly packed groups surface
in unison
60°

70°

80°

ARNOUX’S BEAKED WHALE 107

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 SATO’S BEAKED WHALE
Berardius minimus Yamada, Kitamuru and Matsuishi, 2019

Hokkaido whalers in Japan have traditionally recognised two different kinds of Baird’s beaked whale: the relatively
common ‘slate-grey’ form (p. 104) and a rarer, smaller ‘black’ form. The existence of the rarer form as a new species has
been debated for decades. It was officially described as a new species, Sato’s beaked whale, in 2019.
IUCN status Near Threatened (2020).
Population Unknown, but probably rare given the small number of standings. Decreasing.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Whalers from the Japanese island of Hokkaido call it kuro-tsuchi (black Baird’s beaked whale); another
name they use is karasu (crow – after its dark colour), though it is unclear whether this refers to the same species, yet
another new species, or one of the Mesoplodon species found in Hokkaido.
DISTRIBUTION Cool temperate and sub-Arctic waters in the North Pacific, between 40°N and 60°N, and 140°E and
160°W. Confirmed records reveal a disjunct distribution with three main centres: in and around Alaska’s Aleutian
Islands, Hokkaido in Japan, and Russia’s southern Sea of Okhotsk. From April to June, Japanese whalers repeatedly
see groups in the Nemuro Strait, near the northern tip of Hokkaido – the only location known to have regular sightings.
There has also been one possible scientific sighting at sea, in 2009, of a pod of three individuals off Rausu, in Nemuro
Strait. Bite scars from cookiecutter sharks (Isistius brasiliensis) – with a northern limit between 30°N and 43°N –
suggest that at least some individuals migrate south to warm temperate or tropical regions for at least part of the
year (it is possible that the species spends considerable time in warmer, more southerly waters). Little is known about
habitat preferences; it might be expected in waters with depths of more than 500m, with primary habitat on or near the
continental slope and on or near oceanic seamounts.
BEHAVIOUR Very little known. As with other beaked whales, it is likely to be found in deep waters offshore and
probably spends little time at the surface. Japanese whalers in Abashiri, on the Sea of Okhotsk coast of Hokkaido, say
that it is difficult to approach within shooting distance.
FOOD AND FEEDING No information but it is probable that, like other beaked whales, it is a deep diver that feeds
primarily on squid (perhaps taking some smaller fish and shrimp species as well).
TEETH Upper jaw 0; lower jaw 4. Teeth presumed to erupt in both male and female (as in Baird’s beaked whale).
GROUP SIZE AND STRUCTURE No information, though Japanese whalers have described ‘small groups’.

ADULT
Crescent-shaped
blowhole (ends point
backward – cf. all other Flukes wide relative
beaked whale genera) to body length

Tail stock laterally

compressed No median
notch

SIZE
L: ♂ 6.3–6.9m; MAX: 6.9m;
WT: unknown, possibly c. 2–3t;
Calf – L: unknown; WT: unknown

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 Medium- Medium size More spindle-shaped ADULT MALE
length beak body than Baird’s
(proportionately Almost uniformly
shorter than Small dorsal fin
black colouring
Pale white on Baird’s) two-thirds of the
Prominent,
colouring way along back
bulbous
on rostrum forehead Relatively taller
dorsal fin than
Baird’s

White cookiecutter
shark bites fairly
conspicuous
Two pairs of teeth Fewer or less
at tip of lower jaw V-shaped throat Relatively intense tooth rake (Appearance of female
(anterior teeth grooves smaller flippers marks than Baird’s currently unknown)
larger) than Baird’s

A NEW SPECIES
The existence of a new species of beaked whale –
closely related to Baird’s beaked whale – has been
speculated for more than 70 years. First, there
was anecdotal evidence from whalers, operating
from the island of Hokkaido, in northern Japan.
Then, in 2004, a mother and calf of a previously
unfamiliar-looking beaked whale were taken as
bycatch and landed at Shibetsu, Hokkaido. In
2009, a pod of three unidentified beaked whales,
resembling small, black Baird’s beaked whales,
was photographed off Rausu, in Nemuro Strait,
Japan. In the years since, 14 confirmed specimens
have been found: 5 in and around the Aleutian
Islands, Alaska, USA; 6 on Hokkaido, Japan; and
3 from Sakhalin and Kunashir Islands, Russia.
Recent DNA studies on some of these specimens
found a significant genetic difference between
them and Baird’s beaked whale – indeed, a greater
difference than between Baird’s and Arnoux’s
beaked whales.

DORSAL FIN VARIATIONS

75°

60°
AT A GLANCE Cool temperate and sub-Arctic
waters of North Pacific • Medium size (60–70
per cent of length of Baird’s beaked whale)
45°
Nemuro Strait • Predominantly black body colour • Less
scarring than Baird’s beaked whale • Dorsal
fin relatively taller than Baird’s beaked whale
30°

15°
and two-thirds of the way along back
0°

15°

locations where confirmed specimens have been found

SATO’S BEAKED WHALE 109

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 CUVIER’S BEAKED WHALE
Ziphius cavirostris G. Cuvier, 1823

Cuvier’s beaked whale is one of the most frequently seen, easily recognisable and widely distributed species of beaked
whale, though it is still poorly understood. It holds the records for the deepest and longest dives of any mammal.
IUCN status Least Concern (2018). Mediterranean sub-population Vulnerable (2018).
Population Likely to be well over 100,000. Global trend unknown (Mediterranean sub-population decreasing).
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies, though there is a genetically distinct population in the Mediterranean
Sea.
Other names Goose-beaked whale, goosebeak whale.
DISTRIBUTION Widely distributed in cool polar to warm tropical waters worldwide, though generally absent from
very high latitudes and water shallower than 200m (uncommon south of the Antarctic Convergence around 60°S).
Present in many enclosed seas, including the Gulf of Mexico, Caribbean Sea, Sea of Okhotsk, Gulf of California and
the Mediterranean Sea (where it is the only species of beaked whale commonly occurring). Seems to prefer deeper
waters over and near the continental slope, or oceanic waters, with a complex seabed topography; it is most common
in canyons along shelf margins or around oceanic islands or seamounts. Usually occurs in water deeper than 1,000m.
Evidence of seasonal movements in some parts of the range, but other populations appear to be resident.

May have dark ADULT FEMALE

Less extensive linear
oval patch amid Highly variable dark crescents scarring than on male
paler coloration on (none, one or two) slightly Colour variable, from
melon, especially in behind blowhole dark grey to reddish
Mediterranean (rare brown, but usually
in males) browner than male

Head may be paler (lightens with Can be extensive white

Less prominent lower age) but usually not as dramatic in spotting from cookiecutter
No erupted shark bites
jaw than adult male contrast or extent as in male
teeth

DIVE Sequence Often exposes beak on surfacing; entire head and part of body may be exposed when swimming
fast or just before long dive (when often lunges out of water); occasionally flukes before a deep dive in some regions
(not north-west Atlantic). • Depth Holds mammalian record for dive depth (2,992m off southern California, USA);
probably physiologically capable of diving much deeper (possibly to 5,000m); believed to dive to more than 1,000m
for 60+ minutes both day and night, non-stop, year-round (indeed, estimated to spend two-thirds of its life at depths
greater than 1,000m). • Duration Holds mammalian record for longest dive duration (3 hours 42 minutes off Cape
Hatteras, North Carolina, USA); same study analysed 3,680 dives by 26 individuals over 5 years: 5 per cent of dives
exceeded 77.7 minutes; foraging dives average 58 minutes (followed by average 1.9 minutes on the surface), often
interspersed with shorter ‘bounce’ dives; non-feeding dives average c. 12 minutes.
BLOW Bushy blow usually about 1m high, projecting slightly forward and to left (but usually inconspicuous or
invisible).

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 Highly variable dark crescents (one ADULT MALE
Extensive linear parallel scarring on head
or two) slightly behind blowhole
and front half of body (teeth rake marks
Melon more Highly variable pair of dark caused by fighting with other males)
bulbous than in crescents around eye
Mesoplodon, Body colour usually dark Spindle-shaped
less bulbous Head and cape brown (but highly variable, body (robust for a
than in usually paler or from grey to almost white) – beaked whale)
Hyperoodon white (may extend all adult males have extensive
to dorsal fin) white on their bodies
Slight Relatively small, falcate
Smoothly concavity Greatest girth (sometimes triangular) dorsal fin
sloping on top of around midpoint
forehead head Dorsal fin approximately two-
thirds of the way along back
Relatively
short,
poorly May be barnacles
defined on dorsal fin (some
beak regions only)

Small, dark
flippers
Beak may
be light on Eye usually surrounded
upperside, dark by darker coloration
on underside
Two throat
grooves (V-shaped)
Two erupted, forward-
Upturned mouthline
pointing, conical teeth at Lower jaw
(giving unique ‘smile’)
tip of lower jaw (visible extends well
when mouth closed) beyond upper jaw
Orange-brown or
greenish-brown patches
caused by films of
diatoms or algae (some
SIZE individuals completely or
L: ♂ 5.3–6m, ♀ 5.5–6m; partially covered)
WT: 2.2–2.9t; MAX: 8.4m, 3t
Calf – L: 2.3–2.8m; WT: 250–300kg
Tailstock
compressed
Usually reversal laterally
of normal
countershading
(i.e. light above,
dark below)
Extensive round or oval white
scarring (healed bites from
cookiecutter sharks and,
sometimes, lampreys) – absent
in some parts of range
Mature males have seven
times more scarring than
females (1–6 new scars
appear per year)

AT A GLANCE Worldwide except High Arctic

and Antarctic • Colour highly variable, from
80°

slate-grey to brown to white • May be covered

70°

60°

50° in round or oval white scars from cookiecutter

shark bites (some regions only – e.g. rare
40°

30°

20°
10° in north-west Atlantic) • Numerous linear
0°
10°
scars (especially on male) • Medium size •
20°

30°
Smoothly sloping forehead with relatively
40°

50°
short beak • Upturned mouthline (giving
60° unique ‘smile’) • Two conical, forward-pointing
70°
teeth at tip of lower jaw of male (visible when
80°
mouth closed) • Dorsal fin approximately two-
thirds of the way along back

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 BEHAVIOUR Occasionally breaches. Lobtailing often observed in some regions, especially when there is more than one
male in a group. Tends to be very sensitive to low levels of human-made noise, typically escaping quickly, and generally
avoids boats; but it can be inquisitive in some areas. Significant scarring suggests males fight for access to females.
FOOD AND FEEDING Primarily deepwater squid; may consume deepwater fish and crustaceans in some parts of range.
Most feeding occurs at or near seabed (though also sometimes in water column). Likely to be a suction feeder.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only; 8cm tall (including portion buried in jawbone).
GROUP SIZE AND STRUCTURE Typically in small fluid groups of 1–4 animals; groups of up to 25 reported, but more
than 10 is rare. Larger groups typically contain at least two adult males and two or three adult females and juveniles.
Lone individuals are usually older males. Rarely interacts with other cetacean species.

ADULT FEMALE
Large, slit-like blowhole
visible at very close range
Flukes wide in relation
to body length

Underside of flukes light

(sometimes white)
Slightly concave
Flippers tuck away in slight May be barnacles
trailing edge
depressions on body called on flukes
‘flipper pockets’ (in both sexes)

Concavity on top of head Head and front half of body OLD ADULT MALE
becomes more obvious in become progressively whiter
older animals (both sexes) in older adults of both sexes
(usually more obvious in males)
White area may extend
Melon more on upperside as far back
prominent (even as dorsal fin
bulbous) in older
males

Teeth may be worn down Underside darkens with Becomes increasingly covered in
as far as gums in older age (both sexes) white spots (healed cookiecutter
animals (sharply pointed shark bites) as it ages (particularly
in younger males) in lower latitudes)

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 ADULT FEMALE
COLOUR VARIATIONS

YOUNG ADULT FEMALE

Uniform grey, bluish-grey

Rounded, more or brownish upperside CALF
dolphin-like head
Pale head No scarring

No erupted
teeth
Darker spot Counter-shaded No cookiecutter
around the eyes (paler below) shark bites

ADULT MALE
White or cream-coloured teeth may
be partially or completely covered
in reddish-brown or purple-brown
stalked barnacles

DORSAL FIN VARIATIONS

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 NORTHERN BOTTLENOSE WHALE
Hyperoodon ampullatus (Forster, 1770)

The northern bottlenose whale is the largest beaked whale in the North Atlantic, one of the best-known members of
the family and one of the few to have been targeted by whalers on a large scale. The heads of males and females look
so different that early anatomists believed them to be two separate species. References to ‘bottlenose whales’ in the
northern North Pacific refer to Baird’s beaked whales.
IUCN status Near Threatened (2020).
Population Probably in the order of 20,000+. Possibly c. 100,000 before intense whaling began in the 1880s, but this
was reduced to a few tens of thousands by the time whaling ceased in the 1970s. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Northern bottle-nosed whale, bottlehead, flathead, steephead, common bottlenose whale.
DISTRIBUTION Cold temperate to Arctic waters in the North Atlantic. Ranges from the ice edge to at least c. 37°N.
However, rarely seen south of c. 55°N – with one notable exception: a well-studied population in a submarine canyon
called The Gully, 200km south-east of the Atlantic coast of Halifax, Nova Scotia, Canada. Usually frequents waters
deeper than 500m (with a preference for 800–1,800m) over the continental slope. Rarely strays over the continental
shelf, except in submarine canyons, and prefers areas with complex seabed topography, such as the continental shelf
edge, oceanic islands and seamounts. Generally occurs in open water.
BEHAVIOUR Males may use their large, bulbous heads to headbutt one other. When resting at the surface, both sexes
and all ages may hang in the water at a 45° angle, with the entire melon and beak above the surface. Breaching and
lobtailing are not uncommon. Can be very curious towards boats and even large ships, and will often approach closely.
FOOD AND FEEDING Mainly deepwater squid; sometimes fish; rarely prawns, sea cucumbers and starfish. Most
feeding appears to be on or near seabed in deep water; probably suction-feeder.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only (up to 5cm tall).
GROUP SIZE AND STRUCTURE Typically 1–10, rarely more than 20. Group size varies according to region. There is
some segregation by age and sex. Females appear to form loose, fluid associations, but pairs of males form long-term
relationships that can last anything from days to 1–2 years.

ADULT FEMALE
Melon bulbous but Much of melon and
relatively smaller and less face slightly paler
square than adult male
No linear scarring

No erupted Beak may be thicker

teeth than on male

DIVE Sequence Rolls forward, sometimes with head, back and dorsal fin visible simultaneously; rarely flukes.
• Depth Routinely dives to 800+m (average in one study 1,065m); maximum recorded 2,339m. • Duration Routinely
30–40 minutes; maximum recorded 94 minutes; possibly capable of 2 hours.
BLOW Low, puffy blow (1–2m), often clearly visible and canted forward.

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 Huge, rounded melon (often overhanging Uniform medium to dark grey, ADULT MALE
rostrum) with very steep (almost square- chocolate- to olive-brown or tan
shaped) forehead as it matures coloration (brownish or yellowish
Two conical teeth tinge may be enhanced by film or
at tip of lower Much of melon and face (back to Robust patches of diatoms)
jaw (rarely visible eyes) white or cream-coloured body
outside closed
Distinct junction Tip usually
mouth and often
between beak and pointed
wear down or
melon (but no crease) Prominent, erect but
fall out)
relatively small, falcate
dorsal fin (up to 30cm)
two-thirds of the way
along back

Small, blunt
Medium-length, thick, well- Two shallow, flippers (fit Little or no May be mottled with
defined beak (highly variable V-shaped throat into ‘flipper linear scarring round or oval white
appearance – beak looks grooves pockets’ on scarring (healed bites from
smaller relative to melon sides) cookiecutter sharks)
growing larger)
ADULT MALE GREY FORM

OLD ADULT MALE

Huge melon with very steep,
flattened forehead (giving
distinctly squarish profile)

SIZE
L: ♂ 7.5–9m, ♀ 7–8.5m; Much of melon and
WT: c. 5–8t; MAX: 9.8m, 10t face very pale (white
Calf – L: 3–3.5m; WT: c. 300kg or cream-coloured)

80°
AT A GLANCE Cold, deep waters of North
Atlantic • Medium size (larger than other
70°
beaked whales in region) • Grey, tan or
brownish coloration • Huge, squared-off,
60°
bulbous white or cream-coloured melon
50°
(especially in male) • Medium-length, thick,
well-defined beak • Prominent falcate dorsal
fin two-thirds of the way along back • Little
40°

30°
or no linear scarring • Male’s teeth not clearly
20°
visible • Often inquisitive and may approach
10°
stationary vessels

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 SOUTHERN BOTTLENOSE WHALE
Hyperoodon planifrons Flower, 1882

The southern bottlenose whale is by far the most common beaked whale sighted in Antarctic waters, though is still
less well known than its slightly larger northern counterpart, the northern bottlenose whale (with which it forms an
antitropical species pair). It has never been hunted commercially.
IUCN status Least Concern (2018).
Population Unknown, though certainly high hundreds of thousands. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Antarctic bottle-nosed whale, bottlehead, flathead, steephead.
DISTRIBUTION Distribution appears to be continuous in cold temperate to Antarctic waters in the southern hemisphere.
Most sightings are from c. 57–70°S, with areas of concentration from 58–62°S in the South Atlantic and eastern
Indian Oceans. There seems to be a north–south migration in winter–summer. During the summer, most frequently seen
within 120km of the ice edge, sometimes reaching the edge itself. Usually in waters deeper than 1,000m, preferring
areas with complex seabed topography, such as submarine canyons, the continental shelf edge, oceanic islands and
seamounts.
BEHAVIOUR Breaching (often many times in rapid succession), lobtailing and other aerial behaviours are not uncommon.
Unlike northern bottlenose whales, males of the southern species seem to fight with their teeth more than headbutt.
FOOD AND FEEDING Mainly oceanic deepwater squid; also krill and occasionally fish (especially Patagonian toothfish).
May compete with sperm whales for same prey species (though bottlenose whales probably take smaller individuals).
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only (up to 5cm tall).
GROUP SIZE AND STRUCTURE Typically 1–5. Usually fewer than 10, with occasional sightings of up to 25. Nothing is
known about group composition.

ADULT FEMALE
Melon bulbous but Much of melon
relatively smaller and face paler
No linear
and less square
scarring
than in male

No erupted Beak may

teeth be thicker

DIVE Sequence Arches steeply before sounding dive; rarely flukes • Depth Unknown. • Duration Dives range
from 15–40 minutes.
BLOW Low, puffy blow (1–2m), often clearly visible and canted forward.

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 Huge, rounded melon (often Some adult males Uniform light greyish to ADULT MALE
overhanging rostrum) with very light in colour brownish coloration (brownish
very steep forehead or yellowish tinge may be
Spinal field (and much enhanced by film of diatoms);
Front of melon may be
of back) may be very can appear golden in sunlight
Distinct flatter in older males, giving
light (almost white)
junction a more squarish profile
between Robust body Tip usually pointed
Much of melon and face
beak and
(back to eyes) white or
melon (but Prominent, erect but
cream-coloured
no crease) relatively small, falcate
dorsal fin (up to 30cm) two-
thirds of the way along back

Small, blunt flippers (fit into

Medium-length, thick, ‘flipper pockets’ on sides) May be mottled with round
well-defined beak Two shallow, V-shaped or oval white scarring (healed
(highly variable length) throat grooves bites from cookiecutter sharks
– more extensive with age)
Two conical teeth at tip of lower jaw
(rarely visible outside closed mouth
and often wear down or fall out) May have white or cream-coloured melon (light ADULT FEMALE
Teeth may have stalked colour extends to just behind blowhole – usually
barnacles attached further back than in Longman’s beaked whale) Flukes wide
relative to
body length Usually
no
median
notch
(or
shallow
Crescent-shaped blowhole one)
Colour highly
(ends point forward)
variable
OLD ADULT MALE
Heavy linear
scarring typical
(from male–male
SIZE combat); increases
L: ♂ 6–7m, ♀ 6.5–7.5m; with age
WT: 6–7.5t; MAX: 7.8m
Calf – L: 2–3m; WT: unknown

80°

70°

60°
AT A GLANCE Cool temperate to Antarctic
50° waters of southern hemisphere • Medium
size • Huge bulbous melon (especially in
40°

30°

male), sometimes white or cream-coloured

20°
10°
0°
10° • Uniform light greyish to brownish coloration
20°

30° • Medium-length, thick, well-defined beak

40°

50°
• Prominent falcate dorsal fin two-thirds of
60° the way along back • Extensive linear scarring
70°
• No clearly visible teeth
80°

SOUTHERN BOTTLENOSE WHALE 117

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 SHEPHERD’S BEAKED WHALE
Tasmacetus shepherdi Oliver, 1937

One of the least known of all cetaceans – and yet among the most distinctive – with males, females and calves sharing
the same contrasting colour pattern. This is also the only beaked whale to have a mouthful of functional teeth (in both
jaws and present in both sexes).
IUCN status Data Deficient (2018).
Population Unknown. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Tasman whale, Tasman beaked whale.
DISTRIBUTION Presumed to have a circumpolar distribution in deep, cool, temperate oceanic waters of the southern
hemisphere. Known from fewer than 50 strandings and a couple of dozen sightings at sea, mostly from 30–46°S.
Apparent hotspots include New Zealand, southern Australia, Shag Rocks near South Georgia, and Tristan da Cunha.
The mean depth of water for sightings in Australia and New Zealand is 1,208m.
BEHAVIOUR Very little known. Some breaching, lobtailing and spyhopping observed. Individuals in a group typically
surface and dive close together; may swim in a moderately spaced ‘chorus line’ for several minutes. Unless travelling,
they often return to the surface within 100–150m of the original dive. No evidence of evasive behaviour in the presence
of boats.
FOOD AND FEEDING Little known, but prey includes fish (especially eelpout), squid and crabs.
TEETH Upper jaw 34–42; lower jaw 36–56.
GROUP SIZE AND STRUCTURE Very little information, but the few reliable sightings suggest that group size ranges
from 2–14 individuals with a mean of 5.4 (3–6 being most common).

ADULT
Readily identifiable from above by
pale creamy-brown melon and clear
Pale colour of May appear
boundary between darker cape and
melon may extend browner due to film
lighter back behind dorsal fin
down rostrum of diatoms on skin
dorsally

No (or
minimal)
median
notch

Lighter pigmentation of
Crescent-shaped Pale 'shoulder blaze' tailstock may continue
blowhole often visible above uninterrupted to
water on surfacing upperside of flukes

DIVE Sequence Typically skims beak through surface (parallel to water) but may lift it entirely out of water (typically
at angle of c. 30–40°); tends to remain below surface between blows; usually little arching of back on sounding dive;
does not show flukes. • Depth No information. • Duration Limited encounters suggest 5–15 minutes, with surface
intervals of 4–17 minutes.
BLOW Bushy blow 1–2m tall and canted forward (often visible from considerable distance).

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 Bulbous, pale melon Unique pale ‘shoulder May have single or paired ADULT MALE
(more prominent than blaze’ 20–30cm wide Dark brownish-grey cape linear scars (presumably (female almost
in mesoplodonts, above flipper and on upperside and sides, from fighting with other identical except
less prominent extending vertically extending from about males or social behaviour teeth at tip of lower
than in Berardius from underside blowhole to mid-dorsal within group) jaw do not erupt)
and Hyperoodon of body (top often fin region (colour may
– comparable to visible above water vary according to lighting Small dorsal fin two-thirds of the
Indopacetus) on surfacing) conditions or activity) way along back (30–35cm high),
variable in shape (usually falcate
Steep Dark brownish- Spindle- but some more triangular)
forehead grey around eye shaped
(giving mask-like body Dorsal fin usually dark brownish-
(with clear
appearance) grey but bicoloured in many
demarcation
individuals (dark brownish-grey in
from beak)
front, pale grey or brownish-grey
behind – changing at point where
dark cape meets lighter tailstock)

Small flippers
Two enlarged Underside mostly
teeth protrude Two shallow creamy white or Tailstock
forward at tip of throat grooves pale grey-brown noticeably
lower jaw (rarely Long, dark paler May be some
Beak more pointed than in Strong contrast between scarring from
visible at sea) brownish-grey, mesoplodonts (with slightly light and dark areas (but cookiecutter shark
dolphin-like beak protruding lower jaw)
(grows relatively separating lines not sharp) bites
longer with age)

Some individuals may have a

DARK GREY TYPE more triangular dorsal fin
SIZE
L: ♂ up to 7m, ♀ up to 6.6m;
WT: c. 2.3–3.5t;
Calf – L: c. 3–3.5m; WT: unknown
o (or
minimal)
median
otch

80°

AT A GLANCE Cool temperate deep waters of

70°

60°

50° southern hemisphere • Medium size (larger

40°

30°
than mesoplodonts) • Pale bulbous melon
20°
10°
with steep forehead • Contrasting dark
0°
10°
brownish-grey cape and much paler tailstock
20°

30°
• Dark mask-like feature over eyes • Top of
40°
pale ‘shoulder blaze’ often visible on surfacing
• Dorsal fin often bicoloured (darker in front,
50°

60°

70° lighter behind) • Mainly small, close-knit

80° groups
presumed range (from limited sightings and strandings)

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 LONGMAN’S BEAKED WHALE
Indopacetus pacificus (Longman, 1926)

Longman’s beaked whale was one of cetology’s great long-standing mysteries. Until 2003, the only evidence for its
existence came from two weathered skulls: one found on an Australian beach in 1882, and the other on the floor of
a Somalian fertiliser factory in 1955. However, live animals are now being seen with some regularity at scattered
locations in the tropical Indo-Pacific and there have been about 20 strandings.
IUCN status Least Concern (2020).
Population Unknown. Despite the paucity of sightings, its wide distribution suggests that it is not rare. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy Originally placed in the genus Mesoplodon, but morphological and genetic studies prove that it should be
in its own genus; no recognised forms or subspecies.
Other names Indo-Pacific beaked whale, tropical bottlenose whale.
DISTRIBUTION Distribution is poorly known, but it appears to be widespread and fairly continuous in the subtropical
and tropical Indo-Pacific. Thought to be more abundant in the western part of its distribution. Sightings tend to be in
areas with surface water temperatures of 21–31°C (mostly in water warmer than 26°C). May push further south or
north with warm currents. Most sightings are oceanic, over or near areas with steep bottom topography, in depths of
250–2,500m or more.
BEHAVIOUR Larger groups tend to be more active at the surface and may ignore or approach boats.
FOOD AND FEEDING Presumed to feed mainly on deepwater squid, with some fish. Feeding techniques unknown.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Tight groups tend to be large (1–110 recorded, with overall average of 18.5) and may
vary regionally.

ADULT FEMALE
Melon less Adult may lose Little or no linear scarring
bulbous than pale melon
in male with age May appear darker
than male (due to lack
of linear scarring)

No erupted May be extensive oval

teeth white scarring from
cookiecutter shark bites
SIZE
L: ♂ 5.9–6.1m (based on two males),
♀ 5.7–6.5m;
WT: c. 6–7.5t (guesstimate);
Calf – L: c. 2.3–2.9m; WT: c. 230kg

DIVE Sequence Swims faster and more ‘aggressively’ than most other beaked whales; when surfacing quickly,
head and beak appear quite high out of water as it throws up rooster-tail of spray; little arching of back (cf. Cuvier’s
beaked whale) before long dive. • Depth Unknown, but probably a deep diver. • Duration 11–33 minutes; one
individual tracked underwater acoustically for 45 minutes (contact lost before it surfaced).
BLOW Low, bushy blow fairly conspicuous and angled slightly forwards.

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 Melon drops steeply to ADULT MALE
Pale tan or lighter-coloured head
beak (average c. 75°
(extends as far back as blowhole –
– cf. 90° in southern Bluish-grey or brownish-grey to olive-brown
cf. southern bottlenose whale)
bottlenose whale) or bronze-brown upperside and tailstock
Dark area behind (highly variable between individuals and
Crease according to light conditions)
between melon forms part of
melon dark patch around eye
Appears greyer in overcast
and beak Bulging conditions, browner in
Beak usually May have paler ‘ear
melon bright conditions
dark above, patch’ in dark area
and white, behind eye Dorsal fin may have
light grey, Spindle-shaped darker leading edge
pale tan or body and pale centre
pink below
(variable) Relatively tall, erect
and falcate dorsal
fin two-thirds of the
way along back

Fairly Small, blunt Pale flanks rise to May be extensive

Moderately straight
long, well- mouthline flippers shoulder (can be linear scarring May be extensive
Two shallow, conspicuous at sea)
defined oval white scarring
V-shaped
dolphin-like Flippers mostly Much lighter sides from cookiecutter
throat grooves
beak dark on upperside, and underside shark bites (often
Single pair of pear-shaped teeth at light on underside (shading to white looks spotty)
tip of lower jaw (male only), barely around genital region)
visible outside closed mouth except
at close range
Coloration ADULT MALE VARIATION
In older males, melon may drop almost variable
perpendicular to beak (sometimes with Many individuals covered
well-developed overhang) in numerous light, oval
cookiecutter shark bites

80°

70°

60°

50°
AT A GLANCE Warm waters of the Indo-
40° Pacific • Medium size • Spindle-shaped body
• Low bushy blow often angled forward •
30°

20°

Conspicuous, bulging pale melon • Distinct,

10°
0°
10°
20° sharply demarcated beak • Falcate, dolphin-
30°

40°
like dorsal fin • Fin two-thirds of the way
50°
along back • Apparent colour varies according
to weather conditions
60°

70°

80°

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 PERRIN’S BEAKED WHALE
Mesoplodon perrini Dalebout, Mead, Baker & van Helden, 2002

Officially named in 2002, this is one of the least-known beaked whales (all information is based on just six stranded
specimens, from southern California). Its external appearance closely resembles Hector’s beaked whale (though the
two species are not closely related). Its nearest relative is the pygmy beaked whale.
IUCN status Endangered (2020).
Population Possibly fewer than 1,000 mature individuals. Decreasing.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names California beaked whale.
DISTRIBUTION Currently known only from southern California, USA. Strandings range from Torrey Pines State Reserve
(32°55’N – just north of San Diego) north to Fisherman’s Wharf, Monterey (36°37’N). It might be expected in waters
with depths of more than 1,000m (mainly beyond the continental shelf but possibly also close to shore where the water
is sufficiently deep). Researchers recorded unique echolocation pulses 350 nautical miles off the coast of California,
believed to have been made by Perrin’s. There have also been probable acoustic detections off Baja California, Mexico.
BEHAVIOUR This species has never been reliably identified at sea, so there is no information on behaviour. Probably
unobtrusive and difficult to spot in anything but calm conditions.
FOOD AND FEEDING Virtually no information, but very limited evidence from stomach contents suggests primarily mid-
and deepwater squid; may also include deepwater fish and shrimps.
TEETH Upper jaw 0; lower jaw 2. Laterally compressed teeth erupt in males only (females have similar teeth, but they
do not erupt); the exposed portion is roughly the shape of an isosceles triangle; up to 64mm long.
GROUP SIZE AND STRUCTURE No information.
ADULT FEMALE

Small head with ADULT FEMALE

relatively straight Predominantly
mouthline dark grey Flukes wide
in relation to
body length

No
median
notch

Broad, crescent-
shaped blowhole Tailstock
Body flecked with oval
(ends point forward) compressed
cookiecutter shark bites
laterally

DIVE SEQUENCE (speculative)

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 Appearance of darker face ADULT MALE
Smoothly sloping, mask (dak grey colour
non-bulbous melon (based on little information)
extending from near gape Dark grey upperside
to eye and to just behind grading to lighter grey
blowhole) or white underside
Melon may be paler Moderate to heavy linear
and/or browner scarring (single, rather than
Small head with parallel, tooth rake marks)
Relatively short beak relatively straight Spindle-
(shorter than other mouthline (does not shaped body Small, triangular or slightly
Mesoplodon beaked rise towards rear) falcate dorsal fin two-thirds
whales except of the way along back
pygmy and
Hector’s) Limited evidence
suggests single scar lines
(cf. two parallel lines
Dark grey typical of most male
rostrum beaked whales)

Small, narrow White patch

Two relatively Two shallow
flippers around Body flecked with
large, laterally throat grooves
umbilicus oval cookiecutter
compressed Light grey to white shark bites
triangular teeth c. throat and lower jaw
1–2cm behind tip of Teeth may be colonised
lower jaw (situated by stalked barnacles
just behind tip in
adult male Hector’s Teeth exposed above FLUKES (UNDERSIDE)
beaked whale) gumline by c. 3cm (visible
outside closed mouth)

Distinctive ‘starburst’
SIZE
light-and-dark patterning
L: ♂ 3.9m, ♀ 4.3–4.4m;
on underside of tail (also
WT: c. 900kg; MAX: 4.53m
Calf – L: c. 2–2.1m; WT: unknown
occurs on Stejneger’s
(based on very few specimens) beaked whale)

50°

AT A GLANCE Eastern North Pacific • Small

40°
to medium size • Generally nondescript
counter-shaded coloration • Small triangular
30°
or slightly falcate dorsal fin two-thirds of the
way along back • Appearance of darker face
mask • Two large triangular teeth near tip of
20°
lower jaw may be visible at close range.

10°

range shows possible distribution strandings

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 PYGMY BEAKED WHALE
Mesoplodon peruvianus Reyes, Mead & Van Waerebeek, 1991

Formally described in 1991, the pygmy or Peruvian beaked whale is the smallest species of beaked whale. It is very
poorly known: for many years the majority of records were curated specimens from Peru, but recently there have been
more live sightings in the Gulf of California, Mexico, and elsewhere.
IUCN status Least Concern (2020).
Population Unknown, but likely tens of thousands (reasonably common within its limited range). Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies. This is the previously unnamed species formerly known as
‘Mesoplodon sp. A’.
Other names Peruvian beaked whale (used interchangeably with ‘pygmy’), lesser beaked whale.
DISTRIBUTION Originally described primarily from freshly captured specimens landed in Peruvian fishing ports.
However, in recent years it has become the most frequently sighted Mesoplodon in deep waters of the sub-tropical and
tropical eastern Pacific Ocean (although this is relative); Mexico’s Gulf of California is a sightings hotspot. A single male
stranded near Kaikoura, South Island, New Zealand (42°31’S), in 1991, is considered to be a vagrant.
BEHAVIOUR Generally unobtrusive and difficult to spot in anything but calm conditions. Groups typically dive for 15–30
minutes, surface again some distance away, breathe half a dozen times and then dive again. Has been known to
approach small boats very closely (albeit briefly) but usually keeps its distance. Breaching, lobtailing and other surface
behaviours have been recorded, but appear to be rare.
FOOD AND FEEDING Limited evidence suggests primarily mid- and deepwater fish, but also probably deepwater squid
and shrimps. Feeding techniques unknown.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only; 31–65mm long.
GROUP SIZE AND STRUCTURE Typically in groups of 2–5 (sometimes 1–8). Groups usually include mixed sex and age
classes.
Some individuals have
More nondescript ADULT FEMALE
darker forehead (giving Dorsal fin often more
than male
mask-like appearance) triangular than in male
Shallow angle
to forehead (less (but variable)
pronounced melon Mostly olive-brown
than in male) to grey-brown
upperside and sides
No significant
scarring

No erupted
teeth Paler grey to
whitish lower SIZE
Mouthline much less sides and L: ♂ 3.4–3.9m, ♀ 3.4–3.6m;
arched than in male underside WT: unknown; MAX: 4.1m
Calf – L: c. 1.6m; WT: unknown

DIVE Sequence Surfaces with shallow roll, showing top of head (sometimes also much of beak) and back; slightly
arches back before diving. • Depth Likely to forage at depths greater than 500m.• Duration Typically 15–30 minutes
(based on limited observations).
BLOW
Indistinct and rarely visible.

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 ADULT MALE

Adult males have broad white or Rest of body dark brown or

Melon bulges slightly cream swathe across back and black (may have brownish
in front of blowhole sides (in front of dorsal fin) patches due to diatoms on skin)
(and more pronounced
than in female) Small triangular or slightly falcate
Slender, spindle- dorsal fin two-thirds of the way along
Melon slopes shaped body back (variable but often similar in
smoothly to Relatively shape to that of harbour porpoise)
short, narrow small head
beak Dorsal fin not as
falcate as in other
mesoplodonts

Wide base

Beak tip Small flippers

may be Moderate to heavy Deep tailstock
brown or Arched Two throat grooves linear scarring (tooth
white mouthline rake marks)
Two small, conical teeth at
apex of arched lower jaw
Teeth perpendicular in (difficult to observe at sea) ADULT MALE
adult male (lean forward Flukes wide in
20–40° in sub-adult) relation to body
length

No median
notch
in most
individuals

DORSAL FIN VARIATIONS Tailstock

compressed
laterally

30°

20° AT A GLANCE Mainly warm waters of eastern

Pacific Ocean • Small size • Adult male has
broad white swathe across back and sides •
10°

0° Small groups • Dorsal fin variable but usually

small, wide-based and roughly triangular •
10°
Difficult to spot except in calm conditions •
20°
Long dives between surfacings

30°

(plus one stranding on South Island, New Zealand)

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 DERANIYAGALA’S BEAKED WHALE
Mesoplodon hotaula Deraniyagala, 1963

Currently known from only 11 confirmed specimens and a handful of likely sightings at sea. This poorly known beaked
whale was originally described in 1963 from a specimen in Sri Lanka, but its status as a new species was disputed for
more than half a century (until DNA analysis of new specimens proved the original declaration right).
IUCN status Data Deficient (2020).
Population Unknown, but presumed to be rare or uncommon. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies. For many years it was considered synonymous with the very similar
ginkgo-toothed beaked whale (and may be indistinguishable from it at sea), but was formally accepted as a separate
species in 2014.
Other names Atoll beaked whale (rarely used since specimens have been found in places other than atolls).
DISTRIBUTION Currently known only from a small number of widely scattered strandings and a few tentative sightings
in the Indo-Pacific. It appears to have a tropical range in the Indian Ocean and at least part of the Pacific Ocean. Little is
known about habitat preferences, but it might be expected in waters with depths of more than 500m.
BEHAVIOUR Virtually nothing is known. There have been a number of possible sightings at Palmyra Atoll, in the central
Pacific, and during two of these distant animals were observed leaping completely out of the water.
FOOD AND FEEDING Like other mesoplodonts, likely to feed primarily on squid; maybe also deepwater fish. Feeding
techniques unknown.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only; 10cm long, 9cm wide (in comparison, ginkgo-toothed beaked
whale teeth are wider than they are long).
GROUP SIZE AND STRUCTURE Most sightings of mesoplodonts likely to have been Deraniyagala’s were of paired
animals (including at least two mother–calf pairs), giving a mean group size of 2.2 individuals (ranging from two to
three).
ADULT MALE
Crescent-shaped blowhole
(ends point forward)
Tailstock laterally
compressed

Upper jaw darker

than lower jaw

ADULT FEMALE FLUKES ADULT MALE FLUKES

(upperside) (upperside)

Median lobe with No median notch

small notch

Flukes wide in relation

Flukes wide in relation
to body length
to body length

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 ADULT MALE
(Based on single adult male from
Shallow the Seychelles, 2009)
forehead merges Predominantly dark
with rostrum (no (Adult females may be more
blue-black pigmentation brownish than bluish)
distinct melon) (grading to slightly
lighter shade on
underside)
Upper ‘lips’ whitish Slender Long, linear scars may heal
(grading to blue- head quickly to skin colour (more
black on rostrum) May be lighter visible on males of other
mottled grey Mesoplodon species)
Tip of Slender, spindle-
on cheek
rostrum shaped body
light grey Falcate dorsal fin two-
No dark thirds of the way along
eye patch back (can be taller in
some individuals)

Small, narrow
flippers

Two shallow throat

Small number of light spots on
grooves
back and sides probably made by
Teeth may cookiecutter shark bites (though may
be broken Lower jaw and throat heal same colour as surrounding skin)
or worn predominantly white

Tip of Small portion of teeth exposed

lower Stalked barnacles on front top edge of raised area
jaw pale may grow on of lower jaw
erupted teeth
(giving a ‘tasselled’
appearance)
Moderately
long beak SIZE
with arched L: ♂ 3.9–4.3m, ♀ 4.5–4.8m;
mouthline WT: unknown; MAX: unknown
Calf – L: unknown; WT: unknown

80°

AT A GLANCE Tropical Indian and west

70°

60°

50° Pacific Oceans • Small to medium size •

40°

30°
Predominantly dark with pale lower jaw and
20°
10°
throat • Possibly no long linear scarring •
0°
10°
Prominent falcate dorsal fin two-thirds of the
20°

30°
way along back • Visible teeth near apex of
40°
moderately arched mouthline • Teeth may
be covered in stalked barnacles • May be
50°

60°

70° indistinguishable from ginkgo-toothed beaked

80° whale at sea

possible range stranding locations

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 GINKGO-TOOTHED BEAKED WHALE
Mesoplodon ginkgodens Nishiwaki and Kamiya, 1958

There are no confirmed sightings of live ginkgo-toothed beaked whales at sea, although there have been several possible
ones. There have been fewer than 30 widely scattered strandings and captures of this poorly known species, spread
across the Pacific and Indian Oceans.
IUCN status Data Deficient (2020).
Population Unknown. Probably uncommon, given the small number of records. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies. For many years Deraniyagala’s beaked whale was considered
synonymous with ginkgo-toothed, but Deraniyagala’s was formally accepted as a separate species in 2014. The two
species may be indistinguishable at sea.
Other names Japanese beaked whale, ginkgo-toothed whale.
DISTRIBUTION Exact distribution is unclear, due to the small number of records. Records are widely scattered, with
most concentrated in the western Pacific (Indian Ocean records are more likely to represent Deraniyagala’s). Mainly
deep tropical to temperate waters, and probably more common in areas of complex seabed topography.
BEHAVIOUR There have been no confirmed sightings of live animals at sea, so there is no information on behaviour.
FOOD AND FEEDING Very little known. Like other beaked whales, it is presumed to eat mainly deepwater squid, and
some fish.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only (6.5cm tall).
GROUP SIZE AND STRUCTURE Nothing known.

ADULT FEMALE
Beak may have May be paler than male
white tip (especially on underside)

Teeth do not
erupt

Lower jaw paler

ADULT MALE
Crescent-shaped blowhole Flukes wide in
(ends point forward) relation to body
length (up to 25
per cent)

No
median
notch

Tailstock
compressed
laterally
Trailing edge may
be slightly convex

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Smoothly sloping, slightly
bulbous melon merges with
rostrum (no crease)
ADULT MALE
Basic dark grey (or brownish)
countershading coloration
(poorly known from few fresh
specimens)

May be lighter
mottled grey Little or no pale
on cheek Spindle-shaped body linear scarring
Flattened tooth near
apex of arch on Small head
each side of lower
jaw (mostly hidden
in gum) Small, falcate dorsal
fin two-thirds of the
way along back

Small, narrow
Dark eye flippers
Moderately
long beak with patch Extensive round or oval white
pale tip (upper scars from cookiecutter shark bites
and lower jaws) (especially in urogenital area)
Two shallow,
V-shaped throat
Prominent arch in grooves Teeth resemble
mouthline slightly behind leaves of ginkgo tree
middle of lower jaw

SIZE
L: ♂ 4.7–5.3m, ♀ 4.7–5.3m;
WT: c. 1–1.5t; MAX: 5.3m, 2t Teeth similar (in size and shape) to those of
Calf – L: 2–2.5m; WT: unknown Deraniyagala’s beaked whale and wider-toothed
variants of Gray’s beaked whale

80°

70°
AT A GLANCE Tropical to temperate waters
60° of Pacific and Indian Oceans • Medium size •
50°

40°
Little or no pale linear scarring • Moderately
30° long beak with white tip • Smoothly sloping,
slightly bulbous melon • Prominent arch in
20°
10°

mouthline slightly behind middle of lower

0°
10°

jaw • Wide tooth near apex of arch on each

20°

30°

side of lower jaw • Small, falcate dorsal fin

40°

50°

60°
two-thirds of the way along back • May be
70°
indistinguishable from Deraniyagala’s beaked
80°
whale at sea
possible range strandings

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 GRAY’S BEAKED WHALE
Mesoplodon grayi von Haast, 1876

Our knowledge of Gray’s beaked whale has improved significantly in recent years, though it is still rarely seen at sea.
However, strandings are fairly common and it is probably quite abundant. On one extraordinary occasion in June 2001,
a mother and calf spent almost five days in Mahurangi Harbour, North Island, New Zealand, giving researchers a rare
opportunity to observe this species closely.
IUCN status Least Concern (2020).
Population Unknown. Appears to be fairly common in at least parts of its range, based on genetic evidence and the
number of records. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Scamperdown beaked whale, southern beaked whale.
DISTRIBUTION Circumpolar in temperate offshore waters of the southern hemisphere, with most records between 30°S
and 45°S. In summer months it appears near the Antarctic Peninsula and along the shores of the continent, even among
sea ice. There appears to be a hotspot in the area between North Island, New Zealand, and the Chatham Islands. Usually
in waters deeper than 2,000m along and beyond the continental shelf edge. Likely to be more common in areas of
complex seabed topography. A single female stranded in the Netherlands, in 1927, which was undoubtedly extralimital.
BEHAVIOUR Breaching (usually at a shallow angle), spyhopping, flipper-slapping and lobtailing have all been observed.
May porpoise out of the water when swimming fast, making low, arc-shaped leaps.
FOOD AND FEEDING Very little known. Probably feeds on small deepwater squid and fish.
TEETH Upper jaw 34–44; lower jaw 2. Unique among mesoplodonts in that adults of both sexes have very small teeth
(each less than 1cm tall) in the upper jaw. It is rare for the lower teeth to erupt in females.
GROUP SIZE AND STRUCTURE Most sightings have been of singles or pairs; New Zealand strandings records suggest
average group size of 3.4 (for those not alone). However, there have been several mass strandings (including one of 28
animals in the Chatham Islands, in 1874).
Both sexes can be more
orangish or yellowish ADULT FEMALE
Paler overall colour (due to film of diatoms)
Longer, more slender
beak than male
Little or no linear
scarring

Teeth rarely erupt

on lower jaw SIZE
L: ♂ 4.7–5.2m, ♀ 4.5–5.3m;
WT: 0.9–1.1t; MAX: 6m, c. 1.5t
Calf – L: 2.1–2.4m; WT: unknown

DIVE Sequence Often surfaces slowly at angle of 45° showing long white beak; typically does not raise flukes before
sounding dive. • Depth Likely to forage at depths greater than 500m.
BLOW May be low, diffuse blow.

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 Smoothly sloping, slightly Predominantly dark grey or black ADULT MALE
bulbous melon merges with (sometimes with yellowish or
rostrum (no crease) orangish tinge from diatoms) Often has long, wide,
Relatively straight single or paired linear
Beak (and
mouthline Slender, spindle- scarring on body
part of melon)
shaped body
usually white
to light grey Relatively small,
robust head Small, falcate dorsal
(variable) fin two-thirds of
the way along back
(variable shape)

Small, narrow
flippers (edges may Wider tail flukes
Extremely long, have light border) White patch in than in female
relatively slender urogenital region
beak (average 38cm Two shallow, V-shaped
– longest of any throat grooves Sometimes May be extensive
mesoplodont), often paler grey on light, oval scars from
visible underwater Two triangular teeth on lower jaw underside cookiecutter shark bites
just behind midpoint of mouthline
(become heavily worn over time)
ADULT FEMALE
Crescent-shaped blowhole
(ends point forward)

May be more
orangish or yellowish
ADULT FEMALE VARIATION (exaggerated by film
of diatoms)

80°

70°

60°

50° AT A GLANCE Temperate waters of southern

hemisphere • Medium size • Spindle-shaped
40°

30°

20°
10° body • Relatively small head • Long, slender
0°
10°
white to light grey beak • Beak appears at
20°

30°
45° angle above water on surfacing • Small
40°

50°
triangular teeth in middle of each side of
60° lower jaw • Possibly in small groups
70°

80°

extralimital record

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 HECTOR’S BEAKED WHALE
Mesoplodon hectori (Gray, 1871)

Known from only a few dozen strandings and just one confirmed sighting at sea (a juvenile off southwestern Australia),
Hector’s beaked whale is one of the least known of all cetaceans.
IUCN status Data Deficient (2020).
Population Unknown. Information is sparse, but it appears to be rare. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names New Zealand beaked whale, skew-beaked whale (after the huge asymmetry of the skull).
DISTRIBUTION Cool temperate waters of the southern hemisphere, between 32°S and 55°S. There are no records
between New Zealand and the Pacific coast of South America – explained either by a break in distribution or simply a
lack of data. It is presumed to occur in deep waters beyond the edge of the continental shelf.
BEHAVIOUR Known primarily from strandings. In the only confirmed sighting of this species alive at sea, a single
3m-long healthy-looking animal was observed in 1999 in shallow water about 50m from shore in southwestern
Australia – almost certainly atypical habitat for the species – and it breached several times near a research vessel. It
remained in the area for two weeks before disappearing.
FOOD AND FEEDING Virtually unknown. It probably feeds in deep water on squid (possibly also fish and invertebrates).
The only confirmed live sighting recorded dive times of up to 4 minutes, but the maximum is likely much longer.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE No information, but likely to occur in small groups.

ADULT FEMALE
Relatively nondescript
Dark grey colouring
upper jaw Less linear
scarring
No erupted
teeth

Pale grey or whitish

blaze behind eyes ADULT FEMALE
Broad flukes in
relation to body
length

No
median
notch

Tailstock
compressed
Crescent-shaped blowhole laterally
(ends point forward) Almost straight
trailing edge

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 ADULT MALE
May be dark area around
eyes (extending to darker Dark grey-brown
Smoothly face mask in some upperside
sloping, slightly individuals) Small triangular or slightly
bulging melon falcate dorsal fin two-thirds of
Spindle-shaped
Small head with the way along back
body
Light grey to relatively straight
white beak mouthline (does not
and front of rise towards rear) Moderate to heavy single
melon and closely paired linear
scarring (tooth rake marks)

Medium-sized
beak Small, narrow
flippers Slightly lighter Body may be flecked
underside with oval cookiecutter
Two relatively large, Two shallow shark bites
laterally compressed throat
triangular teeth less grooves
than 1cm behind tip of
lower jaw
ADULT MALE VARIATION

SIZE
L: ♂ 4–4.3m, ♀ 4–4.4m;
WT: c. 900kg; MAX: 4.5m, c. 1t
Calf – L: 1.8–2.1m; WT: unknown

80°

70°

AT A GLANCE Temperate waters of southern

60°

50°

40°

30°
hemisphere • Small to medium size • Small
20°
10°
triangular dorsal fin two-thirds of the way
0°
10°
along back • Light grey to white beak and
20° front of melon • Smoothly sloping melon •
Moderate to heavy scarring • Two laterally
30°

40°

compressed triangular teeth at tip of lower

50°

60°

70° jaw
80°

possible range

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 HUBBS’ BEAKED WHALE
Mesoplodon carlhubbsi Moore, 1963

Known from fewer than 60 records in the North Pacific – mostly strandings and fishery entanglements, with just a few
reliable sightings at sea – Hubbs’ beaked whale is very poorly understood. It is strikingly similar to Andrews’ beaked
whale, which lives far away in the cold waters of the Southern Ocean.
IUCN status Data Deficient (2020).
Population Unknown. Paucity of sightings suggests that it may be rare but, like all mesoplodonts, it is inconspicuous
at sea and may simply be missed. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Arch-beaked whale.
DISTRIBUTION Distribution is known mainly from strandings. Endemic to deep offshore cold temperate waters of the
North Pacific. The majority of records are from western North America, largely along the path of the south-flowing,
cold-water California Current. The lack of land masses where strandings could be recorded may account for the paucity
of records in the central North Pacific, so it is possible that the distribution is continuous.
BEHAVIOUR There have been only a few confirmed sightings, so virtually nothing is known about behaviour at sea.
FOOD AND FEEDING Limited evidence suggests mainly deepwater squid, and some deepwater fish. Probably suction-
feeder (sucks prey into mouth and swallows it whole).
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Very little information suggests groups of 1–5.

Mid-grey upperside,
fading to light grey or ADULT FEMALE
No white ‘cap’ (though
may be pale) white underside

Less
pronounced
melon No linear scarring

Mouthline far
less arched
May have light-
coloured or whitish No visible
beak (but contrast with teeth SIZE
head colour not as L: 4.7–5.3m; WT: c. 1–1.5t
great as in male) Calf – L: c. 1.7–2.3m; WT: unknown

DIVE Sequence (speculative) Beak and white cap on head visible (one report of lifting head out of water on
surfacing); probably does not show flukes. • Depth Possibly 500–3,000m. • Duration Unknown, but likely up to
one hour.
BLOW Blow indistinct.

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 Forehead rises at steeper Spindle- ADULT MALE
angle than Andrews’ shaped
beaked whale (no crease body Uniformly
Single flattened tooth between melon and beak) dark grey to
at apex of arch on each black
side of lower jaw White ‘cap’ on moderately
bulbous melon (cf. Scars up to
Andrews’ beaked whale) 2m long
Relatively short,
stubby, bright Small, pointed,
white beak moderately falcate
Relatively dorsal fin two-thirds
small head of the way along back
(22–23cm high)

Prominently
arched Small, narrow May have extensive single
mouthline flippers and closely paired linear
scarring (tooth rake marks)
Tips of teeth exposed (to Two shallow
May be some light
tip of rostrum or higher) V-shaped throat
oval scarring from
when mouth closed grooves
cookiecutter shark bites

ADULT MALE
Flippers fit into
‘flipper pockets’
(slight depressions on Flukes wide
sides of body) relative to body

No
median
notch

Crescent-shaped blowhole
(ends pointing forward)

75°

60° AT A GLANCE Cool temperate waters of

North Pacific • Medium size • Uniformly
45°
dark • Bright white ‘cap’ and beak • Heavily
scarred • Small falcate dorsal fin two-thirds
30° of the way along back • Two tusks on arched
lower jaw
15°

0°

15°

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 BLAINVILLE’S BEAKED WHALE
Mesoplodon densirostris (Blainville, 1817)

Blainville’s beaked whale is the most commonly observed Mesoplodon in tropical waters worldwide (though this is
relative) and is the most widely distributed member of the genus. It has a strongly arched lower jaw, with teeth that
protrude like a pair of horns, and its rostrum is formed of the densest bone of any animal.
IUCN status Least Concern (2020).
Population Unknown, but appears to be relatively common in most tropical seas. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Dense-beaked whale, tropical beaked whale.
DISTRIBUTION Tropical to warm temperate waters in both hemispheres. There have been nearly 400 known strandings
worldwide, and Blainville’s beaked whale is seen fairly frequently in a few key hotspots, including Hawaii, the
Bahamas and the Canary Islands. It is one of the most tropical of the mesoplodonts; higher-latitude records are usually
associated with warm-water currents. Occurs in many enclosed seas, including the Gulf of Mexico, the Caribbean Sea
and the Sea of Japan, but is considered a vagrant in the Mediterranean. Seems to prefer waters of intermediate depth
(500–1,500m in Hawaii and the Bahamas) over continental shelf waters, deep submarine canyons and steeply sloping
regions around seamounts. However, it is also known in much deeper waters (at least 5,000m) in the open ocean and
has been reported in waters as shallow as 320m (the mean depth for seven sightings in the Canary Islands). In the few
areas where it has been studied, it shows a high degree of site fidelity (known individuals have been seen repeatedly in
the same area over one or two decades).
BEHAVIOUR Behaviour better known than for any other species of Mesoplodon. It rarely breaches or performs other
aerial behaviours. Not known to occur in mixed aggregations with other cetaceans. Behaviour around boats varies
enormously.
FOOD AND FEEDING Mainly deepwater squid and fish, some crustaceans, with regional differences. Foraging dives
occur day and night; believed to forage along seabed, at least sometimes; suction-feeder.

Crescent-shaped ADULT MALE

blowhole (ends point
forward)
Flukes wide relative
to body length

No
median
notch

Laterally
Wide ‘cheeks’ compressed
tailstock

DIVE Sequence Briefly lifts beak out of water at angle of c. 45° on surfacing (entire head may clear surface); may
be slight pause as it levels and blows; tailstock appears as it arches slightly and rolls forward to dive (arches higher
on terminal dives); rarely, if ever, shows flukes. • Depth Three types of dive: shallow (2–4m); long, deep, foraging
(regularly more than 1,000m for 1+ hours); and intermediate (30–300m) to avoid detection by predators. Record
in Hawaii of adult female diving to 800m accompanied all the way with calf; maximum recorded 1,599m.
• Duration Typically 20–45 minutes; maximum recorded 83 minutes.
BLOW Inconspicuous blow usually low and canted forward.

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 Fairly nondescript ADULT MALE
Small head with dark greyish-brown
flattened melon or brownish-grey
upperside
Spindle-shaped body
Smoothly sloping, non-
bulbous melon merges
with rostrum (no crease) Extensive single and paired
linear scarring (often quite
Yellowish-orange deep), especially on back
Flattened tooth erupts sheen (diatom between blowhole and
from highest point on film) may cover dorsal fin (caused by male–
arch (tip often extends parts of body male combat)
well above rostrum,
though amount
visible highly
Small, slightly falcate
variable)
or triangular dorsal fin
two-thirds of the way
along back

Moderately
long beak
Two shallow, Small, narrow
V-shaped throat flippers (tuck into
grooves 'flipper pockets' on Often heavily
May have black spots sides of body) pockmarked with
in a line around mouth
healed round or oval
(scars from hooks on
Highly arched lower jaw white scars from
squid tentacles)
(arch very wide and often cookiecutter shark bites
towers above melon)
May have slightly
paler underside
Teeth may be covered by tassels
No of dark purple or reddish-brown
median stalked barnacles (sometimes
otch obscuring teeth altogether)

80°

70°
AT A GLANCE Tropical to warm temperate
60°
waters worldwide • Medium size •
50° Mostly nondescript grey-brown colouring •
Pockmarked with healed cookiecutter shark
40°

30°

bites • Tangled web of mainly parallel linear

20°
10°
0°
10° scarring • Very strongly arched lower jaw
20°

30° • Flattened, forward-tilting teeth on jaw

40°

50°
arches • Stalked barnacles on teeth look like
60° pompoms • Small head with flattened melon
70°
• Small, slightly falcate or triangular dorsal fin
80°
two-thirds of the way along back

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 TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Varies geographically, but typically in groups of 3–7 (mean 4.1); the largest groups
observed in Hawaii and the Bahamas each contained 11 individuals; can also be found in pairs or alone. Groups are
usually harems, with a single adult male accompanying several adult females with their calves and/or juveniles. Sub-
adults appear to stay in separate groups, and tend to occur in less productive waters. Occasional sightings of larger
aggregations, including more than one adult male, are probably temporary aggregations of two or more groups.

Little or no ADULT FEMALE

Teeth not visible linear scarring
(do not erupt)

Arched lower jaw (much Pockmarked with

less extreme than in male) Tends to be paler
healed cookiecutter
on underside
shark bites

Nondescript light grey CALF

countershading (though
Darker patch may have coating of
around eye brown diatoms)

Moderately arched
lower jaw (with age, Relatively small
male develops fully dorsal fin may
formed arches before have dark edges
teeth erupt)

Shorter, stubbier beak

DENSE BEAK
The rostrum of Blainville’s beaked whale becomes
secondarily ossified as individuals mature, especially
in males, forming the densest bone currently known.
SIZE
Three possible functions have been proposed: to act as
L: ♂ 4.3–4.8m, ♀ 4.3–4.7m;
ballast (reducing the energetic cost of deep diving); as an WT: 0.8–1t; MAX: 4.9m, 1.03t
adaptation for transmitting sound, during echolocation; Calf – L: 2–2.5m; WT: c. 60kg
or, most likely, as a mechanical reinforcement to prevent
impact damage to the skull during male–male combat.

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 ADULT TWO-TONED MALE

Ochre to gold colour

patches caused by
diatom infestations ADULT TWO-TONED FEMALE

ADULT MALE VARIATION

ADULT FEMALE VARIATION

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 SOWERBY’S BEAKED WHALE
Mesoplodon bidens (Sowerby, 1804)

Sowerby’s beaked whale was the first Mesoplodon beaked whale to be described: a male stranded in 1800, in the
Moray Firth, northeastern Scotland (UK), and the skull was preserved. A few years later, James Sowerby, an English
watercolour artist and naturalist, painted a picture of it and how he imagined the whole animal might have looked.
IUCN status Least Concern (2020).
Population Unknown, but probably fairly common in parts of its range. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms of subspecies.
Other names North Sea beaked whale, North Atlantic beaked whale.
DISTRIBUTION Deep, cold offshore waters of the northern North Atlantic (among the most northerly species of
Mesoplodon). Appears to be considerably more common in the eastern North Atlantic, and the centre of abundance
appears to be northern Europe. The vast majority of strandings have been between 50°N and 60°N. Occurs mainly in
deep waters beyond the continental shelf edge and often associated with areas of complex seabed topography. May be
seen close to shore where deep water approaches the coast.
BEHAVIOUR Small groups typically surface within a couple of body lengths from each other. Breaching, spyhopping and
tail-slapping have been observed.
FOOD AND FEEDING Unusual among beaked whales in taking mainly small mid- and deepwater fish; some squid.
Feeding techniques unknown.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only; both sexes also possess small vestigial teeth, which do not
normally erupt.
GROUP SIZE AND STRUCTURE Very little information, but seems to be 3–10 (8–10 have been recorded on a number
of occasions).

ADULT MALE
Crescent-shaped
blowhole (ends Flippers fit into
point forward) ‘flipper pockets’ Flukes wide relative
to body length

No
median
notch

Tailstock
compressed
laterally

DIVE Sequence Usually surfaces at 30–45° angle, clearly showing beak; melon and much of head may also be
visible; swimming behaviour often described as ‘calm and unhurried’. • Depth Main prey typically occurs at 400–
750m. • Duration Typically 12–28 minutes, but probably capable of c. 1 hour.
BLOW Invisible or inconspicuous (small and diffuse) blow angled slightly forward.

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 Beak can be ADULT MALE
dark grey or Distinctive bulge Linear scarring on many
very light grey on forehead (in individuals (variable – less
front of blowhole) extensive than many other
Long, relatively mesoplodonts – more with age)
slender beak Dark bluish-grey to lighter
(length variable) Relatively slate-grey upperside (may have
small head Fairly streamlined, brownish tinge due to diatoms)
spindle-shaped body
Melon slopes
gently onto May be dark Small, falcate dorsal fin
rostrum (no around eyes two-thirds of the way
crease) along back (variable shape)

Small, narrow
flippers Slightly paler sides May have white or light
Two shallow, V-shaped and underside grey spots (probably
throat grooves cookiecutter shark bites)
Two small teeth
two-thirds of the Mouthline mostly straight
way along lower with slight arch at rear
jaw (visible outside
closed mouth)
Stalked barnacles may
attach to teeth SIZE
Teeth do not rise above level of L: ♂ 4.5–5.5m, ♀ 4.4–5.1m;
upper rostrum, so rake marks WT: 1–1.3t; MAX: 5.5m, 1.5t
mostly single (not paired) Calf – L: 2.1–2.4m; WT: 170–185kg

ADULT FEMALE
No erupted teeth

Little or no
linear scarring

80°

70°

60°

50°

40°
AT A GLANCE Cool waters of North Atlantic
30°

20°
• Nondescript light to dark grey above, lighter
10°
0°
below • White linear scars may be present •
10°
20°
Medium size • Long, slender beak visible on
30°
surfacing • Two teeth two-thirds of the way
along beak • Distinctive bulge on forehead •
40°

50°

60°
Usually unobtrusive/elusive behaviour
70°

80°

Mediterranean records extralimital records

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 TRUE’S BEAKED WHALE
Mesoplodon mirus True, 1913

The taxonomic status of the two disjunct populations of True’s beaked whale – widely separated by the tropics – has long
been a puzzle. However, recent evidence suggests that they may, indeed, represent two different species: True’s beaked
whale in the North Atlantic and the newly proposed Ramari’s beaked whale in the southern hemisphere.
IUCN status Least Concern (2020); assessed as a single species.
Population Unknown, but sightings have become more regular, at least in the North Atlantic. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy See ‘Anti-tropical Species’ box opposite.
Other names Wonderful beaked whale.
DISTRIBUTION Prefers deep, principally warm temperate offshore waters and may favour areas of complex seabed
topography. Rarely occurs between 30°N and 30°S. The apparent gap in distribution between New Zealand and South
America is likely to reflect a lack of survey effort and possible stranding sites more than its absence.
BEHAVIOUR Energetic breaching has been recorded on several occasions. Tail-slapping has also been observed.
FOOD AND FEEDING Probably mostly deepwater squid; some fish. Feeding techniques unknown.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in males only.
GROUP SIZE AND STRUCTURE Typically 1–4 (based on a few rare sightings).

ADULT MALE SOUTHERN HEMISPHERE (POSSIBLY RAMARI’S)

Bluish-grey to dark Dorsal fin usually light Whitish or light grey

Upper jaw grey upperside but variable (all dark, all tailstock distinctly lighter
dark light or in between)

Upperside of flukes
generally dark

Whitish or light
grey underside
and lower jaw Dark streaks radiate out from centre of
trailing edge on underside of flukes

ADULT FEMALE SOUTHERN HEMISPHERE (POSSIBLY RAMARI’S)

DIVE Sequence Surfaces at angle, possibly showing entire beak and head (to just below eye level). • Depth Likely
to forage deeper than 500m (one study suggests main prey found in waters 200–800m deep). • Duration Unknown.

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 Rounded melon Relatively small head ADULT MALE NORTH ATLANTIC (TRUE’S)
slopes steeply Slender, spindle-
onto rostrum May be distinct indentation shaped body Medium grey to brownish-grey
(no crease) behind blowhole upperside and tailstock (posterior
region of tailstock may be darker)
May be pale or whitish blaze covering
melon from behind blowhole Dorsal fin may be darker than back
Two small
teeth at tip May be narrow, dark line Small, moderately falcate to
of lower jaw between eye and top of head triangular dorsal fin two-
visible when thirds of the way along back
Dark patch around
mouth closed eye (variable) Some individuals may
(do not erupt have lighter tailstock
in female) (not unlike southern
hemisphere animals)

Front half
of beak
dark Small, narrow
flippers May be white or light
Mid-length, pink urogenital patch May be closely spaced
stubby, dolphin- Fairly straight or slightly in some animals
curved mouthline parallel scars on some
like beak Lighter grey to slate- mature males – and
Two well-defined grey underside cookiecutter shark bites
May be dark flecking on V-shaped throat grooves
throat and lower jaw
ADULT FEMALE NORTH ATLANTIC (TRUE’S)
(more in older animals)
Some individuals may have
lighter tailstock (not unlike
southern hemisphere animals)

ANTI-TROPICAL SPECIES
Recent genetic and morphological analysis of museum
and archival specimens of True’s beaked whales from the
northern and southern hemispheres reveal very different
genetics and skull shape (it is still unclear if they are
also distinguishable by coloration). There is believed to
have been no gene flow between the populations for at
least 0.35 million years. If the proposed split is formally
accepted, the northern hemisphere animals will remain
True’s beaked whales (Mesoplodon mirus) and the
southern hemisphere animals will become Ramari’s
beaked whales (Mesoplodon eueu). The common name
honours Māori whale expert Ramari Stewart.

80°

70°

60°
SIZE
50°

40°
L: ♂ 4.8–5.3m, ♀ 4.8–5.4m;
30° WT: 1–1.4t; MAX: 5.4m, 1.4t
Calf – L: 2–2.5m; WT: unknown
20°
10°
0°
10°
Females may be slightly larger than males.
20°

AT A GLANCE North Atlantic and southern

30°

40°

50°
hemisphere • Temperate offshore waters •
Medium size • Rounded melon • Mid-length
60°

70°

80°
beak with two small teeth at tip • Closely
True’s beaked whale strandings outside known range spaced parallel scars • Small dorsal fin two-
Ramari’s beaked whale possible Ramari’s range thirds of the way along back

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 STEJNEGER’S BEAKED WHALE
Mesoplodon stejnegeri True, 1885

Known mostly from strandings – predominantly along the west coast of Honshu, Japan, and in the Aleutian Islands,
Alaska (the vast majority in the western Aleutian Islands – at least 74 individuals in 42 separate events by 2020) –
Stejneger’s beaked whale is rarely seen alive at sea. Sometimes called the sabre-toothed beaked whale, the male has
two particularly large teeth, like tusks, that are used for fighting.
IUCN status Near Threatened (2020).
Population Unknown, but given the paucity of records it appears to be rare in most parts of its range. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Bering Sea beaked whale, North Pacific beaked whale, sabre-toothed beaked whale.
DISTRIBUTION Primarily cold temperate and sub-Arctic waters of the North Pacific. Seems to prefer deep areas of
complex seabed topography. Most live sightings have been in the deep waters of the Aleutian Trench, Alaska; however,
it is also known over the shallow northern Bering Sea shelf (30–70m).
BEHAVIOUR Very little information. Known to breach. Appears to be shy and difficult to approach. Reports of roaring
and groaning sounds made at the surface.
FOOD AND FEEDING Mostly deepwater squid, some fish. Suction is main feeding method.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Typically 2–4 but ranges from 1–15. Groups may contain animals of mixed sexes and
ages, or can be segregated. May be tightly bunched together at the surface – sometimes touching or nearly touching –
and typically swim and dive in unison.
May be More obviously ADULT FEMALE
considerably lighter counter-shaded
Teeth do Little or no linear
not erupt scarring

Lighter lower jaw Flippers may

and throat Creamy-white
be dark underside (may be May be dark margin
along trailing edge FLUKES (underside)
pinkish when active)

Underside of flukes may have

SIZE
L: ♂ 4.6–5.7m, ♀ 4.8–5.4m;
WT: 1–1.3t; MAX: 5.7m, 1.6t
Calf – L: 2.1–2.3m; WT: c. 80kg
starburst of light grey or white
concentric lines radiating from
midpoint of trailing edge (tends to be Starburst
more exaggerated in Stejneger’s than intensifies
other mesoplodonts) with age

DIVE • Sequence Tip of beak breaks surface first; blowhole and upperside of head appear briefly; low profile as head
quickly disappears and dorsal fin rolls forward. • Depth Preferred prey suggests at least 200m; presumably capable
of diving much deeper (possibly to 1,500m). • Duration At least 15 minutes, probably much longer.
BLOW Blow indistinct.

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 Dark cranial cap ADULT MALE
Smoothly sloping,
(extending back roughly
non-bulbous melon Uniformly dark bluish-grey to nearly black
to blowhole) extends
merges with (can appear brownish due to diatoms)
downward to surround
rostrum (no crease)
eyes (gives overall Prominent (mostly
appearance of 'hood' or Overall coloration may darken with age paired) linear scarring
'helmet') (making dark cap and flipper pockets (especially on back)
Small head
less clearly demarcated with time)
Two large,
laterally
compressed, Variable pale Spindle-shaped Relatively small, nearly
triangular teeth ‘collar’ behind body triangular to slightly falcate
on leading cranial cap dorsal fin two-thirds of the
edge of arch way along back
near middle
of lower jaw

Medium-
length beak Flipper pockets Underside (including May be
with strongly considerably darker lower jaw and throat) distinct keel
arched than surrounding may be slightly paler on underside
mouthline area (may look like of tailstock
flipper shadows) Extensive round or oval white
Teeth may project scarring from cookiecutter shark Cookiecutter shark scars
higher than top Small, narrow
bites (and sometimes lampreys), may be absent in Sea of
of rostrum when flippers
especially on rear half of body and Japan animals
mouth closed Two shallow V-shaped underside (more in older animals)
Broken teeth may have throat grooves ADULT MALE
stalked barnacles attached
Flukes wide
Crescent-shaped blowhole relative to body
(ends point forward) length
No
median
notch

Tailstock
Teeth point forward and Flippers fit into darkly pigmented compressed
slightly inward (may ‘flipper pockets’ (slight laterally
constrict opening of jaw) depressions on sides of body)
75°

60°

AT A GLANCE Cold offshore waters of

northern North Pacific • Medium size •
45°
Spindle-shaped body • Dark cranial ‘cap’ •
Gently-sloping forehead • Strongly arched
mouthline • Two large, exposed, flattened
30°

15° teeth • Small groups bunched together

0°

15°

2013 stranding of a female in good condition suggests range

may extend further north

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 GERVAIS’ BEAKED WHALE
Mesoplodon europaeus (Gervais, 1855)

Gervais’ beaked whale is known from more than 300 records in the North Atlantic, and just a handful in the South
Atlantic. Most of these are strandings – there have been few reliable sightings at sea – so information on its life and
habits is sparse.
IUCN status Least Concern (2020).
Population Unknown, but believed to be relatively common. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names European beaked whale, Gulf Stream beaked whale, Antillean beaked whale.
DISTRIBUTION Most records are from the western North Atlantic – Florida and North Carolina account for more than
40 per cent of all worldwide records combined – and it is the most commonly sighted mesoplodont off the US Atlantic
coast and in the Gulf of Mexico. Nearly half of all records in the eastern North Atlantic (21 strandings involving 24
individuals) are from the Canary Islands. There are scattered records in the South Atlantic. Seems to prefer deep waters
in the tropics and subtropics, but there are records from warm temperate and even cold temperate waters. Likely to be
more common in areas of complex seabed topography.
BEHAVIOUR Virtually nothing known.
FOOD AND FEEDING Little known. Feeds mainly on deepwater squid, possibly some deepwater fish.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Limited information suggests that Gervais’ beaked whales are usually found alone or in
small, close-knit groups of up to five individuals.

ADULT FEMALE
May have dark band in Often has distinctive dark band down
front of dark eye patch Small pale areas centre of back with a series of dark,
extending upwards and behind and in front of wavy, vertical stripes extending short
backwards to behind dark eye-patch distance down sides (variable)
blowhole (variable)

Paler than male

No erupted
teeth

May be white patch

in genital area

SIZE
L: ♂ 4.2–4.6m, ♀ 4.2–4.8m;
WT: 0.8–1t; MAX: 5.2m, 1.2t
Calf – L: c. 1.7–2.2m; WT: c. 80kg

DIVE Sequence May surface at angle of about 45°; briefly shows beak and much of head; slight pause before rolls
forward; tends to sink below surface rather than arching back; does not show flukes on sounding dive.
BLOW Inconspicuous blow.

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 Single tooth on small arch about one-third of ADULT MALE
the way along each side of lower jaw (7–10cm Dark band down centre of back
from tip), visible outside closed mouth may be partially obscured by
darker upperside (often more
Dark patch around eye often conspicuous in female and calf)
more pronounced than in other Relatively little pale linear
mesoplodonts (variable) scarring (usually single
Smoothly sloping, Spindle- lines when present)
slightly bulbous melon Very small shaped
merges with rostrum head body
(no crease) Small, wide-based, slightly
falcate dorsal fin two-thirds of
the way along back (variable –
shark-like to falcate)

Mid-length beak Medium to dark grey (sometimes

with relatively Two shallow, Small, narrow
brownish) upperside, paler
straight mouthline V-shaped throat flippers (darker than
underside (may become darker
(slightly raised area grooves underside of body)
with age)
around tusks)
ADULT MALE
Crescent-shaped blowhole Flukes wide relative to
(ends point forward) body length (up to 33
per cent of body length)

No
median
notch

Tailstock compressed
Distinctive dark band down centre laterally
ADULT FEMALE of back often has series of dark,
wavy, vertical stripes extending short Dark bands not present in
distance down sides (variable) all Gervais' beaked whales
– but are unique

50°

AT A GLANCE Tropical to warm temperate

40°
waters of Atlantic Ocean • Medium size •
30° Medium to dark grey above, paler below •
20°
Little or no linear scarring • Very small head
with slightly bulbous melon • Dark patch
around eye • Medium-length beak • Two
10°

0°
teeth on small arch one-third of the way along
10°
lower jaw (male only) • Small dorsal fin two-
20° thirds of the way along back • Females and
juveniles may have tiger-like stripes
primary known range strandings and sightings locations
likely range (outside primary known range)

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 ANDREWS’ BEAKED WHALE
Mesoplodon bowdoini Andrews, 1908

Andrews’ beaked whale is one of the least known of all the world’s cetaceans – there has never been a confirmed
sighting at sea and our limited knowledge comes from just over 50 strandings, all in the cooler waters of the southern
hemisphere. It is strikingly similar to Hubbs’ beaked whale, found in the North Pacific, but recent genetic and
morphological studies confirm their specific distinctiveness.
IUCN status Data Deficient (2020).
Population Unknown, but the frequency of strandings around New Zealand and southern Australia, at least, suggests
that it may not be all that rare. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Splay-toothed (or splaytooth) beaked whale, deep-crested beaked whale.
DISTRIBUTION Known only from about 50 strandings in cool temperate to sub-polar waters between 32°S and
approximately the Antarctic Convergence. Most stranding records have come from New Zealand and its surrounding
islands and the southern coasts of Australia. The overall range may be circumpolar in the southern hemisphere,
although there is a large gap in the records between the Chatham Islands and the west coast of South America (which
could represent a break in distribution or, more likely, reflect a general shortage of cetacean records from this part of
the world). Presumably prefers deep offshore waters.
BEHAVIOUR No information.
FOOD AND FEEDING Limited evidence suggests mainly deepwater squid, some deepwater fish. Probably suction feeder.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Very little information but possibly groups of 1–5.

Uniformly slate-grey to
ADULT FEMALE
greyish-brown fading to
Light patch in
paler sides and underside
front of eye
No visible
Little or no
teeth
linear scarring

White lower jaw, dark

upper jaw (variable – Mouthline far Flippers darker than
can be white-tipped) less arched surroundings

OLD ADULT MALE FLUKES (UNDERSIDE) SUB-ADULT MALE FLUKES (UNDERSIDE)

DIVE Depth Possibly 500–3,000m • Duration Unknown, but likely up to 1 hour.

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 Forehead rises at No white ‘cap’ (cf. ADULT MALE
Uniformly dark May have extensive single
shallower angle Hubbs’ beaked whale)
bluish-grey, blue- and closely paired linear
than Hubbs’ (no
Relatively black or dark brown scarring (tooth rake marks)
crease between
melon and beak) small head
Spindle-shaped Small, blunt-tipped,
May be small pale body triangular to moderately
Tips of teeth
exposed (to tip patch in front of falcate dorsal fin two-thirds
of rostrum or eyes (variable) of the way along back
higher) when (variable shape)
mouth closed
May have pale
grey saddle

Two shallow V-shaped

throat grooves Small, narrow
Relatively flippers
Prominently arched
short, stubby May be extensive light
mouthline
beak (anterior oval scarring from
half white, to Single flattened tooth at apex of cookiecutter shark bites
just past teeth – arch on each side of lower jaw
white may extend
Stalked barnacle may
further back on
attach to teeth
lower jaw) ADULT MALE
Flippers fit into ‘flipper
Crescent-shaped blowhole Flukes wide relative
pockets’ (slight depressions
(ends point forward) to body length
on sides of body)

Tailstock
SIZE compressed
laterally No median notch
L: 3.9–4.4m; (but may be slight
WT: possibly c. 1–1.5t
prominence)
Calf – L: c. 2.2m; WT: unknown

80°

70°

60°

AT A GLANCE Cooler waters of southern

50°

40°

30°

20°
hemisphere • Small to medium size •
10°
0°
Uniformly dark • Relatively short, heavy,
10°
20°
white-tipped beak • Heavily scarred • Small,
30°

40°
falcate fin two-thirds of the way along back
50° • Two tusks on arched lower jaw • No white
60°
‘cap’
70°

80°

possible range (no strandings or sightings records)

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 STRAP-TOOTHED BEAKED WHALE
Mesoplodon layardii (Gray, 1865)

The largest of the Mesoplodon whales, the strap-toothed beaked whale is well known from strandings but rarely seen
at sea (though the unique colour pattern and dentition of the male makes it one of the most readily identifiable of all the
beaked whales). The adult male has perhaps the most bizarre teeth in the animal kingdom (with the possible exception
of the narwhal).
IUCN status Least Concern (2020).
Population Unknown. With 200+ documented standings, it is considered uncommon but not rare. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Strap-toothed whale, straptooth beaked whale, Layard’s beaked whale, long-toothed beaked whale.
DISTRIBUTION Believed to have a continuous distribution in deep, cold temperate waters of the southern hemisphere,
mostly between 35°S and 63°S. More than 200 strandings are known worldwide, with approximately half in Australia
and New Zealand. Most sightings of live animals at sea have been in Australia and New Zealand, too, in water deeper
than 2,000m and beyond the continental shelf. Believed to be more common in areas of complex seabed topography.
BEHAVIOUR Several observations of breaching. Difficult to approach: in response to boats and ships, seen sinking
slowly below the surface, or diving sideways (exposing a single flipper as it disappears).
FOOD AND FEEDING Mainly deepwater squid; may also take some fish and crustaceans. Presumably suction feeder;
teeth may act as ‘guard-rails’ to keep food on direct path into mouth.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Few confirmed sightings, but in most cases it has been observed singly, in female–calf
pairs, or in groups of 2–5 (up to 10).
Little or no
No erupted teeth
ADULT FEMALE
linear scarring

Crescent-shaped blowhole ADULT MALE

(ends point forward, cf. Light grey to white
Arnoux’s and Baird’s) outer margin

No
median
notch

Relatively wide,
Teeth may cross over
roughly triangular-
top of upper jaw
shaped flukes

DIVE Sequence Surfaces at angle of 45°; head and beak often lifted right out of water (adult male showing teeth);
does not show flukes. • Depth Forages at depths greater than 500m; vampire squid (found in stomachs of stranded
whales) mainly at 700–1,500m. • Duration Unknown.
BLOW Indistinct blow.

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 May have patches Upperside mostly light ADULT MALE
Steeply rising of yellow-orange grey roughly between
forehead diatoms blowhole and dorsal fin Complex contrasting light grey and black
(variable extent) coloration (highly variable and may have
Two long Black melon extends bronze tinge due to diatom film)
curved, to black eye-mask Spindle-shaped
flattened body Small, falcate dorsal fin two-
white teeth Relatively small head thirds of the way along back
(up to 34cm) with moderately
May have linear
near middle bulbous melon
scarring but superficial
of lower jaw
flesh wounds may heal
same colour as skin
(nearly invisible)

Small, pointed, light

Long, grey spot or ‘arrow’ Tailstock
slender beak Light grey to just behind flipper predominantly
predominantly white throat black
Light grey to
white (black at Two shallow Small, narrow, white patch May be some healed
base) throat grooves black flippers around urogenital (nearly invisible)
Mouthline mostly straight area (with two cookiecutter shark bites
(slight, smoothly rising forward-pointing
arch towards rear) extensions on
either side)

ADULT MALE

Exposed portion of teeth may

host large stalked barnacles
Tusks emerge from and can be covered in
middle of lower jaw, curl greenish-brown diatoms
backward around 45°
and inward, crossing over
top of upper jaw (often
preventing it from opening
more than 3–4cm – cf.
6.5cm in adult female)
SIZE
L: ♂ 5–5.9m, ♀ 5–6.1m;
WT: 1.3–2.7t; MAX: 6.2m, 2.8t
No Calf – L: c. 2.2–3m; WT: unknown
median
notch
80°

70°

60°

50°

40°
AT A GLANCE Cold temperate waters of
30° southern hemisphere • Medium size •
Complex contrasting light grey and black
20°
10°

coloration • Black melon and face ‘mask’

0°
10°

• Predominantly white beak • Faint blow,

20°

30°

angled forward • Strap-like teeth that can

40°

50°

60°
curve over upper jaw in male
70°

80°

extralimital records

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 SPADE-TOOTHED WHALE
Mesoplodon traversii (Gray, 1874)

The spade-toothed whale is one of the least known of all the world’s living cetaceans – and of all large mammals. The
only proof of its existence is from two strandings (a mother and calf pair, and one adult male), two weathered skulls and a
single jawbone with teeth. There has never been a confirmed sighting at sea.
IUCN status Data Deficient (2020).
Population Unknown. Possibly rare, but could be evading observation in remote areas far from land. Trend unknown.
Classification Odontoceti, family Ziphiidae.
Taxonomy No recognised forms or subspecies.
Other names Spade-toothed beaked whale, Bahamonde’s beaked whale.
DISTRIBUTION Very little information, since there is only a handful of records. However, it is possible that the spade-
toothed whale has simply gone unnoticed elsewhere. On 31 December 2010, a mother and male calf stranded on
Opape Beach (38°S), North Island, New Zealand, and subsequently died. On 23 December 2017, an adult male was
seen swimming in Waipiro Bay (38°S), North Island, New Zealand, before stranding and dying soon afterwards. These
three animals have provided all the information available on the likely appearance of this species. Apart from them,
there are just three specimens: a mandible with teeth from an adult male found on Pitt Island (44°S), in the Chatham
Islands, in 1872 (the type specimen); a skull without a mandible found on White Island (37°S), North Island, in the
1950s; and another skull without a mandible found on Robinson Crusoe Island, in the Juan Fernández Islands, off Chile
(33°S), in 1986.
BEHAVIOUR With no confirmed sightings of healthy animals at sea, nothing is known about its behaviour.
FOOD AND FEEDING Very little known. Like other beaked whales, it is presumed to eat mainly deepwater squid, and
some fish.
TEETH Upper jaw 0; lower jaw 2. Teeth erupt in male only.
GROUP SIZE AND STRUCTURE Nothing known.

ADULT FEMALE
Simpler coloration than male
Dark patch (simple counter-shading)
around eye

SIZE
L: ♂ 5.2m (based on single adult male),
♀ 5.3m (based on single adult female);
WT: unknown
Calf – L: unknown; only known juvenile
was 3.5m

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 Pronounced melon (more ADULT MALE
prominent than in Gray’s beaked
whale, most similar to strap- Spindle-shaped body
toothed beaked whale)
Dark grey or
black upperside
Upper ‘lip’ in Relatively
front of tooth small head Dark eye-patch
pale grey (joins dark
upperside)
Small, falcate
Long, dorsal fin two-
slender thirds of the way
beak along back

Beak dark Small, dark,

Two shallow,
grey or narrow flippers Paler underside (may be
V-shaped throat
black grooves ‘Cheeks’ mottled darker patch over belly)
medium grey
Two large, wide-based teeth
in middle of each side of
lower jaw (large enough to
be clearly visible)
Large denticle at tip of
ADULT MALE TEETH tooth (may be worn down
in older animals)

Teeth taller (c. 23–24cm)

than those of strap-toothed
beaked whale (based on
Teeth wider than
two teeth from single
those of strap-toothed
specimen)
beaked whale

80°

70°

60°

AT A GLANCE Temperate (and possibly sub-

50°

40°

30°

20°
tropical) waters of South Pacific (and possibly
10°
0°
other oceans) • Medium size • Long, slender
10°
20°
beak • Pronounced melon • Male has two
30°
large, backward-leaning teeth in middle of
lower jaw • Small, falcate dorsal fin two-
40°

50°

60°
thirds of the way along back
70°

80°

probable range possible range strandings/specimens

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 KILLER WHALE or ORCA
Orcinus orca (Linnaeus, 1758)

Two thousand years ago, Roman scholar Pliny the Elder described the killer whale as ‘an enormous mass of flesh
armed with savage teeth’. Even as recently as the early 1970s, US Navy diving manuals described it as ‘extremely
ferocious’, warning that it ‘will attack human beings at every opportunity’. But the killer whale does not deserve its
killer reputation any more than any other apex predator.
IUCN status Data Deficient (2017). Strait of Gibraltar sub-population Critically Endangered (2019).
Population No reliable global estimate, but likely minimum 50,000 (probably considerably more, especially given the
lack of information from large oceanic areas and the Arctic, and likely underestimates in the Antarctic). The latest
estimate for the Antarctic south of 60°S is c. 25,000–27,000 (some estimates are three times higher, on the basis
that most animals are within the pack ice, where they are difficult to find). There are estimated to be c. 22,100 in the
central and eastern North Atlantic (2020). Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy Several possible contenders for separate subspecies or even species status (Bigg’s and Ross Sea are,
perhaps, the most obvious). In the meantime, the term ‘ecotype’ is used for ecologically distinct populations that
do not interbreed (even if they inhabit the same waters) while recognising scientific uncertainty about killer whale
taxonomy. There could be as many as 40 ecotypes altogether; as well as those illustrated in this guide, they might
include New Zealand coastal, New Zealand pelagic, Eastern Tropical Pacific and ecotypes in Argentina, Papua New
Guinea and other locations around the world.
Other names Blackfish (non-taxonomic group of six dark-coloured members of Delphinidae with ‘whale’ in their
name); historically, grampus.
Moulting skin may ADULT MALE RESIDENT
appear as grey mottling
Variable Upperside of
saddle patch flukes black
(underside white)

Distinct
notch in
middle

Broad flukes
Straight or slightly
convex trailing edge

DIVE Sequence Outline of adult male’s tall dorsal fin unmistakable; tip of dorsal
fin typically breaks surface first (followed by top of head). • Depth Varies with
prey and location; maximum recorded more than 1,000m (near South Georgia),
but potentially even deeper (especially males); foraging residents usually dive
less than 100m. • Duration Varies with prey and location; maximum recorded
16 minutes.
BLOW Fairly tall, columnar blow (up to 5m), bushy at the top and projects
slightly forward.

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 Robust, spindle- Exceptionally tall, erect ADULT MALE RESIDENT
Conspicuous elliptical
shaped body dorsal fin (up to 1.8m)
white patch above Saddle patch often ‘open’
and behind each eye Fin triangular (varies (i.e. with dark incursion)
widely in size and shape)
Dorsal fin may Light to dark grey area of
Predominantly jet- variable shape behind and
black body colour slant slightly
forward in some below dorsal fin (saddle patch)
(often two-tone
Huge conical ecotypes
grey in Antarctic) Saddle patch may be laterally
head with
poorly asymmetrical in some individuals
defined beak
Saddle patch sometimes
non-existent (especially
in tropical populations)

Sharp
White lobe extends demarcation Fluke tips may be
White lower jaw, up sides behind between black curled downward in
throat and underside dorsal fin and white areas some mature males
(cf. female)
Disproportionately large, Dorsal fin, flippers and tail flukes all Scarring usually made by killer whale
oval flippers (grow with substantially larger in proportion to tooth marks during bouts of play (though
age – up to 2m) body size (cf. female) sometimes more serious interactions)

FLUKES

SIZE
L: ♂ 5.6–9m, ♀ 4.5–7.7m;
WT: 1.3–6.6t; MAX: 9.8m, 10t
Calf – L: 2–2.8m; WT: 160–200kg
Body size varies considerably among ecotypes. Highly
sexually dimorphic – mature males up to 17 per cent
longer and 40 per cent heavier than mature females.

80°

70°

60°

AT A GLANCE Worldwide distribution

50°

40°

30°

20°
• Medium size • Two-tone colouring,
10°
0°
predominantly jet black (or grey) and white
10°
20°
• Exceptionally tall dorsal fin of male •
30°

40°
Pronounced sexual dimorphism • White patch
50° above and behind each eye • Usually in family
60°
groups
70°

80°

primary range secondary range particular areas of concentration

KILLER WHALE 155

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 DISTRIBUTION The most cosmopolitan cetacean, with worldwide (though patchy) distribution. Occurs in all oceans
and many enclosed seas. Found in all temperatures and depths, from tropical to polar waters and from the surf zone to
the open sea, though the highest densities are in cold temperate to polar coastal waters with high productivity. Most
abundant in the Southern Ocean south of 60°S. Commonly enters heavy, consolidated ice in the Antarctic, but rarely
does so in the Arctic (though it is becoming seasonally more abundant further north as pack-ice extent and duration
decline with climate change). Widespread but rarer in tropical and offshore waters.
BEHAVIOUR Can be very active at the surface, especially when socialising or after a successful hunt. Often breaches,
flipper-slaps and lobtails. Frequently spyhops; several animals may do so together. Occasionally bow-rides or (more
frequently) wake-rides. Northern residents beach-rub on smooth pebbles in shallow water.
FOOD AND FEEDING Apex predator with extremely diverse diet (c. 150 prey species known globally) but high level
of specialisation depending on ecotype and location. Includes 31 species of cetaceans, 19 species of pinnipeds, 44
species of bony fishes, 22 species of sharks and rays (including great white), 20 species of seabirds, five species of
squid and octopus, two species of sea turtle and two species of terrestrial mammals (sitka black-tailed deer and
moose, swimming between islands). Huge range of feeding strategies, including ‘beaching’ in Punta Norte, Argentina,
and Crozet Island, Indian Ocean, to catch seals on shore; ‘wave-washing’ in Antarctic, to wash Weddell seals from ice
floes; ‘endurance-exhaustion’ in Strait of Gibraltar, to catch bluefin tuna; ‘carousel feeding’ in Iceland and Norway, to
catch schooling herring. Often hunts cooperatively. Will take fish from longline fishing operations.
TEETH Upper jaw 20–28; lower jaw 20–28. Teeth may be worn flat in some ecotypes.
GROUP SIZE AND STRUCTURE Fewer than 20 typical for most ecotypes. Occasionally gather in larger, temporary
groups of up to 150 individuals, which are probably to reinforce social bonds between matrilines and pods, or for
mating. Individuals seen alone are almost always males.

ADULT MALE RESIDENT

Pointed or slightly ADULT FEMALE AND

rounded tip NEWBORN CALF RESIDENT

Smaller, falcate dorsal Often with ‘open’

fin (up to 90cm) saddle patch
Fluke tips rarely
curled down
(cf. male)

Falcate dorsal fin

Large flipper (though May have in both sexes Saddle patch indistinct
proportionately faint cape or absent (becomes
smaller than male’s)
consistently apparent at
two years – once formed,
More muted does not change)
colour pattern

Head slightly more

conical than adult White areas typically have rusty-
orange hue (until 6–12 months old)

156 BLACKFISH

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 KNOWN KILLER WHALE ECOTYPES IN THE NORTH PACIFIC
MALE DORSAL FIN AND SADDLE PATCH COMPARISONS
Dorsal fin continuously
Open saddle patch common Dorsal fin has Saddle patch often rounded over tip (without
(either uniform grey or contains pointed tip larger than in residents sharp angle at rear corner
varying amounts of black) and offshores seen in residents)

Saddle patch
Dorsal fin has never open Saddle patch usually quite
rounded tip (but faint and roughly same
usually with sharp size as in residents
angle at rear corner)
Saddle patch
open or closed

RESIDENT BIGG’S OFFSHORE

RESIDENT (OR FISH-EATING) KILLER WHALE

• ‘Typical’ black-and-white killer whale • Leading edge of dorsal fin tends to be straight or
• Male dorsal fin tip usually more rounded than Bigg’s slightly concave
and ends in pointed trailing tip • Saddle patch usually very open (considerable black
• Typically a few nicks and scars on trailing edge of the incursion in otherwise pale grey) and rarely closed
dorsal fin • Middle of saddle patch rarely extends further forward
• Male dorsal fin may lean forward to varying degrees than midpoint of dorsal fin base
• Fin tip tends to be positioned over front end of base • No obvious dorsal cape
• Dorsal fin often has wavy trailing edge (especially in • White eye patch a medium-sized oval (parallel to
older males) body axis)
L: ♂ 6.9m, ♀ 6m; Max: 7.2m
Distribution Known mainly from the north-east Pacific, ranging from the Aleutian Islands, Alaska (USA), through British
Columbia (Canada) and Washington to Monterey Bay, California (USA). May also occur elsewhere in the North Pacific: a
large population of fish-eating killer whales in the Russian Far East, for example, corresponds in appearance, behaviour,
acoustic activity and genetics with this ecotype (its range includes the central Sea of Okhotsk, southern and central
Kamchatka Peninsula, the Commander and Kuril Islands, and the southern Bering Sea); it is known as Type R. Often
inhabits sheltered coastal waterways and not known to venture far beyond the continental shelf.
Food and feeding Primarily fish. Usually ignores marine mammals, which rarely show avoidance behaviour.
Group size Pod typically composed of three matrilines (ranges from 1–11) with average of 18 whales (typically 10–25,
but ranging from 2–49).
Remarks The term ‘resident’ is rather misleading when describing the site-fidelity and movement patterns of these
whales – so they are often referred to as fish-eating killer whales.

Falcate, female-like dorsal fin in

JUVENILE RESIDENT both sexes (difficult to tell apart)
Dorsal fin starts to grow quickly
in male when c. 15 years old

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 BIGG’S (OR TRANSIENT) KILLER WHALE
• ‘Typical’ black-and-white killer whale • Saddle patch always closed (no black incursion into
• Largest of three North Pacific ecotypes grey)
• Male dorsal fin tip straighter and more pointed than • Middle of saddle patch typically extends further
forward (cf. residents) – often past midline of
resident
dorsal fin
• Typically many nicks and scars on trailing edge of fin • White eye-patch medium-sized oval (slanted very
• Fin tip tends to be positioned over centre of base slightly downwards towards rear)
• Large saddle patch uniformly grey • No obvious dorsal cape
L: ♂ 8m, ♀ 7m; Max: 9.8m
Distribution Known mainly from the Bering Sea through British Columbia (Canada) and Washington (USA) to Baja
California (Mexico). May also occur elsewhere in the North Pacific: a very poorly known population of mammal-eating
killer whales in the Russian Far East, known as Type T, corresponds in appearance, behaviour and acoustic activity with
this ecotype. Movement patterns coincide with seasonally available prey species. Coastal and offshore waters. Travels
over a wider range than residents and rarely keeps to predictable routes or stays in the same place for long.
Food and feeding Primarily mammals, especially cetaceans, pinnipeds and sea otters. Preference varies with location.
Will kill swimming seabirds (usually abandoning the carcasses) and takes some squid. Not known to take fish.
Group size Typically 2–6 animals, consisting of a mother and her offspring. In recent years, large temporary aggregations
of 30 or more have been observed. Lone individuals (usually males) are sometimes seen.
Remarks The consensus among researchers is that the ecotype formerly known as the ‘transient killer whale’ should
be called ‘Bigg’s killer whale’, in honour of the late pioneering killer whale researcher Dr Michael Bigg (and because the
term ‘transient’ is rather misleading when describing the site-fidelity and movement patterns of these whales).

ADULT MALE BIGG'S

ADULT FEMALE AND CALF BIGG'S

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 OFFSHORE
• ‘Typical’ black-and-white killer whale • Saddle patch usually quite faint (normally closed – no
• Overall appearance very similar to resident black incursion into grey – though open in some
• Smallest of three known North Pacific ecotypes individuals)
• Relatively smaller dorsal fin • Regularly slaps tail when swimming
• Male dorsal fin tip continuously rounded over tip • Less sexual dimorphism than in resident or Bigg’s
(more rounded than Bigg’s, without sharp corner of (tendency for sexes to be more similar in size)
resident) • Extreme tooth wear normal even in sub-adults (often
• Usually more nicks and scars on trailing edge of dorsal worn flat to gum line – likely from eating sharks
fin than resident that have abrasive skin); similar tooth wear
• May have oval scars from cookiecutter shark bites unknown in resident or Bigg’s
• Grey saddle patch similar in size to that of resident • White eye-patch a medium-sized oval (parallel to
body axis)
L: ♂ 6.5m, ♀ 5.5m; Max: 7.2m
Distribution The least known of the three North Pacific ecotypes. Ranging between southern California and the
eastern Aleutian Islands, Alaska, offshores travel extensively throughout their range. It is unclear whether they occur
predominantly in the open ocean or over the continental shelf; they will sporadically visit coastal (and occasionally
protected inshore) waters.
Food and feeding Bony and cartilaginous fish, especially sharks (including great white, blue, Pacific sleeper, Pacific
spiny dogfish); also known to take Chinook salmon and Pacific halibut. There is no evidence of mammal-eating.
Group size Normally large (50–100 animals is not unusual); occasionally 200-plus in temporary gatherings (possibly
related to prey density).

ADULT MALE OFFSHORE

ADULT FEMALE AND CALF OFFSHORE

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 KNOWN KILLER WHALE POPULATIONS IN THE NORTH ATLANTIC
Note Possible ecotype delineation is less clear in the North Atlantic, but the best-known populations are described here.

ICELANDIC SUMMER-SPAWNING HERRING-FEEDERS

• ‘Typical’ black-and-white killer whale • Front end of eye patch in front of blowhole
• Smaller than those of West Coast Community • Significant tooth wear (teeth often worn smooth to
• More closely resembles north-east Pacific residents the gum line)
• Medium- to large-sized oval eye patch (parallel to • Conspicuous saddle patch
body axis) • Worn teeth produce wide rake marks
L: ♂ 6.3m, ♀ 5.9m; Max: 6.6m
Distribution Iceland, but c. 5 per cent move to north-east Scotland (UK) in the spring and summer (especially Shetland).
No contemporary movement has been detected between Iceland and Norway.
Food and feeding Primarily schooling herring, moving between the herring wintering, spawning and feeding grounds
around Iceland. Carousel feeding frequently observed during winter and summer: the whales work in groups of 3–9 to
round up herring; they split a group of fish from the larger school, swim in fast circles (blowing bubbles, flashing their
undersides and lobtailing) to herd them into a tighter ball, whip their tails into the ball to stun or kill as many fish as
possible, then pick them off one-by-one. Some Icelandic killer whales appear to specialise on herring and follow it year-
round, while others feed on it only seasonally or opportunistically. At least 12 other prey species have been recorded in
Icelandic waters, including whales, dolphins, porpoises, seals, eider ducks, fish and squid. In Scotland, killer whales feed
on herring (offshore) and harbour and grey seals (inshore), and are known to take porpoises, otters and seabirds. Scottish
individuals return to Iceland for the winter, where they feed on the same herring stocks as those that remain year-round.
Group size 4–6 when feeding on marine mammals; 6–30 when feeding on fish close to shore; up to 300 feeding on fish
along the continental shelf edge.
ADULT MALE NORTH-EAST ATLANTIC
(ICELANDIC HERRING-FEEDERS,
NORWEGIAN HERRING-FEEDERS AND
NORTH-EAST ATLANTIC MACKEREL-FEEDERS)

ADULT FEMALE AND CALF NORTH-EAST ATLANTIC

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 NORWEGIAN SPRING-SPAWNING HERRING-FEEDERS
• No discernible differences from Icelandic herring-feeders
L: ♂ 6.2m, ♀ 5.5m; Max: 6.6m
Distribution Most of Norway’s killer whales follow the movements of its spring-spawning herring. No contemporary
movement has been detected between Norway and Iceland.
Food and feeding Primarily schooling herring; recent evidence suggests that, in addition to feeding on overwintering
herring inside the Norwegian fjords, at least some of the whales follow the herring to their offshore spawning grounds.
Other prey items documented include mackerel, cod, salmon, squid, harbour porpoise and harbour and grey seals. Many
whales in this population may include seals in their diet on a regular basis.
Group size Herring-feeding groups range from 6–30 individuals (median 15), seal-feeding groups from 3–11 (median 5).

NORTH-EAST ATLANTIC MACKEREL-FEEDERS

• Worn teeth (due to suction feeding) smaller and male dorsal fin often more rounded at
• Subtly different features to Icelandic and Norwegian tip (butter knife shape)
herring-feeding killer whales, e.g. eye patch often
L: ♂ 6.3m, ♀ 5.9m; Max: 6.6m
Distribution Ranges throughout the northern North Sea, the Irish Sea and the Norwegian Sea, and into the Arctic. Mostly
offshore, but also coastal. Known mainly between the Northern Isles (UK) and southern Norway in mid- to late autumn;
west of the Hebrides (UK) in winter; and in the Norwegian Sea (including Iceland) up to 72°N during late summer.
Food and feeding Mackerel (for at least part of the year – whether it targets mackerel year-round or switches to other
seasonally available prey is unknown). Frequently feeds around fishing trawlers.
Group size Those feeding in the Norwegian Sea have been observed in groups ranging from 1–40 (average 8); groups
feeding around fishing trawlers range from 1–70 (average 13). Maximum group size 200.

WEST COAST COMMUNITY

• ‘Typical’ black-and-white killer whale • Medium- to large-sized oval eye patch (slanting down
• Larger than Icelandic or Norwegian herring specialists towards rear – a key distinguishing feature)
• More closely resembles NE Pacific Bigg’s and Type A • Faint saddle patch
L: ♂ unknown, ♀ 6.1m; Max: unknown
Distribution The UK and Ireland, patrolling a vast area from Pembrokeshire and the southern Irish Sea, along the west
coast of Ireland, and north to the Outer Hebrides (where they are seen most often).
Food and feeding Little understood, but known to have taken harbour seals and harbour porpoises.
Group size Usually 2–3, but too few animals to be certain (currently only 8 individuals).

ADULT MALE WEST COAST COMMUNITY

(most distinct individual –
adult male called John Coe)

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 STRAIT OF GIBRALTAR BLUEFIN TUNA-FEEDERS
• ‘Typical’ black-and-white killer whale • Conspicuous saddle patch uniformly grey and closed
• Medium- to large-sized oval eye patch (parallel to • Male dorsal fin tip usually rounded and ends in
body axis) pointed trailing tip (similar to NE Pacific resident)
L: ♂ 6m, ♀ 5.3m; Max: 7.3m
Distribution In spring and summer, known mainly from the Strait of Gibraltar, with sporadic sightings in the Gulf of
Cadiz and Alboran Sea plus along the west and north coasts of the Iberian peninsula. They have been sighted in the
Strait in autumn and winter, but it is also possible that they follow bluefin tuna into the eastern Atlantic Ocean.
Food and feeding Atlantic bluefin tuna, which is chased at high speed and for up to 30 minutes at a time during spring
and summer. Approximately half the population (pods A1 and A2) frequently depredates tuna from baited hooks in a
drop longline fishery during summer. It is unknown whether they are dependent on tuna year-round; however, at least
one pod (D) may opportunistically feed on coastal fish species.
Group size The small population is subdivided into five pods (A1, A2, B, C and D) each comprising 7–15 individuals.
Remarks In recent years there have been dozens of apparently aggressive interactions between some individuals in
this population and boats (mainly yachts). The cause is uncertain.

ADULT MALE STRAIT OF GIBRALTAR

BLUEFIN TUNA-FEEDER

ADULT FEMALE STRAIT OF GIBRALTAR

BLUEFIN TUNA-FEEDER

NORTH-WEST ATLANTIC
• ‘Typical’ black-and-white killer whale • Dorsal fin tip usually rounded and ends in pointed
• Medium- to large-sized oval eye patch (parallel to trailing tip (similar to NE Pacific resident)
body axis) • Conspicuous saddle patch uniformly grey and mostly
closed
L: ♂ 6.7m, ♀ 5.5–6.5m; Max: unknown
Distribution Known mainly from northern Newfoundland and Labrador (Canada), and Greenland, especially in summer
(though there is considerably less observer effort in winter, the seasonal arrival of pack ice likely limits distribution).
Fishermen are also reporting killer whales far offshore, on the Grand Banks of Newfoundland.
Food and feeding Takes a variety of seals, other cetaceans, fish, cephalopods and occasionally seabirds. There
appears to be some group-specific prey specialisation (e.g. fish and cephalopods in west Greenland, marine mammals
in east Greenland).
Group size Typically 2–6 (average five); rarely more than 15, occasionally as many as 30; one-quarter of all sightings
are lone individuals.

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 KNOWN KILLER WHALE ECOTYPES IN THE ANTARCTIC AND
ADJACENT WATERS
TYPE A (ANTARCTIC KILLER WHALE)
• ‘Typical’ black-and-white killer whale • Usually no visible dorsal cape
• Possibly largest of Antarctic ecotypes (Large Type B • Saddle patch can be brownish
may be equal in size) • White patches occasionally tinted slightly yellowish
• Medium-sized oval eye patch (oriented parallel to (with diatoms)
body axis) • Black body colour occasionally tinted slightly brownish
• Male saddle patch usually closed, female’s can be (with diatoms)
slightly open
L: ♂ 7.3m, ♀ 6.4m; Max: 9.2m
Distribution During the southern summer, it is circumpolar in Antarctic waters, mostly in offshore, ice-free, open water;
frequently seen around the Antarctic Peninsula. Seasonal movements are poorly understood, but it is known to migrate
away from Antarctica to lower, warmer latitudes, at least for short periods. Killer whales that look like Type A have
been observed in New Zealand, Australia, South Africa, West Africa and Chilean Patagonia, as well as the Crozet
Archipelago, the Kerguelen Islands and Macquarie Island, but whether any of these other whales migrate to Antarctic
waters is unknown. Currently, Type A is a ‘catch-all’ ecotype that includes anything that is not Type B, C or D – and,
ultimately, it could include more than one ecotype.
Food and feeding In Antarctic waters, predominantly Antarctic minke whales and elephant seals, though it may
take calves of other baleen whale species and other seals; it has also been observed chasing (though not catching)
penguins. It is not known what this ecotype feeds on when away from Antarctic waters.
Group size 10–15 (ranges from 1–38).

ADULT MALE TYPE A

(ANTARCTIC)

ADULT FEMALE AND CALF TYPE A

(ANTARCTIC)

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 LARGE TYPE B (PACK ICE KILLER WHALE)
• Two-tone grey and white (not black and white) • Saddle patch almost always closed
• Larger and more robust body than Small Type B • Dark grey dorsal cape (often demarcated by narrow
(Gerlache) killer whale white border originating as an extension of lower
• Often covered in diatoms (turns white areas yellowish, saddle) stretching from forehead to just behind
grey areas brownish) dorsal fin
• Eye patch variable but always larger than any other • Very similar to Small Type B (Gerlache) killer whales
killer whale (at least twice as large as in Type A) but twice the size
• Eye patch oriented parallel to body axis • Often seen spyhopping around ice floes (looking for
• Small oval scars from cookiecutter shark bites seals)
common
L: ♂ unknown, ♀ unknown. Max: 9m; estimated to be at least twice the bulk of the Gerlache killer whale.
Distribution Circumpolar in Antarctic waters during summer, mostly inshore around dense pack ice and, especially, floe
ice. It is common around the northwestern half of the Antarctic Peninsula and retreats south with the summer break-
up of fast ice. Winter distribution is unknown but, while it apparently spends most of the year in Antarctic waters, it
periodically undertakes rapid round-trip migrations to tropical and sub-tropical waters (30–37°S). These are known
as ‘maintenance migrations’, believed to allow skin regeneration without the high cost of heat loss. The Antarctic
Peninsula population migrates north, east of the Falkland Islands and Argentina, to Uruguay and Brazil. It is likely that
‘clean’ grey-and-white individuals have recently returned from the tropics (where they shed diatoms with their skin).
Food and feeding Feeds preferentially on Weddell seals (usually in coordinated groups, by ‘wave-washing’) and will
often ignore crabeater and leopard seals. It occasionally takes Antarctic minke whales and elephant seals, and may
also take humpback whale calves.
Group size Usually fewer than 10.

ADULT MALE LARGE TYPE B

(PACK ICE)
ADULT MALE LARGE TYPE B (PACK ICE),
WITH DIATOMS (not to scale)

ADULT FEMALE AND CALF LARGE TYPE B

(PACK ICE)

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 SMALL TYPE B (GERLACHE KILLER WHALE)
• Two-toned grey and white (not black and white) • Eye patch may be oriented parallel to body axis or
• Smaller and slimmer body than Large Type B (Pack Ice) slightly slanted
killer whale • Dark grey dorsal cape present (though can be
• Often covered in diatoms (turns white areas yellowish, indistinct), stretching from just in front of eye patch
grey areas brownish) backward to just behind dorsal fin, and continuous
• Small oval scars from cookiecutter shark bites common with the lower leading edge of the saddle (often
demarcated by narrow white border originating in
• Eye patch variable but always larger than in any other
saddle)
killer whale (except Pack Ice killer whale)
• Very similar to Large Type B (Pack Ice) killer whales
• Eye patch narrower than Pack Ice killer whale
but half the size
• Saddle patch usually (but not always) closed
L: ♂ unknown, ♀ unknown. Max: 7m; estimated to be roughly half the bulk of Pack Ice killer whales.
Distribution Known mainly from the Antarctic Peninsula (western side and the western Weddell Sea); the Gerlache
Strait and Antarctic Sound are hotspots. Usually in more open water (it tends to avoid pack ice), often near penguin
colonies. Spends much of the year in Antarctica, but periodically undertakes rapid (6–7-week) round-trip ‘maintenance
migrations’ to tropical and sub-tropical waters (30–37°S).
Food and feeding Has only been observed feeding on penguins, especially gentoo and chinstrap (it eats only the breast
muscles and discards the rest), but it probably feeds mainly on fish and squid caught near the ocean floor (it is a deep
diver).
Group size Often 50-plus.
ADULT MALE SMALL TYPE B
(GERLACHE)

ADULT FEMALE AND CALF SMALL TYPE B

(GERLACHE)

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 TYPE C (ROSS SEA KILLER WHALE)
• Two-toned grey and white (not black and white)
• May be covered in diatoms (turns white areas yellow
or orange, black and grey areas brown)
• Small oval scars from cookiecutter shark bites
common
• Smallest killer whale ecotype known
• Dark grey dorsal cape usually visible (often
demarcated by narrow white border originating in
saddle)
• Small, narrow, wispy eye patch (slanted forwards at
45°angle to body axis)
• Saddle patch usually closed and very distinct

L: ♂ 5.6m, ♀ 5.2m. Max: 6.1m; weighs several times less than Type A and Type B (Pack Ice) killer whales (which could
conceivably prey on it).
Distribution Known mainly from East Antarctica, predominantly in the Ross Sea but also west along the Adélie Land to
Wilkes Land coasts, with smaller numbers as far west as Prydz Bay. Commonly reported in McMurdo Sound. It lives in
dense pack ice, polynyas and leads in the fast ice (often many kilometres from open water) and is concentrated where
most sea ice remains year-round. Spends most of the year in Antarctica (and has been recorded in the sea ice during
winter) but the presence of cookiecutter shark bites and sightings off New Zealand and Australia indicate that there are
at least some migrations to lower latitude and even tropical and sub-tropical waters.
Food and feeding Only known to feed on fish, primarily large 2m-long Antarctic toothfish, but also takes at least two
much smaller species of icefish and may take super-abundant (but very small) Antarctic silverfish. Limited evidence of
hunting penguins. It routinely dives to 200–400m, with a maximum of at least 700m.
Group size 10–120 (up to 200); group size appears to have been decreasing in recent years (current average about 14).

ADULT MALE TYPE C

(ROSS SEA)

ADULT FEMALE AND CALF TYPE C

(ROSS SEA)

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 TYPE D (SUB-ANTARCTIC KILLER WHALE)
• ‘Typical’ black-and-white killer whale • Male dorsal fin relatively short, narrow, noticeably
• Distinctive minuscule white eye patch (parallel swept back and with sharply pointed tip (more
to body axis) – sometimes absent – makes falcate and pointed than other Antarctic ecotypes)
identification easy • Moderately conspicuous saddle patch
• Noticeably bulbous melon, cf. other killer whales • No conspicuous dorsal cape
(more like pilot whale in some individuals) • No yellowish or brownish colouring (caused in some
• Marked sexual dimorphism in dorsal fin size and ecotypes by diatoms)
shape (as in other ecotypes)
L: ♂ unknown, ♀ unknown. Max: 7.3m.
Distribution Circumpolar in sub-Antarctic waters, ranging from 40°S to 60°S; sometimes associated with islands.
There have been many live sightings in recent years, at the northern edge of the Southern Ocean (including at Crozet,
South Georgia and New Zealand sub-Antarctic islands); and it is now being seen almost annually in the Drake Passage
and between the Falkland Islands and South Georgia (one particular hotspot is an area along the continental shelf
edge, south of Cape Horn, in Chilean waters).
Food and feeding Little known, but certainly includes fish (it takes Patagonian toothfish from longline fisheries near the
Crozet Archipelago and off Chile).
Group size Range 9–35, average 18 (but based on little information).
Remarks This is the most distinctive-looking killer whale ecotype, immediately recognisable by its extremely small (or
sometimes absent) white eye patch.

ADULT MALE TYPE D (SUB-ANTARCTIC)

ADULT FEMALE AND CALF TYPE D (SUB-ANTARCTIC)

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 SHORT-FINNED PILOT WHALE
Globicephala macrorhynchus Gray, 1846

The short-finned pilot whale is a distinctive-looking animal but, at sea, is virtually impossible to tell apart from its close
relative, the long-finned pilot whale (the main distinguishing feature is a subtle difference in the length and shape of
their flippers). Both species are highly sexually dimorphic in both size and appearance.
IUCN status Least Concern (2018).
Population Unknown. Approximate regional estimates total c. 700,000, but large swathes of the range have not been
surveyed. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised subspecies; there is genetic evidence of three types or forms of undetermined taxonomic
status: one in the Atlantic and two in the Pacific/Indian Oceans. Two forms – exhibiting differences in appearance,
morphology, vocal repertoire, life history and genetics – were first noted in the colder Oyashio Current off northern
Japan (called the shiho type) and in the warmer Kuroshio Current off southern Japan (the naisa type). It is now
believed that these are more widespread: the shiho type occurs throughout the eastern Pacific, and the naisa type
occurs throughout the remainder of the species’ distribution; separated by the East Pacific Barrier, they are likely to be
different subspecies.
Other names Pothead, blackfish (term normally used for non-taxonomic group of six dark-coloured members of
Delphinidae with ‘whale’ in their name).
DISTRIBUTION Widely distributed in deep tropical, sub-tropical and warm temperate waters worldwide. Does not
normally range north of 50°N or south of 40°S. Absent from the Mediterranean Sea (where the long-finned pilot whale is
resident). There are long-term residents in some areas, but other populations may move long distances (individual whales
have been recorded travelling up to 2,400km/month). Seasonal inshore–offshore (winter/early spring–summer/autumn)

May have very square head from ADULT MALE

above (especially older males)

Distinct
median
notch

Pale grey or whitish

Crescent-shaped Pale grey ‘eyebrow’ blazes from saddle patch (highly
Pointed tips
blowhole (ends above eyes towards front of variable)
point forward) dorsal fin (highly variable)

DIVE Sequence Quite leisurely dive sequence; head raised relatively high above
surface (eyes often visible); flukes sometimes raised above surface before deep
dive; several whales typically surface close together. • Depth Foraging dives can
exceed 1,000m, but depth varies greatly with region and time of day; in Hawaii,
typically to 700–1,000m during day, 300–500m at night); in Canary Islands
performs deep dives day and night; maximum recorded 1,552m.
• Duration Typically 12–15 (occasionally 20) minutes (varies with sex, size and
behaviour); maximum recorded 27 minutes (Hawaii).
BLOW Strong, low, shapeless blow (up to c. 1m), quite conspicuous in calm
weather (but tends to dissipate quickly).

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Pale grey or whitish
‘eyebrow’ blaze angled
up and backward (highly
variable – most prominent in
eastern North Atlantic)
ADULT MALE
Fin low in profile and one-
Predominantly dark grey third of the way along back
to black or brown (noticeably further forward
than in other cetaceans)
Long, robust
body Extremely wide, broad-based,
falcate or hooked dorsal

Round, exceptionally Pale grey or whitish saddle

bulbous head patch (highly variable –
(exaggerated in most prominent in eastern
older animals) North Atlantic)

Flukes may
Indistinct develop
beak Pointed tip Most adults have upturned tips
some scarring
Straight
Melon may mouthline Deepened tailstock
protrude beyond, angled upward (more pronounced
or overhang, Long, slender, May have oval scars from in male)
mouth by up to sickle-shaped cookiecutter shark bites (but
10cm (especially in flippers (slightly generally heal to background
older males) longer in male) colour)
ADULT MALE

Light grey to white narrow

midline stripe connects chest
Light grey to white anchor-shaped patch on patch to wider urogenital patch
chest (visible when animal spyhops)

80°

70°

60°

50°

40°

AT A GLANCE Warm waters worldwide

30°

20°

• Medium size • Black, blackish or brownish

10°
0°

• Rounded, bulbous melon • Indistinct beak

10°
20°

• Broad-based, backswept dorsal fin far

30°

40°

50°

60°
forward • Small to large groups
70°

80°

primary range secondary range approx. overlap with

long-finned pilot whale

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 movements in some regions are related to the seasonal spawning migrations of squid. Prefers the continental shelf
break, continental slope and island slope waters, and areas with complex topography such as seamounts and ridges.
Abundance is lower in deep oceanic environments. Will approach nearshore areas where the water is sufficiently deep.
BEHAVIOUR Often observed in aggregations with other cetacean species. There are accounts of pilot whales behaving
aggressively toward some cetaceans and they tend to harass larger whales, but the role is sometimes reversed with
smaller species (such as melon-headed whales), when the pilot whales are the victims. Oceanic whitetip sharks
often follow pilot whales, to scavenge on lost or discarded prey (or they rely on the whales to find prey at depth).
More aerially active than the long-finned pilot whale, breaching occasionally (though not as often as many smaller
delphinids), spyhopping and lobtailing. Spends much of the day logging (resting) at the surface. More prone to mass
strandings than almost any other cetacean (apart from the long-finned pilot whale), probably in part because of its
strong social bonds. Will grieve for dead members of the group and will carry a dead calf around for hours or days.
Reaction to boats varies according to location.
FOOD AND FEEDING Mainly squid; some octopus, and mid- and deepwater fish. While foraging may spread out in
‘chorus line’ up to 3km long.
TEETH Upper jaw 14–18; lower jaw 14–18.
GROUP SIZE AND STRUCTURE Highly social, living in matrilineal groups (consisting of a matriarch with her immediate
kin) similar in structure to killer whale groups (though not quite as stable). Typical group size is 15–50 (average 18
around the main Hawaiian islands, 15 in Madeira), including all ages and sexes (though there tend to be more adult
females). They usually remain in the family group for life. Males will mate during temporary aggregations of separate
family groups. Several family groups may join together to form a pod or school, typically with 30–90 members (up to
several hundred). Rarely seen alone (very occasionally, lone adult males are reported).

ADULT MALE

Head may become

SIZE
L: ♂ 4.2–7.3m, ♀ 3.2–5.1m;
WT: 1–3.5t; MAX: 7.3m, 3.6t
Calf – L: 1.4–1.9m; WT: 40–85kg
more bulbous and
conspicuously flattened
on front in older animals
(especially in western and
central North Pacific)

Less bulbous Significantly smaller

head dorsal fin
Less deepened
tailstock
ADULT FEMALE

Dorsal fin not ADULT MALE SHIHO TYPE

as broad (eastern Pacific and northern Japan)
Adult male has rounder Brighter, broader, more distinct light
head (especially when grey saddle patch behind dorsal fin
viewed from above) (with distinct rear margin)
Larger size (maximum
female 4.2–5.1m,
male 6.2–7.3m)

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 Broader Darker, less distinct ADULT MALE NAISA TYPE
Adult male has flatter,
dorsal fin saddle patch (just visible (throughout range except eastern
squarer or slightly concave
head (especially when in very good light or Pacific and northern Japan)
viewed from above) underwater) that fades
into dark pigmentation of
Smaller size (maximum
tailstock
female 3.2–4.1m, male
4.2–5.3m)

Many individuals
can appear quite ADULT MALE BROWN VARIATION
brown

Both sexes have

more dolphin-like,
Paler than adult falcate dorsal fin
CALF

MALE DORSAL FIN VARIATION Slimmer body shape

Deep concave
trailing edge
Base may be more
than twice as long
Thickened leading as fin is tall
edge in older males

Adult male fin often

very hooked

FLIPPER COMPARISON 14–19 per cent 18–27 per cent

of body length of body length
Strongly angled
Gently curved leading edge
leading edge (forms ‘elbow’)

short-finned pilot whale long-finned pilot whale

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 LONG-FINNED PILOT WHALE
Globicephala melas (Traill, 1809)

With experience, it is possible to tell the sex and approximate age of long-finned and short-finned pilot whales by
looking at their dorsal fins: they change shape as they grow older and are quite different on females and males. The
adult male pilot whale’s fin is like no other: low in profile and exceptionally broad-based (the base can be more than
twice as long as the fin is tall).
IUCN status Least Concern (2018). Mediterranean sub-population Data Deficient (2010).
Population Unknown, a guesstimate of c. 1 million suggested (based on available regional estimates). Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy Two subspecies recognised: North Atlantic long-finned pilot whale (G. m. melas) and southern long-finned
pilot whale (G. m. edwardii). An undescribed subspecies in Japanese waters (now extinct) was informally known as
the North Pacific long-finned pilot whale.
Other names Pothead, caaing whale, blackfish (term normally used for non-taxonomic group of six dark-coloured
members of Delphinidae with ‘whale’ in their name).
DISTRIBUTION The two extant subspecies are widely distributed in deep cold temperate to sub-polar waters of the
North Atlantic and southern hemisphere, but separated by a wide tropical belt. Prefers the continental shelf break,
continental slope and island slope waters, and areas with complex topography such as seamounts and ridges. Most
sightings are in waters deeper than 2,000m.
BEHAVIOUR Often observed in mixed-species aggregations. Less aerially active than the short-finned pilot whale, often
spyhopping and lobtailing but only occasionally breaching. Spends much of the day logging (resting) at the surface. Like
short-finned pilot whale, more prone to mass strandings than almost any other cetacean, probably partly because of its
strong social bonds. Reaction to boats varies according to location.
FOOD AND FEEDING Mainly squid and other cephalopods; some small to medium-sized fish; occasionally shrimps;
however, great variation according to location. Deep foraging tends to be at night in most regions.
TEETH Upper jaw 16–26; lower jaw 16–26.
GROUP SIZE AND STRUCTURE Highly social, living in matrilineal groups similar in structure to killer whale groups
(though not quite as stable). Typical family group size is 8–20, with considerable geographical variation, including all
ages and sexes. They remain in the group for life. Several family groups may join to form a pod or school, frequently
with up to 50 members (sometimes 100+ and occasionally up to 1,200).
ADULT MALE SOUTHERN HEMISPHERE
‘Eyebrow’ streak behind eye
can be long, white and very Saddle patch can be solid white (most
conspicuous other populations show just hint)

Typically
glossy black

DIVE Sequence Quite leisurely dive sequence; head raised relatively high above
surface (eyes often visible); flukes sometimes lifted above surface before deep
dive; several whales typically surface close together. • Depth Most feeding 30–
500m; maximum recorded 828m but likely capable of deeper. • Duration Typically
2–12 minutes (maximum recorded 18 minutes).
BLOW Strong, low, shapeless blow (up to c. 1m height); quite conspicuous in calm
weather, but tends to dissipate quickly.

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 Round, exceptionally bulbous
head (exaggerated and
sometimes with flattened
front in older animals)
Pale grey or whitish
‘eyebrow’ blaze angled
up and backward (highly
variable – often not
visible in North Atlantic)
Predominantly Extremely wide, ADULT MALE NORTH ATLANTIC
dark grey to jet broad-based, falcate
black or brown or hooked dorsal fin
Fin low in profile and one-third of the
Long, robust way along back (noticeably further
body forward than in other cetaceans except
short-finned pilot whale)
Deep, concave
trailing edge
Pale grey or whitish saddle
patch (highly variable –
most prominent in southern
hemisphere)

Pointed tip
Deepened tailstock
Indistinct beak Long, slender, sickle- (more pronounced
Straight mouthline Most adults have
shaped flippers (slightly in male)
angled upward some scarring Flukes may
longer in male)
develop
Melon may protrude beyond, or
upturned tips
overhang, mouth by up to 10cm
(especially in older males)
Significantly smaller, ADULT FEMALE NORTH ATLANTIC
Less bulbous thinner dorsal fin
head
Less deepened
tailstock

SIZE
L: ♂ 4–6.7m, ♀ 3.8–5.7m;
WT: 1.3–2.3t; MAX: 6.7m, 2.3t
Calf – L: 1.7–1.8m; WT: c. 75–80kg

80°

70°

60°

50°

AT A GLANCE Cold waters of North Atlantic

40°

30°

20°
10°
and southern hemisphere • Medium size
0°
10°
• Black, blackish or brownish • Rounded,
20°

30°
bulbous melon • Indistinct beak • Broad-
40°
based, backswept dorsal fin positioned far
forward • Small to large groups
50°

60°

70°

80°

approx. overlap with short-finned pilot whale

LONG-FINNED PILOT WHALE 173

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 FALSE KILLER WHALE
Pseudorca crassidens (Owen, 1846)

Despite its name, the false killer whale belongs taxonomically to the dolphin family, Delphinidae, and it often behaves
more like one of its energetic and sprightly smaller relatives.
IUCN status Near Threatened (2018).
Population Unknown. Considered among the least abundant delphinids, even in locations with the highest densities.
Trend unknown (but the best studied population, around Hawaii, has been declining).
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Pseudorca, blackfish (term normally used for non-taxonomic group of six dark- coloured members of
Delphinidae with ‘whale’ in their name).
DISTRIBUTION Tropical to warm temperate waters worldwide, mainly between c. 50°N and c. 50°S. Density is much
higher in lower latitudes. Sightings in cooler temperate waters (such as the Baltic Sea, off the UK and Canada’s British
Columbia) are generally considered extralimital. Primarily favours deep oceanic waters, particularly those deeper than
1,500m; it also occurs where deep water approaches the coast, especially around oceanic islands. However, certain
populations appear to be more coastal and occur in shallower water.
BEHAVIOUR An exuberant, fast-swimming cetacean. Often leaps clear of the water, especially when attacking
prey; will often breach with prey in the mouth, and may throw it quite high into the air. Mass strandings are fairly
common (likely due to strong social bonds). Regularly associates with other cetaceans and long-term associations with
bottlenose dolphins have been documented. However, it occasionally behaves aggressively towards other cetaceans.
Not shy of boats and will readily bow-ride and wake-ride.
FOOD AND FEEDING Varies by region, but mainly large fish; also takes squid; will attack and eat other small cetaceans.
Cooperative feeder and will share prey; hunts day and night.
TEETH Upper jaw 14–22; lower jaw 16–24.
GROUP SIZE AND STRUCTURE Highly variable depending on location, typically ranging from 10–60 (less commonly
2–100); exceptionally, larger groups have been reported (the largest mass stranding involved at least 835 animals).
Where one small group is present, often other small groups are scattered over a wider area. Smaller groups consist
primarily of closely related individuals and are of mixed age and sex; females (and possibly males) seem to remain
within the social group in which they were born. Larger groups are likely temporary associations of smaller, more stable
groups.
DORSAL FIN VARIATIONS

Fin shape highly

Usually
variable
rounded tip SIZE
L: ♂ 4–6m, ♀ 4–5.1m;
WT: 1.1–2t; MAX: 6.1m, 2.2t
Calf – L: 1.5–2.1m; WT: c. 80kg
Males larger than females; regional
size differences (e.g. 10–20 per cent
larger in Japan than South Africa).

DIVE Sequence When swimming slowly, head and melon break surface (eyes may be visible), and may strongly arch
tailstock but flukes rarely visible; when swimming quickly, may porpoise just clear of water in low, flat arcs, but often
shows little more than splash and dorsal fin. • Depth Most feeding near surface, but will forage along seafloor;
capable of more than 1,000m. • Duration Long dives typically 4–6 minutes; maximum recorded 18 minutes.
BLOW Short and bushy but only sometimes visible.

174 BLACKFISH

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 Small, conical ADULT
Dark grey to black Slightly darker dorsal
head with (may appear slightly
non-bulbous cape (visible only in Relatively narrow-
paler slate-grey in good light) based, falcate dorsal
Front melon bright light) fin midway along back
of head
may be May have
flattened light grey Long, Dorsal fin taller
in older areas on slender than it is long
males sides of body (18–41cm high)
head (highly
variable)
Dorsal fin smaller (in
proportion to amount
of back visible) than on
any other blackfish

No
Narrow flippers
discernible
Long relatively far
beak May be light star-shaped
mouthline forward on body
scars from cookiecutter shark Deep tailstock
Melon overhangs bites (most heal to same
Distinct bulge colour as background)
lower jaw more
on leading edge
in male
(makes S-shape)

ADULT
Narrow,
tapered head Slender, relatively
small flukes

Median
notch

Distinct bulge on leading edge

of flippers (like an ‘elbow’)

80°

70°

60°

50°

40°

30°
AT A GLANCE Warm (mainly offshore) waters
20° worldwide • Medium size • Dark grey to
black • Long, slender body • Relatively
10°
0°

narrow-based, falcate dorsal fin • Small,

10°
20°

conical head with no beak • Distinct bulge

30°

40°

50°

60°
on leading edge of flippers • Small, often
70°
exuberant groups
80°

primary range secondary range

FALSE KILLER WHALE 175

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 PYGMY KILLER WHALE
Feresa attenuata Gray, 1874

Despite its name, the pygmy killer whale belongs taxonomically to the dolphin family, Delphinidae. Until 1952 it was
known from only two skulls, collected in 1827 and 1874. It is still poorly known, but our knowledge has improved
dramatically in recent years.
IUCN status Least Concern (2017).
Population Unknown. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Blackfish (term normally used for non-taxonomic group of six dark- coloured members of Delphinidae
with ‘whale’ in their name).
DISTRIBUTION Tropical and sub-tropical waters worldwide, between 40°N and 35°S, overlapping almost exactly with
the melon-headed whale. Most sightings are in deep waters offshore and around oceanic islands where deep, clear
waters are found near the coast.
BEHAVIOUR Pygmy killer whales can be quite difficult to spot and, while high breaches have been observed, rarely
engage in aerial behaviour. Their reaction to boats is extremely variable, but they will occasionally bow-ride slow-
moving vessels. They spend most of the day travelling slowly, socialising or resting motionless at the surface.
FOOD AND FEEDING Mostly squid and fish; known to attack and possibly eat other dolphins in eastern tropical Pacific.
Most feeding seems to occur at night.
TEETH Upper jaw 16–22; lower jaw 22–26.
GROUP SIZE AND STRUCTURE Usually 12–50, although they have been encountered in pairs and herds of up to several
hundred; the average group size in Hawaii is nine.

Head rounded (not as ADULT

White patches and spotting
narrow or triangular as
typical (cf. melon-headed whale)
in melon-headed whale)

Fairly prominent dark,

narrow dorsal cape
SIZE
L: ♂ 2–2.6m, ♀ 2–2.6m;
WT: 110–170kg; MAX: 2.7m, 228kg
Calf – L: c. 80cm; WT: c. 15kg

DIVE Sequence Surfaces quietly and discreetly (rather a slow, sluggish swimmer), rarely porpoising and keeping low
profile; herd often swims shoulder to shoulder in coordinated ‘chorus line’. • Depth Believed to feed at depth; in
Hawaii, most commonly seen in water 500–3,500m deep. • Duration Unknown.
BLOW Rarely visible.

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 Melon may be more Distinct In poor light Tall, falcate Trailing edge of dorsal ADULT
extensive (even forward- demarcation appears dorsal fin in fin may be damaged
leaning) in some older between darker uniformly dark middle of back
individuals grey or blackish (shape varies Fairly prominent dark,
cape and lighter
with age) narrow dorsal cape
Cape broadens grey sides
to large dark
Moderately Most adults of both sexes have
‘cap’ on crown
robust widely spaced, paired, white
Rounded forebody linear rake marks (made by
to bulbous other pygmy killer whales)
head
Noticeably slimmer
behind dorsal fin

‘Lips’ often Dorsal cape does

pale grey not dip as low below
dorsal fin as on melon- Male has
to white Neck very flexible pronounced ventral
headed whale
Moderately long, keel (not present
No visible beak No face Cookiecutter shark scars on female)
(though upper jaw slender flippers
‘mask’ with rounded tips often present (may be
slightly overhangs tinged pink from blood flow)
tip of lower jaw) especially in older animals

Melon-headed whale
c. 20 per cent FLIPPER COMPARISON
of body length
Slightly convex
leading edge, Acutely pointed tip
straight trailing edge

HEAD VARIATIONS False killer whale

Pygmy killer whale c. 10 per cent
c. 18–23 per cent of body length
of body length

Convex leading
edge, concave Characteristic ‘elbow’ on leading
trailing edge Rounded tip edge gives S-shaped appearance

80°

70°

60°

50°

40°
AT A GLANCE Tropical and subtropical waters
30° worldwide • Small size • Appears uniformly
dark in poor light • Distinct dark dorsal cape
20°
10°

and no face ‘mask’ • Relatively large, broad

0°
10°

dorsal fin in middle of back • Rounded to

20°

30°

40°

50°
bulbous head • Generally slow and lethargic
60° • Typically in small herds of fewer than 50
70°

80°

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 MELON-HEADED WHALE
Peponocephala electra (Gray, 1846)

Despite its name, the melon-headed whale belongs taxonomically to the dolphin family, Delphinidae. It was known only
from skeletons until the 1960s, but nowadays is seen regularly in several parts of the world.
IUCN status Least Concern (2019).
Population Unknown. One guesstimate suggests minimum c. 600,000; considered to be relatively common in some
parts of its range. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Electra dolphin, little blackfish, Hawaiian blackfish (‘blackfish’ is normally used for non-taxonomic group
of six dark-coloured members of Delphinidae with ‘whale’ in their name); sometimes affectionately called a ‘pep’, after
the scientific name.
DISTRIBUTION Tropical and sub-tropical waters worldwide, overlapping almost exactly with the pygmy killer whale.
Most records are between 20°N and 20°S, and it is rarely seen north of 40°N or south of 35°S (rare records from higher
latitudes are usually associated with incursions of warm-water currents). Most sightings are in deep waters offshore
and around oceanic islands (typically in depths of 300–2,000m). There is some evidence of inshore movements during
the day (for resting and socialising) and offshore movements to feed at night.
BEHAVIOUR Usually encountered in large, dense, fast-swimming herds, which are notorious for suddenly changing
direction. They are often in mixed aggregations with Fraser’s dolphins, and have been observed with other cetaceans.
They tend to flee from approaching vessels in the eastern tropical Pacific, but will enthusiastically bow-ride elsewhere.
Breaching and spyhopping are fairly common. During the day in calm seas, large groups often rest by logging or milling
about at the surface.
FOOD AND FEEDING Mainly squid, but also small fish and crustaceans; may also prey opportunistically on dolphins in
some areas. Most feeding seems to occur at night.
TEETH Upper jaw 40-52; lower jaw 40-52.
GROUP SIZE AND STRUCTURE Usually in large, tight-knit groups of 100–500, with exceptional sightings of up to 2,000
individuals.
Fewer white patches and less
spotting than in pygmy killer whale
ADULT
Subtle dark, broad
dorsal cape

Head triangular and SIZE

pointed (less rounded than L: ♂ 2.4–2.8m, ♀ 2.3–2.7m;
in pygmy killer whale) WT: 160–210kg; MAX: 2.8m, 228kg
Calf – L: 1–1.2m; WT: c. 15kg

DIVE Sequence When swimming slowly, head and melon break surface briefly before dorsal fin appears, and
flukes rarely visible; when swimming quickly, porpoises clear of water or skims surface, producing much spray.
• Depth Typically prefers depths greater than 1,000m and feeds deep in water column; maximum recorded dive
472m. • Duration Maximum recorded 12 minutes.
BLOW Rarely visible.

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 Triangular head In poor light Moderately Tall, broad-based dorsal Dorsal fin often more
ADULT MALE
shape (can appears robust body fin in middle of back falcate in older individuals
become more uniformly dark
bulbous in older grey or blackish Trailing edge of
individuals) Subtle dark dorsal
Light grey dorsal fin may be
cape (visible in
crown damaged
good light)
Dorsal cape dips
much lower below
dorsal fin than on
pygmy killer whale

No visible
beak White linear rake Male has pronounced
White or light grey
areas common around Moderately long, marks rare ventral keel (not
throat (may extend sharply pointed, sickle- present in female)
along underside) shaped flippers

Less robust body Adult female has ADULT FEMALE

Less bulbous lower dorsal fin
melon
Narrower flukes

Sometimes a Relatively shorter Less pronounced

hint of beak flippers than male ventral keel

Light grey Dark cap

Head more triangular HEAD COMPARISON on crown
and pointed crown
Subtle dark face Head more bulbous
‘mask’ (more visible and rounded
in good light)
‘Lips’ (and, in some No face ‘mask’
older animals, tip of
lower jaw) often pale ‘Lips’ (and sometimes
grey to white entire chin) pale grey
to white
Melon-headed whale Pygmy killer whale

80°

70°

60°

50°
AT A GLANCE Tropical and subtropical waters
40° worldwide • Small size • Indistinct dark
dorsal cape and face ‘mask’ visible in good
30°

20°

light • Appears uniformly dark in poor light

10°
0°

• Tall, broad-based dorsal fin in middle of

10°
20°

back • Triangular, pointed head • May swim

30°

40°

50°
at high speed • Typically in large herds of
100+
60°

70°

80°

MELON-HEADED WHALE 179

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 RISSO’S DOLPHIN
Grampus griseus (G. Cuvier, 1812)

Risso’s dolphin is the most heavily scarred of all the dolphins and the largest species called ‘dolphin’. There is
huge variation in colour – between individuals, age classes and regions – and this is one of the most distinctive
characteristics of the species. It is quite easy to identify at close range – the only smallish, blunt-headed cetacean that
is typically light in colour.
IUCN status Least Concern (2018). Mediterranean sub-population Data Deficient (2010).
Population Unknown. The sum of existing estimates is 350,000, but this is likely to be a fraction of the actual total.
Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies. Most closely related to the blackfish.
Other names Grampus; in older literature, the killer whale was also called grampus.
DISTRIBUTION Widely distributed in both hemispheres, from the tropics to cool temperate waters, occurring in all
habitats from coastal to oceanic and including many semi-enclosed seas. However, it shows a strong range-wide
preference for mid-temperate waters between c. 30° and 45° latitude. Favours water warmer than 12°C (it is rarely
found in water less than 10°C). This results in seasonal movements in some regions. Prefers deep waters of the
continental shelf break, upper slopes and submarine canyons, especially in areas with steep seafloor topography
(generally 400–1,000m deep). It also occurs in some oceanic areas beyond the continental slope and will enter shallow
coastal waters to feed seasonally on cuttlefish. There is evidence that habitat use is coordinated to avoid spatial and
temporal overlap with other deep-diving odontocetes, including Cuvier’s beaked whales and sperm whales.
BEHAVIOUR During daytime, it is usually socialising, resting or travelling. When socialising it can be aerially active and
will breach, spyhop (often revealing the entire head and body down to the flippers), head-slap, lobtail and flipper-slap.
Commonly associates with other cetaceans. Readily bow-rides, wake-rides and associates with boats in some areas,
but elsewhere it does not approach boats; it is not particularly shy or nervous, but typically maintains a ‘personal space’
and slowly turns away.

ADULT MALE HIGHER LATITUDES

Shallow, vertical
crease on front Appears mostly
of melon greyish-white

Appendages
usually darker

Conspicuous dark
area around base of
Broad head
dorsal fin

DIVE Sequence Usually surfaces slowly at 45° angle; eye usually appears above surface; tall dorsal fin conspicuous
as back arches slightly; members of cluster often travel and surface in synchrony. • Depth Often less than 50m,
but up to 300m; maximum recorded 460m. • Duration Typically 1–10 minutes, likely capable of more (one report –
unverified – of 30 minutes).

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 ADULT MALE HIGHER LATITUDES
Some healed wounds
Robust body shape (appears to
on head may be from
have most of bulk in front of
beaks and suckers of
dorsal fin) Underlying body colour dark to
squid prey
pale grey (highly variable between
Tends to be lighter individuals and regions)
overall in higher
Front of melon has latitudes
Very tall (up to c. 45cm), erect,
distinct vertical Bulbous, squarish
moderately falcate dorsal fin midway
cleft or furrow head with
along back (highly variable shape)
(visible only at indistinct beak
close range) (squarer profile
Dorsal fin usually darker than
than most other
rest of body (and one of tallest
small cetaceans)
in proportion to body length of
any cetacean)
Relatively
narrow base

Relatively
slender tailstock
(especially
Mouthline slopes immediately in
Typically covered with
upward (slightly front of flukes)
white scratches and
downturned Chin
blotches (healed wounds
at corners) often
Long, pointed, sickle- from intraspecific fighting
white May have
shaped flippers and/or play)
dark eye
patch (usually darker than
rest of body)
ADULT MALE

Distinct vertical V-shaped

groove on front of melon
(unique to Risso’s dolphin,
SIZE function unknown)
L: ♂ 2.9–3.8m, ♀ 2.8–3.8m;
WT: 300–400kg; MAX: 4.1m, c. 500kg
Calf – L: 1–1.5m; WT: 20–30kg

80°

70°

AT A GLANCE Worldwide from tropics to cool

60°

50°

40°

30°
temperate waters • Small size • Robust body
20°
10°
• Squarish head (side view) with indistinct
0°
10°
beak • Cleft melon • Extensive linear
20°

30°
scarring • Highly variable coloration within
40° single group • Older animals almost white
• Appendages usually darker than rest of
50°

60°

70° body • Very tall, erect dorsal fin

80°

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 FOOD AND FEEDING Mainly deepwater squid and octopuses, but also some cuttlefish and, rarely, krill. Most feeding
appears to be during late afternoon and at night; evidence of cooperative feeding.
TEETH Upper jaw 0–4 (vestigial – usually unerupted); lower jaw 4–14. Teeth present in both sexes near the front of the
lower jaw (usually 6–8); may be worn down (or missing) in older adults.
GROUP SIZE AND STRUCTURE Typically 5–30, but often up to 100; there are reports of as many as 4,000 together.
There appears to be a ‘stratified’ group structure, with an average of 3–12 individuals in stable clusters grouped by age
and sex classes. Young animals appear to remain in the vicinity of their natal group for some years after being weaned,
then form pods of sub-adults at the age of 6–8 years.
OLDER MALE
Front half typically
whiter than rear half Older male can be nearly
white (gets whiter
with age, due to lost
pigmentation and scarring)

Appendages usually
remain darker

ADULT FEMALE

Tends to have less Generally not as

muscular body white as male (highly
variable)

ADULT MALE LOWER LATITUDES

Tends to be much darker
than in higher latitudes
Darker grey cape
very distinctive

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 Whitish ‘anchor patch’ on ADULT MALE
chest (may be obliterated by
white scarring) Diffuse whitish patch
around urogenital
area (variable)

Anchor patch and

urogenital patch
connected by light streak
Rare browner form in which
scratches repigment

ADULT MALE BROWN FORM

Dark grey or dark brownish-

Distinct darker cape grey (then darkens to near
Beak slightly more
prominent than in adult black before becoming paler
at sexual maturity) SUB-ADULT

Usually silvery-grey upperside,

creamy-white underside (turns
pale grey as juvenile) CALF
Distinct narrow,
Light grey dorsal fin
dark cape extends
(darkens with age)
down over eye
Cape dips slightly
below dorsal fin

‘Face’ region often Pale grey streak Little or no scarring

light olive-brown along side

RISSO’S DOLPHIN 183

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 FRASER’S DOLPHIN
Lagenodelphis hosei Fraser, 1956

For many years, Fraser’s dolphin was known only from a partial skeleton found on a beach in Sarawak, Malaysian
Borneo, some time before 1895. The first official record in the wild was in 1971, but nowadays it is a fairly familiar sight
in several parts of the world.
IUCN status Least Concern (2018).
Population Unknown. Approximate regional estimates total 350,000, but large swathes of the range have not been
surveyed. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Sarawak dolphin, Bornean dolphin.
DISTRIBUTION Tropical, sub-tropical and occasionally warm temperate waters. Recent sightings in the Azores (c. 38°N)
and Madeira (c. 33°N) may reveal the species as a potential bio-indicator of climate change (as it expands its range into
warming waters further north). Mainly oceanic, in water deeper than 1,000m; sometimes close to shore where deep
water approaches the coast.
BEHAVIOUR Active and energetic swimmer, usually in tight, fast-moving schools that whip the sea surface into a froth.
Frequently found in association with other cetaceans. Occasionally performs low, relatively undemonstrative breaches.
Response to boats varies from avoidance to quite approachable (will bow-ride, briefly, in some areas).
FOOD AND FEEDING Mesopelagic fish, cephalopods, crustaceans. Feeding techniques unknown.
TEETH Upper jaw 72–88; lower jaw 68–88.
GROUP SIZE AND STRUCTURE Herds tend to be large, typically with 40–300 animals, but groups as small as 4–15 and
as large as 2,500 are occasionally seen.
Dorsal fin typically
Facial and side less triangular and ADULT FEMALE
stripes faint to more falcate
moderate

Ventral keel usually

Beak may be absent or only
more distinct slightly developed

ADULT FEMALE
VARIATION

DIVE Sequence When swimming slowly, only blowhole, part of back and dorsal fin exposed; rolls forward with
slight arching of back; when swimming quickly, porpoises in long, low-angled, splashy leaps. • Depth Near surface
to c. 600m; physiological studies indicate capable of deep diving. • Duration Unknown.

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 Dark bluish-grey Small dorsal fin ADULT MALE
or brownish-grey (max. height 22cm)
Greyish-white or upperside midway along back
creamy border above Fin typically triangular
Stocky body
dark lateral stripe (or slightly falcate)

Fin tends to be more erect or

Stubby beak, canted forward in adult male
short (3–6cm)
but distinct
Mid-grey sides

Dark ‘bandit
mask’ (highly Dark eye-to-anus side
variable) Very small, slender stripe (highly variable
flippers with Dark stripe may Often has
between individuals pronounced
pointed tips and with age, sex and widen and darken
Tip of beak with age (can be jet ventral keel
and ‘lips’ dark location)
black in older adult
White lower jaw males)
Dark jaw-to-flipper and underside
stripe (may merge (pink when active)
with face mask)

SIZE
L: ♂ 2.2–2.7m, ♀ 2.1–2.6m;
WT: 130–200kg; MAX: 2.7m, 209kg
Calf – L: 1–1.1m; WT: 15–20kg
Dark stripe from
tip of upper jaw to ADULT
apex of melon

Small
median
notch

Very small
flukes Concave trailing
Blowhole very slightly
to left of midline edge

80°

70°

60°

50°
AT A GLANCE Deep tropical and sub-tropical
40° waters worldwide • Small size • Stocky
body • Short but distinct beak • Male often
30°

20°

has dark ‘bandit mask’ and lateral stripe •

10°
0°

Small, triangular dorsal fin, flippers and flukes

10°
20°

• Much individual variation within herds •

30°

40°

50°
Splashy, tight-knit herds leave distinct white
water
60°

70°

80°

FRASER’S DOLPHIN 185

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 ATLANTIC WHITE-SIDED DOLPHIN
Lagenorhynchus acutus (Gray, 1828)

Calling the Atlantic white-sided dolphin ‘white-sided’ is a bit of misnomer – its markings are complex, bold and more
colourful than those of most other dolphins – though the brilliant white patch on either side is one of the most striking
features of this gregarious dolphin.
IUCN status Least Concern (2019).
Population Minimum 200,000–300,000 may be a reasonable guesstimate. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy The genus Lagenorhynchus is under revision, and this species may be put into its own genus
(Leucopleurus). No recognised forms or subspecies.
Other names White-side, springer, jumper; affectionately called a ‘lag’ by researchers (from the generic name).
DISTRIBUTION Cold temperate to sub-Arctic waters of the North Atlantic, typically within a temperature range of
1–16°C (preferring 5–11°C). The extreme limits of the range are poorly known. Prefers fairly deep waters (primarily
100–500m) with high seabed relief of the outer continental shelf and slope, but also occurs in oceanic waters and
will enter fjords and inlets less than 50m deep. There are large-scale seasonal shifts in abundance in some regions:
typically to more northerly latitudes and/or closer to shore during warmer months.
BEHAVIOUR Lively and acrobatic, especially in larger groups. Will often leap and, less often, tail slap. Its leaps are
either simple (no spinning or twisting – clearing and re-entering the water with a smooth arc) or complex (higher,
involving twists and turns in the air). Will associate and feed with large baleen whales and sometimes forms mixed
groups with other cetaceans. Keen bow-rider and wake-rider, and will ride the bow waves of mysticetes.
FOOD AND FEEDING Mainly small schooling fish, squid, shrimps. Known to feed cooperatively on sand lance off New
England, by herding prey into tight ball against surface.
TEETH Upper jaw 58–80; lower jaw 62–76.
GROUP SIZE AND STRUCTURE Small, relatively fluid, sub-groups of 2–10; aggregations of 30–100 typical (though
average off the UK is fewer than 10); large aggregations of up to 500 not uncommon; and exceptional records of up to
4,000. Group sizes tend to be larger during travel and social interaction.

ADULT
Yellow to ochre and
white patches often
visible from above

Distinct
median
notch

Flukes distinctively
swept back

DIVE Sequence Bubbles often appear before head breaks surface; briefly shows much of beak, head and eyes;
yellow and white patches often visible simultaneously; strongly arches back. • Depth Unknown, but likely fairly
shallow. • Duration Usually less than one minute; maximum recorded four minutes.

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 ADULT
Upperside and all
Mid-sides Complex, sharply appendages dark Tall, prominent, falcate dorsal fin
medium to demarcated grey to black (when midway along back (up to 33cm
light grey colour pattern surfacing, appears tall – typically more than 12 per
to have Stenella-like cent of total body length)
Short, stubby cape)
beak (5cm or less) Robust body Tip usually pointed
well defined from
melon by distinct
crease Fin uniformly dark
grey to black

Gently Narrow yellow

sloping Narrow, bright white patch to ochre patch
melon Black eye- (from below dorsal fin to (from below
patch midway along tailstock) trailing edge of
dorsal fin to rear
tailstock)

Moderately broad,
pointed, sickle-
Beak dark grey shaped flippers Distinctively deep tailstock
to black above, with strong dorsal and
white below ventral keels (markedly more
(may be some May be 8–12 small pronounced in adult male)
ochre colouring tubercles along Underside and
on underside) leading edge lower sides
predominantly
white (to
urogenital area)
Thin black line Oblique medium to light
from beak to eye grey stripe from rear
patch (variable) margin of lower jaw to
leading edge of flipper
SIZE
L: ♂ 2.2–2.7m, ♀ 2–2.5m;
WT: 170–230kg; MAX: 2.8m, 235kg
Calf – L: 1–1.2m; WT: c. 24–30kg

80°

AT A GLANCE Cold temperate to sub-Arctic

70°
?
waters of the North Atlantic • Small size
? • Complex, sharply demarcated colour pattern
60° • Yellow to ochre patch on tailstock • Bold,
bright white patch on side • Very tall, pointed,
50°
falcate dorsal fin midway along back • Short,
? ? ?
stubby beak • Distinctive dorsal and ventral
keels on tailstock • Often lively and acrobatic
40°

30°

? uncertain range

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 PACIFIC WHITE-SIDED DOLPHIN
Lagenorhynchus obliquidens Gill, 1865

The Pacific white-sided dolphin is remarkably lively and energetic, repeatedly leaping high out of the water and doing a
variety of somersaults, backflips, spins and cartwheels. A large school of these gregarious dolphins often throws up so
much spray that their splashes can be seen long before the dolphins themselves.
IUCN status Least Concern (2018).
Population Possibly in excess of 1 million. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy The genus Lagenorhynchus is under revision, and this species may be moved to a different genus
(Sagmatias). No recognised subspecies, but there may be as many as six geographical forms (indistinguishable in the
field). There are also a number of uncommon anomalous colour patterns or ‘morphs’, including all-black and largely
all-white (though not albinistic) individuals; the commonest is the ‘Brownell type’.
Other names Affectionately called a ‘lag’ by researchers (from the generic name).
DISTRIBUTION Found in a continuous band across cool temperate waters of the North Pacific and some adjacent seas.
Deep oceanic waters, usually within 200km of the coast, but also in nearshore waters where deeper water approaches
closer to shore.
BEHAVIOUR Highly acrobatic, especially while travelling, with single leaps more common while feeding or socialising.
Breaches may include side-slaps and belly-flops, and it will also flipper-slap and tail-slap. Often seen in association
with other marine mammals. Can be extremely inquisitive and an avid bow-rider and wake-rider.
FOOD AND FEEDING Small schooling fish and cephalopods; occasionally shrimps. Large herds c­ ooperatively corral
schools of fish into bait balls close to the surface.
TEETH Upper jaw 46–72; lower jaw 46–72.
GROUP SIZE AND STRUCTURE Highly gregarious, typically in herds of up to 100, but sometimes up to several thousand.
Large herds often segregate into sub-groups according to age and sex.
Distinctive, light grey ‘suspender’
ADULT stripes particularly conspicuous
Flukes dark grey
on both sides

Median
notch

Slightly concave
trailing edge

Flippers may have

paler patch on SIZE
upper surface L: ♂ 1.7–2.5m, ♀ 1.7–2.4m;
WT: 90–170kg; MAX: 2.5m, 198kg
Calf – L: 90–110cm; WT: c. 15kg

DIVE Sequence Typically surfaces quite fast; may produce Dall’s porpoise-like rooster tail of spray; may cut through
water with just dorsal fin showing (shark-like). • Depth Offshore populations pursue fish found at depths of 500–
1,000m; coastal populations mostly eat surface-schooling prey. • Duration Average 24 seconds, with longer dives
rarely more than 3 minutes; maximum 6.2 minutes.

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 Tall, prominent, strongly falcate ADULT
Complex grey, white to lobate (broadly rounded)
Short, stubby beak well and black colour pattern dorsal fin midway along back
defined from melon by (highly variable)
Dark grey to (up to 28cm – typically up to
shallow crease
black upperside 12 per cent of total body length)

Rostrum, Robust body Fin more hooked Dorsal fin strikingly

beak tip in older males bicoloured (leading edge
and ‘lips’ Mid-grey eye- dark grey, c. two-thirds of
dark grey ring stretches to posterior portion light grey
to black corner of beak to white – highly variable)
Light grey
‘suspender’
stripes Older male may
have extensive
scarring

Narrow, dark
grey to black No significant
stripe from Brilliant white keels
beak to flipper underside (to
Large sickle- Distinct black border
Light grey shaped flippers between white underside urogenital area)
thoracic patch and light grey sides

Wide, pure white stripe

above eye runs back towards ADULT BROWNELL MORPH
urogenital patch (highly variable)

Less distinct (or absent)

Very dark black border between white
thoracic patch underside and light grey sides

75°

AT A GLANCE Cool temperate waters of the

60° North Pacific • Small size • Tall, prominent,
strikingly bicoloured dorsal fin • Complex grey,
45°
white and black coloration • Brilliant white
underside • Pale grey thoracic patch • Light
30° grey ‘suspender’ stripes along back • Short,
stubby beak • Acrobatic and demonstrative
• Tends to approach boats
15°

0°

15°

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 DUSKY DOLPHIN
Lagenorhynchus obscurus (Gray, 1828)

‘Duskies’ are so acrobatic that at least some animals in a large school are likely to be in mid-air at any one time –
performing the extraordinary high leaps and somersaults for which they are so well known. This is the best studied ‘lag’
– an affectionate term used by researchers for the six dolphins in the genus Lagenorhynchus.
IUCN status Least Concern (2018). Chilean/Peruvian subspecies Vulnerable (2019).
Population Unknown, though it appears to be abundant in most parts of its range. Global trend unknown (Chilean/
Peruvian subspecies decreasing).
Classification Odontoceti, family Delphinidae.
Taxonomy The genus Lagenorhynchus is under revision, and this species may be moved to a separate genus
(Sagmatias). Three subspecies are currently recognised (but probably indistinguishable in the field): Argentinian dusky
dolphin or Fitzroy’s dolphin (L. o. fitzroyi); African dusky dolphin (L. o. obscurus); Chilean/Peruvian dusky dolphin (L. o.
posidonia). The New Zealand dusky dolphin may be a fourth, unnamed subspecies (possibly L. o. superciliosis).
Other names Affectionately called a ‘lag’ by researchers (from the generic name).
DISTRIBUTION Seven apparently disjunct populations over a discontinuous range in cool temperate waters of the southern
hemisphere. The populations are: New Zealand (including the Chatham and Campbell Islands); southern and central South
America; southwestern Africa; Amsterdam and St Paul Islands; Prince Edward, Marion and Crozet Islands; Tristan da
Cunha and Gough Island; and southern Australia (including Tasmania). Associated with cool upwelling areas and cold
currents. Mainly coastal, it prefers shallow waters (less than 500m, typically less than 200m) and is found predominantly
over the continental shelf. Prefers sea surface temperatures of 10–18°C, but will venture into colder waters.
BEHAVIOUR Highly acrobatic and frequently leaps high out of the water multiple times. Associates with a wide variety
of other cetaceans. Generally approachable and curious around boats; frequently bow-rides.
FOOD AND FEEDING Wide variety of schooling fish and squid. Large groups cooperate to feed on schooling fish.
TEETH Upper jaw 52–78; lower jaw 52–78.
GROUP SIZE AND STRUCTURE Ranging from 2–1,000 or more (occasionally 2,000), with group size and structure
varying greatly according to season, activity, prey and location. Fission-fusion dynamics is the norm, in which groups
frequently fluctuate in size and composition.

Narrow, light grey tailstock blazes (‘suspenders’)

ADULT stretch to mid-body (highly variable – nearly to
blowhole in some individuals)

Distinct,
narrow
median
notch

Flippers pale grey

with darkened edges Concave trailing edges

DIVE Sequence When swimming slowly, beak tip breaks surface first and tailstock often barely visible; when
swimming quickly, porpoises with neat re-entry or cuts through water with only dorsal fin visible (shark-like). •
Depth Maximum recorded 156m. • Duration Non-foraging dives average c. 21 seconds, foraging dives more than
90 seconds.

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 Predominantly dark grey ADULT
Gently to bluish-black upperside
Tall, prominent, falcate
sloping dorsal fin (up to 28cm)
Short, dark forehead Moderately slightly behind mid-body
grey to black Large, pale grey robust body
beak (clearly thoracic patch extends
demarcated onto most of face Pointed tip
from melon
by shallow Male dorsal fin more curved
Small grey and with broader base
crease) to dark grey
eye patch
Dorsal fin variably bicoloured
(dark grey to black leading edge,
light grey trailing edge)

Pale grey patch on tailstock

splits into two blazes (longer
Beak narrower White underside dorsal blaze variable –
Dark ‘lips’
and slightly extends to just in front of
Small flippers curved
longer than in dorsal fin or beyond)
on leading edge
Pacific white-
sided dolphin
RARE HYBRID WITH SOUTHERN RIGHT WHALE DOLPHIN
(sharing characteristics with both species)
Sharp dividing line between black
upperside and white underside Small dorsal fin
two-thirds of the
way along back

SIZE
Black upperside does not extend L: ♂ 1.7–2m, ♀ 1.7–2m;
below eyes (unlike in southern WT: 70–85kg; MAX: 2.1m, 100kg
right whale dolphin) Calf – L: 80–100cm; WT: c. 9–10kg

80°

70°

60°

50°

40°
AT A GLANCE Cool temperate waters of
30° southern hemisphere • Small size • Complex
black, white and grey coloration • Pale grey
20°
10°

face and thoracic patch • Two forward-

0°
10°

pointing, pale grey side blazes • White

20°

30°

40°
? underside • Tall, prominent, bicoloured dorsal
50°

60°
fin • Gently sloping forehead • Short, dark
70°
beak • Gregarious and highly acrobatic
L. o. fitzroyi L. o. obscurus
L. o. posidonia L. o. posidonia secondary range
unnamed subspecies unnamed subspecies secondary range

DUSKY DOLPHIN 191

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 HOURGLASS DOLPHIN
Lagenorhynchus cruciger (Quoy and Gaimard, 1824)

Uniquely, the hourglass dolphin was formally described – and accepted as a valid species – solely on the basis of rough
drawings made at sea in 1820. It is often seen, particularly in the Drake Passage, but there have been few specimens to
study and it remains one of the least known of all the dolphins.
IUCN status Least Concern (2018).
Population The only abundance estimate is 144,300 south of the Antarctic Convergence. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy The genus Lagenorhynchus is under revision, and this species may be moved to a separate genus
(Sagmatias); no recognised forms or subspecies.
Other names As with all Lagenorhynchus dolphins, affectionately called a ‘lag’ by researchers (from the generic name).
DISTRIBUTION Circumpolar in both Antarctic and sub-Antarctic waters. Most sightings are from 45–65°S. Occurs on
both sides of the Antarctic Convergence. Closely associated with the Antarctic Circumpolar Current, it is most often
seen in areas with turbulent seas. Most sightings are in water colder than 7°C. Found near the ice edge in some areas.
Mainly deep offshore waters but sometimes in depths less than 200m near islands and banks.
BEHAVIOUR Hourglass dolphins often associate with fin whales (to such a degree that whalers used them to find
whales) and, less often, with other cetaceans. Seem to be attracted to ships and will often change course and approach
from a considerable distance to bow-ride enthusiastically, or surf in the wake. They appear so quickly that they are
easily overlooked as they slip onto the bow. Active, generally fast swimmers, especially enjoying wave-surfing in high
seas, they often breach.
FOOD AND FEEDING Small fish, squid and crustaceans. Often forages in association with great shearwaters, black-
browed albatrosses and other seabirds.
TEETH Upper jaw 52–68; lower jaw 54–70.
GROUP SIZE AND STRUCTURE Typically 1–12; groups of up to 100 have been reported.

Dorsal fin rarely has sharp ADULT FEMALE

backward bend (cf. male)
Some individuals (both sexes) have
patches of brownish-grey on sides Dorsal fin has more
as well as black and white pointed tip

Dorsal fin smaller and

more strongly falcate

Keel on underside of tailstock

less prominent than on male

DIVE Sequence When swimming slowly, back and dorsal fin appear while head still visible; gentle roll to dive,
flukes perhaps appearing (or slapped onto surface); when swimming fast and high, leaps at long, low angles then
swims rapidly just below surface (penguin-like); when swimming fast and low, travels very close to surface with
only top of head and dorsal fin visible; produces distinctive ‘rooster tail’ of spray (similar to that made by Dall’s
porpoise). • Depth Prey choice suggests that it feeds mainly in surface waters. • Duration Unknown.

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 Dark 'eye spot' Predominantly black with ADULT MALE
outlined with variable contrasting ‘hourglass’ Sharp backward
thin white line pattern of wide white bend halfway up
lateral streaks (variable) Tall, strongly falcate dorsal
fin midway along back
Short, stubby Forward white (larger and more markedly
but well- streak extends Robust body hooked than in female)
defined black onto face
beak above eye Dorsal fin variable
shape but always black
Melon Rounder tip
black than in female

Flukes black
on both sides

Dark ‘lips’ Underside Keel on

predominantly underside of
Thin black line from Long, curved tailstock
eye to corner of white
Flippers black on flippers with
mouth concave trailing Thin white line Distinctive hook-shaped
upperside (partially
edge between two lateral white mark below rear
white underside)
streaks (variable) lateral patch

ADULT

SIZE
L: ♂ 1.6–1.9m, ♀ 1.4–1.8m;
WT: 70–90kg; MAX: 1.9m, 94kg
Calf – L: c. 0.9–1.2m; WT: unknown

80°

70°

60°

50°

40°
AT A GLANCE Oceanic waters in sub-
Antarctic and Antarctic • Small size • Sharply
30°

20°

demarcated black-and-white coloration •

10°
0°

Tall, falcate, black dorsal fin • Porpoises

10°
20°

like a penguin when swimming fast • Eager

30°

40°

50°

60°
bow-rider
70°

80°

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 WHITE-BEAKED DOLPHIN
Lagenorhynchus albirostris (Gray, 1846)

Despite their name, not all white-beaked dolphins have white beaks – many are actually quite dark or flecked. The
species’ Greenlandic name (aarluarsuk) means ‘killer whale look-alike’.
IUCN status Least Concern (2018).
Population Minimum 100,000 (the sum of available abundance estimates). Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy The genus Lagenorhynchus is under revision: the white-beaked dolphin is the type species and could
ultimately be the only remaining member; no recognised forms or subspecies.
Other names Squidhound, jumper, springer; as with all Lagenorhynchus dolphins, affectionately called a ‘lag’ by
researchers (from the generic name).
DISTRIBUTION Cold temperate to ice-free polar waters of the North Atlantic. More common in European waters
than North American. Four areas of high density have been identified: the Labrador Shelf (including south-west
Greenland); Iceland; Scotland (including the northern Irish Sea and northern North Sea); and the northern coast of
Norway (extending north into the White Sea). Sometimes occurs up to the edge of the pack ice. Prefers sea surface
temperatures of 5–15°C. Mainly coastal, in water less than 200m deep, but also occurs in deeper, offshore waters (up
to 1,000m in the Barents Sea and off West Greenland).
BEHAVIOUR Acrobatic, frequently leaping out of the water and performing a range of aerial behaviours. Can be quite
elusive in some areas, but in others frequently approaches boats from a distance to bow-ride and jump in the wake.
FOOD AND FEEDING Pelagic schooling and benthic fish; may also take squid, octopus and benthic crustaceans. Feeds
alone deep underwater and cooperatively to herd fish against the sea surface.
TEETH Upper jaw 46–56; lower jaw 44–56. First three teeth in each row often concealed within the gum.
GROUP SIZE AND STRUCTURE Usually 5–30; average 9 in Iceland, 6 in Svalbard, 4–6 in Denmark. Rarely seen alone.
Groups of several hundred known, especially offshore.

Rear light band may rise ADULT

to join band on other side Flukes grow
and form pale saddle proportionately
larger with age

Flukes dark grey on both

SIZE sides (may be speckled
L: 2.4–3.1m; with white on underside)
WT: 180–275kg; MAX: 3.2m, 354kg
Calf – L: 1.1–1.3m; WT: c. 40kg

DIVE Sequence When travelling fast, tends not to porpoise cleanly out of water – skims over surface, producing
distinctive ‘rooster-tail’ spray; when swimming slowly: head, back and top of beak appear above surface, then
dorsal fin appears before it rolls gently to dive. • Depth Unknown; one Icelandic individual reached 45m.
• Duration Very little information; average in Iceland 24–28 seconds; maximum recorded 78 seconds.

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 Tall, dark, strongly falcate Tip pointed and ADULT
Very robust body dorsal fin in middle of back often hooked
(especially male) (more prominent in male
Shallow crease and with age) May be off-white
between beak Upperside patch on dorsal fin
and melon mostly black Blazes on sides and
Short, to dark grey saddle vary extensively in
thick beak width and extent
(5–8cm
long) Distinctive band of light grey
along length of body from
behind blowhole to tailstock
(variable, with indistinct border)

Beak Falcate Underside mostly

variable Long, pointed, trailing white to pale grey
Many individuals have
colour dark grey flippers edge
Large dark medium grey patch on
(up to 19 per cent
May have patch above chest and abdomen (with
of body length)
dark or light and behind variable longitudinal white
‘speckles’ May have dark grey flipper line down middle)
between eye stripe between corner
and flipper of mouth and flipper BEAK VARIATIONS
White mottled with dark
grey and with pinkish tip Percentages based on
Dark grey with (32 per cent) studies in Iceland (i.e.
white tip
eastern Atlantic)
(52 per cent)

White mottled
with dark grey
and with white
tip (9 per cent)

Ashy-grey or dark grey (paler

than head) with no white
Pure white
or pinkish tip (especially in
(7 per cent)
western Atlantic)

80°

70°
AT A GLANCE Cool waters of the North
Atlantic • Small size • Complex, diffuse (and
variable) grey, black and white coloration
• Distinctive band of light grey along side
60°

50°
• Very robust body • Greyish-white ‘saddle’
behind dorsal fin • Short, thick beak (often
40° white) • Tall, dark, falcate dorsal fin
30°

primary range possible range

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 PEALE’S DOLPHIN
Lagenorhynchus australis (Peale, 1848)

With complex markings broadly similar to those of dusky and Pacific white-sided dolphins, the poorly known Peale’s
dolphin is best identified by its distinctive dark face.
IUCN status Least Concern (2018).
Population Unknown. Surveys of the Patagonian shelf (c. 1,300km along southern Argentina) and the Falkland Islands
estimated c. 19,924 and 1,896 individuals respectively. Appears to be stable.
Classification Odontoceti, family Delphinidae.
Taxonomy The genus Lagenorhynchus is under revision, and this species may be moved to a separate genus
(Sagmatias); no recognised forms or subspecies.
Other names Plough-share dolphin; as with all Lagenorhynchus dolphins, affectionately called a ‘lag’ by researchers
(from the generic name).
DISTRIBUTION Cold temperate and sub-polar waters along both coasts of southern South America, extending east
to the Falkland Islands. Occurs in two different habitats: predominantly protected bays, channels and fjord entrances
in southern Chile; and open, wave-washed coasts and over shallow continental shelves in northern Chile (north of
Chiloé) and throughout most of its range in Argentina. It occurs in both habitats around the Falkland Islands, where it
frequently forages in coastal kelp beds (but also has a preference for the 100m isobaths further offshore). Frequently
within sight of land and often in less than 20m of water. However, it does occur as far as 300km offshore in some areas
(though rarely in water deeper than 200m – maximum 300m – and abundance decreases with greater depth). Occurs
mainly where the sea surface temperature is c. 5–15°C.
BEHAVIOUR Ponderous and slow-moving much of the time, swimming quite inconspicuously, but prone to bursts
of activity. Often observed in mixed-species groups with other dolphins. Frequently plays in coastal surf and may
breach repeatedly. Often slaps its head, flukes or flippers onto the surface, and sometimes spyhops. Commonly and
energetically bow-rides. Will also wake-ride.
FOOD AND FEEDING Variety of fish, cephalopods and crustaceans. Forages in kelp beds (where it picks small octopuses
off fronds) and open waters.
TEETH Upper jaw 54–74; lower jaw 54–72.
GROUP SIZE AND STRUCTURE Usually 4–5 (ranging from 1–30); aggregations of up to 100 rarely observed.

Light spinal blaze ADULT

fades into black on
Rounded tips
back near blowhole

Distinct
median
notch

Flukes dark
on both sides

DIVE Sequence When swimming quickly, mostly hidden behind wall of

spray as water splashes up high around face (thus nickname ‘plough-share
dolphin’); groups usually surface simultaneously; may make long, low leaps;
when swimming slowly, only blowhole, small part of back and dorsal fin
exposed. • Depth Unknown, but feeds mostly near bottom in shallow
water. • Duration Average 28 seconds (ranges from 3–157 seconds).

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 Both light-coloured Single pale grey Dark grey to Tall, falcate dorsal ADULT
Predominantly patches may be spinal blaze or black upperside fin (variable)
dark grey to flecked with ‘suspender’ midway along back
black face darker grey
Robust body Pointed tip
(variable)
Darker double Fin somewhat two-toned:
eye-rings (not dark grey to black with
always visible) poorly defined paler grey
trailing edge
Pale grey to
white patch on
tailstock

Short,
stubby Snowy-white
beak underside
Underside white
Recurved, area extends onto
Tip of beak, ‘lips’ Lower edge of
pointed flanks, producing Large pale grey
and lower jaw thoracic patch
flippers white ‘armpit’ thoracic patch
dark grey to black usually bordered
above flippers
with narrow dark
Black throat (extent Flippers dark grey line (variable)
highly variable) to black with lighter
trailing edge
Complex colour pattern
ADULT VARIATION individually variable

DORSAL FIN VARIATIONS

SIZE
L: ♂ 1.4–2.2m, ♀ 1.3–2.1m;
MAX: 115kg (no informaton on
weight range)
Calf – L: 1–1.3m; WT: unknown

10°

0°

10°

20°
AT A GLANCE Shallow waters of southern
South America • Often associated with kelp
30°
• Small size • Stocky body • Indistinct beak
40° • Complex grey, black and white coloration •
Dark mask-like face • Tall, falcate dorsal fin
50°

60°

possible range

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 CHILEAN DOLPHIN
Cephalorhynchus eutropia (Gray, 1846)

Found in shallow waters along the southwestern coast of South America, the porpoise-like Chilean dolphin is poorly
known and seems to be uncommon.
IUCN status Near Threatened (2017).
Population Unknown, but it appears to be relatively uncommon (low thousands at most). Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Black dolphin.
DISTRIBUTION Unevenly distributed along 2,600km of the central and southern Chilean coast; there have also been a
small number of vagrants in Argentina. Most sightings are within 500m of the coast (though there has been little survey
effort in adjacent offshore waters). Found in Chile’s intricate network of fjords, channels and sheltered bays, as well as
estuaries and rivers (as far as 12km upstream). Prefers shallow water of 3–15m (rarely deeper than 30m), particularly
in areas with a high tidal range and rapid tidal flow, near rivers or over shallow banks at fjord entrances. Often seen in
the surf zone.
BEHAVIOUR Can be quite active, porpoising and occasionally breaching. Tends to be shy and elusive around boats in
the southern parts of the range – perhaps a learned response to hunting – but some groups approach boats in the north
and bow-ride or surf in the wake.
FOOD AND FEEDING Small benthic and pelagic schooling fishes, as well as squid, octopuses and crustaceans. Some
evidence of cooperative feeding.
TEETH Upper jaw 58–68; lower 58–68.
GROUP SIZE AND STRUCTURE Usually 2–3, often 4–10, sometimes up to 15; occasionally as many as 50 (probably the
merging of several small groups) especially in the north. Exceptionally large aggregations of c. 400 have been reported
along the open coast north of Valdivia.

Main white patch on belly of ADULT

male splits into three arms on
White chin tailstock, similar to that of killer
whale (female mostly white)

Both sexes have a thin, oval,

dark grey patch in urinogenital
White patches separated by area (broader in female)
medium to dark grey band
stretching across chest
between flippers
SIZE
L: ♂ 1.2–1.7m, ♀ 1.2–1.7m;
WT: 30–60kg; MAX: 1.7m, 63kg
Calf – L: c. 0.9–1m; WT: c. 8–10kg

DIVE Sequence Tip of rostrum appears first, then melon; body rolls relatively high as rounded dorsal fin appears;
flukes rarely visible; also makes vertical leaps, usually re-entering water headfirst, with little splash.
• Depth Probably less than 30m. • Duration Usually less than three minutes.

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 ADULT
Slightly pointed, conical Mostly mid-grey to
Stocky body (girth
head (more pointed than brownish-grey (may
can be nearly two- Distinctly rounded dorsal
other Cephalorhynchus seem black at distance)
thirds of body length) fin just behind middle of
dolphins)
back (variable shape)

Dark grey band Dorsal fin blackish

from blowhole to (contrasts with
above eye lighter body)
Light to
mid-grey Dorsal fin relatively
‘cap’ on low and leans
melon backwards

May have white ‘armpits’

just behind flippers (not
Short, always present or visible Male has finger-shaped white
stubby White throat patches (separated by grey
from side)
beak bands) on lower rear body

Small, dark, paddle-

Wide, dark grey shaped flippers with
Mid-grey lower jaw band from around rounded tips
joins dark band to eye to flipper
ADULT
flippers
Pointed tips

Distinct
median
notch

Dark bands from behind

blowhole to eye and Flukes concave
onto rostrum on trailing edge

10°

0°

10° AT A GLANCE Mainly central and southern

Chile • Shallow coastal waters • Small size
20°
• Prominent, dark, distinctly rounded dorsal
30° fin • Complex shades of grey and white •
Finger-shaped white band on lower rear body
40° • Indistinct beak • Small groups

50°

CHILEAN DOLPHIN 199

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 COMMERSON’S DOLPHIN
Cephalorhynchus commersonii (Lacépède, 1804)

Commerson’s dolphin has the strangest distribution: the main stronghold is in southern South America and around the
Falkland Islands, but there is also an isolated population in the Kerguelen Islands, 8,500km away in the Indian Ocean.
Suitable habitat is lacking in between, so interchange between them seems unlikely.
IUCN status Least Concern (2017). Subspecies kerguelensis has not been assessed separately.
Population Unknown, but probably one of the most abundant Cephalorhynchus species. Estimated at least 40,000 in
Argentina and c. 22,000 for the Patagonian shelf. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy Two subspecies are recognised: South American Commerson’s dolphin (C. c. commersonii) in southern
South America, and Kerguelen Islands Commerson’s dolphin (C. c. kerguelenensis) in the French Southern and Antarctic
Territory of Kerguelen; there is growing evidence that animals in the Falkland Islands may be another subspecies.
Other names Black-and-white dolphin, piebald dolphin, skunk dolphin.
DISTRIBUTION The nominate subspecies (commersonii) prefers sheltered habitats such as fjords, narrow passages,
bays, harbours and river estuaries, and where there are strong currents and/or a great tidal range. Nearly always seen
in green water within sight of land, typically in depths less than 200m (sometimes inside coastal breakers, as shallow
as 1m – at home in turbulent waters close to shore). Often seen in the surf zone. Attracted to kelp beds. The subspecies
kerguelenensis is restricted to the immediate vicinity of the Kerguelen Islands, and prefers open waters, kelp-lined
coastlines and protected areas between islets.
BEHAVIOUR Quick, active and playful, Commerson’s dolphin seems to delight in surfing and darting through breakers,
and often swims upside down or spins on its longitudinal axis underwater. Strongly attracted to boats, it readily bow-
rides and wake-rides.
FOOD AND FEEDING Wide variety of fish, squid, octopuses, small crustaceans, marine worms, other benthic
invertebrates; kerguelenensis has a more restricted diet – mostly fish (especially mackerel icefish). Single animals
forage on seabed, in kelp forests and in tidal areas; large groups coordinate to herd schooling fish against surface or
shore.
TEETH Upper jaw 56–70; lower jaw 56–70.
GROUP SIZE AND STRUCTURE Usually 1–10; sometimes up to 15; aggregations of 100+ tend to be short-lived (10–30
minutes). Solitary individuals and small groups (2–4) are more common in the north, larger groups more common in the
south.

Tips rounded
ADULT

Small
median
notch

‘Widow’s peak’ (fraying of border between

black and white behind blowhole) highly Concave trailing
variable (poorly defined or absent in Kerguelen) edge

DIVE Sequence When swimming slowly, rolls forward leisurely as dorsal fin appears, then head submerges and
fin disappears; when swimming rapidly, porpoises out of water with low, horizontal leaps. • Depth Unknown.
• Duration Typically 20–30 seconds.

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 Basically white with Low, fairly large, rounded dorsal ADULT SOUTH AMERICAN
Dorsal fin shape
black face, flippers, fin (sometimes described as like
highly variable
Gently dorsal fin, narrow cape, a ‘Mickey Mouse ear’) slightly
sloping rear tailstock and flukes Robust body behind middle of back
Dorsal fin rises
forehead at shallow angle
Conical and leans slightly
head backward

Slightly convex
trailing edge

Small flippers with

Indistinct White rounded tips White areas in Falkland
beak throat animals frequently
May have saw-toothed serrations on leading more dusky grey
Relatively straight edge of flippers (especially left flipper)
mouthline slants up invisible at sea (not present in all individuals)
towards eye
ADULT KERGUELEN
Markedly Dark areas tend to be dark
Coloration may grey (rather than black)
resemble young South larger
American animals Light areas with greyer tinge (rather
May be subtle, than white) but highly variable
diffuse paler
patch on top Dark areas may be
of melon more extensive
Borders between
dark and light
less distinct

Slightly more Underside

well-defined Flippers more tapered
pure white
beak
White throat SIZE – SOUTH AMERICAN SIZE – KERGUELEN
tends to be L: ♂ 1.2–1.4m, ♀ 1.3–1.5m; L: ♂ 1.4–1.7m, ♀ 1.5–1.7m;
less extensive WT: 25–45kg; MAX: 1.5m, 66kg WT: 30–50kg; MAX: 1.74m, 86kg
and more Calf – L: 65–75cm; WT: 4.5–8kg Calf – L: 65–75cm; WT: 4.5–7kg
asymmetrical

70°

60°

50°

40°

30°

20°
AT A GLANCE Southern South America
10° and Falkland Islands • Small population in
Kerguelen Islands • Shallow, coastal waters
0°

10°

20°

30°
• Small size • Porpoise-like appearance •
40° Low, rounded dorsal fin • Sharply demarcated
50°
black and white • Fast and active • Likely to
60°
approach boats
70°

South American subspecies secondary range

Kerguelen Islands subspecies

COMMERSON’S DOLPHIN 201

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 HEAVISIDE’S DOLPHIN
Cephalorhynchus heavisidii (Gray, 1828)

Endemic to the Benguela ecosystem along the southwestern coast of Africa, this beautiful small dolphin is fond of
surfing and can often be seen playing in the waves close to shore.
IUCN status Near Threatened (2017).
Population Unknown, but total of two key regional estimates (6,345 from Table Bay to Lambert’s Bay, southern South
Africa, and 1,594 in the Namibian Islands’ Marine Protected Area, southern Namibia) suggests c. 10,000. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Haviside’s dolphin, Benguela dolphin.
DISTRIBUTION Restricted to cool waters of the Benguela ecosystem along 2,500km of coastline in southern Angola,
Namibia and South Africa, ranging from at least 16°30’S (southern Angola) to 34°20’S (South Africa). Tends to stay in
water less than 100m deep, but regularly ranges up to 30km from shore. It is generally closest to shore between sunrise
and midday – preferring areas with sandy shores and big swells (especially in the breakers) – and furthest offshore
between mid-afternoon and sunrise.
BEHAVIOUR Energetic and sometimes boisterous. Fond of surfing and can often be seen playing and jumping in the
waves. Performs a variety of leaps (often in competitive pairs or small groups). Readily approaches boats, frequently
bow-rides and will sometimes play in the wake.
FOOD AND FEEDING Mainly demersal (some pelagic) fish; will take squid and octopus. Most feeding at night.
TEETH Upper 44–56 jaw; lower jaw 44–56.
GROUP SIZE AND STRUCTURE Usually 2–3, but typically ranges from 1–10; larger aggregations of multiple groups
numbering as many as 100 individuals can occur in some high-density areas.

ADULT
Narrow band from
dark cape extends
as far as blowhole
Pointed tips

Distinct
median
notch

SIZE
L: 1.2–1.7m; Flukes may be slightly
WT: 50–75kg; MAX: 1.8m, 75kg concave on trailing edge
Calf – L: 80–85cm; WT: unknown,
but possibly c. 10kg

DIVE Sequence Head appears first, closely followed by tip of dorsal fin; body rolls relatively high in water; head
submerges and dorsal fin disappears; flukes rarely visible; when swimming quickly, often clears surface at low
angle; also makes vertical leaps, usually with headfirst re-entry (little splash). • Depth Forages mostly on or near
seabed in relatively shallow water (less than 100m). • Duration Unknown.

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 Leading edge usually ADULT
longer and more convex
Stocky body Large, broad-based,
than trailing edge
Light grey to (girth can be roughly triangular
mid-grey head nearly two-thirds dorsal fin in middle
and thorax of body length) of back
Distinctive dark
Darker around grey to bluish-
eyes (may black cape (extent
extend over variable)
much of face)

Cone-shaped
head

White ‘armpits’ just Areas between

behind flippers (not finger-shaped
always visible from side) marks light grey
Short, to bluish-black
Finger-shaped
stubby White patch white band on
beak on chest Irregular serrations Small, slender flippers lower rear body
between on leading edge of with blunt tips
Lower jaw flippers (invisible
projects flippers
at sea)
slightly beyond ADULT
tip of snout
White ‘armpits’ just
behind flippers

Shapes of pure white patterns vary Main white patch on underside

White patch on chest slightly between sexes (in male, central splits into three arms or
between flippers finger of white is pointed and extends 'tridents' on tailstock (similar
beyond anus – in female, it is shortened/ to that of killer whale)
squared off and stops at anus)

30°

20°

AT A GLANCE Atlantic coast of southern

Africa • Small size • Complex black, grey and
10°

0°
white body pattern • Finger-shaped white
marks on lower rear body • Large, wide-
10° based triangular dorsal fin in middle of back
• Indistinct beak • Small group size • Very
20° energetic

30°

HEAVISIDE’S DOLPHIN 203

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 HECTOR’S DOLPHIN
Cephalorhynchus hectori (Van Bénéden, 1881)

One of the smallest dolphins in the world, Hector’s dolphin is found only in New Zealand. Its numbers and range have
declined dramatically in the past 30 years, and now one of the two subspecies – Māui dolphin (previously known as
North Island Hector’s dolphin) – is on the brink of extinction.
IUCN status Endangered (2008). Māui subspecies Critically Endangered (2008).
Population South Island Hector’s dolphin: c. 15,000; Māui dolphin: c. 60. In the mid-1970s there were c. 50,000
Hector’s dolphins altogether (including c. 2,000 Māui dolphins). Populations of both subspecies declining.
Classification Odontoceti, family Delphinidae.
Taxonomy Two subspecies are recognised: South Island Hector’s dolphin (C. h. hectori) and Māui dolphin (C. h. maui);
they look identical (Māui is marginally larger) but are genetically distinct.
Other names New Zealand dolphin.
DISTRIBUTION Endemic to New Zealand: mainly South Island with a very small population on the west coast of North
Island (mainly between Manukau Harbour and Port Waikato). There are occasionally sightings off the east coast of
North Island (subspecies unknown). Typical habitat is in murky water (less than 4m visibility) and in depths of less than
40m (though up to 100m), and within 40km of shore (the distribution of both subspecies is related to water depth, rather
than distance from shore). It is often seen just beyond the surf line or inside harbours. Recent research reveals some
movement of South Island individuals into the Māui range, but the current genetic isolation of the two subspecies is
largely maintained by geographical isolation and their typically small home ranges.
BEHAVIOUR Quite aerially active. Large groups are particularly boisterous, with lots of chasing, leaping, bubble-
blowing and vigorous tail slapping. Will also spyhop. Frequently surfs, especially in rough weather. Strongly attracted
to small boats and readily bow-rides.
FOOD AND FEEDING Wide variety of small fish; also squid and octopus. Feeds throughout water column, from surface
to seafloor; strongly attracted to inshore trawlers (groups of up to 50 may follow trawlers for several hours).
TEETH Upper jaw 48–62; lower jaw 48–62.
GROUP SIZE AND STRUCTURE Typically 2–10. Frequently several small groups form large, temporary aggregations of
c. 25 (sometimes up to 50 or more), mingling and separating again within 10–30 minutes – often swapping members in
the process. Individuals in groups are not related, and long-term associations are rare.
ADULT
Dark grey to black crescent-
shaped band arches across Dark grey to
head just behind blowhole black flukes

Small
median
notch

Deeply concave
Grey forehead trailing edge

DIVE Sequence Typically surfaces slowly and leisurely, with little or no splash; rolls slowly forward, then dorsal
fin disappears; in occasional fast travel, actively surfaces and splashes. • Depth Probably to 50m or more.
• Duration Longest dives typically c. 90 seconds or less.

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 May be single linear Low-profile, fairly large, broad- ADULT
scarring (tooth based, rounded, dark grey to black
Dark grey to black band Robust body
rake marks from dorsal fin (sometimes described as
extends forward from (girth can be up Leading
conspecifics) shaped like a ‘Mickey Mouse ear’)
flippers to encircle eyes to 68 per cent edge rises
Predominantly of total length) at shallow Rounded, convex trailing
light grey angle edge makes fin appear to
Conical head upperside lean backwards
and sides
Fin slightly behind
Indistinct middle of back
beak Dark grey to black area
around base of dorsal fin
(possibly remnant of what
was once a cape)

Sharp
Dark grey
demarcation
to black Small white patch Mainly white or creamy- between
beak tip behind flipper in white lower jaw, throat markings
Saw-toothed ‘armpit’ area and underside
Slender, finger-shaped
serrations on leading Rounded, dark (difficult to see) white projections
edge of flippers grey flippers Faint band
across mid-body extend from ventral
(difficult to see at sea) with blunt tips
(variable) patch onto lower sides

Dark grey area ADULT MALE

between flippers
(highly variable shape)
Mainly white Large, dark grey urogenital
underside patch on male (reduced or
absent on female) Female may be more
white on belly and
tailstock

White or creamy-white ventral area splits into

three finger-shaped projections behind umbilicus

SIZE – SOUTH ISLAND SIZE – MĀUI Underside of

L: ♂ 1.2–1.4m, ♀ 1.3–1.5m; L: ♂ 1.3–1.5m, ♀ 1.3–1.6m; flukes dark grey
WT: 35–50kg; MAX: 1.5m, 50kg WT: 40–60kg; MAX: 1.6m, 65kg to black
Calf – L: 60–70cm; WT: 8–10kg Calf – L: 60–75cm; WT: 8–10kg

North Island (Māui) subspecies

main concentration
isolated groups

AT A GLANCE Shallow coastal waters of

New Zealand • Small size • Mainly light grey
with dark appendages • White on belly •
Mostly dark face • Rounded dorsal fin leans
backward • Small groups
South Island subspecies
highest abundance
moderate abundance
isolated groups

HECTOR’S DOLPHIN 205

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 NORTHERN RIGHT WHALE DOLPHIN
Lissodelphis borealis (Peale, 1848)

With its striking black-and-white markings, slender body and no dorsal fin, the northern right whale dolphin is easily
identifiable within its range. The two species of right whale dolphins may look superficially similar, but their markings are
strikingly different and they are widely separated geographically.
IUCN status Least Concern (2018).
Population Believed to be low hundreds of thousands; one of the most abundant oceanic dolphins in the North
Pacific. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies; some populations are characterised by a ‘swirled’ colour morph
(which may resemble the southern right whale dolphin).
Other names Affectionately called a ‘lisso’ by researchers (from the generic name).
DISTRIBUTION Cool to warm temperate waters in the North Pacific, mainly from 31–50°N in the east and 35–51°N in
the west. Favours deep oceanic waters from the outer continental shelf and beyond, but also occurs where deep waters
approach the coast. Most abundant where sea surface temperatures are 8–19°C.
BEHAVIOUR Fast swimmer, capable of bursts of speed up to 34km/h. Frequently associates with at least 14 other
cetacean species. May erupt into bouts of high excitement, with much aerial activity such as breaching, spyhopping,
belly-flopping, and side- and fluke-slapping, and bursts of energetic swimming. Response to boats varies enormously: it
will bow-ride (especially in the presence of other bow-riding dolphins) but can be skittish and easily startled.
FOOD AND FEEDING Mainly fish, but some squid. Feeding techniques unknown.
TEETH Upper jaw 74–104; lower jaw 84–108.
GROUP SIZE AND STRUCTURE Highly gregarious: 100–200 common, but sometimes up to 2,000–3,000. Rarely seen
alone.
White band runs from
behind throat to fluke notch In female, white band
ADULT and widens on chest broadens slightly
around genitals

Flukes mostly white on underside

except for black trailing edge and
central patch around white band

SIZE
L: ♂ 2.2–2.6m, ♀ 2.1–2.3m;
WT: 60–100kg; MAX: 3.1m, 113kg
Calf – L: c. 0.8–1m; WT: unknown

DIVE Sequence When swimming quickly, makes graceful, ‘bouncing’, low-angled leaps (up to 7m) or splashy belly
flops, creating much surface disturbance; can appear almost eel-like; when swimming slowly, makes low-profile roll,
barely breaking surface to breathe (easy to miss). • Depth Probably capable of 200+m. • Duration 10–75 seconds;
maximum recorded 6.2 minutes.

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 Shallow crease Primarily Extremely ADULT
between beak jet black slender body
No dorsal fin Sharp
and melon
Black face or ridge demarcation
and beak Gently sloping between black
forehead and white

Extremely
narrow
tailstock

Straight Small, narrow,

Small but Irregular white band
mouthline White chest recurved flippers
distinct on underside (varies
with pointed tips considerably in extent)
beak Small white patch
just behind tip of
lower jaw

ADULT ‘SWIRLED’ COLOUR MORPH

Lower sides of White extends above More extensive

face may be and onto upper base white band rising
white of flippers onto lower sides

ADULT
Flippers black
Tiny flukes with
pointed tips

Median
notch

Crescent-shaped
patch of light grey
Concave trailing edge

75°

AT A GLANCE Deep temperate waters of

the North Pacific • Small size (appears even
60°

smaller at sea than measurements suggest)

45°
• No dorsal fin • Mainly black with white
band on underside • Extremely slender body
• Small but distinct beak • Low-angled leaps
30°

15° • Usually in sizeable groups

0°

15°

NORTHERN RIGHT WHALE DOLPHIN 207

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 SOUTHERN RIGHT WHALE DOLPHIN
Lissodelphis peronii (Lacépède, 1804)

Dolphins living in high latitudes tend to be just black and white, and the southern right whale dolphin is no exception.
With its striking black-and-white markings, slender body and complete lack of a dorsal fin, it is easy to recognise.
IUCN status Least Concern (2018).
Population Unknown, though considered fairly common throughout its range. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Mealy-mouthed porpoise.
DISTRIBUTION Cool temperate to sub-Antarctic waters in the southern hemisphere, mainly from c. 25–61°S. The range
extends furthest north along the west coasts of continents (due to the cold counterclockwise currents of the southern
hemisphere). The southern limit appears to be the Antarctic Convergence. Mainly in deep oceanic waters from the
outer continental shelf and beyond, but also where water deeper than 200m approaches the coast.
BEHAVIOUR Relatively fast swimmer. May erupt into bouts of high excitement, with much aerial activity such as
breaching, spyhopping, belly-flopping, and side- and fluke-slapping. Frequently in mixed groups with other cetaceans.
Response to boats varies, but it does not appear to be particularly attracted and only rarely bow-rides.
FOOD AND FEEDING Mainly fish and squid; possibly some krill. Feeding techniques unknown.
TEETH Upper jaw 78–98; lower jaw 78–98.
GROUP SIZE AND STRUCTURE Highly gregarious. Often in hundreds, sometimes up to 1,000.

ADULT Tiny flukes with

pointed tips

Median
notch

Flippers mostly
white (may have Concave
dark trailing edge) Variable, mostly white trailing
flukes, fading to black edge
along trailing edge
SIZE (white underside)
L: ♂ 2.2–2.9m, ♀ 2.1–2.6m;
WT: 60–100kg; MAX: 3m, 116kg
Calf – L: c. 1m; WT: unknown

DIVE Sequence When swimming quickly, makes graceful, ‘bouncing’, low-angled leaps or splashy belly flops,
creating much surface disturbance; rather penguin-like; when swimming slowly, makes low-profile roll, barely
breaking surface to breathe. • Depth Probably capable of 200m-plus (main prey at 200–1,000m). • Duration 10–75
seconds; maximum recorded 6.4 minutes.

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 ADULT
Sharply demarcated
jet black and white White extends high up sides
(producing S-shape (varies considerably in extent)
Shallow crease
along sides) Very slender body and may appear yellowish
between beak
(slightly more robust
and melon
than northern right
whale dolphin) No dorsal fin
Gently
sloping or ridge
Beak and Extremely
lower forehead
narrow
forehead tailstock
white

Small, narrow,
Straight
recurved flippers
mouthline
Small but with pointed tips
distinct May have dark
beak spots on beak FLIPPER VARIATIONS

RARE HYBRID WITH DUSKY DOLPHIN

(sharing characteristics with both species)

Sharp dividing line

between black upperside Small dorsal fin
and white underside two-thirds of the
way along back

Black upperside does not extend

below eyes (unlike in southern
right whale dolphin)

80°

70°

60°

50°

40°
AT A GLANCE Deep cold waters of the
southern hemisphere • Small size • No dorsal
30°

20°

fin • Strikingly black-and-white coloration

10°
0°

• Slender body • Small but distinct beak

10°
20°

• Mostly white face and beak • Low-angled

30°

40°

50°

60°
leaps • Usually in sizeable groups
70°

80°

SOUTHERN RIGHT WHALE DOLPHIN 209

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 AUSTRALIAN SNUBFIN DOLPHIN
Orcaella heinsohni Beasley, Robertson and Arnold, 2005

Until recently, the Australian snubfin dolphin was regarded as a form of Irrawaddy dolphin, but the two were separated
in 2005 – largely on the basis of skull morphology and genetics, but also external characters such as coloration, height
of the dorsal fin, and presence or absence of a dorsal groove.
IUCN status Vulnerable (2017).
Population Probably fewer than c. 10,000 mature individuals. Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names None.
DISTRIBUTION Poorly documented range in a narrow strip of shallow, protected coastal and estuarine tropical and
sub-tropical waters of the Sahul Shelf (part of the continental shelf extending from the northern coast of Australia to
New Guinea). Occurs throughout northern Australia – although the distribution is fragmented – and in the Kikori Delta
in the Gulf of Papua, southern Papua New Guinea. Most sightings are within 6km of shore, though up to 20km where
the water is shallow. Typically in water less than 18m deep, and some populations regularly enter water as shallow as
2m. Unlike the Irrawaddy dolphin, it does not regularly inhabit freshwater, but has been recorded up to 50km upstream
in some larger tidal rivers (and is rarely more than 20km from the nearest river mouth).
BEHAVIOUR Not particularly acrobatic and usually makes only low leaps when disturbed, socialising or swimming
against a strong current. Spyhopping, body-rubbing, rolling sideways and tail-slapping are sometimes observed. Groups
may suddenly become energetic, performing playful, splashy leaps. Can spit a narrow, well-directed jet of water 1–2m
into the air, usually in association with feeding (perhaps to herd small fish). Usually shy of vessels and not known to
bow-ride.
FOOD AND FEEDING Wide variety of fish, squid, octopus, cuttlefish and crustaceans. Feeds throughout water column to
seafloor; known to hunt cooperatively.
TEETH Upper jaw 27–42; lower jaw 29–37.
GROUP SIZE AND STRUCTURE Typically 2–6, but sometimes alone or in aggregations of up to 25.
Crescent-shaped blowhole ADULT
slightly to left of midline No shallow groove
(ends point forward) along back
(cf. Irrawaddy dolphin)

Median
notch

Neck more flexible than

in most other dolphins SIZE
(first two neck vertebrae L: ♂ 2.1–2.7m, ♀ 1.9–2.3m;
unfused) WT: 114–130kg; MAX: 2.7m, 133kg
Calf – L: c. 1m; WT: c. 10–12kg

DIVE Sequence Surfaces inconspicuously with little splash; rolls forward into dive, showing little or nothing of
tailstock; occasionally lifts flukes out of water before steep dive; rarely porpoises with low, horizontal, splashy
leaps. • Depth Unknown. • Duration Typically 30 seconds to three minutes; maximum recorded 12 minutes
(when disturbed).

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 ADULT
Subtle darker slate-grey
to brown dorsal cape (can
Small, triangular to
appear reddish in certain
slightly falcate dorsal
lighting), cf. Irrawaddy
fin (variable)
dolphin Rounded or slightly
Blunt, bulbous
Margin of cape pointed tip (variable)
head (may be
fairly straight
more bulbous in
older males)
Fin c. 10cm high
Distinct neck (c. 4 per cent of
No total length)
crease (behind
discernible
blowhole)
beak
Fin three-fifths of
the way along back

Lighter grey to
brownish sides
Convex
Older individuals may
leading Exceptionally Pale grey, brown,
Straight mouthline have extensive scarring
edge broad flippers whitish or pinkish
(angled up toward (from encounters with
eye – appears to underside Australian humpback
Rounded tip
be ‘smiling’) dolphins and large
sharks)

Body colour Fin shape

variable variable ADULT VARIATION

edian
ch

10°

0°

AT A GLANCE Coastal northern Australia and

10°
southern Papua New Guinea • Small size
• Subtle three-toned grey to brown coloration
20°
• Bulbous head with no beak • Small dorsal
fin behind midpoint of back • No dorsal
30°
groove • Cryptic, low surfacing • Usually in
small groups
40°

primary range possible range

AUSTRALIAN SNUBFIN DOLPHIN 211

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 IRRAWADDY DOLPHIN
Orcaella brevirostris (Owen in Gray, 1866)

Recently separated from the similar Australian snubfin dolphin, the Irrawaddy dolphin is superficially like a finless
porpoise with a dorsal fin. It is not classified as a ‘river dolphin’, yet is found in three large river systems (as well as
coastal waters).
IUCN status Endangered (2017). Mekong River sub-population Critically Endangered (2004). Malampaya Sound
sub-population Critically Endangered (2004). Iloilo-Guimaras sub-population Critically Endangered (2018). Mahakam
River sub-population Critically Endangered (2008). Ayeyarwady (Irrawaddy) River sub-population Critically Endangered
(2004); Songkhla Lake sub-population Critically Endangered (2004).
Population Regional estimates total c. 7,000. Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies (although freshwater sub-populations are possibly disjunct and new
subspecies designations may be warranted).
Other names Mahakam River dolphin, pesut.
DISTRIBUTION Poorly documented range in shallow, protected coastal, estuarine and fresh tropical and sub-tropical
waters of the Sunda Shelf (part of the Southeast Asian continental shelf). Prefers low-salinity waters. Fragmented into
relatively small populations, and absent from long stretches of coastline. Also occurs in three brackish water bodies
(Songkhla Lake in Thailand, Chilika Lake in India and Malampaya Sound in the Philippines) and three large river systems
(Myanmar’s Irrawaddy/Ayeyarwady, Indonesian Borneo’s Mahakam and the Mekong of Lao PDR and Cambodia). May
enter the lower reaches (up to c. 86km) of other rivers. Riverine populations are concentrated in relatively deep pools
(10–50m) at confluences or above and below rapids. Coastal populations are typically within a few kilometres of shore
and of river estuaries, in water a few metres deep; also in large lagoons and mangrove forests.
BEHAVIOUR Not particularly acrobatic and usually makes only low leaps when disturbed, socialising or swimming
against a strong current. Spyhopping, body-rubbing and tail-slapping are sometimes observed. Can spit a narrow,
well-directed jet of water 1–2m into the air, usually in association with feeding. Remarkable habit in the Irrawaddy
(Ayeyarwady) River is fishing cooperatively with cast-net fishermen. Usually shy of vessels and not known to bow-ride.
FOOD AND FEEDING Wide variety of fish, squid, octopus, cuttlefish and crustaceans. Primarily a diurnal feeder; known
to hunt cooperatively.
TEETH Upper jaw 16–38; lower jaw 22–36. Teeth may not erupt in some populations (e.g. Mahakam River).
GROUP SIZE AND STRUCTURE Typically 2–6, but varies with location; up to 20 when two or more groups come
together, and as many as 30 in some deepwater riverine pools during the dry season.

ADULT COLOUR VARIATION

DIVE Sequence Surfaces inconspicuously with little splash; blowhole and

only uppermost body (including dorsal fin) visible; may lift flukes out of
water before steep dive, or flip to one side, showing single fluke; rarely
porpoises (low, horizontal, splashy leaps). • Depth Unknown. • Duration
Typically 30 seconds to three minutes; maximum recorded 12 minutes
(when disturbed).

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 Indistinct neck ADULT
crease in some Uniformly dark Small, triangular to slightly
individuals slate-grey to bluish- No darker cape falcate dorsal fin (variable)
grey upperside and (cf. snubfin three-fifths of the way
sides (variable) dolphin)
Blunt, bulbous along back
head (more Moderately
bulbous in robust body
Rounded (rarely
older males) pointed) tip

Fin c. 3 per cent

of total length

Exceptionally Paler grey underside

No Animals in captivity
broad flippers (may appear almost
discernible often lose much of
beak Convex Rounded tip white in muddy water)
pigmentation (may
Straight mouthline leading
become whitish)
(angled up towards edge
eye – appears to
be ‘smiling’)
SIZE
L: ♂ 1.7–2.7m, ♀ 1.7–2.2m;
WT: 115–130kg; MAX: 2.8m, 130kg
Calf – L: c. 1m; WT: c. 10–12kg

Crescent-shaped Shallow groove along ADULT

blowhole slightly to back (from just behind
left of midline (ends neck crease to just Broad flukes (up
point forward) before dorsal fin) to one-quarter of
body length)

Median
notch

Neck more flexible than in

most other dolphins (first Narrow tailstock Shallowly concave
two neck vertebrae unfused) trailing edge

30°

20°
AT A GLANCE Tropical and sub-tropical Indo-
Pacific • Coastal, brackish and fresh waters
10°
• Small size • Two-toned grey coloration •
Bulbous head with no beak • Small dorsal
0°
fin behind midpoint of back • Shallow dorsal
groove • Cryptic, low surfacing • Usually in
10°
small groups
20°

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 ROUGH-TOOTHED DOLPHIN
Steno bredanensis (Lesson, 1828)

Unmistakable at close range, with its smoothly sloping forehead, the rough-toothed dolphin has been described as
looking more like an extinct ichthyosaur (a marine reptile from the age of the dinosaurs) than a cetacean.
IUCN status Least Concern (2018).
Population Minimum c. 250,000. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Slopehead.
DISTRIBUTION Tropical to sub-tropical (and some warm temperate) waters in the Atlantic, Pacific and Indian Oceans,
mainly from c. 40°N to c. 35°S. Also in many semi-enclosed seas; formerly considered a vagrant in the Mediterranean,
but a small, relict population is present in the far eastern corner. Prefers deep offshore waters beyond the continental
shelf – usually deeper than c. 1,000m – and rarely close to land except around islands with steep drop-offs (though
found in shallow coastal waters in some areas). Shows fidelity to at least some oceanic islands.
BEHAVIOUR Can appear quite lethargic and inactive, and it is not wildly acrobatic, but does breach fairly regularly and
often multiple times in succession (although not particularly high). Spyhopping, surface-slapping and low-angled arced
leaping are quite common. Well known for swimming shoulder to shoulder in a synchronised ‘chorus line’. Reaction to
boats varies. In most places, quite easy to approach as long as it is not actively foraging. Often bow-rides and wake-
rides.
FOOD AND FEEDING Fish, squid and octopus. Sometimes hunts cooperatively; thought to feed primarily on near-surface
species, some evidence of food sharing.
TEETH Upper jaw 38–52; lower jaw 38–56.
GROUP SIZE AND STRUCTURE Most common group size is 10–20, though occasionally smaller groups or alone; up
to 50 not uncommon in the eastern tropical Pacific and central Atlantic; some reports of more than 300 (probably
aggregations of sub-groups).
Dorsal cape narrowest
between blowhole ADULT
and dorsal fin

DIVE Sequence When swimming slowly, surfaces unobtrusively (although dorsal fin striking), with slight arching
of back as it dives; at moderate speed, skims along with head and chin just above surface, forming distinctive
walls of spray (looks rather like surfing). • Depth Deepest recorded 399m (though morphologically capable
of deeper); dives deeper at night. • Duration Varies with location; average 4–7 minutes in Hawaii; maximum
recorded 15 minutes.

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 Sides Linear scarring from bites Fairly large, slightly falcate ADULT
Melon smoothly intermediate of other rough-toothed or slightly triangular dorsal
tapers into grey dolphins (especially older fin in middle of back (more
moderately long animals) falcate with age) Wide base to
beak (producing dorsal fin
Stocky forebody
cone-shaped head)
(narrower tailstock) Narrow, dark grey dorsal
Large, slightly cape dips slightly onto
protruding eyes flanks below dorsal fin
No demarcation often in dark
between beak patch
and melon Border between
dark sides and
lighter underside
very irregular

Female has May have Slight keel on

proportionately Wide eye-to- remoras underside of
Unusually long
longer beak flipper stripe attached tailstock (absent
(17–19 per cent
(barely visible) of body length), in female)
slender flippers
Underside (including Sides and underside often covered
throat, lower jaw, part in white or pinkish oval scars
of upper jaw and tip of Flippers further back
Female has fewer from cookiecutter sharks (more
beak) often light grey, than in most other
linear scars spotted with age – when scars
whitish or pinkish small cetaceans
blend together, throat and belly can
appear overall white or pinkish)

Slightly paler, more

muted colour pattern
CALF

SIZE
L: ♂ 2.2–2.7m, ♀ 2.1–2.6m; Usually lacks
WT: 90–155kg; MAX: 2.8m cookiecutter
Calf – L: c. 1–1.2m; WT: c. 15kg shark scars

80°

70°

60° AT A GLANCE Offshore waters in tropics

50°

40°
and sub-tropics • Small size (but chunky) •
30°
Complex three-toned coloration • Prominent,
slightly falcate dorsal fin • Cone-shaped head
20°
10°

• Melon slopes smoothly into moderately

0°
10°

long beak • Unusually large flippers • May

20°

30°

be covered in pink or white blotches • Almost

40°

50°

60°
reptilian in appearance • Often ‘skims’ along
70°
surface
80°

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 ATLANTIC HUMPBACK DOLPHIN
Sousa teuszii (Kükenthal, 1892)

The Atlantic humpback dolphin lives up to its name: its dorsal fin sits on top of an extraordinary elongated hump. It is
a very rare dolphin, with a fragmented distribution along the west coast of Africa, and immediate action is required to
save it from extinction.
IUCN status Critically Endangered (2017).
Population Probably fewer than c. 3,000. Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Atlantic hump-backed dolphin.
DISTRIBUTION Mainly shallow, nearshore tropical and sub-tropical waters along the west coast of mainland Africa, in
the eastern Atlantic. Confirmed in 13 countries, from Western Sahara to southern Angola, but is not yet documented in
a further six within the likely range (Sierra Leone, Liberia, Côte d’Ivoire, Ghana, Equatorial Guinea or the tiny coastline
of the Democratic Republic of Congo). However, with long stretches of coastline without any reported sightings,
distribution may not be continuous. It does not occur around offshore islands. Prefers soft-sediment bottoms and
habitats strongly influenced by tidal and wave motion, such as surf zones, estuaries, channels, mudflats and sandbars,
including mangroves and exposed open coasts; often ‘patrols’ beaches just beyond the breaking waves. Typically in less
than 20m of water (frequently as shallow as 3m). Usually within 1–2km of shore and frequently within 100m; recorded
as far as 13km from shore where the water is sufficiently shallow. Occasionally occupies rivers where there is tidal
influence, but there is no evidence of separate freshwater populations.
BEHAVIOUR A slow, deliberate swimmer and generally inconspicuous. Not as aerially active as most other dolphins,
but it does occasionally leap. Will sometimes hang vertically in the water, with the head out (spyhopping). Reaction to
boats varies, but if it is approached cautiously there is usually just a subtle avoidance response maintaining a ‘personal
space’ of 15–20m from the boat). It may approach closer if the engine is switched off. Does not bow-ride.
FOOD AND FEEDING Nearshore coastal, estuarine and reef fish. Sometimes forages cooperatively to herd mullet, but
may also spread out over area and forage individually.
TEETH Upper jaw 54–64; lower jaw 52–62.
GROUP SIZE AND STRUCTURE The typical group size is 1–10 (accounting for 65 per cent of sightings), but as many as
45 have been seen in the Gulf of Guinea.

ADULT Broad flukes

Deep
median
notch

Subtle darker cape

DIVE Sequence Beak (and often much of head) lifted clear of

water on surfacing, often rising at angle of more than 45°;
flukes often lifted above surface before deep dive. • Depth
Details unknown, but usually shallow (probably less than
20m). • Duration Typically 40–60 seconds.

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 Uniform slate- Relatively small, slightly ADULT
grey to black falcate, backswept dorsal
upperside and fin midway along back
Melon and sides (may Robust body (variable shape – can be Much more rounded tip than
beak can be appear brownish (male may be nub-like or more triangular) on similar Indian Ocean
conspicuously in some lighting) more robust than humpback dolphin
paler female)
Fin sits on prominent, wide-
Light grey eye- based dorsal hump (connective
Shallow crease patch in some
between beak individuals Darker dorsal tissue, present in all age
and melon cape visible in classes and both sexes)
(especially good light
younger
animals) Dorsal hump more exaggerated in
Moderately males (entire structure 26–32 per
rounded cent of total body length), producing
melon ‘lumpy’ overall appearance

Moderately
Moderately deepened
long, slender tailstock
beak (relatively Fades to lighter Mid- to lower tailstock
Broad flippers
shorter Very subtle light grey or whitish paler grey and ‘spotty’
with straight
than other Relatively grey ‘anchor patch’ underside (may (with dark spots and
trailing edge
humpback straight on underside (apex have subtle pink flecks) in some regions
and rounded
dolphins – mouthline continues as distinct flush during intense (e.g. Senegal and
tips
up to 31cm) pale line between activity) Guinea) but uniformly
May be pale grey throat and navel) dark in other regions
‘cheek’ below eye in (e.g. Angola)
some individuals
May be whitish scarring on dorsal fin
and ridge (especially individuals with
most pronounced humps) DORSAL VARIATIONS

SIZE
L: 2.3–2.8m;
WT: c. 140–280kg; MAX: 2.85m, 166kg
Calf – L: c. 1m; WT: c. 10kg

30°

AT A GLANCE Tropical and sub-tropical West

20° Africa • Nearshore waters • Small size •
Dorsal fin sits on distinctive elongated hump
10° • Dorsal profile often appears uniform brown,
grey or black (depending on light) • Robust
0° body with deepened tailstock • Moderately
long, slender beak exposed on surfacing •
10°
Often indifferent or shy towards boats

20°

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 INDO-PACIFIC HUMPBACK DOLPHIN
Sousa chinensis (Osbeck, 1765)

Previously, all humpback dolphins from South Africa to China and Australia were classified as the Indo-Pacific
humpback dolphin (Sousa chinensis). But in 2014 this was split into three distinct species: Indo-Pacific, Indian Ocean
and Australian. There may be yet another species (tentatively S. lentiginosa – see illustration on p. 220) along the
eastern coast of India, and in Bangladesh and Myanmar but, until the taxonomic status is clarified, this is currently
treated as S. chinensis.
IUCN status Vulnerable (2015); Taiwanese subspecies Critically Endangered (2017).
Population Probably fewer than c. 16,000. Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy Two subspecies are recognised: Chinese humpback dolphin (S. c. chinensis) and Taiwanese humpback
dolphin (S. c. taiwanensis). As well as the form known as lentiginosa, there is also uncertainty about the exact
taxonomic status of animals in mainland Malaysia and Borneo (‘borneensis-type’).
Other names Indo-Pacific hump-backed dolphin, Chinese white dolphin, Borneo white dolphin, Taiwanese white dolphin.
DISTRIBUTION Found in shallow coastal waters – typically less than 20–30m deep – and rarely more than a few
kilometres from shore. Highest densities occur in and around estuaries, but it also occurs along open coasts, over rocky
reefs, in bays, coastal lagoons and mangrove swamps, and in areas with sandbanks and mudbanks. It will sometimes
enter rivers and inland waterways, but remains within the range of tidal influence. The distribution is discontinuous.
BEHAVIOUR Tends to swim slowly and deliberately. Moderately acrobatic – breaches, acrobatic leaps and spyhops are
not uncommon. Reaction to boats varies with region. Rarely bow-rides or wake-rides.
FOOD AND FEEDING Wide variety of nearshore, estuarine and reef fish; occasionally cephalopods and crustaceans. In
Hong Kong sometimes seen with mud on body, suggesting bottom feeding.
TEETH Upper jaw 64–76; lower jaw 58–76.
GROUP SIZE AND STRUCTURE Typically in small groups of 2–6, sometimes up to 10; larger groups in some regions.

ADULT (SOUTHEAST ASIA –

Dorsal fin more triangular and BORNEENSIS-TYPE)
even lower and wider-based More extensive
dark coloration

May have slight Typically has dark

pinkish tinge spots and blotches
over much of body

SIZE
L: ♂ 2–2.6m, ♀ 2–2.6m;
WT: 200–240kg; MAX: 2.7m, 240kg
Calf – L: c. 1m; WT: c. 10–12kg

DIVE Sequence Beak (and often much of head) lifted clear of water on surfacing. • Depth Shallow diver, typically
to maximum 30m. • Duration Typically 40–60 seconds; maximum c. 5 minutes.

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 Background mostly white or ADULT MALE (CHINA, TAIWAN AND HONG KONG)
pinkish-white (highly variable
according to age and sex) Relatively low, wide-based,
slightly falcate dorsal fin Male tends to retain
Moderately rounded Pinkish tinge caused by midway along back (15–21 some dark spotting (older
melon with steep blood flushing to surface per cent of total body length) animals typically have
forehead (not pigmentation) small dark spots on back
No distinct and/or dorsal fin)
Distinct junction Robust body
between beak dorsal hump
No dorsal or ventral
and melon (but
ridges on tailstock
no crease)
(present on Indian
Ocean humpback
dolphin)

Older animals may

Moderately Dark blotches, spots and flecks have wrinkled skin
long, well- can be extensive, minimal
Relatively straight Broad flippers (usually (especially on tailstock)
defined beak or absent (highly variable
(up to 34cm) mouthline with rounded tips) according to age and sex)

ADULT FEMALE VARIATIONS

Female loses pigmentation
earlier than male (older (CHINA, TAIWAN AND HONG KONG)
female often pure white
or bright pink)

30°

AT A GLANCE Tropical to warm temperate

20°
waters in South Asia and Southeast Asia •
Small size • Nearshore waters, usually near
freshwater inputs • Mostly white coloration
10°

(often with pinkish tinge) • Often with dark

blotches, spots and flecks • No distinct dorsal
0°

10°
hump • Low, wide-based, slightly falcate to
triangular dorsal fin midway along back •
20°
Moderately long, well-defined beak

Indo-Pacific humpback dolphin

taxonomically uncertain Sousa sp. (lentiginosa) – see p. 220

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 INDIAN OCEAN HUMPBACK DOLPHIN
Sousa plumbea (G. Cuvier, 1829)

The Indian Ocean humpback dolphin was recognised as a distinct species in 2014; previously, all humpback dolphins from
South Africa to Australia were classified as Indo-Pacific humpback dolphins (Sousa chinensis). The taxonomic status of
humpback dolphins in the Bay of Bengal (‘lentiginosa’) remains uncertain (they are currently included under S. chinensis).
IUCN status Endangered (2015).
Population Possibly c. 10,000 (certainly no more than low tens of thousands; fewer than 500 in South Africa).
Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Indian humpback dolphin, plumbeous dolphin.
DISTRIBUTION Found exclusively in tropical to warm temperate waters in the Indian Ocean, along a narrow, shallow,
coastal strip. Occurs in semi-enclosed seas such as the Gulf of Aden, Red Sea and Persian Gulf, and around several
offshore islands (including the Andamans, Mayotte, the Bazaruto Archipelago and Zanzibar). Large portions of the range
have not been surveyed, but its distribution appears to be discontinuous. Strong preference for protected areas, such as
sandy bays, coastal lagoons, rocky reefs, river estuaries and mangroves. Rarely occurs more than 3km from shore (often
only a few hundred metres) or in waters deeper than 25m (sometimes as shallow as 2m).
BEHAVIOUR Tends to be a slow, deliberate swimmer. Occasionally breaches and performs other aerial manoeuvres.
Tends to be quite shy of boats and very rarely bow-rides.
FOOD AND FEEDING Wide variety of nearshore, estuarine and reef fish; occasionally squid, octopuses and crustaceans.
Typically forages near seabed in shallow, murky water; in the Arabian Gulf and Mozambique’s Bazaruto Archipelago,
deliberately strands on exposed sandbanks in pursuit of fish.
TEETH Upper jaw 66–78; lower jaw 62–74.
GROUP SIZE AND STRUCTURE Typically fewer than 10 individuals, though larger groups in some regions (30–100 is not
unusual in Arabian waters).
ADULT (BAY OF BENGAL –
May be pink patches Wide-based
LENTIGINOSA-TYPE)
on dorsal fin and dorsal fin Slight evidence of
Upperside other parts of body dorsal hump (especially
mid-grey behind dorsal fin)
Shows characteristics
associated with
both Indo-Pacific
and Indian Ocean
humpback dolphins

May have some Overall appearance is much

Sides of dark flecking lighter than western Indian
head paler Ocean individuals, especially
on belly and sides

DIVE Sequence Beak (and often much of head) lifted clear of water on surfacing; body arched (showing much of
back, hump, dorsal fin and tailstock) in a high, tight roll; flukes often lifted above surface before deep foraging dive.
• Depth Shallow diver, typically to maximum 25m. • Duration Typically 40–60 seconds; maximum c. 5 minutes.

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 May be loss of pigmentation or Uniform dark grey ADULT
scarring on head, beak, dorsal hump or brownish-grey
and dorsal fin (pinkish or whitish – (variable but Fin sits on prominent, wide-based dorsal
generally much Much more hump (connective tissue, present in all
varies with age, sex and locality)
darker than in Bay pointed tip age classes and both sexes) that gets
of Bengal) than on similar proportionately larger in older animals
Moderately Atlantic
rounded melon humpback Dorsal hump more exaggerated in
with steep dolphin males (entire structure 23–38 per
forehead Small, highly cent of total body length), producing
falcate dorsal overall ‘lumpy’ appearance
Distinct Robust
junction fin midway
body
between along back May be well-developed dorsal and
beak and ventral ridges on tailstock (variable
melon (but by region), especially in male
no deep
crease) Spotting (if present)
limited to small
areas of tailstock
and dorsal hump

Long, slender beak

(up to 38cm) may
Relatively Broad flippers Slightly Deepened tailstock
be pinkish
straight (usually with paler belly (dorsal and ventral)
mouthline rounded tips)

SIZE
L: ♂ 1.8–2.7m, ♀ 1.7–2.5m;
WT: 200–250kg; MAX: 2.8m, 260kg
Calf – L: c. 1–1.1m; WT: 14kg ADULT
Relatively
broad flukes

Median
notch

Rounded tips

40°

30°

20° AT A GLANCE Tropical to warm temperate

waters of western Indian Ocean • Small
size • Robust body • Uniform dark grey or
10°

0° brownish-grey coloration • May be some

10°
loss of pigmentation • Small, pointed dorsal
fin sits on exceptionally large hump • Long,
20°
slender beak • Moderately rounded melon
30°
with steep forehead

Indian Ocean humpback dolphin

taxonomically uncertain Sousa sp. (lentiginosa) extralimital records

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 AUSTRALIAN HUMPBACK DOLPHIN
Sousa sahulensis Jefferson and Rosenbaum, 2014

Split from the Indo-Pacific humpback dolphin (Sousa chinensis) in 2014, the Australian humpback dolphin is now
recognised as a distinct species and differs from other members of the genus in genetics, morphology, coloration and
range.
IUCN status Vulnerable (2015).
Population Probably fewer than c. 10,000 mature individuals. Decreasing.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Sahul dolphin.
DISTRIBUTION Tropical and sub-tropical Australia and southern New Guinea. Distribution throughout the range is
mostly sparse. Shallow and protected coastal habitats such as inlets, shallow bays, sandy-bottomed estuaries, major
tidal rivers (up to 50km upstream), reefs, coastal archipelagos, seagrass meadows, mangroves and occasionally in
dredged channels; rarely in open stretches of coastline. Generally within c. 10km of shore and often much closer; little
survey work has been done further offshore but it has been recorded up to 70km from land (especially in sheltered and
protected waters such as around the Great Barrier Reef). Depth preference varies with location, but rarely deeper than
20m and most commonly shallower than 10m (sometimes in 1–2m).
BEHAVIOUR Tends to be a slow, deliberate swimmer. Moderately acrobatic, capable of making high leaps and
somersaults. Males are regularly observed carrying marine sponges, which they appear to use for posturing in sexual
displays.
FOOD AND FEEDING Mostly nearshore, estuarine and reef fish; rarely cephalopods or crustaceans. Solitary feeding
mainly near seafloor, during long dives; large groups usually feed close to surface.
TEETH Upper jaw 62–70; lower jaw 62–68.
GROUP SIZE AND STRUCTURE Typically 1–5 (up to 10); as many as 30–35 have been observed in feeding aggregations,
especially while following trawlers.
Broad flukes with
rounded tips
ADULT
Deep
median
notch

Very limited loss of

pigmentation on leading ADULT FEMALE
edge or upper half of
dorsal fin (cf. male) Fewer nicks
and notches
on dorsal fin
SIZE
L: ♂ 2.1–2.6m, ♀ 2–2.6m;
WT: c. 240kg; MAX: 2.7m
Calf – L: c. 1m; WT: c. 10–12kg

DIVE Sequence Beak (and often much of head) lifted clear of water on surfacing; body arched (showing much
of back, hump and dorsal fin) in a high, tight roll; flukes often lifted above surface before deep foraging dive.
• Depth Details unknown, but mainly shallow (usually less than 20m). • Duration Typically 40–60 seconds;
maximum c. 5 minutes.

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Slightly curved, diagonal
separation between
dark upperside and
light underside
Moderately rounded
melon (slightly Robust body
depressed in profile)
Dark grey upperside
(resembling a ‘cape’) and
tailstock (varies between
individuals and with age)
ADULT MALE
May exhibit loss of pigmentation on
leading edge and upper half of dorsal
fin (varies with age, sex and locality)
Low, triangular to slightly falcate,
wide-based dorsal fin midway along
back (base ranges from 14–24 per
cent of total body length)
Little or no
dorsal hump

Variable amounts of dark

Shallow crease and light flecking, white
between beak scarring and white or
and melon pinkish spotting

Lighter grey
May have invasion of light
Moderately long, Relatively straight or whitish
Broad flippers (usually colour from underside into
slender beak (up mouthline underside
with rounded tips) lower cape (below and
to 35cm) behind dorsal fin)

OLDER ADULT MALE

Significant pigmentation loss (perhaps
due to intraspecific aggression) leaves
More nicks and notches on
large pinkish-white areas on dorsal fin
trailing edge of dorsal fin
(especially upper half and leading edge)
Some older animals
Beak and very light (though not
front of melon completely white)
pinkish white

10°

0°

AT A GLANCE Coastal tropical and sub-

10° tropical Australia and southern New Guinea
• Small size • Robust body • Broadly two-
20°
toned grey coloration • Diagonal dorsal cape
• Little or no dorsal hump • Low, very wide-
30° based dorsal fin • Moderately long, slender
beak • Usually shy and difficult to approach
40°

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 COMMON BOTTLENOSE DOLPHIN
Tursiops truncatus (Montagu, 1821)

The common bottlenose dolphin is the quintessential dolphin and, thanks to its coastal habits, prevalence in captivity
and frequent appearances on television, one of the best-known cetaceans. But its taxonomy is still in dispute – due
to huge geographical variation in size, shape, skull morphology and coloration. Over the years, more than 20 nominal
species have been proposed, but only two are currently recognised (common bottlenose and Indo-Pacific bottlenose).
Others could be accepted in the future.
IUCN status Least Concern (2018). Lahille’s subspecies (T. t. gephyreus) Vulnerable (2019). Black Sea subspecies
(T. t. ponticus) Endangered (2008). Fiordland (New Zealand) sub-population Critically Endangered (2010).
Mediterranean sub-population Vulnerable (2009).
Population Minimum 750,000 (based on available abundance estimates – but much of the range has not been
surveyed). Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy Three subspecies are recognised: common bottlenose dolphin (T. t. truncatus), found in tropical to
temperate waters worldwide; Lahille’s bottlenose dolphin (T. t. gephyreus), a larger, coastal form found in the western
South Atlantic (which has been proposed as a separate species); and Black Sea bottlenose dolphin (T. t. ponticus),
known only from the Black Sea, Kerch Strait (and connecting part of the Azov Sea) and the Turkish Straits system.
The ‘Burrunan bottlenose dolphin (Tursiops australis)’, found off southern and southeastern Australia, was proposed
as a new species in 2011, but more research is needed. In the North Atlantic, there appear to be two ‘ecotypes’ – a
smaller, coastal form and a larger, more robust offshore form – that may be separated in the future (but differences are
subtle and vary according to location).
Other names Bottlenose dolphin, bottle-nosed dolphin.
DISTRIBUTION Widespread in tropical to temperate waters worldwide. Most abundant between 45°N and 45°S, except
in northern Europe (with significant numbers around the United Kingdom and as far north as the Faroe Islands at 62°N).
Most often seen in shallow coastal waters and around oceanic islands, but also out to the continental shelf edge and
most abundant in deep offshore waters. Frequently in bays, lagoons, channels and around harbours, and ventures into
rivers for brief periods. Frequently near population centres. In the Indian Ocean, where Indo-Pacific bottlenose dolphins
can be prevalent along the coast, common bottlenose dolphins tend to live mainly offshore.
ADULT Distinct crease
between beak
and melon

Dark ‘bridle’ down melon

(between blowhole and
beak) often bordered with
lighter stripes

Extensive tooth rake marks

from intraspecific fighting

DIVE Sequence When swimming slowly, tip of beak often breaks surface first; when swimming quickly, porpoises
with neat re-entry. • Depth Highly variable depending on location and prey; typically up to 70m, but offshore often
several hundred metres; maximum c. 1,000m. • Duration Offshore average c. 1–5 minutes (maximum recorded 13
minutes); inshore typically 30–120 seconds (maximum recorded 8 minutes).

224 LONGER-BEAKED OCEANIC DOLPHINS

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Generally darker than Indo-Pacific
bottlenose (but highly variable)
Three-toned coloration
(can be subtle – often
appears uniform)
Moderately bulbous
melon (slightly more
bulbous than in Indo-
Pacific bottlenose)
Tall, falcate dorsal fin ADULT
midway along back Fin relatively smaller,
narrower-based and slightly
more falcate than in Indo-
Robust body (slightly
Pacific bottlenose
more robust than
in Indo-Pacific Light grey to very dark grey
bottlenose) upperside (may be visible as
dorsal cape at close range)
May have pale spinal blaze
(faint to distinct) and/or pale
brush strokes on side

Some populations
may appear brownish
in bright sunlight

Short to
moderate
length, stubby Large, recurved,
slightly pointed Mid-tone light grey
beak separated
flippers to mid-grey sides,
from melon by
fading to dark above,
distinct crease
Whitish, cream-coloured light below
or pale grey belly (may
Gently curving have pinkish hue)
mouthline Generally no spotting but occasionally
Wide beak-to-flipper has small dark flecks on belly and sides
(ostensibly like stripe (often very faint)
a ‘smile’) (especially in Atlantic – possibly due to
occasional hybridisation with sympatric
Atlantic spotted dolphin)

SIZE
L: ♂ 1.9–3.8m, ♀ 1.8–3.5m;
WT: 136–600kg; MAX: 3.9m, 635kg
Calf – L: 1–1.5m; WT: c. 15–25kg
Wide variation in size between
populations.

80°

70°

60°

50°
AT A GLANCE Tropical to temperate waters
40° worldwide • Small size • Robust body •
Short, stubby beak • Archetypal dolphin •
30°

20°

Ostensibly ‘smiling’ mouthline • Three-toned

10°
0°

coloration (subtle to distinct) • Rarely has

10°
20°

small dark flecks on underside • In coastal

30°

40°

50°
waters usually in small groups • Often bow-
rides
60°

70°

80°

COMMON BOTTLENOSE DOLPHIN 225

004 WDP FG.indd 225 29/11/2021 12:29

 BEHAVIOUR Tends to be active much of the time and will leap, lobtail, porpoise, body surf and perform other aerial
behaviours. Keen bow-rider and wake-rider. It will bow-ride any vessel, from a small motor boat to a large oceanic
cargo vessel or cruise ship, and will even ride the bow-waves in front of large whales (‘snout-riding’). Known to attack
and kill harbour porpoises in Scotland and Wales (UK) and California (US); the reasons for this behaviour are unclear,
and it is more widespread than simply the aberrant behaviour of one or two individuals. Some wild common bottlenose
dolphins have become ‘friendly’ – solitary individuals, not part of a social group, which hang around harbours and
befriend people, apparently preferring to associate with boats, divers and swimmers rather than with other dolphins.
Some stay for weeks or months, others for years.
FOOD AND FEEDING Generalist feeder overall, but specialisation within populations and among individuals; wide
variety of fish, cephalopods and crustaceans; will attempt to swallow absurdly large prey. Wide variety of techniques,
depending on prey and location, including high-speed chasing, bubble-blowing to herd prey towards surface, ‘fish-
whacking’ (knocking fish out of water with flukes – sometimes catching them in mid-air), ‘strand-feeding’ (sending
wave of water that pushes fish onto mudbanks, then temporarily beaching themselves to grab the fish), ‘kerplunking’
(scaring fish out from seagrass beds and other vegetative cover with bubble-forming tail slaps), and ‘mud-ringing’ (one
dolphin creates ring-shaped mud plume, then others catch fish in mid-air as they leap out of ring). Will feed behind
shrimp trawlers (to eat discarded fish and steal fish from fishing gear). In Mauritania and Brazil, regularly drive mullet
towards fishermen holding nets in shallow water.
TEETH Upper jaw 36–54; lower jaw 36–54. Teeth often worn down or missing in older animals.
GROUP SIZE AND STRUCTURE Typically 2–15, but offshore sometimes in large herds of several hundred; groups of
more than 1,000 have been reported. Generally, groups are smaller close to shore. Group structure varies greatly, but
tends to be relatively fluid.

ADULT (BURRUNAN)
Intermediate grey Dark bluish-grey
sides (dip below upperside
Tall, falcate dorsal fin
dorsal fin)
midway along back
Smaller size
(2.3–2.8m)
Distinct three-toned
colour pattern

Pale spinal blaze may Off-white underside

Short, stubby beak extend from thoracic (often extends over eye
area towards base of No ventral and above flipper)
dorsal fin (variable) spotting

CALF

Similar coloration Slightly

to adult slimmer body

Relatively shorter,
stubbier beak

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 ADULT VARIATIONS

COMMON BOTTLENOSE DOLPHIN 227

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 INDO-PACIFIC BOTTLENOSE DOLPHIN
Tursiops aduncus (Ehrenburg, 1833)

Split from the common bottlenose dolphin in 2000, the slightly smaller Indo-Pacific bottlenose dolphin is the only
cetacean known to use tools. In Shark Bay, Western Australia, more than 70 individuals have learned to ‘wear’ marine
sponges on their beaks as protective ‘gloves’ or shields when foraging for prey along the seafloor.
IUCN status Near Threatened (2019).
Population Minimum c. 40,000. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names None.
DISTRIBUTION Widespread but discontinuous distribution throughout coastal tropical to temperate waters in the
Indian and western Pacific Oceans. Found throughout the islands and peninsulas of the Indo–Malay archipelago,
around some oceanic islands (such as the Maldives, Seychelles, Réunion and Madagascar) and in some semi-enclosed
seas (including the Gulf of Thailand, Red Sea and Persian Gulf). Almost exclusively over the continental shelf, especially
in shallow coastal waters (less than 100m deep) in areas with sandy or rocky bottoms, as well as reefs and seagrass
beds. Will concentrate in and around estuaries.
BEHAVIOUR Generally less acrobatic than the common bottlenose dolphin, but still capable of some high leaps. Can be
a keen bow-rider and wake-rider. In Australia, there are several locations where Indo-Pacific bottlenose dolphins come
into shallow water to be fed by people.
FOOD AND FEEDING Predominantly benthic and reef-dwelling fish, and cephalopods; occasionally small benthic
sharks; little overlap with preferred prey of common bottlenose dolphin. Wide variety of techniques, depending on prey
and location, including ‘bottom-grubbing’, ‘sponging’, ‘shelling’ (lifting large shells out of water to dislodge fish hiding
inside), ‘strand-feeding’, ‘snacking’ (chasing fish ‘belly up’ near surface), ‘octopus tossing’ (throwing octopuses in the
air before consumption, to avoid risk of suffocation) and ‘kerplunking’.
TEETH Upper jaw 46–58; lower jaw 46–58.
GROUP SIZE AND STRUCTURE Typically 6–60, but sometimes in large groups of several hundred.

ADULT Dark dorsal cape Relatively

broad flukes

Median
notch

Dark stripes from

blowhole to apex of
melon may be visible
SIZE
L: ♂ 1.8–2.6m, ♀ 1.8–2.6m;
WT: 120–200kg; MAX: 2.7m, 230kg
Calf – L: 0.8–1.2m; WT: 9–21kg
Wide variation in size between populations.

DIVE Sequence When swimming slowly, tip of beak often breaks surface first; when swimming quickly, porpoises
with neat re-entry. • Depth Highly variable depending on location and prey, but usually quite shallow; maximum
200m. • Duration Typically 30–120 seconds (maximum recorded 10 minutes).

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 Moderately robust body Mostly uniform mid-grey to dark ADULT
(slightly more slender than grey upperside (may be visible as
in common bottlenose) dorsal cape at close range)
Moderately bulbous
melon (slightly Three-toned coloration Generally lighter than
Prominent, tall,
less bulbous (can be subtle – often common bottlenose
falcate dorsal fin
than in common appears uniform in poor (but highly variable)
midway along back
bottlenose) light or from a distance)
Beak Fin relatively larger, Some populations
separated May have dark wider-based and slightly may appear
from ring around eye less falcate than in brownish in
melon by common bottlenose bright sunlight
distinct
crease

Mid-tone light grey

to mid-grey sides,
fading to dark
Tip of beak Gently curving above, light below
and ‘lips’ mouthline Large, recurved, Typically has May be some pale
tend to get (ostensibly like pointed flippers pale spinal blaze oval or round scarring
paler with a ‘smile’) extending to below from cookiecutter shark
age Wide beak-to- dorsal fin (variable bites (especially around
flipper stripe but often quite oceanic islands)
Moderate-length, stubby (usually very faint) prominent)
beak (slightly longer
Appendages generally Whitish, cream- May have prominent dark flecks
and narrower than in
larger and broader coloured or pale or spots on underside (variable
common bottlenose
(relative to body size) than grey belly (may according to region and
– and length varies
in common bottlenose have pinkish hue) individual, but more intense
according to region)
and darker with age)

ADULT VARIATIONS
(not to scale)

Median
otch

40°

AT A GLANCE Tropical to temperate waters

30°

20°
of Indo-Pacific • Small size • Moderately
10°
robust body • Moderate-length, stubby beak
0°
• Ostensibly ‘smiling’ mouthline • Three-
10° toned coloration (subtle to distinct) • May
20° have dark flecking or spotting on underside
30° • Usually in small- to medium-sized groups
40°
• Occasionally bow-rides
50°

INDO-PACIFIC BOTTLENOSE DOLPHIN 229

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 PANTROPICAL SPOTTED DOLPHIN
Stenella attenuata (Gray, 1846)

The pantropical spotted dolphin is highly variable in appearance – between ages, individuals and regions – from
virtually unspotted to very heavily spotted. Even though it has been severely depleted by tuna purse-seine fishing in the
eastern tropical Pacific, it is still one of the most abundant cetaceans on the planet.
IUCN status Least Concern (2018).
Population Minimum 2.3 million (not including populations yet to be assessed). At least 4 million killed by commercial
tuna fleets. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy Two subspecies are currently recognised: offshore (S. a. attenuata), which is slightly smaller, more slender
and lightly spotted, and coastal (S. a. graffmani), which is slightly larger, stockier and more heavily spotted.
Other names Spotter, bridled dolphin.
DISTRIBUTION Tropical and some sub-tropical waters in the Pacific, Atlantic and Indian Oceans. Within this range,
most abundant in lower latitudes. The offshore subspecies is found mainly in oceanic waters, beyond the continental
shelf edge, and around some oceanic islands, but it does occur nearshore where sufficiently deep water approaches
the coast. Primarily inhabits waters with surface water temperatures above 25°C. The coastal subspecies is usually
within 130km of shore, along the west coasts of Latin America from southern Mexico to northern Peru, often in water
shallower than 50m.
BEHAVIOUR Fast swimmer. Can be highly acrobatic (though it does not spin) and frequently performs breaches and
side-slaps. Readily approaches boats and bow-rides (except on tuna fishing grounds in the eastern tropical Pacific);
females and juveniles are more likely to bow-ride than males. In the eastern tropical Pacific and western Indian
Oceans, it frequently associates with yellowfin tuna and skipjack tuna (perhaps for foraging efficiency or protection
from predators).
FOOD AND FEEDING Offshore subspecies – mainly small epipelagic and mesopelagic fish, squid and crustaceans;
coastal subspecies – possibly mainly larger, bottom-living species. Forages mainly at night; offshore subspecies
exploits deep scattering layer.
TEETH Upper jaw 68–96; lower jaw 68–94.
GROUP SIZE AND STRUCTURE Coastal subspecies generally in groups of 10–20 (ranging from one to c. 100). Offshore
herds can number in the hundreds or thousands, sometimes spread out over several kilometres.
ADULT Pointed tips

Slight
median
notch
White-tipped beak
clearly visible
Slightly concave
SIZE trailing edges
L: ♂ 1.6–2.6m, ♀ 1.6–2.4m;
WT: 90–110kg; MAX: 2.6m, 119kg
Calf – L: 80–85cm; WT: 10–15kg

DIVE Sequence When swimming slowly, tip of beak often breaks surface first; when swimming fast, porpoises
with neat re-entry. • Depth Typically shallower (5–50m) during day, deeper (25–250m) at night (varies according to
region); maximum recorded 342m. • Duration Typically 30–120 seconds; maximum 5.4 minutes.

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 Long, slender Dark dorsal cape sweeps ADULT OFFSHORE
beak (up to 13cm) Typically very little or no very low on side just ahead
separated from white spotting on cape of dorsal fin
melon by distinct (less than in coastal)
Very narrow, falcate dorsal fin
crease (tends to Two-toned (up to 20cm tall) midway along
be slightly more underlying back (variable shape)
slender than coastal Some individuals
colour pattern
subspecies) have small number
Typically pointed tip of cookiecutter shark
More slender
bites (often heal into
Narrow (1–2cm), body than coastal No spinal starburst pattern)
well-defined, dark subspecies blaze
grey ‘mask’ stripe
connects eye patch
to apex of melon Sides and underside
light to medium grey

Small, slender, strongly

recurved flippers
Dark grey May be moderate Two-toned tailstock
Many larger eye-patch ventral keel in (dark upperside,
individuals have Dark grey beak-to-flipper adult male light underside)
brilliant white-tipped stripe (variable)
beak (up to first 4cm)
Typically pale ‘lips’
that gets progressively
(may flush pink)
whiter with age (may Stockier body ADULT COASTAL
flush pink) than offshore
Much more extensive white spotting on subspecies Spotting may extend to
dark dorsal cape (highly variable – may dorsal fin and flippers
be dense enough to obliterate cape)

Two-toned
tailstock

Beak tends to be Male tends

slightly thicker to be more
than offshore Dark spots on underside often fuse and lighten into Typically more heavily spotted
subspecies slightly mottled or (seen from distance) uniform pale spotting than than female
grey colour in older animals (cf. Atlantic spotted dolphin) offshore

80°

70°

60°

50° AT A GLANCE Tropical to sub-tropical waters

worldwide • Small size • Two-toned tailstock
40°

30°

20°
10°
• Dark dorsal cape dips lowest just ahead
0°
10°
of dorsal fin • Highly variable dark and light
20°

30°
spotting • Long, slender beak with white
40°
tip • White or pale grey ‘lips’ • Two-toned
underlying colour pattern • No spinal blaze
50°

60°

70° • Tall, falcate dorsal fin midway along back

80° • Highly variable appearance within group
primary range of offshore subspecies
primary range of coastal subspecies

PANTROPICAL SPOTTED DOLPHIN 231

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 ATLANTIC SPOTTED DOLPHIN
Stenella frontalis (G. Cuvier, 1829)

The Atlantic spotted dolphin is highly variable in appearance – between ages, individuals and regions – from virtually
unspotted to very heavily spotted. Confusion is most likely with common bottlenose dolphins (though heavily-spotted
animals are likely to be Atlantic spotted); look for the more prominent spinal blaze. Pantropical spotted dolphins
generally have a more slender body, a broadly two-toned underlying colour pattern, a tailstock divided into dark upper
and light lower portions, and lack the light-coloured spinal blaze.
IUCN status Least Concern (2018).
Population Probably low hundreds of thousands (83,000 in US waters). Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised subspecies. However, there appear to be two forms: a larger, heavy-bodied, heavily spotted
form occurring mainly over the continental shelf in warmer waters of the western North Atlantic; and a smaller,
slimmer, lightly spotted or unspotted form in more oceanic areas over the continental slope in the Gulf Stream and
the central North Atlantic (and around some offshore islands, such as the Azores).
Other names Spotted dolphin, spotter.
DISTRIBUTION Tropical to warm temperate waters in the Atlantic Ocean. Occurs around some oceanic islands, such
as the Azores and the Bahamas. The heavily spotted form prefers shallow continental shelf waters (typically at least
8–20km offshore). The lightly spotted form occurs over the outer continental shelf, the upper continental slope and in
deep oceanic waters. May prefer shallower waters around oceanic islands, such as over shallow (6–12m) sand banks
in the Bahamas.
BEHAVIOUR Highly acrobatic and capable of some exceptionally high leaps. An avid bow-rider in most of its range.
FOOD AND FEEDING Small to large fish and squid. Offshore form cooperates in herding balls of fish against the surface.
TEETH Upper jaw 64–84; lower jaw 60–80.
GROUP SIZE AND STRUCTURE Generally in small to medium-sized groups of up to 50 animals (occasionally up to 200).
Group sizes tend to be smaller (5–15) nearer shore and larger offshore.
ADULT HEAVILY SPOTTED FORM
Flukes usually
darker with little
or no spotting
Median
notch

Flippers usually
darker with little
or no spotting SIZE
L: ♂ 1.7–2.3m, ♀ 1.7–2.3m;
WT: 110–140kg; MAX: 2.3m, 143kg
Calf – L: 0.8–1.2m; WT: c. 10–15kg

DIVE Sequence When swimming slowly, tip of beak breaks surface first, upperside of melon briefly visible as it
blows, and back and dorsal fin appear briefly (but tailstock often barely visible); when swimming fast, porpoises
with neat re-entry. • Depth Most dives are less than 10m; maximum recorded 60m. • Duration Mostly 2–4 minutes;
maximum recorded 6 minutes.

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 Indistinct three-toned colour Well-developed, lighter ‘FUSED’ ADULT
Slightly pattern (dark grey dorsal spinal blaze sweeps up into HEAVILY SPOTTED FORM
bulbous melon cape, medium grey sides, dorsal cape (distinctiveness
with smoothly white underside) may be varies hugely with amount Tall, falcate dorsal fin (up to 25cm)
sloping visible under spotting of spotting) midway along back (variable shape)
forehead Light dorsal spots become
Body shape variable extensive and coalesce (some
according to region (tends Rounded tip individuals are so heavily
to be moderately robust spotted that background
Distinct
in heavily spotted form) colour is totally obliterated)
crease
between
Underlying colour of
beak and
tailstock monotone
melon
grey (cf. pantropical
spotted dolphin)

Many
individuals Recurved
flippers Heavy, dark ventral
have white- Dark eye-to-flipper Dark ventral spots remain spotting, light dorsal
tipped beak and or beak-to-flipper distinctly defined, with
Appendages spotting (highly variable –
pale ‘lips’ stripe (variable) underlying white underside
proportionately larger more spotting with age)
visible between them (cf.
Moderately long, thick beak (up than in pantropical
pantropical spotted dolphin)
to 13cm) intermediate between spotted dolphin
those of common bottlenose and
pantropical spotted dolphins
Body shape variable ADULT LIGHTLY SPOTTED FORM
according to region (tends
to be moderately slender
in lightly spotted form) Distinct three-toned colour
pattern (dark grey dorsal
Well-developed, lighter cape, mid-grey sides,
spinal blaze sweeps up white underside)
into dorsal cape
Little or no
spotting

60°

AT A GLANCE Tropical to warm temperate

50°

40°
waters of the Atlantic • Small size
30° • Generally chunkier than pantropical spotted
20° dolphin • Many populations heavily spotted
10° • Highly variable appearance within group
0° • Light diagonal spinal blaze • Three-toned
10° underlying colour pattern • Tall, falcate dorsal
20° fin midway along back • Moderately long,
30° white-tipped beak
40°

ATLANTIC SPOTTED DOLPHIN 233

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 SPINNER DOLPHIN
Stenella longirostris (Gray, 1828)

Named for its spectacular habit of leaping high out of the water and spinning up to seven times longitudinally, before
falling back with a great splash, the spinner dolphin is a familiar sight in many parts of the tropics. There is more
geographical variation in form and colour pattern in this species than in almost any other cetacean.
IUCN status Least Concern (2018). Eastern spinner subspecies (orientalis) Vulnerable (2008).
Population One million+ (based on limited surveys, but most of range unsurveyed). At least 2 million killed by
commercial tuna fleets. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy Four subspecies currently recognised: Gray’s (sometimes ‘Hawaiian’) spinner dolphin (S. l. longirostris), the
‘typical’ spinner dolphin; Central American (previously ‘Costa Rican’) spinner dolphin (S. l. centroamericana); eastern
spinner dolphin (S. l. orientalis); and dwarf spinner dolphin (S. l. roseiventris). There is also a hybrid – called the
whitebelly or white-bellied spinner – that is intermediate between Gray’s and eastern spinner dolphins, found in the
eastern tropical Pacific where these two ‘parent’ subspecies meet.
Other names Long-snouted spinner dolphin, longsnout, spinner, rollover; see taxonomy for subspecies common names.
DISTRIBUTION Found in all tropical and most sub-tropical waters in the Pacific, Atlantic and Indian Oceans. Best known
in coastal waters, around oceanic islands and over shallow banks, but also occurs in very large numbers on the high
seas and ranges over vast distances of open water. Coastal spinners – especially around oceanic islands – frequently
move into shallow sandy bays in the morning and rest until late afternoon or early evening; they venture out to deeper
water at night to feed. In some places, they rest actually inside the lagoons of coral atolls.

SPINNING Best known for leaping up to 3m into the air, spinning on its longitudinal axis up to seven times and then
falling back into the water – often up to 14 times in a descending series (each less vigorous than the previous one). It
starts spinning underwater, just before emerging from the surface. Individuals of all ages spin and, once one dolphin
starts, others typically join in. Some other dolphin species spin, but not as many times or with the same frequency.

DIVE Sequence When swimming slowly, tip of beak often breaks surface first;
back and prominent dorsal fin appear briefly – some individuals look back-
to-front (due to forward-leaning dorsal fin); tailstock quickly arches (usually
high), but normally does not fluke; when swimming fast, may porpoise with
clean re-entry; large, travelling schools often churn water into foam.
• Depth Varies with subspecies; offshore, frequently to 200–300m, but
capable of 600m or deeper. • Duration When resting, 1–2 minutes (spends
most of time at surface); when foraging, 3–4 minutes.

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 Three-toned coloration ADULT GRAY’S
(dark grey cape, lighter
Very long, narrow beak, grey sides, white belly)
separated from melon
by distinct crease Dorsal fin relatively small,
Slender body falcate (female) to triangular
(older male – may be canted
slightly forward), midway
Slender head with along body
Beak dark gently sloping
on upperside forehead
(darker with
age), white
on underside

Wide, dark grey

Dark (almost Slender, dark, Belly may flush
eye-to-flipper
black) tip recurved, pink when active May develop
stripe
pointed flippers subtle ventral
Most of lower keel (male only)
jaw white Narrow, dark grey or
black eye-to-beak stripe

Noticeably ADULT GRAY’S

slender body

Distinctive black tip

80°

70°

60°

50° AT A GLANCE Tropical and sub-tropical

waters worldwide • Small size • Usually
40°

30°

20°
10°
slender body • Erect dorsal fin (sometimes
0°
10°
canted forward) midway along back • Huge
20°

30°
variation in appearance according to region
40°

50°
• Long, slender beak • Gently sloping melon
60° • Performs high spinning leaps • Usually
70°
quite gregarious
Gray’s Whitebelly and Whitebelly and eastern
Gray’s overlap overlap
Dwarf Eastern Central American

SPINNER DOLPHIN 235

004 WDP FG.indd 235 29/11/2021 12:29

 BEHAVIOUR One of the most aerial of all dolphins. As well as spinning, it will perform more traditional breaches, as
well as arc-shaped leaps, tail-over-head leaps, side-slaps, fluke-slaps and flipper-slaps. Particularly acrobatic during
the change from rest to foraging. In many parts of the range it readily approaches boats and bow-rides (except on tuna
fishing grounds in the eastern tropical Pacific, where it generally avoids boats).
FOOD AND FEEDING Wide variety of small midwater fish, squid, sergestid shrimps; dwarf spinner feeds on benthic reef
fish and invertebrates. Most feeding at night, resting during day (except dwarf spinners – feed during day).
TEETH Upper jaw 80–124; lower jaw 80–124. There are small differences in tooth count between subspecies.
GROUP SIZE AND STRUCTURE Highly variable according to activity and location, ranging from 10–50 up to several
thousand. The largest schools tend to occur offshore, the smallest in coastal waters.

ADULT MALE CENTRAL AMERICAN

Uniform ‘battleship’
Very long, dark, Triangular or strongly canted dorsal
grey coloration
narrow beak (slightly fin midway along back (may look
longer and narrower as if facing backwards – especially
Slender body
than in eastern prominent in older male)
(proportionately
spinner), separated
slimmer than
from melon by Dorsal fin canted
eastern spinner)
distinct crease Slender head forward slightly less
with gently than in eastern spinner
sloping
forehead
Fluke tips may be
upturned (more
exaggerated
with age)

Dark eye-to-flipper
Slender, dark, May have inconspicuous
stripe (variable)
recurved, pointed whitish patches around Tailstock can become
flippers urogenital area and very deepened (with
‘armpit’ (highly variable) medium to large ventral
keel – generally less well
developed than in eastern
spinner)

SIZE – GRAY’S
L: ♂ 1.6–2.1m, ♀ 1.4–2m;
WT: 55–70kg; MAX: 2.2m, 80kg
Calf – L: 75–80cm; WT: c. 10kg

SIZE – EASTERN
L: ♂ 1.6–2m, ♀ 1.5–1.9m;
WT: 55–70kg; MAX: 2m, 75kg
Calf – L: 75–80cm; WT: c. 10kg SIZE – CENTRAL AMERICAN SIZE – DWARF
L: ♂ 1.9–2.2m, ♀ 1.8–2.1m; L: ♂ 1.4–1.6m, ♀ 1.3–1.5m;
SIZE – WHITEBELLY WT: 55–75kg; MAX: 2.2m, 82kg WT: 23–35kg; MAX: 1.6m, 36kg
L: ♂ 1.6–2.4m, ♀ 1.6–2m; Calf – L: 75–80cm; WT: c. 10kg Calf – L: 50–70cm; WT: 5–7kg
WT: 55–75kg; MAX: 2.4m, 75kg
Calf – L: 75–80cm; WT: c. 10kg

236 LONGER-BEAKED OCEANIC DOLPHINS

004 WDP FG.indd 236 29/11/2021 12:29

 ADULT FEMALE CENTRAL AMERICAN Dorsal fin more falcate or
erect and triangular (not
canted forward)

Fluke tips not

upturned

Tailstock less deepened

(no ventral keel)
ADULT CENTRAL AMERICAN

Three-toned coloration ADULT DWARF

(dark grey cape, lighter Fairly
grey sides, white or slender
pinkish belly) Proportionately large,
body erect to falcate dorsal
Long, narrow fin midway along back
beak separated
from melon by
distinct crease

Dark (almost
black) tip Dark eye-to- White underside
flipper stripe Relatively large, may flush pink
pointed flippers when active
Beak dark on
upperside, white
or pinkish on ADULT DWARF
underside

SPINNER DOLPHIN 237

004 WDP FG.indd 237 29/11/2021 12:29


Long, dark, narrow beak
(slightly shorter and
thicker than in Central
American spinner),
separated from melon
by distinct crease
ADULT MALE EASTERN
Slender body
(proportionately less
slim than Central Very large, triangular or
American spinner) strongly canted dorsal fin
(especially prominent in older
male) midway along back

Uniform ‘battleship’
grey coloration (may Dorsal fin of older male canted
appear purple-grey or forward slightly more than in
bluish-grey in sunlight) Central American spinner

Fluke tips may

be upturned

Tailstock can become very deepened

May have inconspicuous (with exaggerated ventral keel –
Slender, dark, whitish patches around especially in older male – generally
recurved, pointed urogenital area and ‘armpit’ better developed than in Central
flippers (highly variable) American spinner)

ADULT FEMALE EASTERN

Dorsal fin more erect and

slightly falcate or triangular
(not so canted forward)

Fluke tips not

upturned

Tailstock less deepened

(no ventral keel)

ADULT EASTERN

238 LONGER-BEAKED OCEANIC DOLPHINS

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 Coloration two-toned (clear or moderate ADULT MALE WHITEBELLY
darkish grey cape, white lower sides
and belly) or three-toned (with narrow,
medium grey divide between dark
upperside and white underside)
Dorsal fin tends to be
more triangular in male

Robust body Dorsal fin midway

along back

Dark eye-to-flipper White belly may May develop small

stripe flush pink when to moderate ventral
active keel (male only)

ADULT WHITEBELLY

ADULT FEMALE WHITEBELLY

Coloration more
likely to be muted
in female Dorsal fin tends to
be more falcate
than in male

White belly may

flush pink when
active No ventral keel

SPINNER DOLPHIN 239

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 CLYMENE DOLPHIN
Stenella clymene (Gray, 1850)

Molecular studies reveal that the Clymene dolphin (normally pronounced ‘Cly-me-nee’) may have evolved through
extensive hybridisation between spinner and striped dolphins and, in many ways, it appears almost intermediate
between the two species. If so, it would be the first marine mammal known to have arisen in this way. Initially believed
to be a variant of the spinner dolphin, it was not fully accepted as a distinct species until 1981.
IUCN status Least Concern (2018).
Population Unknown, but appears to be relatively common in at least parts of its range. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Short-snouted spinner dolphin, Atlantic spinner dolphin, Senegal dolphin, helmet dolphin.
DISTRIBUTION Tropical, sub-tropical and occasionally warm temperate waters in the Atlantic Ocean, including the
Caribbean Sea and Gulf of Mexico. There are only two mid-Atlantic records, but it is assumed to have a continuous
range. An oceanic species, occurring mainly seaward of the continental shelf (preferring the slope and beyond) and
rarely seen near shore (except where deep water approaches the coast).
BEHAVIOUR Quick, agile and often aerially active. Breaches and spins longitudinally; may spin up to four times (though
the leaps are lower and less frequent, and the spins less elaborate and acrobatic than those of the spinner dolphin –
the only other species that routinely exhibits longitudinal rotations). Response to boats varies from avoidance to quite
inquisitive. Avid bow-rider in some areas and will often approach vessels from a distance.
FOOD AND FEEDING Small mesopelagic fish and squid. Feeds mostly at night; cooperative feeding has been observed.
TEETH Upper 78–104 jaw; lower jaw 76–96.
GROUP SIZE AND STRUCTURE Ranges from one to 1,000, with an overall average of c. 70–80.
ADULT Distinctive dark Slightly concave
dorsal cape trailing edge

Median
notch

Variable amounts
of mid-grey sides
visible from above Fin can be
more triangular
More muted May be paler ADULT VARIATION
facial pattern inner area on
dorsal fin

Dark grey lateral stripe may be

absent, indistinct or bold

DIVE Sequence Porpoises when swimming fast. • Depth Unknown. • Duration Unknown.

240 LONGER-BEAKED OCEANIC DOLPHINS

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 Small and triangular to ADULT
May be paler inner
tall and falcate dorsal fin
Grey cape dips area on dorsal fin
(highly variable)
above eye Dark grey
dorsal cape Fin midway along back
Dark grey to black
‘moustache’ (dark Grey cape dips below dorsal
streaks) on rostrum Stocky body fin (or in some individuals
(variable) further forward, below base
Dark eye-ring of leading edge of dorsal fin)
Distinct crease
between beak May be oval scars
and melon from cookiecutter
shark bites (mainly on
sides and underside)

White or creamy Mid-grey sides

underside, throat (varies in extent
Dark grey and lower jaw and distinctiveness)
stripe Variable dark grey
Beak tip
between eye-to-flipper stripe Distinct dark grey lateral stripe
and ‘lips’
eye and (usually broadens between white underside and mid-
dark grey Slender, curved
beak toward flipper) grey side may be absent, indistinct or
or black flippers with
pointed tips bold (sometimes prompting references
to colour pattern as ‘four-toned’)
Moderately long
Narrow white eye-to-
(less than 12cm)
flipper stripe (above
beak (shorter and
dark stripe) ADULT
thicker than on
spinner dolphin)

Dark grey to black ‘moustache’

(dark streaks) on rostrum visible
on bow-riders (varies in extent
and intensity)

Dark stripe from tip of

beak to base of melon
SIZE (continues to blowhole as
L: ♂ 1.8–2m, ♀ 1.7–1.9m; lighter grey band)
WT: 50–80kg; MAX: 2m, 80kg
Blackish-
Calf – L: 0.9–1.2m; WT: c. 10kg
tipped beak

80°

70°

60°

50° AT A GLANCE Warm waters of Atlantic

Ocean • Small size • Stocky body • Three-
40°

30°

toned colour pattern (highly variable) • Two

20°
10°
0°
10° distinctive dips in the dark cape (giving wavy
20°

30° appearance) • Nearly triangular or slightly

40°

50°
falcate dorsal fin • Medium-length, robust
60° beak • Dark ‘moustache’ on surface of beak
70°

80°

CLYMENE DOLPHIN 241

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 STRIPED DOLPHIN
Stenella coeruleoalba (Meyen, 1833)

Ancient Greeks marvelled at the beautiful ‘brushstrokes’ and colours of striped dolphins, and depicted them in their
frescoes several thousand years ago. Widely distributed in warm waters in both hemispheres, the species is a familiar
sight in many parts of the world.
IUCN status Least Concern (2018). Mediterranean sub-population Vulnerable (2010).
Population Minimum 2.4 million. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Streaker, euphrosyne dolphin, whitebelly.
DISTRIBUTION Widely distributed mainly in tropical to warm temperate waters (roughly between 50°N and 40°S),
though extends into higher latitudes than other Stenella dolphins (it is the only member of the genus that routinely
reaches northern Europe). Typically in water deeper than 1,000m (in many areas sighting rates increase dramatically
with greater depth). Generally occurs outside the continental shelf but also close to shore where waters sufficiently
deep. In the Mediterranean, where it is the most abundant dolphin, it is sometimes in shallower water relatively close
to shore.
BEHAVIOUR Very acrobatic, frequently breaching up to 5–7m high, porpoising upside down and chin-slapping. Performs
a unique behaviour called ‘roto-tailing’, in which it makes a high arcing leap while vigorously whipping its tail in a
circle. Especially nervous of vessels in the eastern tropical Pacific; elsewhere, it will bow-ride and wake-ride, but can
be more easily ‘spooked’ than other tropical dolphins and often dashes away (with low, splashy leaps) for no apparent
reason.
FOOD AND FEEDING Variety of small fish, squid and some crustaceans. Probably mostly nocturnal.
TEETH Upper jaw 78–110; lower jaw 78–110.
GROUP SIZE AND STRUCTURE Typically 10–100 in a dense school, sometimes up to 500, with several thousand
reported occasionally. Great variation between regions. Group size tends to increase the further offshore and deeper
the water.
ADULT

Median
notch

Flukes light grey to

black on both sides
SIZE
L: ♂ 2.2–2.6m, ♀ 2.1–2.4m;
WT: 100–150kg; MAX: 2.6m, 156kg
Calf – L: 90–100cm; WT: 10–15kg

DIVE Sequence Usually seen swimming fast, in long, low arcing leaps; surfaces at shallow angle, sometimes
throwing tail high in air mid-leap; when not clearing surface, typically throws up Dall’s porpoise-like rooster tail
of spray. • Depth May be capable of 700m, but limited information. • Duration Unknown.

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 Slightly bulbous ADULT
melon with smoothly Dark grey, bluish- Moderately tall, falcate
sloping forehead Generally more robust grey or brownish- dorsal fin midway along
than other Stenella grey dorsal cape back (up to 27cm)
Moderately long, species (but ranging
fairly stubby beak to quite slender)
(up to 11cm or
4.5–5.8 per cent of Pale grey or white
body length), with Pale grey spinal blaze extends
distinct crease thorax from thoracic area
between beak towards base of dorsal
and melon fin (highly variable)

May have indistinct

Dark grey to black
eye-to-flipper stripe Pointed tip
Slender, recurved,
light grey to black
flippers
IMMATURE
wounds from
cookiecutter shark
White or pinkish bites (which heal to
belly, throat and background colour)
lower jaw
Short accessory
stripe (not always Dark grey to bluish-black
present) lateral stripe from beak,
through eye, to anus
(widening and slightly
fading at rear)
Dorsal colour
variable (may
be browner)

Lateral stripe and eye-to-flipper

stripes may be more muted

80°

70°

AT A GLANCE Deep tropical to warm

60°

50°

40°

30°
temperate waters worldwide • Small size •
20°
10°
Complex three-toned colour pattern • Long,
0°
10°
dark lateral stripe • Underside bright white or
20°

30°
pinkish • Light grey spinal blaze sweeps back
40° and up towards dorsal fin • Moderately tall,
falcate dorsal fin midway along back • Active,
50°

60°

70° energetic and fast swimmer

80°

STRIPED DOLPHIN 243

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 COMMON DOLPHIN
Delphinus delphis Linnaeus, 1758

Aristotle and Pliny the Elder described the common dolphin in great detail – and it was the first dolphin species to be
scientifically described – yet there has been ongoing debate ever since about whether it should be classified as one,
two or more species. The unique criss-cross or hourglass colour pattern on the sides of all common dolphins (except a
rare dark morph) should distinguish them from other dolphins.
IUCN status Least Concern (2020). Mediterranean sub-population Endangered (2003). Black Sea sub-population
Vulnerable (2008). Gulf of Corinth sub-population Critically Endangered (2019).
Population At least 4–5 million (based on relatively dated regional estimates). Trend unknown, but there have been
dramatic declines in some areas (such as the Mediterranean).
Classification Odontoceti, family Delphinidae.
Taxonomy Much confusion, with more than 20 species described since 1758. Most experts considered it to be a
single species (Delphinus delphis) until 1994, when it was split into two – the short-beaked common dolphin (D.
delphis) and the long-beaked common dolphin (D. capensis). Recent research questioned this split and, since 2016,
it has once again been considered a single species. There are currently four recognised subspecies: common dolphin
(D. d. delphis), Indo-Pacific common dolphin (D. d. tropicalis), Black Sea common dolphin (D. d. ponticus), and Eastern
North Pacific long-beaked common dolphin (D. d. bairdii). It is possible that the Eastern North Pacific long-beaked,
occurring off California, may yet constitute a separate species.
Other names Crisscross dolphin, common porpoise, saddleback dolphin.
DISTRIBUTION Tropical to temperate waters worldwide, roughly from 45°N (North Pacific) and 60°N (North Atlantic) to
50°S. May occasionally follow warm-water currents outside the normal latitudinal distribution. From nearshore waters
to thousands of kilometres offshore, although absent from Gulf of Mexico and the Caribbean and from much of Atlantic
and Indian Oceans. Separate populations occur in some semi-enclosed seas, such as the Mediterranean and Black
Seas.
Dark line along crease ADULT COMMON
Beak light brownish-grey between beak and melon
(often with black tip or (extends posteriorly to
black band near tip) encircle eyes)
Flukes usually dark
brownish-grey above

Deep
median
notch

Dark line from tip of Concave trailing edge

beak to apex of melon

DIVE Sequence When swimming slowly, surfaces at shallow angle, and often begins to blow underwater; when
faster, underside of beak skims along surface; may be faint walls of water on either side of head. • Depth Most
foraging shallower than 50m, but dives to 280m recorded. • Duration Typically c. 10 seconds to 3 minutes; maximum
c. 8 minutes.

244 LONGER-BEAKED OCEANIC DOLPHINS

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 ADULT COMMON
Criss-cross colour pattern on side (where
(Atlantic – including
dark brownish-grey cape, tan to pale yellow
or ochre thoracic patch, light to medium Mediterranean – and Pacific)
Rounded, grey flank patch and white underside meet
bulbous, quite at point below dorsal fin)
steeply rising
melon Tall dorsal fin midway along
Relatively
back (variable shape)
robust body
Beak sharply
demarcated
from melon by Pointed tip
deep crease

Small, slender,
recurved flippers
Black ‘lips’
(pointed at tips)
Black eye-patch continues
as black stripe to crease May have
Flipper-to-anus
Relatively between beak and melon prominent
stripe usually weakly
shorter and (clearly separated from ventral keel
developed or absent
stubbier flipper-to-beak and flipper- (male only)
beak (but still to-anus stripes) Flippers can be dark
moderately long) Slender, dark flipper-to-beak brownish-grey, pale yellow
to ochre, pale grey (or any
stripe joins ‘lip’ patch at varying combination in between)
locations on underside of beak

May have pale grey patch in ADULT COMMON

Dorsal fin highly
variable, from falcate centre (varies from absent to DORSAL FIN VARIATIONS
Deep to erect or triangular covering nearly entire fin)
median
notch

80°

70°
AT A GLANCE Tropical to temperate waters
worldwide • Small size • Criss-cross or
60°

50°

40°
‘hourglass’ colour pattern on sides • Dark
brownish-grey cape dips to ‘V’ under dorsal fin
30°

20°

• Tan to pale yellow or ochre thoracic patch

10°
0°

• Light to medium grey flank patch • White

10°
20°

underside • Details of colour pattern highly

30°

40°

50°

60°
variable • Tall, moderately falcate dorsal fin
70°
midway along back • Often in fast-moving,
80°
splashy groups

COMMON DOLPHIN 245

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 BEHAVIOUR Aerially active – frequently performing a variety of acrobatic leaps and somersaults (sometimes up
to 6–7m high), as well as flipper-slapping and lobtailing. Will also ‘pitch-pole’, a distinctive aerial display in which
the dolphin leaps straight out of the water and slams back lengthwise to create maximum splash. Can also make
clean entries, with no splash. Typically porpoises out of the water when travelling at high speed. An enthusiastic
and energetic bow-rider and wake-rider – even riding the ‘snout waves’ of large mysticete whales; however, not all
individuals in a herd bow-ride. Large, boisterous groups of common dolphins, whipping the ocean’s surface into a froth,
are a common sight in many parts of the world.
FOOD AND FEEDING Wide variety of small schooling fish and squid; some crustaceans. Cooperative feeding techniques
often used to herd fish schools.
TEETH Upper jaw 82–134; lower jaw 82–128.
GROUP SIZE AND STRUCTURE Highly gregarious, in groups of 10–10,000 or more. Group composition is poorly known,
but large herds are believed to be composed of smaller social units of 20–30 individuals that are not necessarily
genetically related. Individuals may swim with preferred associates, but group membership appears to be fluid.

Darker thoracic patch contrasts ADULT EASTERN NORTH PACIFIC LONG-BEAKED

less with cape (border between
them is ‘fuzzier’) Relatively More muted
slender body colour pattern
Light patch in dorsal
Flatter, less bulbous fin usually absent
and less steeply (small and faint if
rising melon present)

Long, slender beak Flipper-to-anus stripe

(6.9–7.6 per cent of Black eye-patch may usually moderately to
total length) be embedded in strongly developed
flipper-to-anus stripe

ADULT EASTERN NORTH PACIFIC LONG-BEAKED

HEAD VARIATION

ADULT COMMON (rare dark morph)

Eye-to-flipper stripe can
be dark and wide (may
Dark dorsal area obliterates Rest of colour form ‘bandit mask’)
characteristic hourglass pattern normal
pattern on side (tan to pale
yellow or ochre colour does
not develop or develops
incompletely)

246 LONGER-BEAKED OCEANIC DOLPHINS

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 ADULT INDO-PACIFIC (Indian Ocean – including Red Sea, Persian Gulf, Gulf of Thailand
– and far western Pacific Ocean)

Relatively
Darker thoracic patch slender body
contrasts less with Light patch in dorsal fin
cape (border between usually absent (small
them ‘fuzzier’) and faint if present)
Flatter, less
bulbous and
less steeply
More muted
rising melon
colour pattern

Extremely long, slender

beak (9.4–9.7 per cent
of total length)

ADULT BLACK SEA

Relatively
Bright yellow slender body
to ochre
thoracic patch Light patch in
dorsal fin often
Moderately present
long, relatively
stubby beak

Black eye-patch continues as black Dark lateral band more

stripe to crease between beak and prominent than in
melon (clearly separated from flipper- other subspecies
to-beak and flipper-to-anus stripes)

SIZE – Black Sea

L: ♂ 1.5–1.8m, ♀ 1.5–1.7m;
WT: c. 150kg; MAX: 2.2m

SIZE – Eastern North Pacific long-beaked

L: ♂ 2–2.6m, ♀ 1.9–2.2m;
WT: 150–235kg; MAX: 2.6m

SIZE – Common SIZE – Indo-Pacific

L: ♂ 1.7–2.5m, ♀ 1.6–2.4m; L: ♂ 2–2.6m, ♀ 1.9–2.2m;
WT: 150–200kg; MAX: 2.7m, 235kg WT: 150–235kg; MAX: 2.6m
Larger in the North Atlantic than the North Pacific. Calf – L: 80–93cm; WT: c. 7–10kg

COMMON DOLPHIN 247

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 TUCUXI
Sotalia fluviatilis (Gervais and Deville in Gervais, 1853)

The tucuxi (pronounced ‘too-koo-shee’) is the only member of the dolphin family Delphinidae to be found exclusively in
freshwater – and one of two dolphins inhabiting the Amazon River basin (the other being the Amazon River dolphin). It
was recently separated from the very similar and closely related Guiana dolphin.
IUCN status Endangered (2020).
Population Unknown. Decreasing (an overall decline of at least 50 per cent during the period 2000–2047 is plausible).
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies. In 2007, S. fluviatilis was formally separated into two distinct species
– the tucuxi (S. fluviatilis) in the Amazon River basin, and the Guiana dolphin (S. guianensis) in coastal waters of
Central and South America and the Orinoco River system.
Other names Brazilian dolphin.
DISTRIBUTION Throughout the Amazon River basin. It appears to be continuous along the Amazon River and is found
in most of its tributaries. There is possible overlap between the two Sotalia species in the mouth of the Amazon River.
Frequently inhabits lakes and oxbows connected to the main system. Strongly prefers areas with reduced current:
waterway junctions, within 50m of riverbanks (rather than in mid-stream), and in lakes. Recent genetic research confirms
that the Sotalia species living in the lower reaches of the Orinoco River system is the Guiana dolphin, not the tucuxi.
Largely sympatric with the Amazon River dolphin, but the two species rarely interact.
BEHAVIOUR Can be quite aerially active and performs a variety of leaps and somersaults. Frequently spyhops, rolls at
the surface, lobtails and flipper-slaps. Generally shy and more difficult to approach than the Guiana dolphin, and not
known to bow-ride.
FOOD AND FEEDING At least 27 species of fish, particularly toothless characins, croakers, catfish. Feeds individually
and in groups, sometimes appearing to use cooperative feeding techniques.
TEETH Upper jaw 56–70; lower jaw 52–66.
GROUP SIZE AND STRUCTURE Typically up to 4 (occasionally 6) in mixed groups of adults and calves. As many as 30
have been reported on occasion.

ADULT MALE Flukes grey on

both sides

SIZE
L: ♂ 1.4–1.5m, ♀ 1.3–1.5m;
WT: c. 35–45kg; MAX: 1.5m, 53kg
Calf – L: 71–83cm; WT: c. 8kg

DIVE Sequence Head and beak appear at c. 45° angle (eye often visible); surfaces quickly (typically spends less than
1 second at surface); when travelling quickly, may porpoise. • Depth Unknown. • Duration Typically 20–120 seconds,
separated by shorter dives of 5–10 seconds.

248 LONGER-BEAKED OCEANIC DOLPHINS

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 Short, wide-based, triangular ADULT
Medium to dark bluish- dorsal fin midway along
or brownish-grey back (narrower base than in
upperside (variable) Guiana dolphin)
Bulbous melon merges
smoothly with rostrum
(no crease) Rather stocky
body May have appearance
of darker cape
Moderately
long, slender
beak Tailstock often
light grey

Light grey to pinkish

underside and sides
Mouthline Broad, pointed (pinker in warm waters
curves gently flippers (grey and when active – result
upward both sides) Variable broad, medium of vascularisation rather
to light grey band than pigmentation)
Lower jaw Variable broad, dark roughly from above
flipper to anus Belly tends to be
light grey band from eye to
lighter than in
or creamy- flipper (more clearly
Guiana dolphin
coloured to defined than in
pinkish Guiana dolphin)

May be hooked or
recurved at tip DORSAL FIN COMPARISON
Smooth
leading edge

Small hump on
leading edge
Wider base

Tucuxi Guiana dolphin Amazon River dolphin

10°
VENEZUELA
ATLANTIC
OCEAN
GUYANA
SURINAME
FRENCH GUIANA
AT A GLANCE Amazon River basin • Small
COLOMBIA
size (smaller than Guiana dolphin) • Dark grey
nco
Bra

above, pinkish-white below • Bulbous melon

0°
zon
Am a
Ne

ECUADOR imoes
gr

Sol
merges smoothly with rostrum (no crease) •
o

Maranón a jós
eir pa
Ta
Moderately long, slender beak • Wide-based,
ad
M

10° triangular dorsal fin midway along back • Can

PERU be aerially active
BOLIVIA
PACIFIC BRAZIL
OCEAN

Amazon basin

TUCUXI 249

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 GUIANA DOLPHIN
Sotalia guianensis (Van Bénéden, 1864)

Separated in 2007 from the closely related and very similar freshwater species, the tucuxi, the Guiana dolphin lives
mainly in warm coastal waters (although its presence has recently been confirmed in the Orinoco River). At first glance,
it resembles a small bottlenose dolphin.
IUCN status Near Threatened (2017).
Population Probably minimum c. 10,000. Trend unknown.
Classification Odontoceti, family Delphinidae.
Taxonomy No recognised forms or subspecies.
Other names Costero, estuarine dolphin, marine tucuxi.
DISTRIBUTION Range appears to be discontinuous along the tropical and sub-tropical Caribbean and Atlantic coasts
of Central and northern South America from the mouth of the Layasiksa River, in northern Nicaragua (14°N), to
Florianópolis, in southern Brazil (27°S). A population has also been described in Maracaibo Lake, a large semi-enclosed
estuarine system in northwestern Venezuela. There is possible overlap between the two Sotalia species in the mouth
of the Amazon River, but only Guiana dolphin occurs in the lower Orinoco. Mostly favours shallow nearshore marine
waters (within 100m of shore), especially in estuaries, bays, large gulfs and other protected coastal waters. Found in a
wide range of water depths, temperatures, salinity and turbidity, but mostly in habitats less than 5m deep.
BEHAVIOUR Can be quite aerially active and performs a variety of leaps and somersaults. Frequently spyhops, rolls at
the surface, lobtails and flipper-slaps. Typically indifferent to quiet boats, and easier to approach than the tucuxi, but
often avoids approaches by boats with running engines. Not known to bow-ride, but it may surf in the waves and wakes
produced by passing boats.
FOOD AND FEEDING At least 70 species of fish; sometimes cephalopods; occasionally crustaceans. Feeds individually
and in groups, using wide variety of feeding techniques, sometimes cooperatively.
TEETH Upper jaw 60–72; lower jaw 56–64.
GROUP SIZE AND STRUCTURE Typically up to 10 (4–6 most common) in mixed groups of adults and calves, with
up to 50–60 not uncommon. Group sizes may be larger, on average, in the wet season. Larger, cooperative feeding
aggregations are most common in southern Brazil with up to 400 observed together.

ADULT Flukes grey on

both sides

Fast swimming

DIVE Sequence Head and beak appear at c. 45° angle; surfaces quickly (typically spends less than one second
at surface); when travelling quickly, may porpoise. • Depth Unknown. • Duration Typically 20–120 seconds,
separated by shorter dives of 5–10 seconds.

Slow swimming

250 LONGER-BEAKED OCEANIC DOLPHINS

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 Medium to dark Small hump on Fin may be ADULT
Bulbous melon bluish- or brownish-grey leading edge of dorsal hooked at tip
merges smoothly upperside (variable) fin (cf. tucuxi)
with rostrum
Short, wide-based, triangular
(no crease)
dorsal fin midway along back
Rather stocky body (wider base than in tucuxi)
Moderately
long, slender May have grey blaze
beak below dorsal fin and
on tailstock (highly
variable)

Broad, pointed Belly tends to

Underside of flippers (grey
lower jaw light Light grey to be darker than
Variable broad, poorly both sides) in tucuxi
grey or creamy- pinkish underside
coloured to defined dark band from and sides
pinkish eye to flipper (less well
Mouthline curves defined than in tucuxi)
gently upward

ADULT VARIATION

Pinker in warm waters and when

SIZE active – result of vascularisation
L: ♂ 1.6–1.9m, ♀ 1.6–2m; rather than pigmentation
WT: c. 50–80kg; MAX: 2.2m, 121kg
Calf – L: 90–106cm; WT: 12–15kg

20°

AT A GLANCE Atlantic coasts of Central and

10°
northern South America (and lower Orinoco
0°
River) • Small size (larger than tucuxi) • Dark
grey above, pinkish-white below • Bulbous
10° melon merges smoothly with rostrum (no
crease) • Moderately long, slender beak •
20° Wide-based, triangular dorsal fin midway
along back • Can be aerially active
30°

GUIANA DOLPHIN 251

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 INDUS RIVER DOLPHIN
Platanista minor Owen, 1853

Thanks to decades of strenuous conservation efforts, the Indus River dolphin population has increased in recent years,
but it is still teetering on the brink. In particular, it is threatened by dams and irrigation barrages (which severely
fragment the population and reduce river flow), fisheries bycatch and pollution.
IUCN status Still listed as South Asian river dolphin: Endangered (2017). Indus subspecies (i.e. this taxon) Endangered
(2004).
Population c. 2,000 (2,000 in Pakistan, c. 7–10 in India). Increasing (up from 132 in 1972).
Classification Odontoceti, family Platanistidae.
Taxonomy Indus and Ganges River dolphins were classified as belonging to a single species from the 1880s until
the 1970s. Then they were separated into two distinct species until the late 1990s, when they were downgraded to a
single species (‘South Asian river dolphin’) with two subspecies. In 2021, however, strong evidence was presented for
a return to separate species status. They are now estimated to have diverged c. 550,000 years ago.
Other names Bhulan, blind river dolphin.
DISTRIBUTION In the 1870s, the Indus River dolphin was widely distributed throughout 3,500km of the Indus River
and its five large tributaries (Jhelum, Chenab, Ravi, Sutlej and Beas), in Pakistan and north-western India. However,
this historical range has declined by 80 per cent in the past century. Development of the vast Indus Basin Irrigation
System has severely fragmented the population within a network of irrigation barrages. The current range comprises
only a 1,154km stretch of the Indus River in Pakistan, between the Chashma and Kotri barrages, plus a very small
surviving sub-population in the Beas River, north-west India. Geographically isolated from the Ganges River dolphin.
Usually favours turbid, fresh (sometimes brackish) water that is relatively shallow (often only a few metres deep). Most
common in areas that create eddy countercurrents, such as small islands, sand bars, river bends or meanderings, and at
the confluences of rivers and tributaries, where it finds refuge in slower water and where prey is abundant.
BEHAVIOUR Generally cryptic and difficult to observe well, especially if there is any water disturbance. Frequently lives
near humans, but does not bow-ride. Rarely breaches, though it sometimes surfaces energetically showing most of the
head and body. Sometimes swims on its side.
FOOD AND FEEDING Wide variety of fish and invertebrates (including prawns, snails and clams). Feeds from surface to
riverbed, using long beak to probe mud.
TEETH Upper jaw 62–74; lower jaw 60–72.
GROUP SIZE AND STRUCTURE Usually solitary or in mother-calf pairs, but may be in fluid groups of up to 10 animals;
loose aggregations of as many as 30 have been reported.

ADULT SIDE-SWIMMING

SIZE
L: ♂ 1.6–2.2m, ♀ 1.6–2.2m;
WT: 70–85kg; MAX: 2.41m, 120kg
Calf – L: 60–70cm; WT: 4–7.5kg

DIVE Sequence Several types of surfacing (largely age-dependent), with varying parts of head, rostrum, dorsal fin,
back and tailstock visible; more energetic surfacing involves beak breaking surface at 45° angle and most or all of
body visible before bending sharply as beak re-enters water first. • Depth Unknown, but usually in shallow water
(less than 30m). • Duration Average dive time 30–90 secs (maximum 8 minutes 24 seconds).

252 RIVER DOLPHINS

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 External ear larger
Fairly uniform May become
Extremely small eyes than eye (located just
brownish-grey or blotchy with age ADULT
barely visible externally above eye – unique in
greyish-brown
(pinholes located just odontocetes)
(variable – and
above upturned corners Low, broad-based,
colour may fade
of mouth) triangular dorsal fin
with age)
(variable – sometimes
Very steep Stocky body more like a hump or
forehead dorsal ridge)

Dorsal fin about

No crease two-thirds of the
between way along back
beak and
forehead

Distinctly Trailing edge of

Very long, flippers usually Sometimes slightly
upturned corner lighter (typically
narrow beak of mouth scalloped but may
(similar length be straight pinkish) on underside
in both sexes) and around rostrum
Anterior teeth longer, Visible ‘finger’
Teeth wear with age visible outside closed Very large, bones along
(become broad and mouth, and interlock broad flippers surface of flippers
flattened)

Distinct (but low)

longitudinal ridge on ADULT
Beak laterally
compressed forehead (in front of
blowhole) Broad flukes
with pointed tips
Widens
at tip
Distinct
median
notch

Blowhole unusual in being single Flippers do not fold flat against

slit (instead of crescent-shaped) body (cf. most marine dolphins)
running along long axis of body and remain perpendicular to it Rear margin
(in ridge on forehead) concave

AFGHANISTAN current range

historical range
Rasul
AT A GLANCE Indus River and tributaries in
Chashma Jinnah Marala
lum Madhopur
Jhe Khanki Beas
na
Qbad
Nehar Pakistan and Beas River in north-west India
b

Che lej
Trimmu Sut

Taunsa
Ghazi Ghat Sidhani
Rav
i Harike Ropar
Hussainiwala • Predominantly brownish-grey colour • Small
size • Very long, narrow (vaguely gharial-like)
Suleimanki

PAKISTAN S ut lej
Islam
Panjnad
beak • Long teeth visible at front end of
Guddu
beak when mouth closed • Low, broad-based
Sukkur INDIA
triangular dorsal fin • Surfaces quietly,
Indus

cryptically, unpredictably and rapidly

Kotri
dam
barrage

INDUS RIVER DOLPHIN 253

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 GANGES RIVER DOLPHIN
Platanista gangetica (Lebeck, 1801)

Living in muddy, riverine waters, the endangered Ganges River dolphin is functionally blind (at most, it can probably
detect changes in light intensity and direction) and relies almost entirely on echolocation to navigate and find food.
IUCN status Still listed as South Asian river dolphin: Endangered (2017). Ganges subspecies (i.e. this taxon)
Endangered (2004).
Population Possibly as few as 3,500. Declining (and waterways development under India’s National Waterways Act
2016 has the potential to wipe out most of its habitable range).
Classification Odontoceti, family Platanistidae.
Taxonomy Indus and Ganges River dolphins were classified as belonging to a single species from the 1880s until
the 1970s. Then they were separated into two distinct species until the late 1990s, when they were downgraded to a
single species (‘South Asian river dolphin’) with two subspecies. In 2021, however, strong evidence was presented for
a return to separate species status. They are now estimated to have diverged c. 550,000 years ago.
Other names Susu, Ganga river dolphin, Gangetic dolphin, Indian river dolphin, blind river dolphin, shushuk, suongsu,
soos, shus and many other vernacular names.
DISTRIBUTION Historically, the Ganges River dolphin occurred throughout the Ganges–Brahmaputra–Megna and
Karnaphuli–Sangu river systems, in the main rivers as well as small and large tributaries. It still has a relatively broad
range in north-east India and Bangladesh, with a small, isolated sub-population in the Karnali River and tributaries
(south-west Nepal) and occasional reported sightings during the monsoon season in Bhutan. However, it has
disappeared from many upstream areas, in particular. Ganges River dolphins have been reported moving along the
coast of the Bay of Bengal when monsoons flush freshwater out along the east coast of India. Distribution tends to
be quite patchy and is highly variable according to season (though, in many places, dams and irrigation barrages now
block these seasonal movements). Geographically isolated from the Indus River dolphin. Usually frequents turbid, fresh
(sometimes brackish) water that is relatively shallow (often only a few metres deep). Most common in areas that create
eddy countercurrents, such as small islands, sand bars, river bends or meanderings, and at the confluences of rivers and
tributaries, where it finds refuge in slower water and where prey is abundant.
BEHAVIOUR Generally cryptic and difficult to observe well, especially if there is any water disturbance. Frequently lives
near humans, but does not bow-ride. Rarely breaches, though it sometimes surfaces energetically showing most of the
head and body. Sometimes swims on its side.
FOOD AND FEEDING Wide variety of fish and invertebrates (including prawns, snails and clams). Feeds from surface to
riverbed, using long beak to probe mud.
TEETH Upper jaw 50–62; lower jaw 52–62.
GROUP SIZE AND STRUCTURE Usually solitary or in mother-calf pairs, but may be in fluid groups of up to 10 animals;
larger, loose aggregations have been reported.

SIZE
L: ♂ 1.5–2.0m, ♀ 1.8–2.5m;
WT: 70–90kg; MAX: 2.67m, 114kg
Calf – L: 60–70cm; WT: 4–7.5kg

DIVE Sequence Several types of surfacing (largely age-dependent), with varying parts of head, rostrum, dorsal
fin, back and tailstock visible; more energetic surfacing involves beak breaking surface at 45° angle and most or
all of body visible before bending sharply as beak re-enters water first. • Depth Unknown, but usually in shallow
water (less than 30m). • Duration Average dive time 2 minutes (maximum 7 minutes 45 seconds).

254 RIVER DOLPHINS

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 Fairly uniform grey or ADULT MALE
Extremely small eyes barely visible greyish-brown (variable
externally (pinholes located just above – and colour may fade Low, broad-based,
upturned corners of mouth) with age) triangular dorsal fin
(variable – sometimes more
External ear larger like a hump or dorsal ridge)
than eye (located just Stocky body
Very steep above eye – unique in
forehead odontocetes) Dorsal fin about two-thirds
of the way along back
No crease
between beak May become
and forehead blotchy with age

Anterior teeth Very large,

longer, visible broad flippers Sometimes
outside closed slightly lighter
mouth, and Distinctly upturned Trailing edge usually (typically pinkish)
interlock corner of mouth scalloped but may be on underside or
straight around rostrum
Teeth wear with age Very long, narrow beak
(become broad and Visible ‘finger’ bones
(not upturned, cf. female) along surface of flippers
flattened)

Colour highly
variable
Beak tends to be
ADULT FEMALE
proportionately
longer in female

Substantially longer
beak may curve upwards
and sideways (only in
female)

AT A GLANCE River systems in India,

Bangladesh, Nepal and, rarely, Bhutan
• Predominantly grey colour • Small size
• Very long, narrow, (vaguely gharial-like)
beak • Long teeth visible at front end of
beak when mouth closed • Low, broad-based
triangular dorsal fin • Surfaces quietly,
cryptically, unpredictably and rapidly

present present (seasonal) absent

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 YANGTZE RIVER DOLPHIN or BAIJI
Lipotes vexillifer Miller, 1918

The Yangtze River dolphin or baiji (pronounced ‘by-gee’ and used interchangeably) has the unfortunate distinction of
being the first cetacean to have been driven to extinction by human activity (although, officially, it is still classified
as ‘possibly extinct’). The last authentic sightings were of a stranded pregnant female in 2001 and a live animal
photographed in 2002. Claimed sightings since are most likely to have been Yangtze finless porpoises.
IUCN status Critically Endangered (Possibly Extinct) (2017).
Population Considered functionally extinct since 2007 (even if a few individuals survive – which is unlikely – there is
no hope of recovery). Genetic analysis estimates an effective population size of more than 100,000 about 1,000 years
ago.
Classification Odontoceti, family Lipotidae.
Taxonomy No recognised forms or subspecies.
Other names Chinese river dolphin; rarely – Changjiang dolphin, whitefin dolphin, whitefag dolphin, ji, peh ch’i
(merely an older way of transliterating the Chinese – i.e. the same as baiji).
DISTRIBUTION The species’ recent historical range was a 1,700km stretch of the middle and lower reaches of the
Yangtze River, China, and smaller connecting rivers, from the Three Gorges area to the river mouth near Shanghai. It
also occurred in the Qiantang (Fuchun) River, south of the Yangtze River mouth, until construction of a hydroelectric
power station began in 1958; in two large lakes connected to the Yangtze (Dongting and Poyang); and in the Yangtze
estuary. The preferred habitat was anywhere near interrupted water flow (such as below bends in the river, mud banks
and sand bars) providing plentiful food and refuge from the strong current.
BEHAVIOUR Elusive and difficult to approach, typically making a long dive and changing direction underwater. Most
active during the day, resting at night in areas where the current is slow. No evidence of aerial behaviour such as
breaching. Used to associate with Yangtze finless porpoises.
FOOD AND FEEDING Freshwater fish. Feeding techniques unknown.
TEETH Upper jaw 62–68; lower jaw 64–72.
GROUP SIZE AND STRUCTURE There were typically 3–4 animals in a group (ranging from 2 to 6); the largest number
recorded together was 16 (no more than 10 after 1980).

ADULT

Distinct
notch

Longitudinally
oval blowhole

SIZE
L: ♂ 1.8–2.3m, ♀ 1.85–2.5m;
WT: 40–170kg; MAX: 2.6m, 240kg
Calf – L: 80–95cm; WT: 2.5–4.8kg

DIVE Sequence Surfaces slowly and smoothly, with typically only top of head, dorsal fin and small part of back
visible. • Depth Unknown. • Duration Typically 10–30 seconds, maximum recorded 3 minutes 20 seconds.
BLOW Blow barely visible but audible (like high-pitched sneeze) at close range.

256 RIVER DOLPHINS

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 Light bluish-grey to Low, broad-based triangular ADULT
Tiny eyes set unusually brownish-grey above
high on sides of head dorsal fin (set two-thirds of
(functional but poor vision) body length from snout)

Fairly stocky body

Pale ‘brush
Rounded melon strokes’ on sides
of head and neck Subtle, light flaring
with steep
on sides of tailstock
forehead

Bulging Broad, rounded Flukes rarely seen

‘cheeks’ flippers above surface
Long, narrow, slightly White to ashy-
upturned beak Lower jaw and white below
(lengthens with age – lower margin of
longer in female) upper jaw white

THE DEMISE OF THE YANGTZE RIVER DOLPHIN

The Yangtze River was once described as ‘teeming’
with dolphins but, by the 1950s, the population
had shrunk to low thousands. There were fewer
than 100 by 1990 and as few as 13 (including 1
calf) by 1997. There have been several intensive
visual and acoustic surveys in recent years, but
they failed to find any evidence of survivors.
Fishing was a major cause of the Yangtze River
dolphin’s decline, but it was also hunted for its
oil and skin. Other threats included depletion of
prey through overfishing, collisions with vessels,
noise pollution, agricultural run-off, industrial and
domestic pollution, riverbank development and
riverbed dredging. Meanwhile, dam construction
fragmented dolphin populations and dramatically
changed water levels, stratification, currents and
sandbanks. The Yangtze River dolphin’s fate was
sealed by an outrageous lack of funds, ineffective
project management, and a lack of coordination and
agreement between conservation groups, western
scientists and Chinese authorities. The result was
too little too late.

YELLOW SEA

AT A GLANCE Probably extinct • Historical

Yangtze range in lower and middle reaches of Yangtze
River, China • Small size • Light bluish-grey to
River
SHANGHAI

Three Gorges Dam

brownish-grey above • Long, narrow, slightly
Three Gorges Dam EAST CHINA SEA
upturned beak • Low, triangular dorsal fin
Reservoir
• Yangtze finless porpoise only other species
Qiantang River within range
Dongting Lake
Poyang Lake

Historical distribution

YANGTZE RIVER DOLPHIN 257

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 AMAZON RIVER DOLPHIN or BOTO
Inia geoffrensis (Blainville, 1817)

The Amazon River dolphin can be brilliant pink in colour and, uniquely among cetaceans, has two types of dentition:
conical grasping front teeth and crushing molar-like rear teeth (for crushing hard-bodied prey). The tucuxi is the only
other dolphin that overlaps in range.
IUCN status Endangered (2018).
Population Unknown. One guesstimate is 15,000, but possibly tens of thousands. Decreasing.
Classification Odontoceti, family Iniidae.
Taxonomy Two subspecies are recognised: Amazon River dolphin or common boto (I. g. geoffrensis) in the Amazon
and Orinoco drainage systems of Brazil, Peru, Ecuador, Colombia and Venezuela; and Bolivian river dolphin or Bolivian
bufeo (I. g. boliviensis) in the upper Madeira River drainage of Bolivia and along the Bolivia–Brazil border; some claim
a third, the ‘Orinoco river dolphin (I. g. humboldtiana)’ in the Orinoco basin of Venezuela and Colombia. There is also
a proposal for a second species, the ‘Araguaian river dolphin (I. araguaiaensis)’ of the Araguaia River and Tocantins
River basin, but current morphological and genetic evidence is considered insufficient to justify such treatment.
Other names Boutu (incorrect phonetic spelling of boto), pink river dolphin, Bolivian bufeo, Bolivian river dolphin, tonina.
DISTRIBUTION Endemic to the Amazon and Orinoco drainage basins in northern South America – a vast area of about
7 million sq km in Brazil, Bolivia, Colombia, Venezuela, Peru and Ecuador (and possibly some rivers in southern Guyana).
Occurs almost everywhere it can physically reach, without venturing into brackish or marine waters. Concentrated in
the main river channels during the dry season, but disperses into the complex mosaic of flooded forests, lakes, channels
and floodplains during the wet season. Prefers highly productive turbid water the colour of white coffee; however, it
also occurs in water the colour of black coffee (even though it is less productive).
BEHAVIOUR Truly a dolphin of the flooded forest, twisting and bending to navigate partially submerged trees and roots,
moving its large flippers independently for greater manoeuvrability; it can even swim backwards. Typically slow-moving
– though capable of short bursts of speed – it will wave a flipper above the surface, spyhop, lobtail and occasionally
breach (young tend to breach more than adults). The ‘chubby’ cheeks are believed to obstruct downward vision, which
may be why it is often observed swimming upside down. Responses to people range from shy to curious and playful.
The species does not bow-ride but often approaches boats, floating houses and people standing on riverbanks.
FOOD AND FEEDING Mainly fish, some crabs and molluscs; occasionally turtles. Usually a solitary feeder, but may work
cooperatively to herd and trap fish in shallow water.
TEETH Upper jaw 46–70; lower jaw 48–70.
GROUP SIZE AND STRUCTURE Mostly solitary, though frequently 2–3. Loose aggregations of up to 19 occur where
there is a rich food supply.

WHY PINK?
Many Amazon river dolphins – especially older males – (which makes the blood underneath the skin more
are bright pink in colour. This is not pigmentation, but is visible). The degree of pinkness varies greatly, according
caused by a progressive loss of the natural grey colour to age, water temperature and clarity, geographic
and the build-up of heavy scar tissue from fighting location and, in particular, the level of activity.

DIVE Sequence Two main types of surfacing: ‘sneak’ (above), which is most common and involves surfacing
slowly at shallow angle; and ‘arch and roll’ (below) when entire beak often exposed, then shows full length of
dorsal ridge before a high-arching roll to dive. • Depth Unknown, but probably shallow. • Duration Usually less
than one minute (30–40 seconds typical); maximum 110 seconds.
BLOW Blow can be explosively loud (like a snort or sneeze) and visible (up to 2m), or quiet and indistinct.

258 RIVER DOLPHINS

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 Bulging melon relatively small Pink or whitish to ADULT MALE
and soft (capable of complex blue-grey or dark grey
shape change – to alter frequency, (usually greyer above, ‘Lumpy’ body
beam width and directionality of paler or pinker below) quite different to
echolocation clicks) streamlined body
Moderately of most oceanic Low, wide-based
Small eyes with robust, dolphins dorsal fin leading to
Sparse, functional lenses flexible body long dorsal ridge
bristly hairs (larger than those of
(mainly on other river dolphins) Gently
upper jaw) sloping peak

Very long, Heavy scarring (tooth

robust beak rake marks caused by
‘Cobblestone’ skin
male–male fighting),
Beak may be scarred lesions common on
especially in older
(especially older Mouthline angles flippers and dorsal and
animals
animals) upwards in corner ventral ridges of tail

May have ‘chubby- In older males, flippers Pink colour varies in

cheeked’ appearance Large, broad, thick, may become upturned extent and intensity
almost triangular, highly on trailing edge
manoeuvrable flippers

Significantly shorter Less ‘ragged’ ADULT FEMALE

Less bulbous appearance
and lighter than male
melon than male Light to moderate scarring
(significantly less extensive
than on male)

Typically grey above,

pink below (variable
– but usually less SIZE
L: ♂ 2.2–2.5m, ♀ 1.8–2.3m;
pink than male) No ‘cobblestone’ WT: 70–185kg; MAX: 2.7m,
skin lesions 207kg
Calf – L: 75–90cm; WT: 10–13kg

10°
VENEZUELA
oco ATLANTIC
Orin
OCEAN
GUYANA FRENCH GUIANA
Casiquiare
SURINAME
AT A GLANCE Rivers and lakes in northern
Canal
COLOMBIA
nco
Bra

South America • Exclusively freshwater •

0°

azon
Ne

ECUADOR imoes Am
gr

Small size • Mid-grey to brilliant pink colour

Sol
o

Tucurui Rapids
Tucurui Dam
Maranón a jós
eir pa

• Very long, slender beak • Low, wide-based

ad Ta
M
Jirau Dam Santo Antonia Dam

dorsal fin leading to dorsal ridge • Large

ia

tins
gua

10° Teotõnio
Toca
A ra

Rapids
PERU
flippers • Bulbous forehead • Alone or in
BOLIVIA
PACIFIC
OCEAN
BRAZIL small groups
Amazon basin Amazon River dolphin Bolivian river dolphin
Region where type Orinoco river dolphin Araguaian boto
is uncertain

AMAZON RIVER DOLPHIN 259

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 FRANCISCANA
Pontoporia blainvillei (Gervais and d’Orbigny, 1844)

Despite being classified as a river dolphin, the franciscana is primarily a marine species. One of the smallest dolphins
in the world, it is known to fishermen as the ‘white ghost’, because it is often light in colour and tends to disappear
when it sees humans.
IUCN status Vulnerable (2017). Rio Grande do Sul/Uruguay sub-population Vulnerable (2003).
Population No overall figure, but probably fewer than 60,000. Four management units (Franciscana Management
Areas) recognised (although FMA–I is split into two sub-units), with abundance estimates ranging from a few hundred
to tens of thousands (FMA–III). Decreasing.
Classification Odontoceti, family Pontoporiidae.
Taxonomy No recognised subspecies, although recent genetic and morphological analyses suggest that franciscanas
inhabiting FMA–I likely represent a distinct subspecies. FMA–II and FMA–IV each comprise two distinct populations
that should probably be divided into FMA–IIa and FMA–IIb and FMA–IVa and FMA–IVb, respectively, for management
purposes.
Other names La Plata River dolphin, La Plata dolphin, toninha.
DISTRIBUTION Endemic to tropical and temperate waters along the east coast of South America (Brazil, Uruguay and
Argentina), the franciscana prefers shallow, turbid water in a narrow strip beyond the surf to about 30m depth. It
tends to avoid deep, clear and cold waters and, beyond 30m, density declines dramatically with distance from shore.
However, there have been sightings in water deeper than 50m and as far as 55km offshore (especially in the northern
part of the range). Range discontinuous, with major gaps. It sporadically enters the La Plata River estuary and other
estuaries.
BEHAVIOUR Generally cryptic and difficult to observe well in the wild, so little is known about its behaviour. Usually
shy, it avoids vessels and does not bow-ride. It does not appear to be aerially active.
FOOD AND FEEDING Wide variety of small marine fish; also takes squid, octopuses and crustaceans. Feeds mostly near
seabed, but uses entire water column and also takes some pelagic prey.
TEETH Upper jaw 106–116; lower jaw 102–112.
GROUP SIZE AND STRUCTURE Usually solitary or in small groups of 2–5, but aggregations up to 50 recorded.

Dorsal ridge
ADULT continues to
tailstock

Deep
median
notch

Crescent-shaped
blowhole set
slightly to left Broad, spatulate
flippers

DIVE Sequence Surfaces quietly and cryptically, with little or no splash; individuals in same group often surface and
breathe in synchrony. • Depth Unknown, but less common in water deeper than 30m. • Duration Foraging dives
average 22 seconds (range 3–83 seconds).

260 RIVER DOLPHINS

004 WDP FG.indd 260 29/11/2021 12:30

 Skin may lighten with Unmarked body ADULT MALE
age (some older animals (no distinctive
predominantly or scarring)
entirely white)
Low to moderately
Brownish to dull tall dorsal fin with
Steep, rounded grey upperside broad base
forehead (variable)
Relatively
Extremely long, small eyes Faint dorsal cape
slender beak may be visible
15–19cm in
male (17–22cm
in female)

Flexible neck Broad, spatulate Lighter brown Broad flukes

(unfused cervical flippers (sometimes yellowish)
vertebrae) to light grey lower
flanks and underside
Irregular and
Beak 12–15 per cent of
undulating
total length in adults Other than length, no known obvious
trailing edge
(longest relative to external morphological differences
body size of any living between northern and southern forms
cetacean) Visible ridges (like
finger bones) along
surface of flippers

DORSAL FIN VARIATION

SIZE
L: ♂ 1.2–1.4m, ♀ 1.5–1.6m; Dorsal fin highly variable (triangular to slightly Some fins can
WT: 20–40kg; MAX: 1.8m, 53kg falcate) but often appears backswept and always resemble those
Calf – L: 70–80cm; WT: 6–8.5kg has bluntly rounded tip of small sharks.

10°
(FMA: Franciscana
Management Area)
0°

AT A GLANCE Shallow tropical and temperate

10°
waters along east coast of South America •
Simple pale counter-shaded colour pattern
FMA-Ia
20°
FMA-Ib • Small size • Very long beak, often lifted
FMA-II into air on surfacing • Broad-based, rounded
30°

FMA-III
dorsal fin • Surfaces quietly and cryptically
with little or no splash
40° FMA-IV

50°

FRANCISCANA 261

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 DALL’S PORPOISE
Phocoenoides dalli (True, 1885)

Dall’s porpoise is probably the fastest small cetacean, typically seen as a splashy blur when it breaks the surface at
high speed. Unlike other porpoises, it often approaches boats and readily bow-rides and wake-rides.
IUCN status Least Concern (2017).
Population Minimum 1.2 million. Trend unknown.
Classification Odontoceti, family Phocoenidae.
Taxonomy Two subspecies are recognised (based on body colour pattern): P. d. dalli (‘dalli-type’ – the nominate form)
and P. d. truei (‘truei-type’). The dalli-type also has two minor colour morphs, distinguished by the size of their white
flank patch (larger in North Pacific–Bering Sea populations and smaller in Sea of Japan–Sea of Okhotsk populations).
Other names True’s porpoise.
DISTRIBUTION Deep cool temperate to sub-Arctic waters of the northern North Pacific and adjacent seas. Prefers
water colder than 17°C, with peak abundance below 13°C. Mainly offshore, but also in coastal areas where the water
is deeper than 100m. Dalli-type rarer where it overlaps with the truei-type (accounting for 4–20 per cent of individuals,
according to location).
BEHAVIOUR An energetic porpoise that can be almost hyperactive, darting jerkily and zigzagging around at high speed
(up to 55km/h). Over short bursts, it may be the fastest small cetacean. A keen bow-rider – indeed, the only porpoise
that often bow-rides – it prefers fast-moving vessels and will lose interest in slower ones. It will also ride the stern
waves of fast boats. Aerial behaviour such as breaching, tail-slapping or porpoising is extremely rare.
FOOD AND FEEDING Range of surface to mid-water fish and squid; may rarely take krill, shrimps and other crustaceans.
Mostly at night; recent research suggests daytime feeding in some areas.
TEETH Dalli-type: upper jaw 46–56; lower jaw 48–56. Truei-type: upper jaw 38–46; lower jaw 40–48.
GROUP SIZE AND STRUCTURE Usually in fluid groups of 2–10 (usually fewer than 5). Larger temporary aggregations
around prey concentrations (but these lack the cohesion of dolphin schools); largest groups occur in oceanic populations.

ADULT MALE DALLI-TYPE Small flukes have convex trailing Tips may become
edges (giving backward appearance) very rounded in
in male (more pronounced with age older animals
and straighter in female)

Prominent
median
notch

Rest of flukes and

Small head appears tailstock usually
White side patches visible black (can be pale White to light grey trailing
triangular from above on bow-riding animals edges on upperside
grey or even white)

Fast swimming

DIVE Sequence Typically swims fast, slicing through surface and creating characteristic V-shaped ‘rooster tail’ of
spray (which, in calm seas, enables positive ID even at great distance); when swimming slowly, barely disturbs
water, surfacing with slow, rolling motion like harbour porpoise but lifting deep tailstock higher. • Depth Most
foraging in less than 100m; however, capable of deep diving and may forage at 500+m. • Duration Typically 1–2
minutes, but some feeding dives may last more than five minutes.
Slow swimming

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 Striking black body with bright May have Fin often canted forward ADULT MALE DALLI-TYPE
white side patches extending pronounced (mostly in large, older
across underside and about hump in front males – not in females) Wide-based
halfway up each flank of fin triangular dorsal fin
Robust body Hooked tip
Relatively (less robust (variable)
small head in female) Light grey to white ‘frosting’
with steeply on upper and rear part of fin
sloping (becomes whiter with age)
forehead
Female has
less deep
tailstock

Short beak
Moderate to large keel
(with no clear White flank patch extends
Some White flank patch makes tailstock appear
demarcation forward roughly to level of
individuals smaller than in truei- exceptionally deep
from melon) leading edge of dorsal fin
Small may have type and does not (more pronounced in
(to midpoint in Sea of Japan males and with age)
flippers flecks of black extend as far forward
population)
near head on white patch

Slightly slimmer but longer ADULT MALE TRUEI-TYPE

body than dalli-type (at least
in Japanese coastal waters) Larger flank patch extends
further forward (at least to
level of flippers)

minent
dian SIZE
ch L: ♂ 1.8–2.4m, ♀ 1.7–2.2m;
WT: 135–200kg; MAX: 2.4m, 218kg
Calf – L: 0.9–1.2m; WT: c. 11kg

75°

AT A GLANCE Cool, deep waters of the

North Pacific and adjacent seas • Striking
black body with prominent white side patch
60°

• Highly distinctive ‘rooster tail’ of spray

45°
common on surfacing • Can be energetic and
almost hyperactive (behaviour more dolphin-
30°
like) • Wide-based, triangular two-toned
15° dorsal fin • Small size • Exceptionally deep
tailstock
0°

15°

dalli-type dalli- and truei-types

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 HARBOUR PORPOISE
Phocoena phocoena (Linnaeus, 1758)

The harbour porpoise may be the most widespread and commonly seen of all the porpoises, but it can be surprisingly
difficult to observe properly. It normally surfaces briefly, shows little of itself and rarely approaches boats, so a typical
sighting is not much more than a fleeting glimpse.
IUCN status Least Concern (2020). Black Sea subspecies Endangered (2008). Baltic Sea sub-population Critically
Endangered (2008).
Population Minimum c. 1 million. Trend unknown globally, but many populations decreasing.
Classification Odontoceti, family Phocoenidae.
Taxonomy Three subspecies are recognised: Atlantic harbour porpoise (P. p. phocoena), Black Sea harbour porpoise
(P. p. relicta) and Pacific harbour porpoise (P. p. vomerina). There may be two more (as yet unnamed): Western Pacific
harbour porpoise, and Afro-Iberian harbour porpoise (from the southern Iberian Peninsula and Mauritania).
Other names Harbor porpoise (American spelling), common porpoise; rarely – herring hog (especially Maine, USA),
puffing pig (especially Atlantic Canada) or puffer, after its sneeze-like blow.
DISTRIBUTION Discontinuous range in cool temperate and sub-Arctic waters of the northern hemisphere. Favours
coastal waters and frequents relatively shallow bays, estuaries, fjords, tidal channels and even harbours (and will also
swim a considerable distance upriver in some areas). Rarely in seas deeper than 200m – although it is known in deep
waters in some inshore regions and was recently found in deep North Atlantic waters during winter. Favours areas of
strong tidal currents, usually near islands or headlands.
BEHAVIOUR Usually avoids boats, or is indifferent, so it can be difficult to approach and follow (although it is more
approachable in some areas, such as the San Francisco Bay area and in the Bay of Fundy, eastern Canada). Most
approachable during extended periods of inactivity, especially on calm days. Very rarely bow-rides or wake-rides.
Acrobatics are uncommon, although it sometimes makes arc-shaped leaps when chasing prey and very occasionally
tail-slaps when socialising.
FOOD AND FEEDING Mainly small schooling fish; some squid and octopuses; calves will eat small crustaceans during
early phase of weaning. Opportunistic, taking prey mainly from near seabed, but will also forage in water column and
close to surface.
TEETH Upper jaw 38–56; lower jaw 38–56. Teeth are spatulate, as in all porpoises.
GROUP SIZE AND STRUCTURE Usually in mother–calf pairs or loose, fluid groups of 1–3 (larger groups of 6–10 are not
uncommon in some areas); several hundred have been observed at good feeding grounds.

ADULT Concave trailing

edges

Prominent
notch in
middle

Flukes all dark

DIVE Sequence Surfaces with slow, forward-rolling motion (as if dorsal fin is mounted on revolving wheel and
lifted briefly above surface, then withdrawn), and dives with little or no splash; when feeding (swimming fast and
erratically) may produce distinctive spray (known as ‘pop-splashing’) – very different to ‘rooster’s tail’ produced by
Dall’s porpoise. • Depth Typically 20–130m (maximum 410m). • Duration Most dives c. 1 minute; maximum
6 minutes.
BLOW Indistinct, but on calm days a sharp, sneeze-like puffing sound may be audible.

264 PORPOISES

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 Overall impression of ADULT
nondescript two-toned Longer leading dorsal fin
pigmentation edge may have 12–19 small
Many subtle bumps (called tubercles or
Varying degree of differences in denticles), possibly serving
bilateral asymmetry pigmentation hydrodynamic function
Small, conical in pigmentation between
head (usually individuals Low, triangular dorsal fin
slightly larger (appears large relative
in Atlantic Great variation
to visible portion of back
than Pacific) in colour pattern
when surfacing)
between individuals
Indistinct beak
(usually slightly Medium to Fin dark grey
shorter in dark grey to black
Atlantic than upperside Concave trailing
Pacific) edge (variable)
Robust
body
Broad base

Fin centrally
placed
Slight dorsal ridge
from dorsal fin to
tailstock

White or light grey

Dark grey Small, dark flippers Body colour underside and chin
or black located in white merges from
‘lips’ area of body dark to light
through flecking
Straight mouthline or streaking,
slopes upward with various
towards eye Dark grey stripe shades of grey
(variable width)
from mouth to
May be 1–3 dark streaks on flipper (almost SIZE
chin (variable length, from L: ♂ 1.2–1.8m, ♀ 1.5–1.9m;
impossible to see
‘lips’ sometimes as far back WT: 45–70kg; MAX: 2m, 75kg
in the field)
minent as area between flippers) Calf – L: 70–90cm; WT: c. 5–6kg
ch in Substantial geographical variation.
dle
80°

70°

60°

50°

40° AT A GLANCE Cool temperate and sub-Arctic

waters of the northern hemisphere • Small
30°

20°

size and robust body • Dark back and low,

10°
0°

triangular dorsal fin • No beak • Usually shy

10°
20°

and undemonstrative • Usually alone or in

30°

40°

50°
small, loose groups • Slow, forward-rolling
motion on surfacing
60°

70°

80°

primary range secondary range

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 VAQUITA
Phocoena sinus Norris and McFarland, 1958

In imminent danger of extinction, the vaquita is the most endangered marine mammal in the world. The biggest threat
for decades has been entanglement and accidental drowning in near-invisible gillnets; these are set for a variety of
species, but the main concern recently has been those set illegally for a 2m-long sea bass-like fish, the totoaba (highly
prized for its swim bladder, used in traditional Chinese medicine). Unless last-ditch conservation efforts are successful,
it will not survive for much longer. Very little is known about its life and habits.
IUCN status Critically Endangered (2017).
Population Just 6–19 (probably 10) in 2018. A 2019 survey estimated a population figure of 9.7. Calculated to have
sustained a population of 5,000 for more than 250,000 years, until relatively recently. Decreasing.
Classification Odontoceti, family Phocoenidae.
Taxonomy No recognised forms or subspecies.
Other names Gulf of California harbour (or harbor) porpoise, cochito (‘little pig’), Gulf porpoise, desert porpoise.
DISTRIBUTION Extreme northern end of the Gulf of California (Sea of Cortez), western Mexico (mainly north of 30°45’N
and west of 114°20’W). Prefers shallow, murky, sediment-laden offshore waters with strong tidal mixing. In modern
times, mostly in water 10–30m deep (rarely deeper than 40m). This is the most restricted distribution of any cetacean,
and there is no evidence to suggest that it has retracted in historical times. The entire range is less than 65km across
and is centred around Roca Consag, a 90m-high granite outcrop 27km east-north-east of San Felipe. Most recent
sightings have been between Roca Consag and San Felipe (almost all within sight of the outcrop) and less than 25km
from shore.
BEHAVIOUR Shy and retiring, typically surfacing away from vessels. Tends to avoid large motorised boats, but may
occasionally approach quiet drifting boats. Does not bow-ride, and aerial displays such as breaching are unknown.
Most sightings are fleeting and once only.
FOOD AND FEEDING Some 21 small (mainly bottom-feeding) fish species known; also takes squid and some
crustaceans. Feeding techniques unknown.
TEETH Upper jaw 32–44; lower jaw 34–40. Teeth are spatulate, as in all porpoises.
GROUP SIZE AND STRUCTURE Most sightings are of 1–3 individuals together, but up to 10 were observed in short-
lived, loose aggregations; small groups often consisted of several mother–calf pairs.
ADULT
Pointed tips

Median
notch

Slightly
pointed tips

DIVE Sequence Surfaces slowly and inconspicuously (almost impossible to see in anything but flat-calm
conditions), making slow, arching roll; typically surfaces 3–5 times, followed by longer dive of 1–3 minutes.
• Depth Shallow (rarely in water deeper than 40m). • Duration Maximum at least 3 minutes.

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 Upper half of leading edge of dorsal fin ADULT
Overall impression has small tubercles (bumps) in mature
Small size (one
dark, but in some adult (begin as whitish spots and
of the smallest
light conditions become more prominent with age)
cetaceans)
can appear olive or
tawny-brown Tall, roughly triangular, falcate dorsal
fin (proportionately taller than in
Pale other porpoises – up to 15cm)
face Stocky,
compact Dorsal fin often has
Melon has slight bulge in middle
distinct body
of leading edge
bulge
Fin halfway
along back Pointed
Blunt tips
head

Underside of
Virtually Whitish tailstock may be
Black or grey Proportionately underside
no beak large, broad-based darker than belly
patch around
each eye flippers
No sharp demarcation
Dark grey-black Paler grey on between darker upperside
stripe from lower sides and paler underside
beak to flipper
Blackish-
(variable)
grey ‘lips’
CALF

Tends to be darker
than adult

SIZE
L: ♂ 1.25–1.45m, ♀ 1.35–1.5m;
WT: 30–48kg; MAX: 1.5m, 55kg
Calf – L: 70–78cm; WT: 7.5kg

MEXICO
er

Tijuana UNIT
Riv

ED
do

MEX STATE
ora
Col

ICO S
Ensenada
El Golfo de Santa Clara
BAJA
CALIFORNIA
AT A GLANCE Extreme northern end of Gulf of
Puerto Peñasco
San Felipe Roca Consag California (Sea of Cortez) • Very small size •
SONORA
Appears all grey or grey-brown in good light
• Prominent dorsal fin • Lack of prominent
Gulf of California beak • Dark ‘lips’ and eye patch • Group size
usually 1–3 • Typically surfaces slowly and
PACIFIC
OCEAN
inconspicuously

primary range secondary range

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 BURMEISTER’S PORPOISE
Phocoena spinipinnis Burmeister, 1865

Burmeister’s porpoise is inconspicuous and easy to overlook. There have been relatively few scientific observations of
live animals, yet it may be fairly common and widespread along the coasts of South America.
IUCN status Near Threatened (2018).
Population Unknown. Trend unknown.
Classification Odontoceti, family Phocoenidae.
Taxonomy No recognised subspecies, but recent genetic studies indicate two sub-populations: Peruvian and Chilean–
Argentinian (which differ genetically and in body size).
Other names None.
DISTRIBUTION Coastal waters of South America, over the continental shelf. Possibly more common in the Pacific than
the Atlantic. Frequents inshore bays, channels and fjords; occasionally observed inside the kelp line. Most sightings in
southern Chile have been in water deeper than 40m (favouring deeper channels) and at least 500m from shore. May
also be present further offshore – up to 50km recorded in Argentina and one 2002 sighting of 150 individuals 20km
offshore in Peru.
BEHAVIOUR Tends to be inconspicuous, and rarely engages in aerial activity such as porpoising or breaching (although
some individuals have been reported riding coastal breakers and occasionally leaping as part of this surfing behaviour).
Almost impossible to see in rough weather. Bursts of speed associated with feeding are common.
FOOD AND FEEDING Mainly fish; also some squid, shrimp and krill. Feeding techniques unknown.
TEETH Upper jaw 20–46; lower jaw 28–46. Teeth are spatulate, as in all porpoises. Young tend to have higher tooth
counts than adults.
GROUP SIZE AND STRUCTURE Typically seen alone, in pairs or in small groups (1–4, occasionally up to 8), though
temporary larger aggregations have been reported.
Tubercles (bumps)
ADULT may extend along
midline of back

Distinct
notch in
middle

May have dark grey

blowhole-to-apex stripe
(usually joins ‘lip’ patch)

Series of small tubercles (bumps) in 2–7

ADULT FIN VARIATIONS
Leading edge long
and straight or
distinctly concave
Dorsal fin shape broadly rows along entire leading edge of dorsal fin Tubercles become
triangular but variable (and often along midline of back) sometimes larger and sharper
(no sexual dimorphism – visible at sea (broadens from one row to with age
though sometimes larger multiple rows at wider part of fin)
in males)
Trailing edge
usually straight or
slightly convex

DIVE Sequence Surfaces causing little disturbance, showing dorsal fin but little of body; slow, forward-rolling
motion, not unlike harbour porpoise; overall impression may be like an undulating sea lion. • Depth Usually in
water shallower than 200m. • Duration Limited records average 1–3 minutes.

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 Blunt, conical head Medium to dark grey or brown ADULT left side
(slightly less bulbous Some individuals upperside and sides (turns jet
in female) have brownish hue black soon after death)
Unique backward-leaning
Small, dorsal fin (rises at very
Well-defined darker eye stocky shallow angle)
patch (surrounded by subtle body
light grey to white halo)
Dorsal fin further
back than on
most other small
cetaceans

Indistinct beak
with slightly Male has slightly
darker ‘lips’ Proportionately deeper tailstock
Forward projection Light to dark grey stripe large, broad-based Lighter grey than female
of eye patch (like from flipper to ‘lip’ patch, flippers to whitish
a ‘tear’) may reach wider than on right side underside
‘lips’ in some (hard to see in the field)
individuals Flipper-to-‘lip’ stripe
well defined by thin,
light grey ‘borders’ ADULT right side

Stripe narrows
Dark grey stripe from
as it curves
flipper to ‘lip’ (narrower
downward
than on left side)
Stripe extends
SIZE further forward
L: 1.4–2.0m; on right side
WT: 70–80kg; MAX: 2m, 105kg
Calf – L: 80–90cm; WT: 4–7kg

10°

primary range
0°
range likely
10°
only under AT A GLANCE Coastal waters of South
certain
oceanographic
America • Small size • Stocky body •
20°
conditions Appears very dark at sea • Unique backward-
30°
leaning dorsal fin well behind midline • Tends
to be inconspicuous, with little disturbance
40°
of water
50°

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 SPECTACLED PORPOISE
Phocoena dioptrica Lahille, 1912

The spectacled porpoise is instantly recognisable, with its striking black-and-white colour pattern and the male’s
enormous dorsal fin. However, it is rarely seen and is one of the least known of all cetaceans.
IUCN status Least Concern (2018).
Population Unknown, but the high genetic diversity and abundance of standings suggests a reasonable population
size.
Classification Odontoceti, family Phocoenidae.
Taxonomy No recognised forms or subspecies; briefly (1985–95) placed in its own genus, Australophocaena, but
genetic and morphometric studies returned it to Phocoena.
Other names None.
DISTRIBUTION Possibly circumpolar in cool temperate to polar waters (mostly, but not exclusively, north of the
Antarctic Convergence). Known mainly from strandings, especially along the east coast of Tierra del Fuego and
southern Argentina, but also from widely scattered offshore islands of the Antarctic Circumpolar Current. Preferred
water temperature is 0.9–10.3°C (most sightings in 4.9–6.2°C). Known mainly from coastal waters (also in some rivers
and turbid channels near the shore) but the primary habitat is believed to be deep offshore waters.
BEHAVIOUR There are only a few dozen confirmed sightings at sea. It is not known to be acrobatic and does not bow-
ride. Usually avoids boats (though it has approached research vessels).
FOOD AND FEEDING Little information, but feeds on anchovies and other small schooling fish, mantis shrimps and squid.
TEETH Upper jaw 32–52; lower jaw 34–46. Teeth are spatulate, as in all porpoises, or peg-shaped; they are often
hidden in the gums.
GROUP SIZE AND STRUCTURE Typical group size is 1–3 (average 2), but as many as 5 have been seen together; most
strandings are of solitary animals. Mother–calf pairs are often accompanied by one or two adult males (unlikely to be
biological fathers of calves – more indicative of a mate-guarding mating system, as seen in Dall’s porpoises).

ADULT MALE May have pale Some individuals have Upperside of flukes
saddle on back relatively light (grey/ usually grey but may
around dorsal fin brown) dorsal colour be white (underside
usually white but
may be grey)

Distinct
median
notch

May be no, one

or two light grey
stripes running
from blowhole to SIZE Small flukes with
apex of melon L: ♂ 1.9–2.2m, ♀ 1.3–2m; fairly straight
WT: 85–115kg; MAX: 2.2m, c. 120kg trailing edge
Calf – L: 0.9–1.2m; WT: probably 10–15kg

DIVE Sequence Usually surfaces inconspicuously with slow forward roll, not unlike harbour porpoise; strongly
arches back, possibly showing top of white sides, before diving; capable of porpoising fast (sometimes clearing
the water). • Depth Unknown. • Duration Unknown.

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 Glossy blue-black, Some individuals have Dorsal fin grows much larger ADULT MALE
grey-black or jet small tubercles on and more rounded in male Dorsal fin may
black upperside leading edge of fin (can be disproportionately be 25cm tall
Characteristic black huge and highly distinctive) Convex trailing edge
eye-patch with fine (uncommon in small cetaceans)
white ‘spectacles’ Convex
(present in most leading May have pale saddle on back
individuals) edge around dorsal fin that may
Extremely
extend up base of fin (believed
May have Robust broad base
Small, to fade rapidly after death)
bulge behind body
blunt
blowhole May show white line
head
along upperside of
tailstock

Slight Very small Brilliant white White sweeps

hint of Black ‘lips’ flippers far Sharp demarcation underside (to upwards at end
a beak surrounded forward on body between black and midway up flanks) of tailstock
by white white along a line level
Flippers can be dark, with the eye (though
medium grey or white may be feathering on
Flippers may have darker (sometimes with grey some individuals)
bands in ‘hand-like’ pattern margins) on upperside
ADULT FEMALE
Usually lighter on back than Usually slightly
male (more noticeable under convex on leading
good lighting conditions) and trailing edges
Dorsal fin smaller (up to
12cm), lower and more
triangular (cf. male)

The only strongly

sexually dimorphic
Both sexes may have one or two indistinct grey stripes (or wider stripe) leading porpoise species
from around ‘lips’ to flippers (may fade with age and absent in some individuals)

80°

70°

60°

AT A GLANCE Cool waters of the southern

50°

40°

30°

20°
hemisphere • Two-toned: black upperside,
10°
0°
brilliant white underside (with sharp
10°
20°
demarcation) • Small size • Absurdly large,
30°

40°
rounded dorsal fin of male • Strong sexual
50° dimorphism • White ‘spectacles’ distinctive
at close range
60°

70°

80°

probable range possible range

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 NARROW-RIDGED FINLESS PORPOISE
Neophocaena asiaeorientalis (Pilleri and Gihr, 1972)

In 2009 it was officially agreed that, instead of one species of finless porpoise, there are actually two – now called the
narrow-ridged finless porpoise and the Indo-Pacific finless porpoise – which are reproductively isolated and can look
sufficiently different to be told apart in the wild.
IUCN status Endangered (2017). Yangtze subspecies Critically Endangered (2012).
Population Unknown. Rough estimates exist for populations in the Yangtze River (c. 1,000), Japan (c. 19,000 in five
different sub-populations) and South Korea–Yellow Sea (21,500+ offshore; 5,500+ inshore). Decreasing.
Classification Odontoceti, family Phocoenidae.
Taxonomy Two subspecies are recognised: East Asian finless porpoise or sunameri (N. a. sunameri), and Yangtze
finless porpoise (N. a. asiaeorientalis). Recent genetic research strongly suggests that the latter is genetically and
reproductively isolated from its marine counterpart, and should be considered a separate species.
Other names Yangtze finless porpoise, East Asian finless porpoise, sunameri; black finless porpoise (resulting from
descriptions of dead animals – which darken after death).
DISTRIBUTION The subspecies sunameri frequents shallow, cool temperate coastal and estuarine waters of the
western North Pacific. Greatest densities tend to be in shallow bays and near the estuaries of large rivers (with a strong
preference for sandy or soft seafloors); may also enter mangrove swamps. Normally in water less than 50m deep, but
it has a greater tendency than the Indo-Pacific finless porpoise (with which it overlaps only in the Taiwan Strait) to
occur offshore. The nominate subspecies asiaeorientalis is the only porpoise restricted to freshwater and (assuming
that the Yangtze River dolphin is extinct) is the only extant cetacean in the Yangtze River, China. It occurs in the middle
and lower reaches of the Yangtze River (although its range has contracted dramatically and it no longer occurs beyond
Yichang, about 1,000km upstream), including Poyang and Dongting Lakes and Gan and Xiang Rivers.
BEHAVIOUR The East Asian subspecies tends to avoid boats in most areas, while the Yangtze subspecies is less shy and
more used to heavy boat traffic. Does not bow-ride and rarely breaches (although individuals in the Yangtze have been
observed leaping from the water and performing ‘tail stands’). Generally unobtrusive.
FOOD AND FEEDING Wide variety of fish, squid, cuttlefish and crustaceans. Feeding techniques unknown.
TEETH Upper jaw 32–42; lower jaw 30–40. Teeth are spatulate, as in all porpoises.
GROUP SIZE AND STRUCTURE Usually seen as singles, pairs (mother and calf or two adults) or in groups of up to 20.

Most individuals have single Dorsal ridge Tubercled patch very ADULT EAST ASIAN
central groove roughly from starts in narrow (0.2–1.2cm –
thoracic region to mid-body central groove sometimes to 2.4cm at
(depth highly variable) widest point)

Relatively
broad flukes

Central groove often splits into two lateral grooves 1–10 (usually 3–8) loosely
longitudinal rows of Trailing edge
running along each side of ridge (lateral grooves
tubercles at widest point concave
usually absent from individuals in Japanese waters)

DIVE Sequence Surfaces briefly and quietly, causing little disturbance of water; shows little of body and may
appear rounded as it rolls into the dive • Depth Most populations rarely in water deeper than 50m. • Duration
Maximum 4 minutes.

272 PORPOISES

004 WDP FG.indd 272 29/11/2021 12:30

 Overall impression Tubercles vary greatly, Body colour in most ADULT EAST ASIAN
of back is deep from pointed prickles populations light grey or
central crease in (sometimes with spine- creamy-white (but some
front of long, high like appearance) to individuals dark grey) Highest point of dorsal ridge
dorsal ridge blunt bumps nearer mid-point of body (cf.
Indo-Pacific finless porpoise)
No dorsal fin
Rounded head Slender body
Dorsal ridge usually obvious
with slightly from thoracic region to tailstock
bulbous melon
(may overhang
Dorsal ridge longer, higher
upper ‘lip’)
(1.2–5.5cm) and narrower
(0.2–1.2cm) than in Indo-
Pacific finless porpoise

No beak Relatively large Japanese colour morph

flippers (found in Japan and northern
China) often creamy-white
May be Neck more
lighter area flexible than in
around throat May have slightly
other porpoises
darker broad band
from mouth to
flipper
Entire dorsal
surface of ridge SIZE
ADULT YANGTZE L: 1.6–2.3m;
Body colour medium has 1–5 loosely
to dark grey (mostly longitudinal rows WT: 40–70kg; MAX: 2.27m, 110kg
of tubercles (East Asian), 1.77m (Yangtze)
darker than East Asian)
Calf – L: 75–85cm; WT: 5–10kg

Dorsal ridge lower (up

to 1.5cm) and narrower
(0.2–0.8cm at widest
point) than in East Asian

AT A GLANCE Shallow coastal waters of

40°
western North Pacific and Yangtze River
• Small size • Great individual, geographical
and age-related differences in colour •
Distinctive high dorsal ridge instead of dorsal
fin • Lighter individuals superficially resemble
30°

extralimital record from Okinawa

small belugas • Rounded head with no beak
• Nearly invisible if water is rough • Usually
20° alone or in small groups

Yangtze subspecies overlap with Indo-Pacific finless porpoise

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004 WDP FG.indd 273 29/11/2021 12:30

 INDO-PACIFIC FINLESS PORPOISE
Neophocaena phocaenoides (G. Cuvier, 1829)

The Indo-Pacific finless porpoise is cryptic by nature. Its range overlaps with the similar narrow-ridged finless porpoise,
but only in the region in and around the Taiwan Strait, where the two species have been observed within tens of metres
of each other.
IUCN status Vulnerable (2017).
Population Likely minimum 10,000. Decreasing.
Classification Odontoceti, family Phocoenidae.
Taxonomy No recognised forms or subspecies.
Other names Wide-ridged finless porpoise.
DISTRIBUTION Wide (albeit discontinuous) range in a narrow strip of shallow tropical to warm temperate coastal
waters in the north-eastern Indian Ocean and Southeast Asia. More tropical and wide-ranging than the narrow-ridged
finless porpoise. The greatest densities tend to be in shallow bays and the lower reaches of some large rivers, usually
in water less than 50m deep. Will enter mangrove swamps and major river systems (up to 60km upstream). Appears to
show a strong preference for sandy or soft seafloors.
BEHAVIOUR Tends to avoid boats in many areas and does not bow-ride (though it will occasionally ride in the wake of
fast vessels). Rarely breaches. Generally unobtrusive, but when startled it may escape with splashes not dissimilar to
those produced by Dall’s porpoise.
FOOD AND FEEDING Wide variety of fish, squid, cuttlefish and crustaceans. Feeding techniques unknown.
TEETH Upper jaw 30–44; lower jaw 32–44. Teeth are spatulate, as in all porpoises.
GROUP SIZE AND STRUCTURE Usually seen as singles, pairs (mother and calf or two adults) or in groups of up to 20
(2–5 is most typical).

Tubercled patch wide (4.8–

ADULT 12cm) and roughly rhomboidally Central groove usually
shaped (but varies greatly in poorly developed or
Relatively
size, shape and position) absent (replaced by
broad flukes
flattened or concave area
upon which tubercled
patch sits)

Lateral grooves
9–25 (usually 10–17) usually poorly
loosely longitudinal rows developed or
of prickly tubercles at absent Trailing edge
widest point concave

DIVE Sequence Surfaces briefly and quietly (though for slightly longer than narrow-ridged finless porpoise),
causing little disturbance of water and showing little of body. • Depth Rarely in water deeper than 50m.
• Duration Typically less than 1 minute, maximum 4 minutes.

274 PORPOISES

004 WDP FG.indd 274 29/11/2021 12:30

 Body colour dark grey to ADULT
Dorsal ridge (if present)
nearly black (but substantial No dorsal fin shorter and broader
geographical variation) (3.1–6cm wide) than in
Rounded head with
narrow-ridged finless
slightly bulbous
porpoise
melon (may
Slender body
overhang upper ‘lip’)
Highest point of
dorsal ridge further
back on body (cf.
narrow-ridged finless
porpoise)

May be lighter
area around
No beak throat and chest
in Indian Ocean
individuals
May be slightly Neck more flexible
darker broad band than in other Relatively SIZE
from mouth to porpoises large flippers L: 1.4–1.7m;
flipper WT: 45–50kg; MAX: 1.71m, 60kg
Calf – L: 75–85cm; WT: 5–10kg

FINLESS PORPOISES’ FINLESS BACKS

The dorsal surface of finless porpoises is unique. Instead of a dorsal fin, there is a structure on the back that
comprises three key features.
1. Dorsal ridge – a long, thin, almost fin-like projection running along the centre of the back.
2. Dorsal grooves – consisting of a ‘central groove’ running along the centre of the back (in front of the dorsal
ridge) that may split into separate ‘lateral grooves’ running along either side of the ridge.
3. Tubercled patch – the skin along the centre of the dorsal surface may be covered in an extensive area of
wart-like protuberances called tubercles (which vary greatly from blunt bumps to pointed prickles and sometimes
prickles with a spine-like appearance).
The purpose of these structures is unclear and they are highly variable between species, populations and
individuals (and sometimes absent).

50°

extralimital record from the

Yellow Sea/Bohai Sea region
40°
AT A GLANCE Shallow, warm coastal and
estuarine waters of north-eastern Indian
Ocean and Southeast Asia • Overall
30°

20° coloration usually darker than narrow-ridged

finless porpoise • Small size • No dorsal fin
• Overall impression of back is flattened area
10°

0° in front of short, low dorsal ridge • Rounded

head with no beak • Nearly invisible if water
is rough • Usually alone or in small groups
10°

20°

overlap with narrow-ridged finless porpoise

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 CARING FOR WHALES,
DOLPHINS AND PORPOISES
Human impact has now reached every square kilometre of
the Earth’s oceans. In particular, commercial whaling and
other forms of hunting, entanglement in fishing nets and
myriad other conflicts with fisheries, overfishing, pollution,
habitat degradation and disturbance, underwater noise,
ingestion of marine debris, ship strikes and climate change
are among the main threats faced by whales, dolphins and
porpoises worldwide.
The IUCN Red List gives no fewer than 23 species – a
quarter of all cetaceans – a threatened rating of Critically
Endangered, Endangered or Vulnerable. But that reveals
only part of the picture, because we simply do not know
enough about most species to judge whether or not they
are in trouble (many haven’t even been evaluated). At the
same time, a frightening number have all but disappeared
from many of their former haunts.
To the best of our knowledge one cetacean
has become extinct in modern times: the Yangtze
River dolphin, or baiji, from China (despite IUCN’s
precautionary Critically Endangered (Possibly Extinct)
classification). The vaquita, a tiny porpoise from western
Mexico, could be the next to go; probably fewer than 10
survivors cling on against all the odds. And several other
species may not be far behind.
We nearly lost many of the larger whale species – if
we had not stopped killing them at the 11th hour, we
could have driven grey whales, blue whales, right whales,
bowhead whales and several other species to extinction.
It is estimated that nearly 2.9 million great whales
were killed during 1900–1999 alone (North Atlantic:
276,442; North Pacific: 563,696; and southern hemisphere:
2,053,956). By the time the worst of the slaughter was
over, we were left merely with the tattered remains –
in many cases, barely 5–10 per cent of their original
populations. Some may never recover. There are still only
about 360 North Atlantic right whales, for example, and
recent population trends and ongoing threats are so dire
that the species could become functionally (reproductively)
extinct within 20 years.
Protecting cetaceans is no easy task: they are mobile
and ignore political boundaries, and they face quite
complex, insidious and cumulative threats. These days
the odds are stacked against them and, for some, without
more help, the future is undoubtedly bleak.
HOW TO HELP
Volunteer Most conservation groups could not survive
without the invaluable help of dedicated volunteers. If
you have spare time, there are many ways to lend support
276 CARING FOR WHALES, DOLPHINS AND PORPOISES
005 endmatter WDP FG.indd 276
– perhaps helping your favourite charity answer letters
from enthusiastic children, training as a marine mammal
medic who helps rescue stranded, injured or lost whales,
dolphins or porpoises, or helping to clear a beach of
rubbish? Or, if you are a photographer, for example, donate
pictures to save the cost of buying them commercially.
Raise urgently needed funds Every conservation group
is short of funds. With more money they could do more
good work. You can raise funds to go into the general pot
or for one particular species, place or project that you
feel passionately about. Alternatively, why not consider
adopting a whale or dolphin through various excellent
schemes, or make regular donations by standing order?
Campaign You could support the conservation campaigns
of your favourite charities. They might invite you to write
a letter or sign a petition, or you could take action on any
other issue close to your heart.
Raise awareness The more people who are aware of
conservation issues – and ultimately care about them
– the better. It takes only one individual to affect the
way hundreds of people see the world. You could: write
letters to the national press about environmental stories,
write articles for your local newspaper, offer yourself for
interviews on local radio, or give talks to local clubs and
organisations.
Live a green life There are many relatively easy ways
to reduce your impact on the world’s oceans. For example,
ensure that any fish you eat comes from a sustainable
fishery with minimal bycatch, reduce the amount of plastic
you use, resist buying overpackaged goods and avoid
supporting companies involved in harmful practices.
Get involved with research You could identify
opportunities to help directly with research work. One way
is to help researchers by sending suitable photographs for
their photo-identification catalogues (such as the Antarctic
Humpback Whale Catalogue, which relies on images
taken by tourists on expedition cruise ships). Fieldwork
is expensive, and the larger the catalogues, the more
information they can provide. A brilliant place to submit
ID photographs is happywhale.com. Images are forwarded
to relevant catalogues, and photographers are informed
of any matches made. Check how the species needs to
be photographed – the underside of its tail, for example,
or one particular side of its body – and do not forget to
include your contact details, sighting date and location
(ideally, the exact coordinates), vessel name and any other
relevant information.
Whatever you are able to do, it all helps.
29/11/2021 12:35
 GLOSSARY
abyssal plain Ocean floor beyond the continental shelf.
ambergris Dark greyish waxy substance formed in the
intestines of sperm whales, once widely used in
perfumes.
amphipod Small shrimp-like crustacean – a food source
for some whales.
anchor patch Variable grey or white anchor- or W-shaped
patch on the chest of some smaller toothed whales.
Antarctic Convergence Natural oceanic boundary in the
Antarctic, where cold, less saline waters from the
south sink below warmer, more saline waters from
the north; considered the northern limit of ‘biological
Antarctica’. Otherwise known as the ‘Polar Front’.
balaenopterid Member of the baleen whale family
Balaenopteridae, otherwise known as ‘rorqual’.
baleen plate Dense comb-like structure hanging down
from the upper jaws of most large whales (baleen
whales, Mysticeti); formerly known as ‘whalebone’.
Hundreds of baleen plates are packed tightly together
to form a giant sieve for filter-feeding small prey.
baleen whale Member of the Mysticeti; a predominantly
large whale with baleen plates instead of teeth.
bathypelagic Inhabiting the deepwater portion of the
open ocean, 1,000–4,000m below the surface.
beak Elongated snout of many cetaceans; anterior portion
of the skull that includes both upper and lower jaws.
benthic Living in, on or just above the ocean floor.
blackfish Colloquial term for six superficially similar
members of the dolphin family (killer whale, false
killer whale, pygmy killer whale, melon-headed whale,
short-finned pilot whale and long-finned pilot whale).
blaze Light streaking, usually grey or white, on the side of
a cetacean’s body, usually starting below the dorsal
fin and pointing up into the cape.
blow Refers to the act of breathing – the explosive
exhalation followed immediately by an inhalation –
and to the visible misty cloud of water droplets formed
when a whale breathes; also known as a ‘spout’.
blowhole Respiratory opening, or nostril, on the top of the
head; baleen whales have two, toothed whales one.
blubber Layer of fatty tissue between the skin and
underlying muscle of most marine mammals;
important for insulation.
bow-riding Swimming or ‘riding’ on the pressure wave
created in front of a vessel or large whale.
breaching Leaping completely (or almost so) out of
the water. Officially, if more than 40 per cent of
the whale’s body leaves the water, it is a ‘breach’;
otherwise it is a ‘lunge’.
bubble-netting Feeding technique used by humpback
whales, producing ‘fishing nets’ by blowing bubbles
underwater.
bycatch Animals caught accidentally or incidentally during
fishing operations (when not the target species).
callosity Area of roughened, keratinised tissue on the
head of a right whale, inhabited by whale lice.
005 endmatter WDP FG.indd 277
caudal fin Tail fin.
caudal peduncle See ‘tailstock’.
cephalopod Member of a group of benthic or swimming
molluscs, including squid, cuttlefish and octopuses.
cetacean Any mammalian member of the Cetacea, a
group of aquatic mammals that includes all whales,
dolphins and porpoises.
chevron V- or U-shaped light-coloured marking on the
back or sides of a cetacean.
continental shelf Area of the seafloor around the edge
of a continent; it slopes gently from the coastline
to a drop-off point called the ‘shelf break’ or ‘shelf
edge’; from there the seafloor drops steeply, via the
‘continental slope’, to the ocean bottom.
continental slope Stretch of the ocean floor that drops
steeply between the ‘shelf break’ and the ‘abyssal
plain’.
cookiecutter shark Small shark (up to 50cm) normally
found in sub-tropical and tropical waters that takes
bites out of marine mammals; the resulting wounds
appear as round or oval craters (each about the size
of an ice cream scoop).
copepod Minute shrimp-like crustacean (usually
planktonic) that occurs in great abundance in the sea
and is an important food source for some whales.
crustacean Member of a group of nearly 70,000
invertebrates (animals without backbones), including
lobsters, crabs, shrimps and barnacles. Mostly
aquatic, crustaceans are an important food source for
many marine animals.
delphinid Member of the oceanic dolphin family
Delphinidae.
deep scattering layer (DSL) A dense layer up to
200m thick of huge numbers of small fish, squid,
crustaceans and plankton, in open oceans worldwide,
which migrate up and down the water column within
a single 24-hour period.
demersal Lives near the seafloor.
diatom Microscopic single-celled algae, abundant in
marine and freshwater environments; often form
a film that coats cetacean bodies, producing a
yellowish, brownish or even greenish tinge.
dorsal Pertaining to the upper surface or the back (or the
upper surface of any body part).
dorsal cape Distinct dark region on the backs of some
toothed whales, dolphins and porpoises, mostly in
front of the dorsal fin (sometimes stretching behind it).
dorsal fin Raised structure on the back of most cetaceans;
not supported by bone.
dorsal ridge Ridge on the back as well as, or instead of,
a dorsal fin; may also refer to ridges on the top of the
rostrum in many baleen whales.
driftnet Fishing net that hangs in the water vertically,
virtually unseen and undetectable, and is carried
freely with the ocean currents and winds; a gillnet
that is not anchored. Notorious for catching
GLOSSARY 277
29/11/2021 12:35
 everything in its path, from seabirds and turtles to
whales and dolphins.
drive fishery Technique used to capture dolphins and
other small toothed whales, usually using speedboats
to herd them into bays or shallow water, where they
are killed.
eastern boundary current Found on the eastern side
of oceanic basins, adjacent to the western coasts of
continents, and flowing from high latitudes towards
the tropics; relatively cold, shallow, broad and slow-
flowing. Usually with more nutrient-rich upwellings
than western boundary currents.
echolocation Process of sending out high-frequency
sounds and interpreting the returning echoes to
build up a ‘sound picture’, as in sonar; used by many
cetaceans to orientate, navigate and find food.
ecotype Term that recognises scientific uncertainty
with regard to systematics and speciation in killer
whales (and, to a lesser degree, some other species);
it recognises a work in progress, avoiding the
immediate need to declare subspecies or species.
El Niño Complex global weather pattern marked by rising
sea temperatures in the central and eastern tropical
Pacific Ocean.
epipelagic Living within 200m of the surface in pelagic
waters.
euphausiid Any member of the order Euphausiacea (86
known species of shrimp-like creatures called krill).
extralimital Occurrence outside the normal range.
falcate Sickle-shaped or back-curved; often used to
describe the shape of a dorsal fin with a concave
rear margin.
flipper Variably shaped, flattened, paddle-like forelimb
of a cetacean (also known as the ‘pectoral fin’ or
‘pec fin’).
flipper-slapping Lying on the back or side, raising one
or both flippers out of the water and then slapping
it/them onto the surface. Also known as ‘flipper-
flopping’, ‘flippering’, ‘pectoral-slapping’ or ‘pec-
slapping’.
fluke (noun) Horizontally flattened tail of a cetacean; two
flukes comprise a cetacean’s tail.
fluke (verb), fluking Natural extension of a deep or
sounding dive – the whale bends its body towards
the seabed and, as it rolls forward and down, the
flukes rise above the surface; larger, more rotund
whales fluke regularly; slimmer whales rarely or
never fluke.
flukeprint A circular swirl of smooth water that resembles
a sheen of oil, made by the downward movement of
the tail creating a vortex, and left on the surface after
a whale has dived.
head-slap When a whale lunges partially out of the water
and forcefully slaps its throat onto the surface with a
large splash; also known as a ‘chin-slap’.
home range Area that an animal patrols regularly.
intraspecific Between individuals of the same species
(‘interspecific’ means between individuals of different
species).
278 GLOSSARY
005 endmatter WDP FG.indd 278
isobath Imaginary line on a chart connecting all the points
with the same depth underwater (rather like an
underwater contour line).
IWC (International Whaling Commission); in 1946,
whalers adopted the International Convention for the
Regulation of Whaling for the ‘orderly development
of the whaling industry’; the IWC was established
as the convention’s decision-making body and has
attempted to regulate whaling (and, more recently,
conserve whales) ever since.
krill About 86 species of small shrimp-like crustaceans
(8–60cm long), comprising much of the ocean’s
zooplankton; a major food source for many large
whales. Also known as a ‘euphausiid’.
lamprey Primitive jawless, eel-like fish with a
permanently open mouth bearing many teeth. Of
43 species, 18 are parasitic (boring into the flesh of
cetaceans and other animals and sucking their blood);
mainly in temperate waters.
La Niña Complex global weather pattern marked by
falling sea temperatures in the central and eastern
tropical Pacific Ocean; often follows an ‘El Niño’.
lobtailing Lifting the tail clear of the water then slapping
it down onto the surface, usually repeatedly and
often forcibly; also known as ‘tail-lobbing’ or ‘fluke-
slapping’.
logging Lying at or just below the surface, inactive and
usually horizontally, to rest.
longline Very long (sometimes tens of kilometres) fishing
line, armed with multiple baited hooks (sometimes
thousands) on shorter branch lines, set to catch large
pelagic fish. Responsible for significant bycatch of
dolphins and other species.
lunging Officially, if less than 40 per cent of the whale’s
body leaves the water, it is termed a ‘lunge’,
otherwise it is a ‘breach’; also known as a ‘half-
breach’ or ‘belly-flop’.
mandible Entire lower jaw.
melon Bulging fatty tissue forming the ‘forehead’ of
toothed cetaceans, thought to focus and modulate
sounds for echolocation.
mesopelagic Inhabiting the intermediate depths of the
open ocean, typically at 200–1,000m.
mesoplodont Beaked whale belonging to the genus
Mesoplodon.
mysid Small shrimp-like crustacean in the order Mysida (c.
1,200 species); mostly benthic.
Mysticeti One of two major groups of cetaceans,
containing all the toothless or baleen whales (known
as mysticetes).
oceanic Open sea environment beyond the edge of the
continental shelf.
Odontoceti One of two major groups of cetaceans,
containing all the toothed whales, dolphins and
porpoises (known as odontocetes).
offshore Well away from the coast.
pectoral fin See ‘flipper’.
peduncle See ‘tailstock’.
pelagic Inhabiting offshore waters of the open ocean
29/11/2021 12:35
 beyond the continental shelf; usually used to describe
animals and plants living in upper portions of the
water column; neither near to the coast, nor close to
the bottom.
photo-identification (photo-ID) Technique for studying
cetaceans using photographs as a permanent record
of identifiable individuals.
plankton Passively floating, or weakly swimming, plant
(phytoplankton) and animal (zooplankton) life that
occurs in swarms, usually near the surface of open
waters.
pod Coordinated group of closely related killer whales or
any group of socially affiliated, medium-sized toothed
whales.
polynya Russian word for ‘open water surrounded by ice’.
porpoising When members of the dolphin family (and,
less commonly, some other cetaceans) travel at high
speed and make low, arcing leaps clear of the water
when they take a breath, before re-entering headfirst;
sometimes called ‘running’.
purse-seine fishing Vertical curtain of netting set around
a shoal of fish, then gathered at the bottom and
drawn in to form a ‘purse’ to prevent the fish from
escaping; responsible for killing more dolphins in the
past 50 years than any other human activity (though
new regulations have reduced bycatch substantially).
rake marks Scarring produced by teeth during
intraspecific fighting (usually between males) or from
attacks by killer whales.
remora Type of fish that has modified its dorsal fin into a
sucker (thus the alternative names ‘whalesucker’ or
‘suckerfish’) to attach onto large marine animals such
as whales and dolphins.
rorqual Baleen whale in the family Balaenopteridae,
characterised by a variable number of expandable
pleats or grooves that run longitudinally from the
chin towards the navel; the name comes from the
Norwegian word rørkval, meaning ‘the whale with
pleats’ (after the grooves); otherwise known as a
‘balaenopterid’.
rostrum Beak-like projection at the front of a cetacean’s
head; also used specifically to describe the upper jaw.
saddle patch Light-coloured, more-or-less saddle-shaped
marking that straddles the back behind the dorsal fin
of some cetaceans.
school Term for a coordinated group, normally used in
association with dolphins, that swims and socialises
together; often used synonymously with ‘herd’.
seamount Underwater mountain, typically rising more
than 1,000m above the surrounding deep-sea floor;
usually an extinct volcano, with the summit some
way beneath the surface; attracts an abundance of
marine life.
sexually dimorphic When males and females of the
same species differ in size or appearance.
shelf break Drop-off point at the edge of the continental
shelf (from there, the seafloor drops steeply, via the
‘continental slope’, to the ocean bottom); also called
the ‘shelf edge’.
005 endmatter WDP FG.indd 279
skim-feeding Technique used by some baleen whales
that involves swimming slowly with the mouth open
and constantly filtering food from the water, typically
along or just below the surface.
sounding dive Deep (and usually longer) dive after a series
of shallow dives; also known as the ‘terminal dive’.
splashguard Elevated fleshy ridge immediately in front
of the blowholes of a baleen whale, which helps to
prevent water from pouring in when the blowholes
are open.
spout See ‘blow’.
spyhopping When a whale raises its head vertically out
of the water, usually exposing the eyes to the air,
before sinking smoothly below the surface without
much splash. Also known as a ‘head rise’ or an
‘eye-out’.
stranding The act of coming ashore, intentionally or
accidentally, alive or dead.
submarine canyon Underwater canyon; deep, narrow,
steep-sided valley cut into the seabed.
tail breaching Throwing the rear portion of the body,
including the flukes, high out of the water and
sideways across the surface, creating a huge splash;
other­wise known as a ‘peduncle throw’ or ‘peduncle
slap’.
tail-slapping Like ‘lobtailing’, but normally associated
with smaller cetaceans.
tailstock Muscular region of the tail between the dorsal
fin and the flukes; also called the ‘caudal peduncle’
or ‘peduncle’.
throat grooves V-shaped grooves (deep folds in the skin
and blubber) on the throat, characteristic of beaked
whales and grey whales.
throat pleats Longitudinal parallel furrows or grooves
on the underside of many baleen whales (backward
from the chin) that allow the throat to expand when
engulfing huge quantities of water to capture prey;
also known as ‘ventral pleats’.
toothed whale See ‘Odontoceti’.
tubercle Circular raised protuberance, or bump, found on
some cetaceans (usually along the edges of pectoral
and dorsal fins, but also on a humpback whale’s
head).
upwelling Process by which ocean water rises from
the depths, forced up by currents, winds or density
gradients; this brings nutrients to the surface.
wake-riding Swimming in the frothy wake of a boat or
ship.
water column Anywhere between the surface and the
seafloor.
western boundary current Found on the western side
of oceanic basins, adjacent to the eastern coasts
of continents, and flowing from the tropics to high
latitudes; relatively warm, deep, narrow and fast-
flowing; tends to have fewer nutrient-rich upwellings
than eastern boundary currents, making it less
productive.
whalebone See ‘baleen plate’.
whale louse An amphipod crustacean (not an insect)
adapted to living on the skin of cetaceans.
GLOSSARY 279
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 SPECIES CHECKLIST
MYSTICETI (BALEEN WHALES)
Right and bowhead whales (family Balaenidae)
❑ North Atlantic right whale (Eubalaena glacialis)
❑ North Pacific right whale (Eubalaena japonica)
❑ Southern right whale (Eubalaena australis)
❑ Bowhead whale (Balaena mysticetus)
Pygmy right whale (family Neobalaenidae)
❑ Pygmy right whale (Caperea marginata)
Grey whale (family Eschrichtiidae)
❑ Grey whale (Eschrichtius robustus)
Rorquals (family Balaenopteridae)
❑ Blue whale (Balaenoptera musculus)
❑ Fin whale (Balaenoptera physalus)
❑ Sei whale (Balaenoptera borealis)
❑ Bryde’s whale (Balaenoptera edeni)
❑ Omura’s whale (Balaenoptera omurai)
❑ Rice’s whale (Balaenoptera ricei)
❑ Common minke whale (Balaenoptera acutorostrata)
❑ Antarctic minke whale (Balaenoptera bonaerensis)
❑ Humpback whale (Megaptera novaeangliae)
ODONTOCETI (TOOTHED WHALES)
Sperm whale (family Physeteridae)
❑ Sperm whale (Physeter macrocephalus)
Pygmy and dwarf sperm whales (family Kogiidae)
❑ Pygmy sperm whale (Kogia breviceps)
❑ Dwarf sperm whale (Kogia sima)
Killer whale, small type B with diatoms
False killer whale, underside
280 SPECIES CHECKLIST
005 endmatter WDP FG.indd 280
Narwhal and beluga (family Monodontidae)
❑ Narwhal (Monodon monoceros)
❑ Beluga (Delphinapterus leucas)
Beluga with diatoms
Beaked whales (family Ziphiidae)
❑ Baird’s beaked whale (Berardius bairdii)
❑ Arnoux’s beaked whale (Berardius arnuxii)
❑ Sato’s beaked whale (Berardius minimus)
❑ Cuvier’s beaked whale (Ziphius cavirostris)
❑ Northern bottlenose whale (Hyperoodon ampullatus)
❑ Southern bottlenose whale (Hyperoodon planifrons)
❑ Shepherd’s beaked whale (Tasmacetus shepherdi)
❑ Longman’s beaked whale (Indopacetus pacificus)
❑ Perrin’s beaked whale (Mesoplodon perrini)
❑ Pygmy beaked whale (Mesoplodon peruvianus)
❑ Deraniyagala’s beaked whale (Mesoplodon hotaula)
❑ Ginkgo-toothed beaked whale (Mesoplodon ginkgodens)
❑ Gray’s beaked whale (Mesoplodon grayi)
❑ Hector’s beaked whale (Mesoplodon hectori)
❑ Hubbs’ beaked whale (Mesoplodon carlhubbsi)
❑ Blainville’s beaked whale (Mesoplodon densirostris)
❑ Sowerby’s beaked whale (Mesoplodon bidens)
❑ True’s beaked whale (Mesoplodon mirus)
❑ Ramari’s beaked whale (Mesoplodon eueu)
❑ Stejneger’s beaked whale (Mesoplodon stejnegeri)
❑ Gervais’ beaked whale (Mesoplodon europaeus)
Female Shepherd’s beaked whale
White-beaked dolphin
29/11/2021 12:35
 ❑ Andrews’ beaked whale (Mesoplodon bowdoini)
❑ Strap-toothed beaked whale (Mesoplodon layardii)
❑ Spade-toothed whale (Mesoplodon traversii)
Dusky dolphin – African
Marine dolphins (family Delphinidae)
❑ Killer whale or orca (Orcinus orca)
❑ Short-finned pilot whale (Globicephala macrorhynchus)
❑ Long-finned pilot whale (Globicephala melas)
❑ False killer whale (Pseudorca crassidens)
❑ Pygmy killer whale (Feresa attenuata)
❑ Melon-headed whale (Peponocephala electra)
❑ Risso’s dolphin (Grampus griseus)
❑ Fraser’s dolphin (Lagenodelphis hosei)
❑ Atlantic white-sided dolphin (Lagenorhynchus acutus)
❑ Pacific white-sided dolphin (Lagenorhynchus obliquidens)
❑ Dusky dolphin (Lagenorhynchus obscurus)
❑ Hourglass dolphin (Lagenorhynchus cruciger)
❑ White-beaked dolphin (Lagenorhynchus albirostris)
❑ Peale’s dolphin (Lagenorhynchus australis)
❑ Chilean dolphin (Cephalorhynchus eutropia)
❑ Commerson’s dolphin (Cephalorhynchus commersonii)
❑ Heaviside’s dolphin (Cephalorhynchus heavisidii)
❑ Hector’s dolphin (Cephalorhynchus hectori)
❑ Northern right whale dolphin (Lissodelphis borealis)
❑ Southern right whale dolphin (Lissodelphis peronii)
❑ Australian snubfin dolphin (Orcaella heinsohni)
❑ Irrawaddy dolphin (Orcaella brevirostris)
❑ Rough-toothed dolphin (Steno bredanensis)
❑ Atlantic humpback dolphin (Sousa teuszii)
❑ Indo-Pacific humpback dolphin (Sousa chinensis)
❑ Indian Ocean humpback dolphin (Sousa plumbea)
Female rough-toothed dolphin
005 endmatter WDP FG.indd 281
❑ Australian humpback dolphin (Sousa sahulensis)
❑ Common bottlenose dolphin (Tursiops truncatus)
❑ Indo-Pacific bottlenose dolphin (Tursiops aduncus)
❑ Pantropical spotted dolphin (Stenella attenuata)
❑ Atlantic spotted dolphin (Stenella frontalis)
❑ Spinner dolphin (Stenella longirostris)
❑ Clymene dolphin (Stenella clymene)
❑ Striped dolphin (Stenella coeruleoalba)
❑ Common dolphin (Delphinus delphis)
❑ Tucuxi (Sotalia fluviatilis)
❑ Guiana dolphin (Sotalia guianensis)
South Asian river dolphins (family Platanistidae)
❑ Indus River dolphin (Platanista minor)
❑ Ganges River dolphin (Platanista gangetica)
Yangtze River dolphin (family Lipotidae)
❑ Yangtze River dolphin or baiji (Lipotes vexillifer)
Amazon River dolphin (family Iniidae)
❑ Amazon River dolphin or boto (Inia geoffrensis)
Franciscana (family Pontoporiidae)
❑ Franciscana (Pontoporia blainvillei)
Male Dall’s porpoise, dalli-type
Porpoises (family Phocoenidae)
❑ Dall’s porpoise (Phocoenoides dalli)
❑ Harbour porpoise (Phocoena phocoena)
❑ Vaquita (Phocoena sinus)
❑ Burmeister’s porpoise (Phocoena spinipinnis)
❑ Spectacled porpoise (Phocoena dioptrica)
❑ Narrow-ridged finless porpoise (Neophocaena
asiaeorientalis)
❑ Indo-Pacific finless porpoise (Neophocaena phocaenoides)
Commerson’s dolphin, underside
SPECIES CHECKLIST 281
29/11/2021 14:08
 SOURCES AND RESOURCES
I am very grateful to the innumerable scientists who have
spent countless hours, days, months and years at sea, and
in the laboratory, progressing our knowledge of cetaceans.
I have read many thousands of their scientific papers
during the preparation of this field guide, which represents
the combined work of all these remarkable people. A
comprehensive catalogue of so many references would
more than double the size of the book so, unfortunately, it
is impossible to list them all. But here is a small but varied
selection of good websites, books and journals that might
be of interest.
WEBSITES
American Cetacean Society – acsonline.org
European Cetacean Society – europeancetaceansociety.eu
Happy Whale – happywhale.com
International Whaling Commission (IWC) – iwc.in
IUCN Red List – iucnredlist.org
IUCN – Species Survival Commission (SSC) Cetacean
Specialist Group – iucn-csg.org
Marine Conservation Society – mcsuk.org
NOAA Fisheries – fisheries.noaa.gov/whales
ORCA – orcaweb.org.uk
Porpoise Conservation Society – porpoise.org
Sea Watch Foundation – seawatchfoundation.org.uk
Society for Marine Mammalogy – marinemammalscience.
org (Committee on Taxonomy 2020 List of Marine
Mammal Species and Subspecies)
Whale and Dolphin Conservation – uk.whales.org
BOOKS
Baird, R. W. 2016. The Lives of Hawai’i’s Dolphins and
Whales: Natural History and Conservation. University
of Hawaii Press.
Berta, A., J. L. Sumich and K. M. Kovacs. 2015 (Third
Edition). Marine Mammals: Evolutionary Biology.
Academic Press.
Bortolotti, D. 2009. Wild Blue: A Natural History of the
World’s Largest Animal. Thomas Allen Publishers.
Brakes, P. and M. P. Simmonds. 2011. Whales and
Dolphins: Cognition, Culture, Conservation and
Human Perceptions. Earthscan.
Burns, J. J., J. J. Montague, and C. J. Cowles (eds). 1993.
The Bowhead Whale. Society for Marine Mammalogy.
Carwardine, M. 2016 (Second Edition). Mark Carwardine’s
Guide to Whale Watching in Britain and Europe.
Bloomsbury.
Carwardine, M. 2017. Mark Carwardine’s Guide to Whale
Watching in North America: USA, Canada, Mexico.
Bloomsbury.
282 SOURCES AND RESOURCES
005 endmatter WDP FG.indd 282
Carwardine, M. 2020. Handbook of Whales, Dolphins and
Porpoises. Bloomsbury.
Darling, J. 2009. Humpbacks: Unveiling the Mysteries.
Granville Island Publishing.
Ellis, R. 2011. The Great Sperm Whale: A Natural History
of the Ocean’s Most Magnificent and Mysterious
Creature. University Press of Kansas.
Ellis, R. and J. G. Mead. 2017. Beaked Whales: A
Complete Guide to their Biology and Conservation.
Johns Hopkins University Press.
Fitzhugh, W. W. and M. T. Nweeia (eds). 2017. Narwhal:
Revealing an Arctic Legend. IPI Press & Arctic
Studies Center, National Museum of Natural History,
Smithsonian Institution.
Ford, J. K. B. 2014. Marine Mammals of British Columbia.
Royal BC Museum.
Heide-Jørgensen, M. P. and K. Laidre. 2006. Greenland’s
Winter Whales. Greenland Institute of Natural
Resources.
Hoyt, E. 2011 (Second Edition). Marine Protected Areas for
Whales, Dolphins and Porpoises: A World Handbook
for Cetacean Habitat Conservation and Planning.
Earthscan.
Jefferson, T. A., M. A. Webber and R. J. Pitman. 2015
(Second Edition). Marine Mammals of the World:
A Comprehensive Guide to their Identification.
Academic Press.
Kraus, S. D. and R. M. Rolland (eds). 2007. The Urban
Whale: North Atlantic Right Whales at the
Crossroads. Harvard University Press.
Laist, D. W. 2017. North Atlantic Right Whales: From
Hunted Leviathan to Conservation Icon. John Hopkins
University Press.
McLeish, T. 2013. Narwhals: Arctic Whales in a Melting
World. University of Washington Press.
Reynolds, J. E. III, R. S. Wells and S. D. Eide. 2000. The
Bottlenose Dolphin: Biology and Conservation.
University Press of Florida.
Ridgway, S. H. and R. Harrison (eds). 1985. Handbook of
Marine Mammals, Vol. 3: The Sirenians and Baleen
Whales. Academic Press.
Ridgway, S. H. and R. Harrison (eds). 1989. Handbook of
Marine Mammals, Vol. 4: River Dolphins and the
Larger Toothed Whales. Academic Press.
Ridgway, S. H. and R. Harrison (eds). 1994. Handbook of
Marine Mammals, Vol. 5: The First Book of Dolphins.
Academic Press.
Ridgway, S. H. and R. Harrison (eds). 1999. Handbook
of Marine Mammals, Vol. 6: The Second Book of
Dolphins and the Porpoises. Academic Press.
29/11/2021 12:35
 Ruiz-Garcia, M. and J. M. Shostell (eds). 2010. Biology, JOURNALS
Evolution and Conservation of River Dolphins within Aquatic Mammals – aquaticmammalsjournal.org
South America and Asia. Nova Science Publishers.
Endangered Species Research – int-res.com/journals/esr/
Sumich, J. 2014. E. robustus: Biology and Human History esr-home
of Gray Whales. Whale Cover Marine Education.
Frontiers in Marine Science – frontiers.org/journals/
Swartz, S. L. 2014. Lagoon Time: A Guide to Gray Whales marine-science
and the Natural History of San Ignacio Lagoon. The
International Whaling Commission Journal of Cetacean
Ocean Foundation.
Research and Management – journal.iwc.int
Turvey, S. 2008. Witness to Extinction: how we failed to
Journal of Mammalogy – academic.oup.com/jmammal
save the Yangtze River Dolphin. Oxford University
Marine Mammal Science – marinemammalscience.org/
Press.
journal
Whitehead, H. 2003. Sperm Whales: Social Evolution in
Nature Scientific Reports – nature.com/scientific reports
the Ocean. University of Chicago Press.
Polar Biology – springer.com/journal/300
Wilson, D. E. and R. A. Mittermeier. 2014. Handbook of the
Mammals of the World, Vol. 4. Sea Mammals. Lynx PLOS ONE – journals.plos.one.org/plosone
Edicions. Royal Society Open Science – royalsocietypublishing.org/
Würsig, B., J. G. M. Thewissen and K. M. Kovacs (eds). journal/rsos
2018. Encyclopedia of Marine Mammals. Academic
Press.
Würsig, B. and M. Würsig (eds). 2010. The Dusky Dolphin:
Master Acrobat of Different Shores. Academic Press.

ARTISTS’ BIOGRAPHIES
Martin Camm, from Bedfordshire, UK, is one of the world’s most renowned illustrators of aquatic life, specialising in
cetaceans. He has contributed to hundreds of books, magazines and journals, and his work is widely published by many
organisations, including the United Nations, the BBC, Greenpeace, International Fund for Animal Welfare (IFAW), Whale
and Dolphin Conservation (WDC), and The Wildlife Trusts.
Toni Llobet, from Catalonia, Spain, thinks of himself more as a naturalist than an artist. He is rigorous in both disciplines,
as his intricate work shows. He has worked as a wildlife illustrator for some 20 years, and his work includes massive
projects such as illustrating the prestigious Handbook of the Mammals of the World (published by Lynx Edicions).
Rebecca Robinson, from Tasmania, Australia, graduated with a BSc in Zoology. In a bid to marry her passion for art and
nature, she completed a Bachelor of Fine Arts degree (with Honours) in Wildlife Illustration. She then worked in the design
department at the Marine Science Institute at the University of California Santa Barbara, and has been a freelance natural
science illustrator ever since.

IMAGE CREDITS
With the exception of the artworks listed on the page below, all artworks in this book remain © Martin Camm
(www.markcarwardine.com).

© Rebecca Robinson (www.markcarwardine.com): all dive sequences and whale blows.

© Toni Llobet: p12, p13, p43, p79 (top two), p81, p82, p83, p152, 153 (top), p214, p215, p232 (bottom); flukes on p8, p14,
p15, p34, p39, p42 (right), p47, p53, p59, p63, p88, p93, p155.
© Toni Llobet from: Wilson, D.E. & Mittermeier, R.A. eds. (2014). Handbook of the Mammals of the World. Vol. 4. Sea
Mammals. Lynx Edicions, Barcelona: p8 (second from top), p41 (top), p43 (top), p65, p66 (bottom), p67, p102, p103 (top),
p181 (top), p182 (middle), p215 (top), p219 (top), p249 (top), p251 (top), p259.

ARTISTS’ BIOGRAPHIES AND IMAGE CREDITS 283

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 ACKNOWLEDGEMENTS
Many wonderful people have helped with this book – answering my endless questions, providing as yet unpublished
information and commenting on early tentative drafts – and it couldn’t have been written without them. Their generous
contributions have helped me to make fewer mistakes than I would have made on my own and I’d like to thank them all
for their time, enthusiasm, generosity and support. It is very much appreciated. All these whale biologists – friends and
colleagues – are listed separately.
Special thanks go to the brilliant wildlife artists Martin Camm, Toni Llobet and Rebecca Robinson, who worked tirelessly
on so many illustrations. Martin and I have collaborated on more whale books than either of us can remember, over more
years than we dare admit, and it’s always a great pleasure. And it has been wonderful working with Toni and Rebecca for
the first time. A very big thank you to Rachel Ashton, my outstandingly efficient manager, for her never-ending patience,
perseverance, encouragement, enthusiasm and general brilliance. I couldn’t do it without you, Rachel. Ahha Actadpyoba
very kindly helped with research and, with her outstanding detective work, discovered all sorts of previously unpublished
snippets of information. My literary agent, Doreen Montgomery, was a huge support, as always over the past quarter
of a century; very sadly, she passed away while I was writing, and I will miss her terribly. Doreen’s daughter, Caroline
Montgomery, kindly took the reins and carries on the Montgomery tradition with aplomb. I am particularly indebted to
Jim Martin at Bloomsbury (without whom this book wouldn’t have happened at all) for his unwavering belief in the whole
idea and for his genuine love and passion for the natural world; even when the project took many years longer than we
had all anticipated, and became two books instead of one, he never stopped smiling and (at least seemingly) took it all
in his stride. Thanks for everything, Jim. Thank you, also, to Alice Ward (Commissioning Editor, Bloomsbury Wildlife),
who smoothed the way so calmly during production of the original Handbook of Whales, Dolphins and Porpoises; it was
a pleasure working with you, Alice. The wonderful Jenny Campbell (Senior Editor, Bloomsbury Wildlife) took over for
this updated and abbreviated version – the Field Guide to Whales, Dolphins and Porpoises – and was a fantastically
understanding and encouraging support and help throughout; thank you for everything, Jenny. I was very lucky, too, to have
James Lowen as the magnificently thorough copy-editor and the brilliant Nigel Redman as proof-reader. And we were
very fortunate to have the incredibly talented Julie Dando as designer – an outstanding professional who went above and
beyond the call of duty; thank you – I really appreciate it.
I’d like to thank my dear friends and family, who have enthused with me when things have been going well and cheered
me on when I’ve been flagging: in particular, Peter Bassett and John Ruthven, for those welcome interludes over coffee in
the Clifton Lido; John Craven, Nick Middleton and Marc Riley for putting up with my relentless chatter about field guides,
whales, dolphins, porpoises, distribution maps, illustrations, scientific papers, etc., etc.; and Roz Kidman Cox and Michaela
Strachan, for their knowing sympathy and good counsel. I’d also like to give my wonderful, kind, encouraging parents,
David and Betty, a special mention: without their never-ending support and encouragement, life would have been so
different. My brother Adam, sister-in-law Vanessa, nieces Jessica and Zoe, Beryl, Al and Jude, Florence and Miller have all
kindly put up with my mind-bendingly long hours and preoccupation with what became known as ‘the-project-that-never-
seems-to-end’. Last, but by no means least, a huge heartfelt thank you for everything to my co-conspirator in life, Debra
Taylor, who always makes things better.
At the end of the day, despite a phenomenal effort to make this field guide as accurate and complete as possible, I
take full responsibility for any mistakes, oversights and inconsistencies that may have crept in. I’d welcome any thoughts,
comments and suggestions – via www.markcarwardine.com – to incorporate in new, improved future editions. Thank you
very much.

284 ACKNOWLEDGEMENTS

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 WITH SPECIAL THANKS TO: Kristin Laidre
Robert L. Pitman Jack Lawson
Charles Anderson Rob Lott
Donald McAlpine
… AND ALL THE SCIENTISTS WHO KINDLY
Colin D. MacLeod
ADVISED ON INDIVIDUAL SPECIES
Amanda Madro
Àlex Aguilar
Anisul Islam Mahmud
Wojtek Bachara
Silvia S. Monteiro
Robin W. Baird
Hilary Moors-Murphy
Isabel Beasley
Dirk R. Neumann
Chiara Giulia Bertulli
Stephanie A. Norman
Arne Bjørge
Giuseppe Notarbartolo di Sciara
Nancy Black
Moira Brown Gregory O’Corry-Crowe
Salvatore Cerchio William F. Perrin
William Cioffi Cindy Peter
Phillip J. Clapham Róisín Pinfield
Diane Claridge Andrew J. Read
Rochelle Constantine Victoria Rowntree
Barbara E. Curry Filipa Samarra
Merel Dalebout Jarrod A. Santora
Jim Darling Marcos César de Oliveira Santos
Natalia Dellabianca Richard Sears
David M. Donnelly Keiko Sekiguchi
Simon H. Elwen Tammy L. Silva
Ruth Esteban Tiu Similä
James Fair Ravindra Kumar Sinha
Ivan D. Fetudin Elisabeth Slooten
Andrew Foote Kate Rose-Ann Sprogis
R. Ewan Fordyce Steven Swartz
Ari Friedlaender Jessica K. D. Taylor
Sonja Heinrich Outi Tervo
Denise Herzing Kirsten Thompson
Sascha Hooker Paul Thompson
Erich Hoyt Fernando Trujillo
Miguel Iñíguez Grigory A. Tsidulko
Maria Iversen Samuel Turvey
Thomas A. Jefferson Koen Van Waerebeek
Eve Jourdain Caroline R. Weir
Catherine Kemper Hal Whitehead
Iain Kerr Tonya Wimmer
Jeremy Kiszka Bernd Würsig

ACKNOWLEDGEMENTS 285

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African dusky dolphin 190
Afro-Iberian harbour porpoise 264
Amazon River dolphin 258
Andrews’ beaked whale 148
Antarctic blue whale 54
Antarctic bottle-nosed whale 116
Antarctic minke whale 80
Antillean beaked whale 146
Araguaian River dolphin 258
Arch-beaked whale 134
Arctic right whale 44
Arctic whale 44
Argentinian dusky dolphin 190
Arnoux’s beaked whale 106
Atlantic harbour porpoise 264
Atlantic humpback dolphin 216
Atlantic hump-backed dolphin 216
Atlantic right whale 32
Atlantic spinner dolphin 240
Atlantic spotted dolphin 232
Atlantic white-sided dolphin 186
atoll beaked whale 126
Australian humpback dolphin 222
Australian snubfin dolphin 210
Bahamonde’s beaked whale 152
baiji 256
Baird’s beaked whale 104
Balaena mysticetus 44
Balaenoptera acutorostrata 76
Balaenoptera bonaerensis 80
Balaenoptera borealis 64
Balaenoptera edeni 68
Balaenoptera musculus 54
Balaenoptera omurai 72
Balaenoptera physalus 60
Balaenoptera ricei 74
beluga 102
beluga whale 102
Benguela dolphin 202
Berardius arnuxii 106
Berardius bairdii 104
Berardius minimus 108
Bering Sea beaked whale 144
bhulan 252
Black-and-white dolphin 200
black Baird’s beaked whale 108
black dolphin 198
black finless porpoise 272
blackfish 154, 168, 172, 174, 176
Black Sea bottlenose dolphin 224
Black Sea common dolphin 244
Black Sea harbour porpoise 264
Blainville’s beaked whale 136
286 INDEX
005 endmatter WDP FG.indd 286
INDEX
blind river dolphin 252, 254
blue whale 54
Bolivian bufeo 258
Bolivian river dolphin 258
Bornean dolphin 184
Borneo white dolphin 218
boto 258
bottlehead 114, 116
bottle-nosed dolphin. 224
bottlenose dolphin 224
boutu 258
bowhead whale 44
Brazilian dolphin 248
bridled dolphin 230
Bryde’s whale 68
Burmeister’s porpoise 268
Burrunan bottlenose dolphin 224
caaing whale 172
cachalot 90
California gray whale 50
Caperea marginata 48
Central American spinner dolphin
234
Cephalorhynchus commersonii 200
Cephalorhynchus eutropia 198
Cephalorhynchus heavisidii 202
Cephalorhynchus hectori 204
Changjiang dolphin 256
Chilean blue whale 54
Chilean dolphin 198
Chilean dusky dolphin 190
Chinese humpback dolphin 218
Chinese river dolphin 256
Chinese white dolphin 218
clymene dolphin 240
coalfish whale 64
cochito 266
Commerson’s dolphin 200
common boto 258
common bottlenose dolphin 224
common bottlenose whale 114
common dolphin 244
common minke whale 76
common porpoise 244, 264
common rorqual 60
Costa Rican spinner dolphin 234
costero 250
crisscross dolphin 244
Cuvier’s beaked whale 110
Dall’s porpoise 262
deep-crested beaked whale 148
Delphinapterus leucas 102
Delphinus delphis 244
dense-beaked whale 136
Deraniyagala’s beaked whale 126
desert porpoise 266
devilfish 50
dusky dolphin 190
dwarf Bryde’s whale 72
dwarf fin whale 72
dwarf minke whale 76
dwarf sperm whale 96
dwarf spinner dolphin 234
East Asian finless porpoise 272
Eastern North Pacific long-beaked
common dolphin 244
eastern spinner dolphin 234
Eden’s whale 68
electra dolphin 178
Eschrichtius robustus 50
estuarine dolphin 250
Eubalaena australis 40
Eubalaena glacialis 32
Eubalaena japonica 36
euphrosyne dolphin 242
European beaked whale 146
false killer whale 174
Feresa attenuata 176
fin whale 60
finback 60
finfish 60
finner 60
Fitzroy’s dolphin 190
flathead 114, 116
four-toothed whale 104, 106
franciscana 260
Fraser’s dolphin 184
Ganga River dolphin 254
Ganges River dolphin 254
Gangetic dolphin 254
Gervais’ beaked whale 146
giant bottlenose whale 104, 106
ginkgo-toothed beaked whale 128
ginkgo-toothed whale 128
Globicephala macrorhynchus 168
Globicephala melas 172
goose-beaked whale 110
goosebeak whale 110
grampus 154, 180
Grampus griseus 180
Gray’s beaked whale 130
Gray’s spinner 234
Gray whale 50
grayback 50
Greenland right whale 44
29/11/2021 12:35
 Greenland whale 44
grey whale 50
Guiana dolphin 248, 250
Gulf of California harbour porpoise
266
Gulf of Mexico Bryde’s whale 74
Gulf of Mexico whale 74
Gulf porpoise 266
Gulf Stream beaked whale 146
harbor porpoise 264
harbour porpoise 264
hardhead 50
Haviside’s dolphin 202
Hawaiian blackfish 178
Hawaiian spinner dolphin 234
Heaviside’s dolphin 202
Hector’s beaked whale 132
Hector’s dolphin 204
helmet dolphin 240
herring hog 264
herring whale 60
hourglass dolphin 192
Hubbs’ beaked whale 134
hump-backed whale 84
humpback whale 84
Hyperoodon ampullatus 114
Hyperoodon planifrons 116
Indian humpback dolphin 220
Indian Ocean humpback dolphin
220
Indian river dolphin 254
Indo-Pacific beaked whale 120
Indo-pacific bottlenose dolphin 228
Indo-Pacific common dolphin 244
Indo-pacific finless porpoise 274
Indo-pacific humpback dolphin 218
Indo-Pacific hump-backed dolphin
218
Indopacetus pacificus 120
Indus River dolphin 252
Inia araguaiaensis 258
Inia geoffrensis 258
Irrawaddy dolphin 212
Japanese beaked whale 128
ji 256
jumper 186, 194
karasu 108
Kerguelen Islands Commerson’s
dolphin 200
killer whale 154
Bigg’s 158
fish-eating 157
Icelandic summer-spawning
herring-feeders 160
large type B (pack ice killer
whale) 164
005 endmatter WDP FG.indd 287
North-east Atlantic mackerel-
feeders 161
North-west Atlantic 162
Norwegian spring-spawning
herring-feeders 161
offshore 159
resident 157
small type B (Gerlache killer
whale) 165
Strait of Gibraltar bluefin
tuna-feeders 162
transient 158
type A (Antarctic killer whale)
163
type C (Ross Sea killer whale)
166
type D (sub-Antarctic killer
whale) 167
west coast community 161
Kogia breviceps 94
Kogia sima 96
kuro-tsuchi 108
lag 186, 188, 190, 192, 194
Lagenodelphis hosei 184
Lagenorhynchus acutus 186
Lagenorhynchus albirostris 194
Lagenorhynchus australis 196
Lagenorhynchus cruciger 192
Lagenorhynchus obliquidens 188
Lagenorhynchus obscurus 190
Lahille’s bottlenose dolphin 224
La Plata dolphin 260
La Plata River dolphin 260
Layard’s beaked whale 150
least rorqual 76
lesser beaked whale 124
lesser cachalot 94
lesser finback 76
lesser fin whale 64
lesser rorqual 76
lesser sperm whale 94
Lipotes vexillifer 256
lisso 206
Lissodelphis borealis 206
Lissodelphis peronii 208
little blackfish 178
little finner 76
little piked whale 76
long-beaked common dolphin 244
long-finned pilot whale 172
Longman’s beaked whale 120
longsnout 234
long-snouted spinner dolphin 234
long-toothed beaked whale 150
Mahakam River dolphin 212
marine tucuxi 250
Māui dolphin 204
mealy-mouthed porpoise 208
Megaptera novaeangliae 84
melon-headed whale 178
Mesoplodon bidens 140
Mesoplodon bowdoini 148
Mesoplodon carlhubbsi 134
Mesoplodon densirostris 136
Mesoplodon eueu 143
Mesoplodon europaeus 146
Mesoplodon ginkgodens 128
Mesoplodon grayi 130
Mesoplodon hectori 132
Mesoplodon hotaula 126
Mesoplodon layardii 150
Mesoplodon mirus 142
Mesoplodon perrini 122
Mesoplodon peruvianus 124
Mesoplodon stejnegeri 144
Mesoplodon traversii 152
Monodon monoceros 98
mud-digger, 50
mussel-digger 50
narrow-ridged finless porpoise 272
narwhal 98
narwhale 98
Neophocaena asiaeorientalis 272
Neophocaena phocaenoides 274
New Zealand beaked whale 106,
132
New Zealand dolphin 204
New Zealand dusky dolphin 190
North Atlantic beaked whale 140
North Atlantic fin whale 60
North Atlantic long-finned pilot
whale 172
North Atlantic minke whale 76
North Atlantic right whale 32
North Pacific beaked whale 144
North Pacific bottlenose whale 104
North Pacific fin whale 60
North Pacific minke whale 76
North Pacific right whale 36
North Sea beaked whale 140
northern blue whale 54
northern bottle-nosed whale 114
northern bottlenose whale 114
northern fourtooth whale 104
northern Indian Ocean blue whale
54
northern minke whale 76
northern right whale 32, 36
northern right whale dolphin 206
northern rorqual 64
northern sei whale 64
Omura’s whale 72
INDEX 287
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 orca 154
Orcaella brevirostris 212
Orcaella heinsohni 210
Orcinus orca 154
Orinoco river dolphin 258
Owen’s pygmy whale 96
Pacific gray whale 50
Pacific harbour porpoise 264
Pacific right whale 36
Pacific white-sided dolphin 188
Pantropical spotted dolphin 230
Peale’s dolphin 196
peh ch’i 256
pep 178
Peponocephala electra 178
Perrin’s beaked whale 122
Peruvian beaked whale 124
Peruvian dusky dolphin 190
pesut 212
Phocoena dioptrica 270
Phocoena phocoena 264
Phocoena sinus 266
Phocoena spinipinnis 268
Phocoenoides dalli 262
Physeter macrocephalus 90
piebald dolphin 200
pikehead 76
pink river dolphin 258
Platanista gangetica 254
Platanista minor 252
plough-share dolphin 196
plumbeous dolphin 220
pollack whale 64
Pontoporia blainvillei 260
pothead 168, 172
pseudorca 174
Pseudorca crassidens 174
puffer 264
puffing pig 264
pygmy beaked whale 124
pygmy blue whale 54
pygmy Bryde’s whale 72
pygmy fin whale 60
pygmy killer whale 176
pygmy right whale 48
pygmy sperm whale 94
Ramari’s beaked whale 142, 143
razorback 60
Rice’s whale 74
Risso’s dolphin 180
rollover 234
rough-toothed dolphin 214
sabre-toothed beaked whale 144
saddleback dolphin 244
Sahul dolphin 222
Sarawak dolphin 184
288 INDEX
005 endmatter WDP FG.indd 288
sardine whale 64
Sato’s beaked whale 108
scamperdown beaked whale 130
scrag whale 50
sea canary 102
sea unicorn 98
Sei whale 64
Senegal dolphin 240
sharp-headed finner 76
Shepherd’s beaked whale 118
short-beaked common dolphin 244
short-finned pilot whale 168
short-headed cachalot 94
short-headed sperm whale 94
short-snouted spinner dolphin 240
shus 254
shushuk 254
Sibbald’s rorqual 54
sittang 68
skew-beaked whale 132
skunk dolphin 200
slopehead 214
small-form Bryde’s whale 72
snub-nosed cachalot 96
soos 254
Sotalia fluviatilis 248
Sotalia guianensis 250
Sousa chinensis 218
Sousa plumbea 220
Sousa sahulensis 222
Sousa teuszii 216
South American Commerson’s
dolphin 200
South Asian river dolphin 252, 254
southern beaked whale 106, 130
southern bottlenose whale 116
southern fin whale 60
southern long-finned pilot whale
172
southern minke whale 80
southern right whale 40
southern right whale dolphin 208
southern sei whale 64
South Island Hector’s dolphin 204
Sowerby’s beaked whale 140
spade-toothed beaked whale 152
spade-toothed whale 152
spectacled porpoise 270
sperm whale 90
spinner 234
spinner dolphin 234
splaytooth beaked whale 148
splay-toothed beaked whale 148
spotted dolphin 232
spotter 230, 232
springer 186, 194
squidhound 194
steephead 114, 116
Stejneger’s beaked whale 144
Stenella attenuata 230
Stenella clymene 240
Stenella coeruleoalba 242
Stenella frontalis 232
Stenella longirostris 234
Steno bredanensis 214
strap-toothed beaked whale 150
strap-toothed whale 150
straptooth beaked whale 150
streaker 242
striped dolphin 242
sulphur-bottom 54
sulphur-bottomed whale 54
sunameri 272
suongsu 254
susu 254
Taiwanese humpback dolphin 218
Taiwanese white dolphin 218
Tasmacetus shepherdi 118
Tasman beaked whale 118
Tasman whale 118
tonina 258
toninha 260
tropical beaked whale 136
tropical bottlenose whale 120
tropical whale 68
‘true’ blue whale 54
True’s beaked whale 142
True’s porpoise 262
tucuxi 248
Tursiops aduncus 228
Tursiops truncatus 224
unicorn whale 98
vaquita 266
Western Pacific harbour porpoise
264
white-beaked dolphin 194
white-bellied spinner 234
whitebelly 242
whitebelly spinner 234
whitefag dolphin 256
whitefin dolphin 256
white-side 186
white whale 102
wonderful beaked whale 142
Yangtze finless porpoise 272
Yangtze River dolphin 256
Ziphius cavirostris 110
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