Distribution, abundance and biology of Group V humpback whales ...

Distribution, abundance and biology
of Group V humpback whales
Megaptera novaeangliae: A review
Liz Vang
Conservation Sciences Unit
Forestry and Wildlife Division
Conservation management report
ISSN 1037-4701
August 2002

Cover photograph: Humpback whales observed in Hervey Bay
Marine Park. Photo: Laura Pitt
Acknowledgements: The assistance of Dr Peter Corkeron and
Dr Robert Peterson in providing constructive criticism of the draft
report is gratefully acknowledged.
Distribution, abundance and biology of Group V humpback whales
Megaptera novaeangliae: A review
ISSN 1037-4701
© The State of Queensland, Environmental Protection Agency 2002
Copyright protects this publication. Except for purposes permitted by
the Copyright Act, storage, transmission or reproduction of all or any
part by any means is prohibited without the prior written permission
of the Environmental Protection Agency.
RE438 August 2002
Produced by the Environmental Protection Agency
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Contents
1 Introduction......................................................... 2
2 Conservation status ............................................ 2
2.1 Status in Queensland............................................... 2
2.2 Status in Australia .................................................... 2
2.3 International status................................................... 3
3 Classifi cation and description ........................... 3
3.1 Taxonomic classifi cation........................................... 3
3.2 Species description .................................................. 4
4 Distribution .......................................................... 5
4.1 Global distribution..................................................... 5
4.2 Distribution of southern hemisphere stocks ............. 5
4.3 Distribution of Group V population............................ 5
4.4 Distribution in Queensland ....................................... 5
4.4.1 Moreton Bay Region....................................... 6
4.4.2 Swain Reef Complex and Bell Cay................. 7
4.4.3 Whitsunday Region ........................................ 7
4.4.4 Cairns Region ................................................ 8
4.4.5 Hervey Bay Marine Park................................. 8
5 Migration and site fi delity................................... 9
5.1 Group V migratory route........................................... 9
5.2 Herd structure and behaviour................................... 9
5.3 Site fi delity and habitat preference ......................... 10
6 Individual identifi cation .................................... 11
6.1 Non-manipulative sampling techniques.................. 11
6.1.1 Photo-identifi cation....................................... 11
6.1.2 Sloughed skin sampling ............................... 12
6.1.3 Acoustic tracking .......................................... 12
6.2 Manipulative sampling techniques ......................... 12
6.2.1 “Discovery tags”............................................ 12
6.3 Biopsy sampling ..................................................... 13
6.4 Telemetry................................................................ 13
7 Reproduction ..................................................... 13
7.1 Age and length ....................................................... 13
7.2 Female reproduction and behaviour....................... 13
7.3 Maternal condition and sex ratio ............................ 14
7.4 Male reproduction and behaviour........................... 14
8 Population estimates & rates of increase ....... 15
8.1 Southern hemisphere population estimates........... 15
8.2 Group V population estimates ................................ 15
8.3 Group V rates of increase....................................... 16
9 References......................................................... 18
1 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

1.Introduction
In 1997, the Department of Environment published the
Conservation and management of whales and dolphins
in Queensland 1997–2001 to guide their conservation
and management. The publication included a review of
humpback whale scientifi c literature and a management
plan that sought to ensure the protection of the species.
The aim of this report is to provide information for current
managers and staff involved in the conservation and
management of humpback whales in Queensland. The
report is primarily based on but not restricted to the key
scientifi c literature published since 1997. For conciseness,
reviews have been cited when possible. Papers presented at
the Humpback 2000 Conference, Brisbane, are not included.
Information relating to the distribution, biology and
abundance of Group V humpback whales is summarised
to provide information about the coastal resources required
by the species especially when in Queensland waters. The
distribution of the species is described at international,
national and regional scales. Summaries of the Group
V migration route, site fi delity, herd structure and habitat
preferences are presented. Details of individual identifi cation
techniques using both invasive and non-invasive methods
are provided with information relating to the impact of the
methodologies. To illustrate the recovery of the species, the
population estimates and rates published by various groups
are summarised and presented graphically.
The references cited are stored in the Environmental
Protection Agency’s marine mammal bibliographic database.
The database is a Procite application and currently contains
560 records. Each record includes a copy of the abstract,
contact details for requesting reprints and reprints held by
the Agency. Staff licensed to use the software Procite can
access a copy of the database from the Central Offi ce
L drive path L:/whale database/MAR-MAM bibliography.
The records can be searched using a variety of
management-related keywords.
2 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002
2 Conservation status
2.1 Status in Queensland
The humpback whale Megaptera novaeangliae is protected
under Queensland legislation out to 3 nautical miles offshore
and under Australian legislation within the Australian
Exclusive Economic Zone (offshore to 200 nautical miles).
In Queensland, the humpback is a protected species under
the Nature Conservation Act 1992 and is listed as vulnerable
under the Nature Conservation (Wildlife) Regulation 1994.
Figure 1 shows the locations of Queensland marine parks
declared under the Marine Parks Act 1982.
Eastern Hervey Bay was the fi rst site to be specifi cally
declared for the protection of humpback whales. The
marine park, declared in 1989, has a single “general use
zone” which is also a Designated Whale Management and
Monitoring Area. The objective of this area is to manage
human activities in the vicinity of humpback whales
Megaptera novaeangliae and to monitor the effects of such
activities to ensure the protection of whales (Department of
Environment and Conservation 1989). Hervey Bay Marine
Park and Moreton Bay Marine Park are the only two areas
where the Queensland Government permits commercial
whale watching.
Since 1997, the humpback has been managed in
Queensland waters under the Nature Conservation
(Whales and Dolphins) Conservation Plan 1997 and the
approved management program for the conservation
of whales and dolphins in Queensland 1997–2001. The
humpback whale is identifi ed as a priority species for
management and declares four areas of special interest for
whales (Department of Environment 1997). Each area of
interest has specifi c management approaches that provide
for the protection of humpback whales.
In the Great Barrier Reef Marine Park, the humpback is
managed under the Commonwealth policy Whale and
Dolphin Conservation in the Great Barrier Reef Marine
Park 2000. The policy intends to complement and reinforce
other State, Commonwealth and internal conservation and
management initiatives. The policy details the Whitsunday
and Cairns Regions as areas specifi cally managed to
minimise the impacts of commercial whale watching.
2.2 Status in Australia
Under Commonwealth legislation, Environment Protection
and Biodiversity Conservation Act 1999, the humpback is
listed as a vulnerable migratory and threatened species.
In 1999, the Environment Protection and Biodiversity
Conservation Act 1999 replaced three Commonwealth
legislative instruments protecting the humpback — the
National Parks and Wildlife Act 1975, the Whale Protection
Act 1980 and the Endangered Species Protection Act 1992.
In 1998, the humpback was down-listed from endangered
to vulnerable under the Endangered Species Protection Act
1992. The Action Plan for Australian Cetaceans 1996 listed
the humpback as vulnerable.

Figure 1. Queensland marine parks and other areas of importance for humpbacks
2.3 International status
The humpback is protected throughout the New Zealand
Exclusive Economic Zone under several pieces of legislation
the Marine Mammals Protection Act 1978 (MMPA);
the Marine Mammals Protection Regulations 1992; the
Treaty of Waitangi Act 1978; and the Fisheries Act 1996.
The International Union Conservation of Nature specialist
Group (IUCN) listed the humpback as vulnerable to
extinction, due to the observed population reduction of more
than 20 percent in the last three generations, due principally
to previous commercial whaling activities (IUCN 1996).
Other international organisations responsible for the
conservation of the humpback include The Convention on
the International Trade of Endangered Species (CITES),
the Convention for Migratory Species (Bonn Convention)
and the International Whaling Commission (IWC).
3.Classifi cation and description
3.1 Taxonomic classifi cation
The extant members of the Order Cetacea are divided into
two sub-orders toothed whales (Odontoceti) and baleen
whales (Mysticeti) to which humpback whales belong
(Figure 2). Baleen whales possess plates of hair like
structures used to sieve prey from water taken in whilst
slowly swimming and have a paired blowhole on top of
head. Species are found in all oceans of the world usually
in high latitudes and most species migrate between summer
feeding and winter areas.
Baleen whales are divided into the following four
families, all of which have living members, right whales
(Balaenidae), pygmy right whale (Neobalaenidae), gray
whales (Eschrichtidae), and rorquals (Balaenopteridae).
The rorquals, to which humpback whales belong, have up
to 100 throat grooves or pleats that extend from underneath
the lower jaw to behind the fl ippers in all members of the
family. The humpback whale has between 12 and 36 throat
grooves that expand whilst feeding but are otherwise
rarely seen.
The humpback is classifi ed as part of the Order Cetacea,
sub-order baleen whales (Mysticeti) and Family rorquals
(Balaenopteridae). The humpback long went under the
name Megaptera nodosa Bonnaterre, 1789, but Kellogg
(1932) showed that Borowski’s name has priority
(Rice 1998). The humpback was placed in its own genus
to emphasise its discrete morphology. There are no
taxonomically valid subgroups currently recognised but
there are several separate populations described based
on the species geographical wintering and/or feeding
separations (ANCA 1996).
3 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

Figure 3. Humpback distribution and migration routes (Anderson et al.1995)
4 Distribution
4.1 Global distribution
The humpback has a cosmopolitan distribution throughout
the world’s oceans (Figure 3). However, despite the species’
unlimited migratory potential, they exhibit signifi cant genetic
worldwide partitioning among oceanic populations and
among sub-populations or “stocks” within these oceanic
populations, and among seasonal habitats within stocks
(Baker et al.1994).
During the summer, the humpback can be found on high
latitude feeding grounds. During the winter, some individuals
migrate to low latitude temperate, tropical or sub-tropical
waters to mate and breed. However, some populations of
humpbacks are non-migratory (for example, animals in the
northern Indian Ocean).
4.2 Distribution of southern hemisphere
stocks
Southern hemisphere baleen whale stocks were defi ned in
relation to their Antarctic summer feeding concentrations
(Paterson and Paterson 1984). These feeding
concentrations correspond with areas defi ned for fi sheries
purposes; the areas are described numerically as Area I, II,
III, IV, V and VI (Figure 3). Tynan (1998) provided support
for the historical description of feeding stocks in Antarctica
through a description of the Antarctic Circumpolar Current
and an associated food web that includes humpback whales.
4.3 Distribution of Group V population
The humpbacks that migrate either side of the Australian
continent feed in Area IV and Area V; the populations are
described as Group IV and Group V respectively (Figure
4). Data gathered on commercial whaling vessels supports
the historical division of the Group V from the Group IV
population (Chittleborough 1965; Dawbin 1966; Paterson
and Paterson 1984; Findlay 1998; Mikhalev 2000).
Further support for the division of the Group V population
from the Group IV and VI is provided from a number of
contemporary studies from different disciplines such
as photo-identifi cation studies (Garrigue and Gill 1994;
Garrigue et al.2000); genetic studies (Baker et al.1994);
and acoustic studies (Gill et al.1995).
Due to some interchange between the Group IV and V
populations while on feeding grounds, the geographical
extremities separating the populations on the feeding
grounds cannot be rigidly defi ned but are considered as
being 130ºE–170ºW (Chittleborough 1965; Paterson 1991).
The southern latitudinal boundary of the wintering ground
is more diffi cult to defi ne as females are thought to calf en
route during migration (Chittleborough 1965). However,
the main calving ground for the Group V population is
considered to be the warmer lagoonal waters of the Great
Barrier Reef in an area between Port Douglas (16°S) and
Whitsunday Islands (21°S) (Paterson & Paterson 1984;
Dawbin 1956; Chittleborough 1965; Chaloupka & Osmond
1999; Garrigue et al.2000)
Several authors (Chittleborough 1965; Dawbin 1966;
Chaloupka and Osmond 1999; Garrigue et al.2000) support
the hypothesis that the Group V population separates into an
eastern group (New Zealand and the Pacifi c Islands) and a
western group (east Australian coast). Garrigue et al.(2000)
used a photo-identifi cation study to determine that there
was no interchange between individuals from the Group V
and Group VI populations found on the New Caledonia and
Tongan wintering grounds.
4.4 Distribution in Queensland
Sheltered waters of the Great Barrier Reef region are
described as an important calving area for the east
Australian humpback whale stock (Paterson and Paterson
1984; 1989; Simmons and Marsh 1985; Chaloupka and
Osmond 1999) (Figure 5). However, studies and anecdotal
evidence indicate that parturition occurs at higher latitudes
and at dispersed sites throughout Queensland waters.
Craig and Herman (2000) presented data on the preference
of sites by female humpbacks dependent upon their
reproductive status. The study hypothesised that undersea
topography has some infl uence on habitat choice made by
females on their wintering grounds.
5 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

Figure 4. Group IV and V distribution and migratory routes (Anderson et al.1995)
The southern Great Barrier Reef has shallow (15–60m)
warm waters characteristic of humpback wintering areas
in Hawaii (Herman and Antionoji 1977) and the West
Indies (Whitehead and Moore 1982). Simmons and Marsh
(1985) concluded from their observations that sea surface
temperature is not a determining factor for the distribution of
humpbacks.
Two factors indicate that the humpback calving grounds
are dispersed throughout Queensland waters, and that
parturition occurs along the migratory route. Paterson and
Paterson (1989) reported a number of sightings of females
with new calves migrating north past North Stradbroke
Island; and newly-born calves have been recorded stranded
at sites such as Moreton Island (Paterson et al.1993; Haines
et al.2000) and Fraser Island (Paterson and Van Dyke 1991;
Haines et al.2000). Parturition has been observed at a few
locations within the Great Barrier Reef at Little Trunk Reef,
Charity Reef and Little Broadhurst Reef (Paterson and
Paterson 1989).
Chaloupka and Osmond (1999) concluded that humpbacks
are present in all months of the year throughout the Great
Barrier Reef Marine Park (GBRMP). This study, and
earlier study by Simmons and Marsh (1985) supported
the hypothesis that humpbacks are resident year round in
the northern Great Barrier Reef. However Chaloupka and
Osmond (1999) found that most pods (75 percent) were
sighted in southern GBR waters below 19ºS (Townsville)
and mainly during winter and spring (July to September).
Humpbacks migrating north are sighted in Queensland
waters from late May. Paterson (1984) recorded a distinct
peak in the abundance of humpbacks migrating north
during late June and July at North Stradbroke Island.
However, there is no corresponding distinct peak that occurs
during the southern migration that occurs during August,
September and October (Paterson 1984).
Winter breeding grounds
(presumed)
Summer feeding grounds
6 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002
During August and September, Chaloupka and Osmond
(1999) reported low sightings of mother-calf pairs in most
areas below 21ºS for the period 1982–1999. They suggested
that the main calving grounds for the east Australian Group
V population occur in the extensive southern GBR lagoonal
waters defi ned in the north by the Whitsunday Group of
islands and reefs and in the east by the Pompey/Swain
Reefs complex. In recent years, anecdotal information
suggests that humpbacks are abundant further north
in areas offshore of Cairns and as late in the year as
November.
4.4.1 Moreton Bay region
Historically, a whaling station was based on Moreton Island.
Paterson and Paterson (1984) summarised the history of
exploitation in relation to a discussion of the patterns of
migration. They concluded that the migratory patterns had
not altered as a result of the commercial exploitation.
The Moreton Bay region is best described as a migratory
corridor for humpbacks although other components of their
biology may also occur here. Paterson (1994) observed
parturition on one occasion and, stranded newly-born calves
have been recorded on several occasions (Paterson et
al.1993; Haines et al.2000; 2001).
For the past 20 years, two independent land-based
censuses have been conducted from Point Lookout,
North Stradbroke Island, or Moreton Island, Queensland
(Bryden et al.1996 and Paterson, Paterson and Cato 1994).
In 1998, a brief overview of the population estimates and
rates of population increase were presented at a workshop
(Department of Environment and Heritage 1999).

Figure 5. Distribution of humpbacks in the Great Barrier Reef 1982–1996. (Chaloupka and Osmond 1999).
4.4.2 Swain Reefs complex and Bell Cay
The Queensland Parks and Wildlife Service conducts
routine bi-annual boat patrols to monitor seabirds at the
Swain Reefs complex and Bell Cay. During winter patrols,
pods of two or three humpbacks are regularly recorded
throughout the Swain Reefs complex. However, during
July of the years 1999, 2000 and 2001, staff recorded and
photographed a large pod of humpbacks in the lee of Bell
Cay, close to the Swain Reefs complex. Pods comprised
of up to 20 individuals and appeared to be stationary, not
migrating north or south (pers. comm. B. Knuckey 2001).
The signifi cance of these aggregations remains unclear.
4.4.3 Whitsunday region
The Whitsunday region is the remains of a coastal mountain
range that was submerged at the end of the ice age. A total
of 76 islands and reefs make up the region, which is situated
between mainland Australia, and the Great Barrier Reef at
latitude of 20·5°S–21·5°S. The area surrounding the island
chain is shallow and sheltered and is reported to be an area
preferred by mother and calf pairs. Bait, Hook and Hardy
Reefs are situated further offshore and are separated from
the outer Great Barrier Reef by a deep-water channel of up
to 60 metres in depth.
7 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

The main calving ground of the east Australian humpback
whale stock is defi ned to the north by the Whitsunday
Islands (Chaloupka and Osmond 1999). Simmons and
Marsh (1985) reported that calves tend to be seen north of
the latitude 21°S early in the migration during the months of
June and July.
Since 1993, the Pacifi c Whale Foundation has undertaken a
mark-recapture study using photo-identifi cation techniques.
Their study (Pacifi c Whale Foundation 1997) describes the
distribution of mother-calf pairs as being in the sheltered
waters both around and inshore of the Whitsunday island
group. However, the whales are widely dispersed with no
obvious pockets of concentration. They also conclude that
whales do not appear to be staying in the region for any
great length of time, except for mother-calf pairs.
Malcom and Duggan (1997) undertook aerial survey of
the Whitsunday Islands to determine the distribution and
abundance of humpback whales. They concluded that most
of the whales were observed south of Hook and East Black
Reef, an area were there is likely to be less disturbance of
whales because it is south of the main vessel transit route.
However, it is uncertain if this is an artefact of the small
sample size.
4.4.4 Cairns region
Anecdotal information suggests that the numbers of
humpbacks are increasing in the inshore waters of the
Cairns region.
4.4.5 Hervey Bay Marine Park
Hervey Bay (latitude 25°S) is a large shallow embayment
of approximately 4000 square kilometres, bound to the
west by the mainland and by Fraser Island to the east.
Most of the Bay is less than 18m deep and has a sand or
mud fl oor. Fraser Island, the world’s largest sand island, is
126 kilometres long. The island extends north and south
bridging the continental shelf. Smith (1997) produced a
comprehensive summary of the research undertaken and
management of Hervey Bay Marine Park.
Based on aerial and boat-based studies, humpbacks
have been found to favour the eastern part of Hervey Bay,
especially Platypus Bay, and tend to aggregate in shallow
water close to the western coast of Fraser Island (Corkeron
et al.1994; Pacifi c Whale Foundation 1993; The Oceania
Project 1999). The area offshore from the Wathumba Creek
is identifi ed as being an area that is favoured by humpbacks
(Corkeron et al.1994; The Oceania Project 1999). It is
uncertain what contributes to this aggregation but it could be
the freshwater infl uence of the creek.
Humpbacks have been recorded in Hervey Bay during the
months of July–November (The Oceania Project 1999;
Pacifi c Whales Foundation 2001). Temporal procession
of whales entering the bay is similar to that described for
the whole Group V population (Chittleborough 1965 and
Dawbin 1966). Immature humpbacks enter the Bay fi rst in
late July and mothers and calves in September, but there
is no data indicating that the Bay is of importance to any
particular age-class of the Group V population (Corkeron et
al.1994). However, the use of the Bay by mothers and calves
in September is coincident with school holidays and an
associated increase in recreational boating activity.
8 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002
Bryden and Corkeron (1989) estimated that approximately
30 percent of the Group V population enters Hervey
Bay each year. Corkeron et al.(1994) concluded that the
similarity between the proportion of calves observed in
Hervey Bay and the estimates of the rate of increase for the
Group V population suggests that females with calves do not
preferentially use the Bay. During September and October,
mother/calf pods move through the Bay in waves or pulses
and often stay for prolonged periods.
There are no published sightings of newborn calves in
Hervey Bay. Rather than being described as a winter
ground, Hervey Bay is considered a resting site for
humpbacks migrating south to Antarctica. There is no
information to suggest that commercial whale watching has
or has not altered the distribution of whales in Hervey Bay.
The pre-whaling distribution is unknown for this localised
area.

5 Migration and site fi delity
The migratory potential of humpback whales is almost
unlimited and individuals may migrate up to 10,000 km
each year between summer feeding and winter grounds
(Baker et al.1990). The humpback migration could be driven
by a breeding population exhibiting strong site fi delity to
natal wintering grounds and which then disperses to access
prey that is only adequately abundant in Antarctic waters
(Gaskin 1982).
It has been hypothesized that not all Group V humpbacks
migrate each year either from the breeding or feeding
grounds. Occasional sightings of humpbacks in northern
GBR waters (above 16°S Cairns) in summer supports
previous claims of a sub-stock resident year round in
northern Australian tropical waters (Chaloupka and Osmond
1999; Simmons and Marsh 1985). Biopsy studies (Brown et
al.1995) and whaling data (Chittleborough 1965) indicate
that some females may remain on the feeding grounds
during Austral summers.
The trigger to commence migration has been attributed to
reduced light conditions (Dawbin 1966), prey availability
(Clapham 1996), breeding condition and water temperature.
Corkeron and Connor (1999) discuss several hypotheses
to explain the migration to winter breeding grounds. They
conclude that the northward migration is undertaken by
pregnant females to reduce the risk of killer whale Orcinus
orca predation on newborn calves in low-latitude waters.
In the absence of experimental evidence, it is impossible
to establish which factor or factors is/are responsible for
migration.
The trigger to leave tropical wintering grounds and return
to feeding grounds may be related to reproductive status or
hunger, although Paterson (1994) suggested that breeding
behaviour was a stronger urge than feeding. Dawbin (1997)
stated that the fl uctuation in water temperature was too
small for it to act as a trigger for migration.
From the observations of a single humpback offshore
of North Stradbroke Island, Paterson (1994) concluded
that humpbacks migrated at a similar rate during the day
and night. This is one of the assumptions used to model
population estimates by two separate groups based on
North Stradbroke Island (Bryden et al.1996; Paterson et
al.1994). Other assumptions are detailed in Brown (1997).
5.1 Group V migratory route
The wintering grounds of humpbacks are typically located in
tropical, shallow lagoonal waters such as Hawaii, Bahamas,
West Indies, the South Pacifi c islands (Dawbin 1959;
Garrigue et al.2000) and the Great Barrier Reef (Simmons
and Marsh 1985; Paterson and Paterson 1984; Osmond and
Chaloupka 1999).
Northward Southward
Lactating females accompanied by weaning yearlings.
Immature male and female.
Mature males together with resting females.
Pregnant females.
Figure 6. Summary of Group IV population herd structure (Dawbin 1997)
The east Australian Group V population migrates northward
along the continental shelf hugging the coast until it reaches
southern Queensland and then disperses into the lagoonal
waters of the GBR (Paterson et al.1994). Bryden and Griffi th
(1980) conducted aerial surveys to determine the width of
the migratory corridor off North Stradbroke Island, southern
Queensland. The surveys were conducted to the continental
shelf edge but no humpbacks were sighted. However, the
migratory corridor was determined to be no more than
10km wide.
Due to research being focused at the ends of migratory
routes (Corkeron et al.1994), we can only speculate on
the navigation tools used by migrating humpbacks. The
humpback, like the closely related gray whale, may follow
bottom topography parallel to the coast during migration
and “spy-hop” to orient themselves when they reach deep
trenches (Gaskin 1982).
Other unsurveyed migratory routes are possible. A chain of
sea mountains lie parallel to the east Australian continental
shelf, the closest approximately 200 nautical miles off
south-east Queensland. This is defi ned northward by the
Fraser Sea Mountain and south by the Taupo Sea Mountain.
These sea mountains lead into the Coral Sea platform that
is approximately 250 nautical miles from the Queensland
coast at 17°S.
The second chain of sea mountains is further offshore and
lies approximately 560 nautical miles off the south-east
Queensland coast. This chain is defi ned northward by the
Chesterfi eld Reefs and southward by Lord Howe Island.
Reports of sightings at the Chesterfi eld Reefs indicate that
the Group V population at least occasionally visits the area
(Gill et al.1995).
The migratory stream of Group V is described as spreading
out as individuals move northward from Antarctica, and
perhaps divide into two sub-populations, one travelling up
the East Australian coast and another travelling into the
eastern Coral Sea around New Caledonia (Chittleborough
1965; Dawbin 1966; Chaloupka and Osmond 1999). Craig
and Herman (2000) suggest that undersea topography has
some infl uence on habitat choice for breeding females in
Hawaii. It may be that the Group V population separates
into three or more substocks and each group uses these
topographical features to migrate north and south.
5.2 Herd structure and behaviour
There is a distinct temporal separation of the migrating
herd dependent upon maturity and reproductive status.
Chittleborough (1965) and Dawbin (1997) describe the
age class of migrating individuals from Group IV and V
respectively. The herd structure described is similar for
both the northward and southward journeys, although the
individuals differ because of the change in reproductive
status of females (Dawbin 1997).
Mixed females (including those in early pregnancy)
and immature males and females.
Mature males.
Females in early lactation.
9 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

During the northern migration there is a distinct peak in
abundance during late June and early July observed off
North Stradbroke Island (Paterson 1984) but no such peak
is observed during the southern migration that occurs
in late August off the Whitsunday Islands (Malcom and
Duggan 1997). In Hervey Bay, the Pacifi c Whale Foundation
(1997) reported humpbacks appearing at earlier dates
in successive years. The peak abundance of humpbacks
in Hervey Bay varies from days to weeks (Corkeron et
al.1994). Dawbin (1956) observed a variation in the peak
abundance of migrating humpbacks past New Zealand to be
as much as fi ve-and-a-half weeks.
The structure of pods was identifi ed by a genetic study by
Brown et al.(1995). They noted the northward migrating
pods were signifi cantly smaller than southward migrating
pods and that more male humpback whales were found in
the larger pods than females. The most common pod type
observed during the study was the male-female pair, which
is suggestive of either mating on migration and/or mate
guarding.
There is growing support for the hypothesis that not all
females migrate annually to winter grounds but remain on
the feeding grounds during rest years, or produce offspring
in the higher latitudes of Antarctica (Chittleborough 1958a;
Brown et al.1995; Craig and Herman 1997; Mikhalev 2000).
Chittleborough (1965) and Brown et al.(1995) documented
the ratio of migrating Group V males to females, sampled in
south-east Queensland, and is approximately 2:1. This is a
similar ratio for other humpback stocks in the North Pacifi c
(Medrano et al.1994; Calambokidis 2000; Craig and Herman
2000). However, researchers from the North Atlantic argue
that there is no evidence for a male-biased sex ratio in
wintering humpbacks in the North Atlantic.
The migration southward may be delayed by some
individuals due to their mating drive being stronger than
hunger as individuals and groups of humpback whales have
been observed to continue to seek mates, at least in the
early stage of the southern migration (Paterson 1984).
5.3 Site fi delity and habitat preference
Humpback whales exhibit a high degree of site fi delity
both towards feeding and wintering grounds (Clapham et
al.1993). Urban et al.(2000) determined that the migratory
movements of eastern Pacifi c humpbacks were clearly
non-random. Photographic comparisons between areas
showed clear evidence for preferred migratory destinations.
Kaufmann (1990) describes the movements of an adult
humpback whale sighted in Antarctic Area V that was
resighted 19 months later in Platypus Bay, Queensland,
Australia. The re-sightings were verifi ed using both tail-fl uke
and lateral body markings. The resighting of this animal
is the fi rst photographic documentation of movement
between the described areas, and provides support for the
assumption, based upon discovery tags, that whales
found along the east cost of Australia migrate from
Antarctic Area V.
Genetic studies demonstrate that, despite the nearly
unlimited migratory potential of the species, there is a
striking degree of genetic structure both within and
between oceanic populations of humpback whales
(Baker et al.1994). Several authors have concluded that
site fi delity in humpbacks is a maternally driven trait
(Baker et al.1994; Palsbøll et al.1995; Urban et al.2000).
Photo-identifi cation studies are not consistent in their
fi ndings in relation to sex biases in site fi delity. Garrigue
et al.(2000) and Sladen et al.(1999) concur in their
conclusions of male exhibiting less fi delity than females. In
New Caledonia Garrigue et al.(2000) found that there is an
interchange of males between winter grounds. In Hawaii
males are more likely to wander between wintering grounds
than females (Sladen et al.1999). However, a photoidentifi
cation study by Craig and Herman (1997) suggests
males show greater site fi delity than females in Hawaiian
wintering grounds.
Baker et al.(1994) concluded from their genetic data that
there was no obvious evidence of sex biases in site fi delity
and that humpbacks used a strategy that is a combination of
imprinting and genetic traits to return to the winter grounds.
Craig and Herman (2000) used photographic techniques
to investigate sex differences in site fi delity and migration.
They concluded that the wintering areas utilised by breeding
females is dependent upon their reproductive status.
Gregr and Trites (2000) used positional information and
oceanographic data (bathymetry, temperature, and salinity)
to predict critical habitat off the coast of British Columbia
for fi ve whale species including humpback whales. Using
six predictor variables (month, depth, slope, depth class,
and sea surface temperature and salinity) the model
developed identifi ed critical habitat for sei, fi n, and male
sperm whales. However, due to the small sample size, the
model was relatively insensitive to the predictor variables
for humpback whales. However, the habitat predictions
did identify humpback whale habitat in sheltered bays
and straits throughout the coast. The application of this
model could be considered for identifying humpback whale
habitat in Queensland. The high degree of site fi delity
exhibited by humpbacks, especially females, is an important
consideration for the management and planning of marine
protected areas.
The high degree of site fi delity exhibited by humpbacks
is likely to have implications for breeding success. It has
been suggested site fi delity is maternally driven and that
females with calves favour habitats that provide protection
from predators and rough sea conditions. If this is the case,
there are only limited areas breeding females can utilise.
If females have already expended huge energy budgets
during migration, the energy required to search for other
suitable breeding habitat may be enough to cause a decline
in breeding success over a long period.
10 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

6 Individual identifi cation
The ability to recognise individual animals has substantially
increased our knowledge of the biology and behaviour of
many taxa (Palsbøll et al.1997). A wide-ranging transient
species such as humpbacks can be tracked using a
number of techniques. The techniques can be divided into
non-manipulative and manipulative. Photo-identifi cation,
sloughed skin sampling and acoustic tracking are nonmanipulative
techniques, whereas conventional tagging
and biopsy sampling are manipulative techniques. Each
technique has its cost and benefi t; the advantages and
disadvantages of photo-identifi cation versus genetics were
debated in depth at the Humpback Whale Research and
Conservation Seminar, Brisbane 1998 (Department of
Environment and Heritage Proceedings 1999).
Some of the conclusions from the seminar were that genetic
techniques are technically superior to photo-identifi cation
as a tool. However, its practical constraints limit its exclusive
use. The photo-identifi cation and genetic techniques should
be integrated to target specifi c issues such as: long-term tag
changes; survivorship (especially with calves); discreteness
of stocks; relatedness of individuals; gender determination of
individuals; abundance; determining the sex composition of
a population and standardising genetic markers to facilitate
the comparison of data across jurisdictions.
Genetic tagging is an effective method for individual
identifi cation even in a large population of wide-ranging
and inaccessible mammals such as cetaceans. The
data obtained from genetic tags can be used to address
evolutionary, demographic and behavioural questions to
which traditional tagging methods are unsuited. All living
organisms possess genetic material that in principle enables
individuals within any taxon to be identifi ed reliably from
minute quantities of tissue. Such tissue is commonly derived
from biopsies, but can also come from sloughed skin, shed
hair or faecal material, thus potentially allowing genotyping
and individual recognition even of unobserved animals
(Palsbøll et al.1997).
Contemporary genetic techniques are capable of extracting
DNA from samples that have been preserved for 10 years
unless they have been fi xed in formalin. As large numbers
of samples exist from commercial whaling operations for a
number of species including humpbacks, genetic typing of
these samples would provide an invaluable context in which
to study extant populations (Valsecchi et al.1997).
Stevick et al.(2001) undertook a double-marking experiment
that compared photographic and genetic techniques
for identifying individuals. His study concluded that
photographic techniques are a reliable method of identifying
humpback whales on a large scale. The study is the fi rst
large-scale test of errors in individual identifi cation by natural
markings for any species.
6.1 Non-manipulative sampling techniques
Non-manipulative techniques have the advantage of minimal
disturbance to an individual as samples are collected
remotely. The two non-manipulative techniques provide very
different types of information; photo-identifi cation relies upon
the repeated recognition of physical features to identify an
individual. Sloughed skin sampling is a genetic technique
that examines the genetic make-up of an individual that
does not vary over time.
The disadvantages of non-manipulative techniques
include high costs in time and data collection and analysis,
diffi culties in using appropriate mathematical analytical
techniques, weather dependence and the certainty of which
individual the sample originated from.
6.1.1 Photo-identifi cation
Photo-identifi cation relies on the ability to identify an
individual based upon the level of pigmentation (black and
white markings and patterns) and the shape of the dorsal
fi n. The photo-identifi cation technique is validated and its
success is reliant upon the quality and content of each
photograph (Hammond 1986). A series of photographs
may be required to clearly identify an individual. Several
authors describe protocols and standards for the taking
of photographs and the evaluation of photographs for
identifi cation purposes (Hammond et al.1990; Mizroch
et al.1990; Friday et al. 2000).
Hammond (1986) reviews the method of estimating
population size from the capture and recapture of animals
with respect to population of naturally marked whales.
The paper is divided into three main sections dealing with
(i) using natural markings to “mark” an animal, (ii) the
basic models, and (iii) the effect on population estimates
of variation in the characteristics of individual animals.
Suggestions are made concerning the sampling of naturally
marked whale populations and the analysis of data from
such experiments.
Traditionally, photo-identifi cation techniques have relied
upon an individual humpback revealing the underside of its
the tail or fl uke, long enough for a good quality photograph
to be taken of the unique pattern of pigmentation on the
underside of its fl uke. However, pigmentation is variable with
age and calves have the greatest variation in pigmentation
during their fi rst two years (Clapham 1993). It is unlikely
that a calf can be identifi ed at a young age using photoidentifi
cation techniques and tracked to maturity.
Research programs using photo-identifi cation have provided
substantial information about the life histories of individuals
on the feeding grounds in Maine, Norway and Antarctica,
and the wintering grounds of Hawaii, Bahamas/West Indies.
Research groups that have developed photo-identifi cation
catalogues of individual humpbacks for the Group V
population include Pacifi c Whale Foundation (1987–2000),
The Oceania Project (1992–2001), The University of Sydney
(Corkeron 1992), Bryden et al.(1996) and Garrigue
(1995–2001).
A recent study by Blackmer et al.(2000) demonstrated
that dorsal fi n shape and the confi guration of the peduncle
knobs, found just behind the dorsal fi n, are highly stable
over time and are easy to capture on fi lm each time a whale
is sighted. They suggest that a protocol should be adopted
of taking photographs based upon an existing protocol used
by fi n whale researchers, which involves taking identifying
photographs from the right side of humpbacks.
Australia has yet to develop a common protocol that
encourages that sharing of photo-identifying data between
researchers and management agencies. Queensland Parks
and Wildlife Service, in a pilot project, investigated the
viability of an existing electronic catalogue that is a sightings
database, a storage facility for data and photography used
to identify individuals; and that assisted in the matching
of photographs to identify individual humpback that return
to locations over long periods. The project has not been
developed beyond the pilot stage but identifi ed the need for
the implementing standard methodologies for data collection
for photo-identifi cation and the resources required for
developing an electronic catalogue.
11 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

6.1.2 Sloughed skin sampling
Sloughed skin sampling is a technique used for collecting
genetic material from samples such as skin, hair and faeces
from an individual at the surface (Palsbøll 1997). Sloughed
whale skin contains enough DNA for genetic analysis,
and offers a non-intrusive method for collecting tissue
(Valsecchi et al.1998). A study undertaken by Valsecchi
(1998) demonstrated that sloughed skin sampling is a viable
alternative to genetic sampling and is particularly effective
when applied to active groups. The sloughed skin sampling
technique offers a viable alternative to biopsy darting in
regions where darting is either not permitted or otherwise
undesirable.
Amos and Hoelzel (1992) described a method for preserving
skin tissue for DNA-based analysis. The collection of skin
tissue for genetic analysis is preferable because skin is
easily accessible and is rich in DNA. Skin from stranded
whales can remain usable for at least a week, and probably
longer depending upon ambient conditions (Amos and
Hoelzel 1992).
6.1.3 Acoustic tracking
Male humpbacks sing while migrating to and from their
wintering grounds and while they are present at the
wintering grounds. However, it is uncertain whether or
not song functions to maintain a space between pods or
to attract females (Tyack 1981). All males in a population
produce the same song, which changes over time and with
increasing distance (Cato 1991).
Noad et al.(1998) undertook an acoustic and visual tracking
study in south-east Queensland and concluded that acoustic
tracking may be especially useful for the study or survey
of whale movements beyond visual range of shore-based
observation points. The information gathered is dependent
upon the number of hydrophones used. Three hydrophones
provided positional information on singing humpbacks and
the paths they took. Three hydrophones are required to
make a direct comparison between visual and acoustic
tracking methods. The study has implications for the
interactions between singers and non-singing humpbacks;
interactions between individual singers; migratory behaviour
and travel rates; pod composition; habitat use; song function
and boat-whale interactions. Although acoustic tracking
gathers information from singing individuals, it cannot be
used as a technique to identify individuals for long-term
studies.
For other innovations in monitoring humpbacks out of visible
range of ships, acoustics have been used with other species
and may eventually prove feasible for monitoring humpback
occurrence at remote areas such as outer reef complexes
(Clark et al.1996 and 2002; Clark and Fristup 1997;
Clark and Ellison 2000; Frankel et al.1995).
6.2 Manipulative sampling techniques
A close approach to a whale is required for the attachment
of a conventional tag, the taking of samples for genetic
analysis and the taking of identifying photographs. The
attachment of tags and biopsy sampling are considered
as manipulative research (Queensland Department
of Environment 1997). However, the certainty with
which a sample is taken and the information provided
is considerable, especially when combined with photoidentifi
cation data.
In general biopsy samples have not been taken from calves
due to cultural sensitivities and the perceived impacts of
the technique. In 1992, a study undertaken by Valsecchi
et al.(2002) targeted mother and calves pairs for genetic
analysis;
biopsy samples were taken from the mothers and when
conditions were favourable sloughed skin samples were
taken from the accompanying calf. The study supports the
notion that mothers travel with their offspring for the fi rst
year of the calf’s life.
6.2.1 “Discovery tags”
Chittleborough (1959a) and Dawbin (1959) used “discovery
tags” to provide distributional information on the group IV
and V populations during the commercial whaling period.
Discovery tags were conventional metal tags inserted
sub-dermally into the humpback. The tags were retrieved
when the harvested whale carcass was processed. The
discovery tags were successfully used to reveal that some
intermingling was occurring on the feeding grounds between
the group IV and V populations; and the migratory routes,
either side of the Australian continent, to the wintering
grounds of the group IV and V populations
(Chittleborough 1965).
6.3 Biopsy sampling
Biopsy sampling is a technique that involves the taking
of a core of skin using a dart fi red from a crossbow. The
biopsy dart is a cylindrical punch measuring approximately
10mm in diameter and 30mm long. A metal fl ange at the
base controls the depth of penetration by the dart. The
dart has a collar of fl otation behind the tip, and may or
may not be tethered to assist retrieval (Brown et al.1994).
The combination of genetic and photo-identifi cation data
provides a powerful tool for identifying individual humpbacks
(Stevick et al.2001).
The reactions of several cetacean species to biopsy
sampling have been investigated (Patenaude and White
1995; Gauthier and Sears 1999; Weinrich et al.1992).
Clapham and Mattila 1993b and Brown et al.(1994)
undertook studies on humpbacks and concluded there
was no signifi cant change in behaviour to the biopsy
sampling. However, Weinrich et al.(1992) and other authors
have noted that there is a greater reaction to the human
disturbance caused by the boat and its occupants than to
the sampling technique itself.
Brown et al.(1994) investigated the behavioural responses
of east Australian humpbacks to biopsy sampling and
suggests that female humpback whales maybe particularly
responsive to human disturbances. They concluded that
biopsy sampling has minimal impact on humpbacks.
Hooker et al.(2001) compared the reactions of biopsy
sampling with tag attachment on northern bottlenose
whales Hyperoodon ampullatus. They found the reactions
of various species to biopsy darting is generally mild, the
most common response is a “startle” reaction, although
the level of reaction varies slightly between species, and
also between populations and individuals. In contrast, the
reaction of cetaceans to tagging with suction-cups-attached
tags has been found to vary dramatically.
Biopsy sampling is described as a manipulative research
technique. The technique is neither encouraged nor
excluded by the management approach adopted in
Queensland.
6.4 Telemetry
Telemetry tags methods include the use of satellite-linked
time-depth recorders, satellite-linked position beacons,
time-depth recorders and VHF radio tags. This technique is
a powerful took for investigating aspects of baleen whales’
ranging behaviour, and are likely to become a powerful tool
in the future. Mate et al.(1997, 1999 and 2000) illustrates
the use and application of the tagging techniques on other
baleen species.
12 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

7 Reproduction
Humpback whales are a long-lived, slow breeding species
and are believed to have a life expectancy of about 50 years.
Clapham and Mead (1999) is an excellent review of the
humpback. The following is an extract from their paper.
The average and maximum life expectancies in this species
are unclear, partly because whaling probably removed most
of the oldest animals from the populations in which this
question has been studies. The oldest whale observed by
Chittleborough (1965) out of many thousands examined
in the Australian catches) was one aged 48 years old.
This estimate is dependent upon acceptance of the age
determination technique employed which assumes that
four layers are laid down each year in the laminar ear plug
found in the auditory meatus.
7.1 Age and length
The species exhibits sexual dimorphism with females
usually being larger than males of the same age. Females
from the Group IV and Group V populations appear to
reach sexual maturity at an average age of 5 or 6 years
old (Chittleborough 1955b and Clapham 1992) or a length
of 39·5ft (12 m) and 39·66ft respectively (Dawbin 1960
and Chittleborough 1955b). Males attain sexually maturity
within a simular range to females, 6–7 years, although
they may not be able to successfully engage in intrasexual
competition until later in life (Clapham 1992).
A calf is defi ned as being approximately 63 percent of
the size of its mother while on wintering grounds and is
considered to be independent when it is weaned which is
usually about 11 months old (Clapham et al.1999a). An
immature female whale is one that has not produced a calf
(Chittleborough 1965) whereas a sexually mature female is
one that has produced a calf (Chittleborough 1955b).
Both males and females reach a maximum size at about
20 years of age (Chittleborough 1965). However, there does
not appear to be any real difference between the growth rate
of humpbacks in the northern and southern hemispheres
(Chittleborough 1965).
7.2 Female reproduction and behaviour
Mature females are believed to conceive during the winter
season en route to or from the winter grounds and return
to give birth the following winter after a gestation period of
10–12 months (Chittleborough 1965). Pregnant females
are the last age class to leave Antarctica on migration and
usually the last to return with their new calves
(Dawbin 1966).
A female usually produces one young every two or
three years, although annual calving has been reported
for a number of individual whales by several authors
(Chittleborough 1965; Straley et al.1994; Weinrich et
al.1993; Clapham & Mayo 1987. Since 1992, The Oceania
Project has conducted a research and education program
in Hervey Bay, Queensland. Their program has positively
photo-identifi ed several female humpbacks that have
successfully demonstrated annual calving
(pers. comm. Wally Franklin 2001).
Corkeron and Connor (1999) argue the reasons for
which baleen whales migrate. The paper references the
physiological aspect of thermoregulation in calves and
suggests that pregnant females spend the greatest time
away from Antarctica to enable calves to build up fat layers.
Paterson (1994) describes probable parturition by a
humpback at Moreton Island. A solitary humpback remained
stationary for at least 45 minutes before rising horizontally
(out of the water) as if being infl ated. Approximately onethird
of its body was above the water and its head and
fl ukes were visible. The whale then “subsided” and was soon
accompanied by a small grey-coloured calf that remained
close to the adult’s pectoral region.
7.3 Maternal condition and sex ratio
Feeding is a rare occurrence for humpback whales on
migration (Dall and Dunstan 1957), which is probably due
to the lack of preferred prey. However, there is evidence
that, as in some other taxa, offspring sex ratio is related
to maternal condition (Clapham 1996). In the northern
hemisphere, occasional feeding of humpback whales on
two known wintering grounds has been reported in Samana
Bay and Maui, Hawaii (Salden 1989). Feeding has been
observed on the winter grounds in Queensland (by the
author 1997) and reported off Fraser Island by The Oceania
Project (pers. comm. Wally Franklin 1997).
Age class Size Estimated age
Mean Min. Max.
Calf: 4.35 m (14 ·27ft) 3 3.96m (13ft) 3 4.57 m (15ft) 3 0-3 months 3
Independent/weaned 3 :
Male
Female
Immature (Puberty) 1 :
Male
Female
29.92ft 2
9.70m (31 82ft) 3
9.89m (32 44ft) 3
(36.75 ft) 1
11.73m (38 50ft) 1
Sexually mature
(Upon 1 st pregnancy) 1 :
Male
Female 12.39m (39.66ft) 1
8.00m (26.25ft) 3 10.00m (30.48ft) 3 more than 1
year 2 separation
from mother
(33ft 4in) 1
11m (35ft 3in) 1
(40ft 10in) 1
13.6m (43ft 6in) 1
less than
5 years
more than
5 years
Figure 7. Summary of age class, length and age in years of humpbacks (Chittleborough 1955b1 and 19652 and 19652 and 1965 ; Clapham et al.1999a3 ) 3 3)
13 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

Variations in the abundance of prey on feeding grounds
may be directly related to the year-to-year variation in
reproductive rates (Clapham 1996) and abundance on the
wintering grounds (Chaloupka and Osmond 1999). Wiley
and Clapham (1993) tested the hypothesis of whether or
not maternal condition affects the sex ratio of offspring.
Their investigation concluded that the sex ratio of calves
born to females in “superior” condition, or that had a calving
interval of three years, was biased toward sons.
7.4 Male reproduction and behaviour
Mature male humpback whales depart the feeding grounds
with mature females (Dawbin 1966). Humpback whale
courting behaviour includes elaborate songs sung by males
during their migration to (Charif et al.2001) and on wintering
grounds (Helweg et al.1998b). Helweg and Herman (1994)
found there was no difference between singing behaviour
during day and night in deep water, although Au et al.(2000)
found that in shallow water off Maui there was a change in
the occurrence of singing behaviour with time of day,
with a peak at night.
The song patterns produced by singing humpback whales
depend on where individuals live, with populations inhabiting
different ocean basins normally singing quite distinct songs
(Noad et al.2000). Singing whales have been used to
determine migratory routes (Charif et al.2001), demonstrate
interchange between populations (Helweg et al.1998)
and as an index of abundance (Noad 1998). Mobley et
al.(1988) played a variety of sounds to humpback whales
on wintering grounds and concluded that the different rates
of behavioural response were attributed to the behaviour
of sexually active males seeking to affi liate with sexually
mature females.
A study conducted by Frankel et al.(1995) supported
the hypothesis that song functions to maintain distance
between singers and McCauley et al.(1995) give a succinct
description of the song and hearing capabilities
of humpbacks.
Richardson et al.(1995) is a comprehensive review of
acoustics and how they relate to marine mammals. The
information presented includes specifi c references to
humpback whales. They concluded there is no research
to demonstrate that acoustics adversely affects humpback
whales. However, more recent research demonstrates
that humpbacks alter their behaviour in response to
anthropogenic noise (Miller et al.2000). Williams et al.(2002)
concluded that weakening or not enforcing the minimum
approach distances for the whale watching of orcas in
Canada would result in higher levels of disturbance.
Corkeron (1995) demonstrated that humpbacks in Hervey
Bay altered their behaviour when vessels were within 300m.
This and several other studies show that the behaviour
of humpbacks is altered by the presence of vessels but it
remains uncertain if the effect is detrimental (Bauer and
Herman 1986; Baker and Herman 1989; Bryden et al.1988).
8 Population estimates and rates
of increase
Reviews of international data indicate a strong recovery
in most studied humpback whale populations (Best 1993
and Clapham 1999b). Paterson et al.(1994) suggests that
the level of recovery of the Group V population, in the
post-whaling period, is due to the stock experiencing no
detrimental environmental factors.
8.1 Southern hemisphere population
estimates
During 1933–1939, the estimated total number of
southern hemisphere baleen numbered 220,000–340,000.
Approximately 10 percent of the southern hemisphere
stock of baleen whales was humpback whales, and the
ratio between the groups I–V was considered to be
1:1:2:3:3 respectively (Matthews 1957 cited Chittleborough
1965). Based on these estimates the Group V population
could be estimated at 22,000–34,000 individuals for
1933–1939 (Figure 8).
8.2 Group V population estimates
Figure 8 is a summary of the population estimates and
illegal catches by the Soviet fl eet for Group V humpbacks
during 1930–2005. Chittleborough (1965) estimates the
Group V pre-whaling population numbered approximately
10,000 individuals. By 1960, he estimated a population size
of only 500 individuals. In 1963, the International Whaling
Commission (IWC) imposed a ban on whaling, and in
1964 the Group V population was estimated at just 400
individuals. Until 1963, humpback population estimates
were derived by modelling the fi sheries-based catch per
unit effort data (Chittleborough 1965).
Chittleborough (1965) compared population estimates
derived by two methods and argues that for the estimate
to be correct using the DeLury method, a further 5000
humpbacks would have had to been fi shed during the years
1960 and 1961. Only 302 and 270 whales were reported
to the IWC for each of those years. Although there is some
uncertainty in these estimates, it is clear that during the late
1950s and early 1960s, the Group V population experienced
a signifi cant decline in numbers.
Recent data presented by Mikhalev (2000) supports
DeLury’s population estimate discussed by Chittleborough
(1965). Mikhalev (2000) presented data collected by two
Soviet vessels illegally whaling during the summer season
of 1959 in the Antarctic Area V. The Russian vessels
signifi cantly exceeded the quota for all countries by illegally
fi shing a total of 11,605 individuals from the Group V
population.
Chittleborough (1965) estimated that the Group V population
would take 36–63 years to regain its unfi shed status of
10,000 individuals. However in 1998, 33 years later, the
Group V population was estimated at 4000 individuals
(Paterson cited QDEH 1999). If a constant rate of increase
is assumed, the 1962 population estimate of 500 individuals
would have to have increased at a rate of approximately
8 percent a year to reach its unfi shed state in 36 years.
The rate of increase for the population to reach its unfi shed
state in 63 years is just 4·8 percent a year. However,
Mikhalev (2000) details the large-scale illegal Soviet catch
data for the late 1960s and illustrates that the catches of
humpback on their feeding grounds during 1963–1968
exceeds the population estimates derived by Chittleborough
(1965). The population estimates for this period are being
revised to account for the illegal catches by the Soviet fl eet
(pers. comm. P Corkeron 2002).
14 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

8.3 Group V rates of increase
In Queensland, three independent groups, using data
collected by two different methods, have modelled the rate
of increase of the population. Paterson et al.(1994) and
Bryden et al.(1997) conducted land-based surveys from
Point Lookout, North Stradbroke Island, since 1978 and
1980 respectively. Paterson (1984), Paterson and Paterson
(1989) and Paterson et al.(1994) estimate the population
is increasing at 11·7 percent a year. Biennial surveys
(1994–1996–1998) undertaken by the Patersons since
1992 confi rm an annual rate of increase of 11·6 percent
(95 CI: ±1 percent). Both sets of shore-based data assume
that whales travel at the same speed throughout the day
and night (Paterson 1994).
Chaloupka and Osmond (1999) modelled data collected
as a secondary consideration by an enforcement agency
undertaking an aerial surveillance program in the Great
Barrier Reef Marine Park. They estimated a much lower
rate of population increase of 3·9 percent a year.
? 34,000 in 1930:
Matthews 1942
10,396 in tropical
waters 1947-61:
Dawbin 1997
10,000:
Chittleborough 1965
Modern studies data:
Bryden et al 1990, 1997; Patterson et al 1989, 1994;
Patterson 1998 (cited QPWS 1998)
4600:
DeLury 1947
500:
Chittleborough 1965
Figure 9. Extrapolated rate of increase for Group V humpback whales 1980–2000
Since the ban on commercial hunting was implemented,
the combined impact of international, national and state
conservation legislation has contributed to an approximately
40 percent recovery in the Group V humpback whale
population. In addition there are no signs that the population
is approaching asymptotic level in response to prey
resources, habitat availability or other population limiting
factors. There are no indications that the introduction of
commercial whale watching in Queensland in 1989, as
managed by QPWS (EPA 2001), has caused any change
in the rate of recovery of this species (Figure 9).
Global ban on commercial
humpback whaling in 1963
11,605 whales fi shed
in Antarctic waters:
Mikhalev 2000
10,000:
Chittleborough 1965
800:
Chittleborough 1965
Modern studies data
Figure 8. Summary of estimated population size, numbers fi shed and associated references for the Group V population 1930–2000
15 • Distribution, abundance and biology of Group V humpback whales Megaptera novaeangliae: A review • August 2002

Appendix 1. Summary of population estimates 1930–2002
Year Pop Estimate (min/max) Reference and notes
1935 22,000-34,000 Matthews 1942 cited Chittleborough 1965
1936
1937
1938
1939 22,000-34,000 Matthews 1942 cited Chittleborough 1965
No data available for this period
1951
1952
1953
1954
1955
1956
1957
1958
10,000 Chittleborough 1965: population remained stable until 1959.
1959 10,000 Chittleborough 1965
1960
1961
DeLury method gives a population estimate of 4600
1962
1963
500 Chittleborough 1965
1964
1965
800
Chittleborough 1965: this is assuming that the mature
stock numbers 400 and equivalent to 52% of the total.
No data available for this period
1977
1978
1979
1980
1981 356 Bryden et al.1990
1982 396 Bryden et al.1990
1983
1984
1985
1986 778 Bryden et al.1990
1987 790 Bryden et al.1990
1988
1989 1100 Paterson and Paterson 1989
1990
1991 1788 (±138) Bryden et al.1997
1992 1900 (±250) Paterson et al.1994
1993 2099 (±152) Bryden et al.1997
1994
1995
1996 3185 (±208) Bryden et al.1996
1997
1998
1999
2000
4000
3500-4000
Paterson 1998 (Cited QDEH 1999)
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• August 2002