REVIEW
published: 14 May 2019
doi: 10.3389/fmars.2019.00247
Common and Antarctic Minke
Whales: Conservation Status and
Future Research Directions
Denise Risch 1* , Thomas Norris 2 , Matthew Curnock 3 and Ari Friedlaender 4
1
Scottish Association for Marine Science, Oban, United Kingdom, 2 Bio-Waves, Inc., Encinitas, CA, United States,
Commonwealth Scientific and Industrial Research Organisation Land and Water, Townsville, QLD, Australia, 4 Institute for
Marine Sciences, University of California, Santa Cruz, Santa Cruz, CA, United States
3
Edited by:
Rob Harcourt,
Macquarie University, Australia
Reviewed by:
Luis Cardona,
University of Barcelona, Spain
Salvatore Cerchio,
New England Aquarium,
United States
*Correspondence:
Denise Risch
denise.risch@sams.ac.uk
Specialty section:
This article was submitted to
Marine Megafauna,
a section of the journal
Frontiers in Marine Science
Received: 07 December 2018
Accepted: 24 April 2019
Published: 14 May 2019
Citation:
Risch D, Norris T, Curnock M and
Friedlaender A (2019) Common
and Antarctic Minke Whales:
Conservation Status and Future
Research Directions.
Front. Mar. Sci. 6:247.
doi: 10.3389/fmars.2019.00247
Minke whales comprise some of the most widely distributed species of baleen
whales, some populations of which are still regularly targeted by commercial whaling.
Here, we review the conservation status of common (Balaenoptera acutorostrata) and
Antarctic (Balaenoptera bonaerensis) minke whale populations, against the backdrop of
ongoing whaling operations and other anthropogenic threats, including climate change,
entanglement in fishing gear, ship strikes, and noise pollution. Although some coastal
minke whale populations have been studied in detail, others, which inhabit remote
and ecologically sensitive locations, such as the Antarctic ice shelf, are among the
least understood populations of marine mammals. The unresolved taxonomy of dwarf
minke whales further highlights some of the existing knowledge gaps concerning these
species. Due to their relatively small size and elusive behaviors, large uncertainties
exist for almost all minke whale populations with respect to behavior, migratory
routes and winter distributions, hindering effective conservation and management.
However, recent advances in research technology, such as passive acoustic monitoring
(PAM), unmanned aerial systems (UAS), multisensor recording tags, and machine
learning assisted photo-identification, are increasingly being applied to study minke
whales and their habitat, and are starting to open new windows into their life history
and ecology. In future research, these non- and less-invasive methods should be
integrated in larger-scale comparative studies aiming to better understand minke whale
behavior, ecological interactions and their varying habitats to drive and support effective
species conservation.
Keywords: population status, minke whales, Antarctic minke whale, conservation, future research
INTRODUCTION
Many baleen whale populations have been reduced extensively by whaling, which makes them more
vulnerable to negative impacts from changes in their habitats (Baker and Clapham, 2004). In the
case of minke whales, several populations are still being impacted by ongoing commercial whaling.
These hunts are carried out against the backdrop of global climate change and other anthropogenic
impacts, such as interactions with fisheries, pollution, as well as ship strike. Given this complex
array of often interacting threats and considering their comparatively long life spans, minke whales,
like other baleen whales, face significant challenges that may lead to severe impacts on populations.
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Risch et al., 2014a). The Gulf of St. Lawrence is a known
summer feeding habitat (Lynas and Sylvestre, 1988; DoniolValcroze et al., 2007). In the central North Atlantic, pronounced
sexual segregation exists on higher latitude feeding grounds, with
females occurring further north, off western Greenland, while
males remain further south and to the east of Greenland (Laidre
et al., 2009). Tracks of satellite-tagged minke whales indicate
a southward movement, following the mid-Atlantic ridge, as
far south as at least 28◦ N starting in autumn (September to
November) (Víkingsson and Heide-Jørgensen, 2015). During
summer and early fall (June to November), minke whales
in the eastern North Atlantic occur around the British Isles
(Macleod et al., 2004; Tetley et al., 2008; Risch et al., 2019)
and Norway (Heide-Jørgensen et al., 2001), while their winter
distribution remains largely unknown. However, minke whales
have been observed year-round around the Canary Islands (van
Waerebeek et al., 1999), but only occasionally off Portugal,
Spain and West Africa (Aguilar et al., 1983; van Waerebeek
et al., 1999). The species is a relatively rare visitor in the
Mediterranean basin with one sighting reported from the Black
Sea (Reeves and Notarbartolo di Sciara, 2006).
In the North Pacific, minke whales are widely distributed
north of 30◦ N, shifting their distribution northward during
summer (Zerbini et al., 2006). However, they also occur
year-round off central California (Dorsey et al., 1990). In
the Central and Eastern Pacific, acoustic data suggest minke
whale presence from approximately 15–35◦ N in deep waters
surrounding subtropical volcanic islands and atolls during
winter and spring (Rankin and Barlow, 2005; Rankin et al.,
2007; Oswald et al., 2011; Norris et al., 2017). Data from
Korea and Japan indicate seasonal southbound migrations in
the western North Pacific, with unknown final destinations
(Ohsumi, 1983).
Minke whales inhabit all oceans, including tropical as well as
polar waters (Figure 1). Since the late 1990s, two different species
have been recognized: the common minke whale (Balaenoptera
acutorostrata) and the Antarctic minke whale (Balaenoptera
bonaerensis). Molecular data suggests that the two species are
sister taxa and their clade is sister to the other Balaenopteridae
and the gray whale (Eschrichtius robustus) (Rychel et al., 2004) but
there is still some ambiguity about the phylogenetic relationships
of minke whales to other species (Perrin et al., 2018). For
the common minke whale three putative subspecies have been
proposed; Balaenoptera acutorostrata acutorostrata in the North
Atlantic, Balaenoptera acutorostrata scammoni in the North
Pacific and the dwarf minke whale, unnamed subspecies, in the
Southern hemisphere (Rice, 1998). Dwarf minke whales were first
recognized as a distinct form of the common minke whale in
the mid-1980s (Best, 1985), and genetic evidence suggests the
existence of at least two different populations in the Southern
Hemisphere (Pastene et al., 2010). The exact placement of the
dwarf minke whales in the taxonomy of this species complex is
still unclear. For example, South Atlantic dwarf minke whales
are more closely related to North Atlantic common minke
whales, than either are to dwarf minke whales from the South
Pacific, indicating that their subspecies status needs further study
(Pastene et al., 2010).
The seasonal distribution and migration patterns of nearly
all populations of minke whales are poorly understood. Their
small size, inconspicuous behavior and seasonal distribution
in remote pelagic and sea ice habitats contribute to this lack
of understanding. Existing knowledge is often obtained from
studying populations that frequent coastal habitats or based on
incomplete historical whaling records. At least some proportion
of the population of minke whales in the western North Atlantic
appear to migrate to the Caribbean during winter (Mitchell, 1991;
FIGURE 1 | Range map of Common (Balaenoptera acutorostrata) and Antarctic (Balaenoptera bonaerensis) minke whales. Source for presented species ranges:
International Union for Conservation of Nature (IUCN) (Cooke et al., 2018a,b).
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understand how large-scale changes in climate and sea ice cover
will affect these ecological relationships, because the linkages
between these predators and their prey remain poorly studied
(Fraser and Hofmann, 2003).
Antarctic minke whales are the largest ice-dependent krill
predators in the Southern Ocean. As their distribution and
ecology are directly tied to sea ice (Herr et al., 2019) and foraging
on krill, any changes that affect the quantity and quality of
their habitat and food availability could be significant. Currently,
Antarctic minke whales and humpback whales are thought to
partition prey resources in this region by feeding in different
habitats (sea ice versus open water) but when the two species
do overlap in time and space, they partition prey vertically
(Friedlaender et al., 2008). With continued sea ice decline, the
potential for overlap, and thus competition for prey, increases as
does the likelihood that krill abundance will decrease due to poor
recruitment, for example (Atkinson et al., 2004). This could lead
to both a limited amount of preferred sea ice habitat and limited
prey availability, increasing the potential for competition with
other krill predators (including humpback whales (Megaptera
novaeangliae), crabeater seals (Lobodon carcinophaga), Antarctic
fur seals (Arctocephalus gazella), as well as a variety of penguin,
seabird, and fish species). In combination, this could mean that
Antarctic minke whales will be displaced to regions that are
currently covered by more sea ice and those that remain may be
at higher predation risk from killer whales in open water (Higdon
and Ferguson, 2009) and be in competition with other predators
for a limited prey resource.
The distribution of dwarf minke whales spans the entire
southern hemisphere (Figure 1). They have been visually
recorded in coastal waters of South America and South Africa
in the Atlantic (Zerbini et al., 1996; Acevedo et al., 2006) and
in waters off both western and eastern Australia, New Zealand,
and New Caledonia in the Pacific (Dawson and Slooten, 1990;
Arnold, 1997; Garrigue and Greaves, 2001; Gedamke et al.,
2001). Although they are less commonly observed south of
65◦ S than Antarctic minke whales (Branch and Butterworth,
2001), dwarf minke whales are sympatric with Antarctic
minke whales in much of their range (Acevedo et al., 2011),
which renders some of the visual based information on the
distribution of both species ambiguous or inaccurate, due
to missing or incorrect identification to species level at sea
(Cooke et al., 2018a).
Antarctic minke whales have a circumpolar distribution south
of 60◦ S during summer (Figure 1), with high densities reported
in parts of the Weddell and Ross Seas (Kasamatsu et al., 1995).
The species is highly associated with sea ice and is generally less
abundant in ice-free waters (Herr et al., 2019). Recent acoustic
data show detections of the species in association with sea ice
for most of the year (Dominello and Široviæ, 2016). However,
this species has also been observed and recorded in sub-tropical
waters of the South Atlantic, South Pacific, and Indian Ocean
during winter (McCauley, 2004; Cerchio et al., 2018). A bi-modal
temporal distribution pattern of Antarctic minke whales has been
visually observed off South Africa, peaking in fall and spring,
suggesting migratory movements by parts of the population to
lower latitude habitats during winter (Best, 1982). There is also
some evidence that Antarctic minke whales occasionally occur
north of the equator (Glover et al., 2010; De Boer, 2015).
With this review, we aim to summarize current knowledge
on the threats global minke whale populations are facing, review
current research and monitoring approaches and discuss data
gaps and future research and conservation priorities.
Whaling
Due to their small size and relatively fast movements, minke
whales only became a target of industrial whaling operations
after the demise of the larger whale species, in the early
1970s (Horwood, 1990). Commercial whaling is carried out
by Norway and Iceland (under objection and reservation to
the 1986 moratorium agreed by the IWC, respectively), and
subsistence hunts carried out by Greenland. Until recently,
when Japan decided to leave the IWC (Normile, 2019), whaling
for minke whales also continued as part of Japan’s special
permit (“scientific”) whaling programme. Overall, since 1985,
43,799 minke whales have reportedly been killed in commercial
(25,583), special permit (15,176) and aboriginal (5,094) whaling
programmes, respectively1 . The scientific whaling by Japan
occurred under Article VIII of the International Convention
for the Regulation of Whaling (ICRW) and had long been
criticized by other member states and scientific expert panels
(Gales et al., 2005; Brierley and Clapham, 2016). The criticism,
with which the authors of this review are in agreement, called
into question the validity of the science being conducted, as
well as made the accusation of the use of minke whaling
and the sale of its products as a cover up for illegal trade
with products from other protected species (Baker et al., 2000,
2010). In 2014 these long-standing concerns led Australia and
New Zealand to bring the issue of whether Japan’s whaling is
legitimately for scientific purposes before the International Court
THREATS
Climate Change
Antarctic minke whales provide a good example of a baleen whale
species that are at risk from multiple environmental pressures
driven by climate change. The marine ecosystem around the
Antarctic Peninsula is experiencing dramatic warming (Morris
and Vaughan, 2003) and significant reductions in the extent
and duration of seasonal sea ice cover (Stammerjohn et al.,
2008, 2011, 2012). These changes affect every component of
the ecosystem (Schofield et al., 2010; Ducklow et al., 2012),
but particularly the demography, behaviors and ecology of
predators that rely on sea ice for foraging habitat and prey,
such as Antarctic krill (Euphausia superba). The life history and
population dynamics of krill are heavily influenced by climate
forcing (Nicol, 2006), so predators of krill must respond to interannual variability in prey availability and alter their behavior
and demography in response to such changes (e.g., penguins
and seals; Fraser et al., 1992; Reid and Croxall, 2001; Fraser
and Hofmann, 2003; Reid et al., 2005). Currently we do not
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of Justice (ICJ), which declared in majority that Japan’s whaling
programme at the time (JARPA II) was not “for the purposes of
scientific research.” Despite this ruling and continued criticism
of lack of scientific justification (Brierley and Clapham, 2016;
Clapham, 2017), Japan’s current whaling programmes in the
Antarctic (NEWREP-A) and North Pacific (NEWREP-NP) were
implemented in seasons 2015/16 and 2017/18, respectively, with
quotas to kill 333 Antarctic minke whales and 174 common
minke whales in the North Pacific2 . With its decision to leave
the IWC, it is anticipated that Japan’s current “scientific”
whaling programmes will stop and be replaced by a commercial
whaling programme in their Exclusive Economic Zone (EEZ)
(Normile, 2019).
therefore modeled or inferred from species-specific vocalizations,
behavioral responses to sounds or anatomical measurements
(Mellinger et al., 2000; Tubelli et al., 2012; Yamato et al., 2012;
McGarry et al., 2017). Estimates of audiograms for baleen whales,
including minke whales, assume highest sensitivities between
200 and 19,000 Hz (National Marine Fisheries Service, 2016).
Simulations of the middle-ear transfer function (METF) of the
minke whale auditory pathway suggested the best frequency
range between approximately 30 and 25,000 Hz (Tubelli et al.,
2012). A recent study in the Northwest Atlantic, investigated the
potential of vessel noise to mask baleen whale vocalizations and
found an ∼80% loss of communication space for minke whale
pulse trains relative to historical “quiet” conditions (Cholewiak
et al., 2018). Off Scotland, minke whale densities have been
observed to decrease during naval exercises, and mortalities of
minke whales occurred during at least two mass stranding events
that have been linked to military sonar exercises (Parsons et al.,
2000; Balcomb and Claridge, 2001). Minke whales have been
observed to respond to mid-frequency active sonar by reducing
or ceasing calling and by exhibiting avoidance behaviors (e.g.,
movement away from the sound source) even at relatively low
received sound levels (Martin et al., 2015; Sivle et al., 2015;
Kvadsheim et al., 2017). Also, results of a recent behavioral
response study indicated that minke whales strongly avoided
ADDs that are used in the context of noise mitigation of
construction activities (McGarry et al., 2017).
Ship Strikes
With the advent of larger, faster moving ships since the 1950s,
there has been a steady increase in ship strikes of baleen whales,
which is likely to continue to increase as ships become even
faster and quieter (Laist et al., 2001; McKenna et al., 2012;
Monnahan et al., 2015). Minke whale mortality events attributed
to ship strikes are usually characterized as blunt trauma (i.e.,
from ships colliding with the animal) or propeller cuts and
lacerations. A study assessing large whale serious injury and
mortality along the east coast of the United States from 1970
to 2009, documented that 17 out of 396 (4.3%) of documented
minke whale mortality events were due to ship strikes (van der
Hoop et al., 2012). Biases and under reporting of ship strike
incidents suggest that many more fatal and non-fatal ship strikes
occur than are documented (Peel et al., 2018). National and
international ship strike databases are now being developed
(Cates et al., 2017). More comprehensive, and in some regions,
mandatory reporting practices are expected to improve our
understanding of this serious problem for whale populations,
including minke whales.
Marine Litter and Chemical Pollution
Human caused pollution is a pervasive and increasing threat
to marine environments (Laist, 1997; Hardesty et al., 2015).
It has been estimated that more than six million metric tons
of plastic enters the oceans each year; this is predicted to
increase 10-fold within the next decade (Jambeck et al., 2015).
Ingestion of plastic marine debris has been reported in nine
species of baleen whales, including minke whales. Effects range
from non-discernible pathological impacts to complete blockage
of the digestive tract resulting in malnutrition, starvation and
in some cases, mortality (Laist, 1997; Baulch and Perry, 2014).
Although levels of persistent organic pollutants (POPs) are
decreasing in some populations (Lebeuf et al., 2014), they are
of concern to many cetacean species, who may accumulate
high concentrations in their blubber (Jepson and Law, 2016).
Baleen whales, including minke whales, typically feeding on lower
trophic levels than toothed whales, have comparatively lower
blubber POP concentrations (Elfes et al., 2010). Nonetheless
POPs may have adverse effects on health and reproduction of
populations of both species groups (Moon et al., 2010; Hall
et al., 2018), especially in combination with other stressors
(Côté et al., 2016).
Underwater Noise
Over the last few decades, the potential impacts of humanmade noise in the marine environment have raised concerns
and received increased attention from scientists, conservationists,
resource managers, policy makers and the public. Sources of
anthropogenic noise are numerous and include military and
scientific sonar, seismic surveys, underwater explosions, offshore
construction and platforms, scientific research, pingers, and
acoustic deterrent devices (ADDs), as well as shipping and
vessel traffic (Prideaux, 2017). Measurements of ambient noise
levels have shown that low-frequency noise has been steadily
increasing, based on recordings analyzed from the 1960s to
2000s in the Northeast Pacific (Andrew et al., 2002; McDonald
et al., 2006a), and from 2002 to 2012 in the Indian Ocean
(Miksis-Olds et al., 2013). These increases in low frequency
noise were primarily driven by shipping noise and noise from
seismic air guns.
Information about minke whale hearing capabilities is
very limited, as audiograms for large whales cannot be
measured directly. Predictions of sensitivity to noise are
2
Entanglement in Fishing Gear
Entanglement in fishing gear is another global problem, affecting
a wide range of marine mammal species (Laist, 1997; Read, 2008;
Baulch and Perry, 2014; Meager, 2016). Partly due to higher
abundance compared to other species in some areas, minke
whales are the species most frequently reported dead due to
entanglement in fishing gear (Ryan et al., 2016). In Scotland,
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for example, evidence of entanglement in static fishing gear was
present in as many as 50% of stranded minke whales examined
from 1990 to 2010 (Northridge et al., 2010). Particularly high
levels of by-catch and entanglement of the J stock population
of minke whales, inhabiting the Sea of Japan, Yellow Sea,
and East China Sea, have raised serious concerns in the IWC
Scientific Committee in recent years (International Whaling
Commission, 2010; Song et al., 2010). In South Korea, by-caught
minke whales can legally be sold and a reporting system has
been in place there since 1994. Although reporting of animals
is not always accurate, these data show that from 1996 to
2014, minke whales by-caught in Korean waters alone exceeded
the estimated level of potential biological removal (PBR) for
this population (Song, 2016). Due to their small size, minke
whales are more likely to die as a result of entanglement than
other species, as they are less likely to be able to reach the
surface when weighted down by fishing gear and more often
sink after death, which may also lead to under-reporting of
entanglement cases for this species (Knowlton et al., 2016). It
has been estimated that 70% of entanglement cases for this
species are fatal (Lien, 1994). More recently, several studies
have highlighted the long-term consequences of entanglement
and animal welfare implications for individuals, often suffering
serious health effects over many years, and urged policy and
management action to be more inclusive of these issues (Cassoff
et al., 2011; Moore and van der Hoop, 2012; Dolman and Moore,
2017; Dolman and Brakes, 2018).
MONITORING AND RESEARCH
APPROACHES
Visual Monitoring and Behavioral Studies
Visual monitoring, from boats, ships or airplanes, is still one
of the most common approaches to study marine mammal
distribution and abundance (Barlow, 2015; Hammond et al.,
2017). Despite recent advances in visual monitoring methods
(Ferguson et al., 2018), these approaches always will be limited
for monitoring animals that spend a large proportion of their
lives under water (Borchers et al., 2013; Thomson et al., 2013).
In addition, visual surveys are dependent on daylight, and
good sighting and weather conditions, and are often spatially
restricted to coastal areas. Therefore, they can only provide a
snapshot of the true distribution, particularly for far-ranging
species such as minke whales (Kaschner et al., 2012). In part
due to such limitations and their small size and inconspicuous
blows and behavior, knowledge on long-term minke whale
distribution and abundance, especially during winter, is still
sparse (Best et al., 2012).
Research on dwarf minke whales has largely been based
on a small population wintering in the Great Barrier Reef,
utilizing dive tourism vessels as platforms of opportunity. Despite
the limitations associated with the use of such platforms (i.e.,
non-systematic survey effort), the ability to study the whales
from in-water, and the collection of long-term datasets (e.g.,
1996 to present) has enabled a diverse and innovative research
programme. Such research has included “interacting population”
size estimation using mark-recapture modeling from photoidentified whales (Sobtzick, 2010), population demography via
underwater videogrammetry (Sobtzick, 2010) behavioral studies
(Mangott et al., 2011), passive acoustic monitoring (PAM)
(Gedamke et al., 2001), social and economic valuation of the
swim-with-whales tourism activity (Stoeckl et al., 2010), applied
research for adaptive management (Curnock et al., 2013), and
recently, genetics, migration and habitat use studies via biopsies
and satellite-tagging (Birtles et al., 2015).
Tourism
Whale watching tourism with a particular focus on minke
whales occurs in several high latitude regions worldwide based
on seasonal feeding aggregations (e.g., West Scotland, Iceland,
Canada’s Gulf of St. Lawrence), as well as in Australia’s Great
Barrier Reef, which provides habitat for a presumed breeding
aggregation of dwarf minke whales (Valentine et al., 2004;
O’Connor et al., 2009). Whale watching of minke whales is
typically vessel-based, with the Great Barrier Reef case being
a unique swim-with activity, that has been managed by the
Great Barrier Reef Marine Park Authority since 2003 (Valentine
et al., 2004; Mangott et al., 2011; Curnock et al., 2013).
To better understand the potential impacts of vessel-based
tourism interactions on minke whales and other baleen whales,
recent research has focussed largely on behavioral responses to
vessels, including changes in swim speed and direction, diving
behavior, and decreased time spent resting or feeding (New
et al., 2015). Relatively few studies have investigated direct
physiological effects associated with disturbance from vessels,
including from direct vessel approaches (Christiansen et al.,
2014), vessel noise (Erbe, 2002), and vessel pollutants such as oil
leaks and exhaust emissions (Lachmuth et al., 2011). While some
effects have been inferred, for example increased stress levels
and energy expenditure among minke whales in the presence
of whale watching vessels (Christiansen et al., 2014), longerterm chronic and cumulative effects of whale watching are of
potentially greater concern, but are difficult to assess and thus
remain largely unstudied and unknown (Bejder et al., 2006;
New et al., 2015).
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Photo-Identification
Photo-identification has been used to recognize and re-sight
individual common minke whales since the late 1970s (Dorsey,
1983). Individual identification from surface photography
typically relies on clear images of dorsal fin profiles, body
scars and pigmentation patterns, and the technique has been
used to establish seasonal and inter-annual site fidelity and
residency times in a number of feeding areas, for example
the east and west coasts of North America (Dorsey, 1983;
Stern et al., 1990), and the Isle of Mull, Scotland (Gill et al.,
2000). Underwater photo-identification of dwarf minke whales
in Australia’s Great Barrier Reef, ongoing since 1996 (Arnold and
Birtles, 1999), is based primarily on lateral pigmentation patterns
which have been shown to have sufficient individual variability
and long-term stability, akin to humpback whale flukes (Arnold
et al., 2005; Sobtzick, 2010). Due to their interactions with
tourists and the growing popularity of underwater photography,
there is a significant “citizen science” component to this
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and even extract DNA (Pirotta et al., 2017). This information
is useful for determining stress and pregnancy as well as the
population of which an individual is a part. UAS can also be
used to survey smaller areas at high-resolution with accurate
geo-referencing. This information is critical when linking the
small-scale movement patterns and behavior of animals to their
environment as such surveys can provide insights as to the
amount of available habitat in an area.
photo-identification programme, to which tens of thousands of
images are now contributed each winter season (Sobtzick, 2010;
Birtles et al., 2014). Mark-recapture modeling has provided an
estimate of the “interacting population” (noting an unknown
proportion of whales that do not interact voluntarily with vessels
and swimmers) over three seasons, consisting of several hundred
animals each year (Sobtzick, 2010). Due to an increasing quantity
of images collected annually, the manual image matching
process is not cost-effective for some larger populations, and
new pattern recognition algorithms may provide a solution
to this problem. Konovalov et al. (2018) report on a “proof
of concept” for recognizing individual dwarf minke whales
using the Deep Learning Convolutional Neural Networks tool.
Further advancements in the use of such technology may
lead to a viable alternative to manual photo analysis in
the near future.
Passive Acoustic Monitoring (PAM)
Minke whales are known to produce a variety of sounds
across their geographic range. In the North Atlantic, frequencymodulated downsweeps and low-frequency pulse trains with
varying inter-pulse interval structure have been attributed to
minke whales (Beamish and Mitchell, 1973; Edds-Walton, 2000;
Mellinger et al., 2000). In the early 2000s, the North Pacific
“boing” sound was identified as a minke whale call (Rankin
and Barlow, 2005), and more recently low-frequency downsweep
and pulse-train sounds were described from recordings made
in British Columbia, Canada (Nikolich and Towers, 2018).
Australian dwarf minke whales produce a complex call,
dubbed the “star wars” vocalization (Gedamke et al., 2001;
Figures 2A–C). In the southern hemisphere, the “bioduck signal”
(Figures 2D,E), which had been recorded in the Southern Ocean
for decades, has recently been attributed to the Antarctic minke
whale (Risch et al., 2014b).
As mentioned above, visual surveys for marine mammals
are carried out mostly during daylight and summer months,
leaving large data gaps for many species’ winter distribution and
daily habitat use. Long-term PAM surveys have been shown to
be capable in filling some of these gaps and are particularly
useful for the study of cryptic species that are difficult to
detect visually, such as minke whales (e.g., Risch et al., 2019).
Recent advances in digital signal acquisition and processing,
increased storage capacity, and reduced power consumption and
instrument size means that PAM is increasingly complementing
visual surveys of marine mammals (Clark et al., 2010; Gerrodette
et al., 2011). Towed hydrophone arrays are now commonly
used on marine mammal surveys, and in some cases, the
passive acoustic data collected have been used for estimates
of relative abundance of minke whales (Barlow and Taylor,
2005; Norris et al., 2017). Large-scale networks of bottommounted hydrophones and autonomous acoustic recorders
have been used to investigate migratory routes, changes in
distribution and to describe previously unknown habitats of
baleen whales (Clark and Gagnon, 2004; Risch et al., 2014a;
Davis et al., 2017). Time-synchronized hydrophone arrays are
now being used in semi-automated approaches to localize and
even track individual animals (Martin et al., 2017). Where
detailed knowledge on the vocal behavior (e.g., calling or
cue rate) of the target population exists, PAM can be used
to estimate densities of animals (Lewis et al., 2007; Efford
et al., 2009; Marques et al., 2013; Harris et al., 2018). For
example, passive acoustic data has been used to estimate relative
abundance of North Pacific minke whales in a small area
using their “boing” vocalizations and applying cue-counting
Satellite Tags, Multisensor Tags, and
Unmanned Aerial Systems (UAS)
Technological advances are improving our ability to measure
and understand the biology of minke whales around the world.
Although not necessarily novel, satellite telemetry is a key tool
for collecting data that improves our understanding of the broadscale movement patterns, behaviors, and associations between
individuals and their environment over times that are relevant
to the biology of the species. Information from satellite tags
can provide insights on critical and preferred habitats. From
foraging to migration, satellite tags offer a unique perspective on
minke whales over extended spatio-temporal ranges that cannot
be easily obtained using other methods (Birtles et al., 2015;
Víkingsson and Heide-Jørgensen, 2015).
The advent of smaller multi-sensor recording tags has
revolutionized our ability to quantify and provide detailed
information on the underwater behavior of diving animals. For
example, researchers are now able to observe and measure
the feeding rates of minke whales (e.g., Friedlaender et al.,
2014). These motion-sensing tags (which can also include pointof-view video) allow for unprecedented measurement of the
kinematics, speed, maneuverability, energy expenditure, and even
information on the habitat of diving animals (Friedlaender et al.,
2009, 2014, 2016; Goldbogen et al., 2015, 2017; Cade et al., 2016).
The ability to measure the fine-scale movement and behavior of
minke whales is critical to providing quantitative information
on their energetic needs, daily behavior, and potential for being
impacted by environmental change.
Unmanned aerial systems (UAS) are an emerging technology
that can be used in a number of different capacities to collect
valuable information on minke whales and their habitats. UAS
have been shown to be able to accurately measure not only the
total length of individual marine mammals, but also their general
health condition (Christiansen et al., 2016; Durban et al., 2016).
This information is critical to being able to extrapolate to the
population-level regarding the age-class distribution, individual
health measures, and overall fitness of a population. UAS are
also being used to collect expired breaths from marine mammals
that can be chemically evaluated to measure hormone levels
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FIGURE 2 | Spectrograms of known minke whale vocalizations from different oceanbasins. (A) North Atlantic slow-down pulse trains, (B) North Pacific “boing”
vocalization, (C) Australian dwarf minke whale “star wars” vocalization, (D) Antactic “bioduck,” (E) Antarctic downsweeps. Note different time and frequency scales.
Spectrogramparameters: (A) Fast Fourier Transform (FFT) = 1024 pt, overlap (ovlp) = 75%,samplerate (SR) = 2000 Hz: frequency resolution (FR) = 1.9 Hz, time
resolution (TR) = 128 ms; (B) FFT = 512 pt, ovlp = 75%, SR = 8000 Hz: FR = 15.6 Hz, TR = 16 ms; (C) FFT = 512 pt, ovlp = 75%, SR = 16,000 Hz: FR = 31.5 Hz,
TR = 8 ms; (D) FFT = 1024pt, ovlp = 75%, SR = 4000 Hz: FR = 3.9 Hz, TR = 64 ms; (E) FFT = 4096 pt, ovlp = 75%, SR = 25,811 Hz: FR = 6.3 Hz, TR = 39 ms.
rate for most marine mammals, including minke whales can be
quite variable (Oleson et al., 2007; Parks et al., 2011b), making it
difficult to estimate accurate and robust cue rates. Techniques to
overcome these challenges include collecting data using acoustic
tags and behavioral focal follow studies (Johnson et al., 2009;
Parks et al., 2011a).
In addition to monitoring population abundance, PAM data
can be used for exploring geographic variability in vocalizations
with the potential to reveal population structure (Rankin and
methods using the calling rate from a single tracked individual
(Martin et al., 2013).
The recent development of these new statistical methods for
density estimation based on PAM data show great promise for
monitoring minke whales in challenging offshore, remote, and
sometimes ice-covered habitats. However, several assumptions
must be met and parameters, such as cue rates, need to be known
in order to reliably estimate animal abundance from passive
acoustic data (Marques et al., 2013). However, the vocalization
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Barlow, 2005; McDonald et al., 2006b), migration routes (Norris
et al., 1995; Risch et al., 2014a), habitat use and environmental
correlates (Baumgartner and Fratantoni, 2008; Stafford et al.,
2009) as well as for the study of behavior and the impact of
anthropogenic activities (Forney et al., 2017).
minimum population estimate for this population is 5,247
animals (Song, 2016).
The abundance of dwarf minke whales in the Southern Ocean
has not been assessed systematically, because in the past most
sighting surveys and whaling records did not distinguish it from
the Antarctic minke whale (Cooke et al., 2018a). For example,
the IWC sightings surveys carried out in the Southern Ocean
(IDCR-SOWER) identified only a small percentage of minke
whale sightings as dwarf minke whale (Branch and Butterworth,
2001). To our knowledge, the only abundance estimate of a dwarf
minke whale population to date is from the Great Barrier Reef,
where photo-ID data collected over 3 years suggested a yearly
fluctuating “interacting population” size of between 342 and 789
individuals (Sobtzick, 2010).
The most recent estimate of overall abundance of Antarctic
minke whales is around 500,000 individuals, down from an
estimated 720,000 in earlier assessments (International Whaling
Commission, 2013). Notwithstanding a lack of confidence in
parts of this assessment, this represents a 31% decline, the
direction and reasons of which currently are unknown. There is
also disagreement as to whether current abundance estimates are
similar to pre-whaling abundance figures. One of the difficulties
in generating abundance estimates for this species is due to its
affinity for sea ice, as has been shown by recent aerial surveys
(Williams et al., 2014; Herr et al., 2019). Despite these logistical
challenges and ongoing discussions, current population estimates
and their trajectory give reason for concern and accordingly have
resulted in the recent classification of Antarctic minke whales as
Near Threatened under the IUCN Red List and under Appendix I
of CITES (Cooke et al., 2018b).
Combining the effects of various anthropogenic pressures
on individuals and populations and addressing them in a
cumulative context still presents a considerable challenge, in
urgent need of new scientific and conservation approaches
(Côté et al., 2016). This is especially true for highly mobile
species and when disturbances change over space and time.
Several frameworks such as the Population Consequences of
Disturbance (PCoD) model (Pirotta et al., 2018) have been
developed in recent years. These models attempt to link health
effects of combined pressures with vital rates that can be scaled
up to population consequences but are still often limited by
our ability to accurately parameterize parts of the model and
deal with uncertainties. While new research and innovations in
methods, tools and techniques are needed to address the many
outstanding information deficiencies, long-term monitoring of
populations and the pressures to which they are exposed,
provides vital feedback on the effectiveness of management
policies and actions. Such monitoring must be integrative,
encompassing not only abundance estimates of the whales, but
also components of the wider system in which they live, including
natural and anthropogenic pressures. In practice, these types of
monitoring programmes and adaptive management approaches
are scarce. However, an example of such a programme has
recently been applied in the Great Barrier Reef, which uses
the Driver, Pressure, State, Impact, and Response (DPSIR)
framework to identify and connect appropriate indicators for
monitoring and enhancing the system-wide understanding
CONSERVATION STATUS AND
MANAGEMENT APPROACHES
Although abundance estimates are missing for large parts of their
known distributional range, common minke whales are currently
listed as a species of Least Concern (LC) under the International
Union for Conservation of Nature (IUCN) Red List (Cooke et al.,
2018a). The Convention on International Trade in Endangered
Species (CITES) lists all common minke whale populations under
Appendix I (threatened with extinction), with the exception of
the West Greenland stock, which is listed under Appendix II (not
currently threatened but trade needs to be strictly regulated).
Latest best estimates by the International Whaling
Commission (IWC) are 90,000 (CI = 60,000–130,000) for
the eastern North Atlantic; 50,000 (CI = 30,000–85,000)
for the central Atlantic, and 17,000 (7,000–40,000) for
West Greenland3 . The most recent abundance estimate
for the Canadian East coast stock is 2,591 (CV = 0.81)
animals (Hayes et al., 2018). It has been suggested, that
there may be two separate breeding populations in the
North Atlantic (Anderwald et al., 2011), but no definite
breeding grounds have yet been confirmed. Other studies have
found no evidence for population structuring in the North
East Atlantic but highlighted the importance of analysis of
more genetic markers to answer the question conclusively
(Quintela et al., 2014).
The North Pacific has generally been surveyed less and
population estimates are thus patchy. The most recent analysis
of inshore ship line transect surveys from the US west coast
(Washington to California) estimated 636 (CV = 0.72) animals
(Carretta et al., 2017). A passive acoustic line-transect survey
of the Northern Mariana Islands estimated between 80 and 91
animals (CV = 0.34) for this area in winter and spring (Norris
et al., 2017). Because acoustic estimates are based on vocalizations
believed to be produced only by males, these are considered
minimum estimates. The latest IWC “best” estimate for the
western North Pacific and Okhotsk Sea is 22,000 (CI under
review) animals3 . At least two sub-populations are currently
recognized in the western North Pacific, although acoustic
records suggest that further population structure might exist
(Rankin and Barlow, 2005). While the sub-population called
“J stock” inhabits the Yellow Sea, East China Sea and the
Sea of Japan, “O stock” inhabits more offshore waters of the
Northwest Pacific and the Okhotsk Sea (Kato, 1992). There
has been serious concern and debate within the IWC about
the status of J stock, which in addition to being targeted by
whaling operations, is also frequently caught incidentally in
fishing gear (Song, 2016; Cooke et al., 2018a). The current
3
https://iwc.int/home (accessed March 29, 2019).
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FIGURE 3 | Example Driver, Pressure, State, Impact, and Response (DPSIR) flowchart identifying system components and relationships relevant to integrated
monitoring of dwarf minke whales in the Great Barrier Reef World Heritage Area (adapted from Curnock et al., 2018, for illustrative purposes).
of knowledge gaps, barriers to effective management and the
prioritization of monitoring effort (see Figure 3).
(Great Barrier Reef Marine Park Authority, 2015). Developed in
the late 1990s by the European Environmental Agency (EEA),
DPSIR provides an integrative model for monitoring and
assessment of linked social-ecological systems at multiple scales
(Gari et al., 2015). Within DPSIR, Drivers are the overarching
causes and trends that influence a range of events/activities
that exert Pressures on parts of the system (e.g., climate change
influences storm intensity, and coastal development influences
ocean pollution). State represents the condition of a value (e.g.,
a characteristic of a whale population, such as abundance, health,
reproduction, etc.) that can/will respond to pressures and change
over time, whereas Impact is considered to be the resultant
effect to human well-being that flows from a change in the
state of a value (e.g., a loss of cultural, economic or heritage
value associated with a whale population). Responses are actions
taken by resource managers and/or communities to influence
drivers, mitigate pressures and/or restore the state of values
(Carr et al., 2007; Tscherning et al., 2012; Gari et al., 2015).
The development of an integrated monitoring programme for
the Great Barrier Reef utilizes DPSIR to identify relationships
and processes between system components (e.g., whales and
shipping, agriculture and corals, etc.) assisting with identification
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CONCLUSION: DATA GAPS, RESEARCH,
AND MONITORING PRIORITIES
The expansive, seasonally changing distribution patterns of
many minke whale populations, means that they are exposed
to a number of anthropogenic threats throughout their range.
Pressures vary in type and severity in different habitats and are
often cumulative, making it difficult to protect such highly mobile
populations, especially when large parts of their year-round
distribution are unknown.
A priority for future research should therefore be to better
understand migratory routes and seasonal habitats, in particular
winter breeding grounds, of minke whale populations worldwide.
Such efforts should focus on populations that are currently
commercially exploited or threatened, like those in the Northwest
Pacific, and those that are currently not being monitored, such
as almost all populations of dwarf minke whales. In the case of
Antarctic minke whales it is important to better understand their
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in-ice distribution and its effects on current population estimates.
Dedicated and opportunistic PAM networks are recommended to
define migration corridors, identify currently unknown offshore
habitats and monitor longer-term changes in distribution (see
Davis et al., 2017). In this context, a better understanding of the
full vocal repertoire, especially behavioral and density dependent
variability in calling rates will also be important.
Both common minke whales in the Arctic and Antarctic
minke whales show sexual segregated migration with females
generally moving toward higher latitudes (Horwood, 1990;
Laidre et al., 2009). A better understanding of such behavior
is important for species monitoring and assessing impacts of
ongoing whaling operations. Non-lethal new technologies such
as PAM, UAS, multi-sensor and satellite tags can help to
address such knowledge gaps even in challenging remote and
offshore habitats.
Entanglements in fishing gear and ship strike are affecting
baleen whale populations worldwide, but the extent of these
impacts and how they relate to other pressures are rarely fully
understood. The case of the J-stock population of common minke
whales in the Northwest Pacific is an example of interactions
of pressures putting a small population at risk for survival.
Japan’s announced withdrawal from the IWC in December 2018
potentially places even more pressure on this endangered (and
other) populations in their EEZ.
Finally, climate change and its effects on global oceanic
ecosystems are changing environmental and anthropogenic
pressures that determine migratory routes and species habitats
at an increasingly faster pace (Tulloch Ayesha et al., 2019),
requiring flexible and adaptable conservation approaches. Across
species, comparative and multi-disciplinary approaches are thus
necessary to conserve minke whale populations worldwide.
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Conflict of Interest Statement: TN was employed by Bio-Waves Inc.
The remaining authors declare that the research was conducted in the absence of
any commercial or financial relationships that could be construed as a potential
conflict of interest.
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May 2019 | Volume 6 | Article 247