Report of the Expert Workshop to
Review the Japanese JARPA II
Special Permit Research Programmej. cetacean res. manage. 16 (suppl.), 2015

369

Report of the Expert Workshop to Review the Japanese
JARPA II Special Permit Research Programme1
The Workshop was held in the Toyomi Center Building,Tokyo from 24-28 February 2014.1
1. INTRODUCTORY ITEMS
1.1 Chair’s opening remarks
The Workshop was chaired by Palka, ex-Chair of the IWC
Scientific Committee. She explained that normally the
Chair of Special Permit Review Workshops was the present
Scientific Committee Chair, who at this time is Kitakado.
However, since Kitakado was a member of the proponents,
the Scientific Committee agreed at their last Annual Meetingthat the previous Committee Chair would be an appropriate
chair for this Review Workshop. Palka then welcomed thePanel members, observers and Japanese proponents to Tokyoand thanked the Fisheries Agency of Japan for hosting theWorkshop. Morishita, the IWC Commissioner for Japan,
also welcomed the Panel and all participants.
The Chair recalled that the JARPA and JARPA II
programmes have been reviewed three previous times: inMay 1997 during an IWC sponsored mid-term review of
the JARPA programme; in January 2005 during a non-IWC
review of the results of JARPA for the period 1987/882003/04; and in December 2006 during an IWC final reviewof the JARPA programme.
Since that time, the Scientific Committee had developedthe ‘Annex P’ process of reviewing special permit
programmes (IWC, 2013b). The Committee had held two
previous review Workshops for special permit programmes
following this process. The first was a review of the ongoing
JARPN II programme in the western North Pacific (IWC,2010) and the second was the final review of the Icelandic
programme (IWC, 2014a).
1.2 Terms of Reference‘Annex P’ contains Terms of Reference for review ofongoing proposals and completed proposals. Those that are
relevant to the JARPA II review can be summarised as:(1) how well the original objectives were met;(2) whether the programme made other contributions toimportant research needs;(3) the relationship of the research to IWC resolutions anddiscussions;(4) utility of lethal and non-lethal techniques; and(5) provide advice on:(a) practical and analytical methods that can improveresearch relative to stated objectives;(b) appropriate sample sizes to meet the statedobjectives;(c) effects on stocks; and(d) when a further review should occur.
1.3 Overview of the process for developing advice for
the Commission
The first component of the process is the present small
specialist Workshop (this has to be held at least 100 days
before the SC meeting), with a limited number of invited
experts chosen by a Standing Steering Group (SSG); alimited number of proponent scientists, primarily to present
Presented to the meeting as SC/65b/Rep02.

1

the proposal and answer points of clarification; and a limited
number of observers. Based on the proponents paper on
analytical methods used in JARPA II, which was submitted
nine months prior the Annual Meeting (e.g. 3 August 2013), in accordance with Annex P the SSG drew up a shortlist of
potential Panel members. Final selection was governed byavailability (many of the potential nominated members wereunable to attend), the need for balance (including between
Scientific Committee and non-Scientific Committee
members) and the available funds. The final Panel is listed
in Annex A and comprised of three scientists who had neverparticipated in the Scientific Committee, two scientists who
have infrequently and not recently participated in ScientificCommittee meetings and three regular members of the
Scientific Committee; this is in addition to the Workshop
Chair and Head of Science. One member, Punt, was also an
author of two ‘O’ papers. Those papers, on statistical catchat age analysis (SCAA) were developed under contract of
the IWC and implemented work directed by the ScientificCommittee and were thus not deemed to include any conflictof interest. His participation in the discussion of that item
within the Panel discussions was to provide information as
required.
Five members of the Scientific Committee attended as
observers and their names are given in Annex A as are theproponent scientists who attended the open sessions.
The report of the Workshop is to be made available for
proponents by 11 March 2014. They are able to commenton the Panel report and to revise documents for submission
to the Scientific Committee in light of comments within
the Panel’s report. The final report, attached commentsfrom proponents and any moderately revised workshop
documents developed as a result of recommendations are to
be made available to the Scientific Committee by 7 April
2014. Major revisions or completely new documents basedon recommendations must meet usual Scientific Committee
document deadlines.
The Scientific Committee may make additional comments
and the report of the Workshop, comments by proponents
and comments by the Committee itself will be sent to the
September 2014 Commission meeting.
1.4 Meeting arrangements and work schedule
The Chair explained that she will follow the previous
Workshop format, with two types of sessions:(1) open sessions where a limited number of scientists
associated with the JARPA II research present the
programme results and answer questions from the Panel– this is also open to observers; and(2) closed sessions where only the Panel members discuss
the proposal and develop their report.
In particular, her general intention was that for themorning sessions on days 1-4, the Japanese scientists andobservers would provide their PowerPoint presentations
for each of Items 3-11, although she recognised that someflexibility would be required and presentations may run into
the early afternoon. Day 5 was for the Panel to discuss its
overall conclusions and develop its report.
In the afternoon, the Panel will hold a closed meetingfor summarising conclusions and recommendations for370

report of the expert workshop to review jarpa ii

each agenda item. During the closed sessions, she notedthat it was possible that the experts may want to ask further
clarifications and questions to the proponent scientists. She
therefore requested that where possible they remain near
the venue in the afternoon; although in this review this was
rarely necessary. The Panel may request working papers
from the proponents on specific topics as was the case inprevious special permit reviews.
She referred to the ‘Annex P’ guidelines which statethat Scientific Committee observers will ‘not normallyparticipate in discussions unless invited to do so by the Chair
under special circumstances’. She was pleased to note thateight papers from Committee members had been submitted
to the Workshop and that most of the authors of these papers
were present. Several of these papers were presented by theauthors and they provided valuable input into the Panel’s
discussions.
1.5 Review of documents and data
SC/F14/J01-38 (papers from proponents), SC/F14/O01-O08(papers from Scientific Committee members), and SC/F14/
R01-R05 (papers from the proponents that were in response
to a SC/F14/O paper) were made available to the Workshop
in accordance with deadlines specified in ‘Annex P’. The listof documents is given as Annex C. A number of background
papers were also available and these are referred to whereappropriate in the text. The proponents, observers andScientific Committee members submitting documents
prepared PowerPoint presentations on each of the major
topics (sometimes these applied to individual papers andsometimes they incorporated the results of several papers).
It was agreed that with permission of the authors, thesepresentations would be made available to observers as well
as Panel members but would be treated in a confidential
manner.
2. ADOPTION OF THE AGENDA
The adopted agenda is given as Annex B.
3. PROPONENTS’ OVERVIEW OF THE RESEARCHOBJECTIVES AND RESULTS (SC/F14/J01-J02)
The overview of objectives and results below is based on
SC/F14/J01 prepared by the proponents. This summary has
not been edited or approved by the Panel but represents theviews of the Proponents. The Panel’s views of individual
items can be found under Items 4 onwards. The Panel’s
conclusions and recommendations are found under Item 12.
3.1 Overview of Objectives
Objectives 1 and 2
The rationale and research needs of JARPA II wereexplained in the original research plan, which was presented
and discussed at the IWC Scientific Committee (IWC SC)
in 2005 (Government of Japan, 2005). The development of
the research objectives of JARPA II took into consideration
the findings of the JARPA. During the 18-year period,
the JARPA research showed evidences of the recovery ofsome whale species such as the fin and humpback whales
which had been depleted by commercial whaling in the past
century. One example is the increase in the abundance ofthe humpback whale in Areas IV and V (Matsuoka et al.,2011). On the other hand the Antarctic minke whales werebroadly stable with at most a slight decline in the same
Areas and period (Hakamada et al., 2013). The increase in
the abundance of some large whale species implied changes

in whale composition and distribution. For example in Area
IV the humpback whale was extending its distribution to thesouth, sharing habit with the Antarctic minke whale along
the ice-edge (Murase et al., 2002). Because whales are top
predators of the Antarctic marine ecosystem, the changes
could have substantial effects on the ecosystem as a whole.
The changes in the ecosystem could also have significant
implications to the IWC conservation and management of
whales.
JARPA research showed that a major transition has
been taking place in recent years, which would affect the
krill availability in the ecosystem and the biology of krill
predators such as the Antarctic minke whale (Fujise et al.,2006). In fact some changes in the biology of the Antarctic
minke whale were reported during the JARPA period
e.g. stable or slight increase in the age at sexual maturity(Zenitani and Kato, 2006) and deterioration in nutritional
condition as revealed by a decrease in blubber thickness,girth and fat weight (Konishi et al., 2008).
The JARPA II research plan therefore emphasised theneed to systematically monitor changes of the Antarctic
marine ecosystem over the long-term as well as changes of
biological parameters in krill predators, and changes in theabundance of cetaceans inhabiting the Antarctic Ocean. It isalso emphasised the need to monitor how cetaceans adapt to
global warming and the changes in the ecosystem structurecaused by human activities so as to provide a scientificbasis for the comprehensive management of whale stocks(Government of Japan, 2005).
It should be noted here that continuous monitoring
programmes form the backbone of all scientific research
programmes which have the aim of providing advice onsustainable levels of catch for marine or terrestrial living
resources. The reason is that population dynamics, and hence
the size of the sustainable yield, can change in a manner that
may not be predictable. Factors indexing these dynamics
must therefore be monitored so that changes can be detected
and important scientific information can be provided to the
IWC as the basis for their conservation and management
decisions.
Objective 1 of JARPA II was therefore the ‘Monitoringof the Antarctic ecosystem’: (i) monitoring of whale
abundance trends and biological parameters; (ii) monitoring
krill abundance and the feeding ecology of whales; (iii)monitoring the effects of contaminants on cetaceans; and(iv) monitoring of cetacean habitat.
Some specific scientific questions to be addressed under
this objective are the following.
Given the results of JARPA that indicated a recent change
in the Antarctic ecosystem, the aim here is the monitoring of
temporal trend of parameters considered as ‘indicators’ ofchanges in the ecosystem.
• Whale abundance based on sighting data:- is the abundance of blue, humpback and fin whales
increasing in the JARPA/JARPA II period?
- is the abundance of Antarctic minke whale still
maintaining a stable trend in the JARPA/JARPA II
period?
• Whale abundance based on SCAA (age and sightingdata):- what is the historical and recent trends in recruitment
and total abundance of Antarctic minke whale?
• Whale distribution based on sighting data:- how the increase in abundance of large whales isaffecting the distribution of Antarctic minke whale?
• Body condition and stomach content weight:

j. cetacean res. manage. 16 (suppl.), 2015- are these indices of nutrition declining in the JARPA/
JARPA II period?
• Biological parameters based on age and reproductivedata:- what is the historical and current trends of age atsexual maturity?
- has the age at sexual maturity increased recently?
- has the pregnancy rate decreased in the JARPA/
JARPA II period?
• Contaminant load based on mercury and organochlorinedata:- what is the pattern of temporal trend of pollutant
levels in Antarctic whale samples?
- what is the accumulation mechanism of pollutants in
the Antarctic ecosystem?
• Cetacean habitat based on oceanographic and marine
debris data:- how have the oceanographic conditions changed
through the JARPA/JARPA II period?
- what is the temporal trend of marine debris in the
JARPA/JARPA II period?
Model building is essential for understanding,interpreting, and predicting the possible change in the
Antarctic ecosystem indicated under the Objective 1.
Models would also contribute to the improvement of the
IWC conservation and management measures, as presented
under Objective 4 of JARPA II (see below).
Objective 2 of JARPA II therefore is ‘Modellingcompetition among whale species (to inform) future371

management objectives’: (i) constructing a model ofcompetition among whale species; and (ii) new management
objectives including the restoration of the cetacean
ecosystem.
The key question for the first part of Objective 2 (i)above is the following:• can the predators-prey interaction alone explain theobserved population trends of whale species and changes
in biological parameters of the Antarctic minke whale
in recent decades, without the need for recourse toenvironmental change hypotheses?
Objectives 3 and 4
The IWC adopted the Revised Management Procedure(RMP), the procedure for the management of commercial
whaling for baleen whales. There is a need for better
estimation of Maximum Sustainable Yield Rate (MSYR) in
order to respond to any concerns over the implementation of
the RMP and to improve its likely deficiencies concerning
utilisation of whale resources. In the past the RMP Small
Areas for Antarctic minke whale in the Antarctic were
established as longitudinal sectors of 10° because there wasno reliable scientific information on its stock structure, but
at the very least, there is a need to redefine appropriate Small
Areas in accordance with research results. Also anotherof the deficiencies of the current RMP is the zero catch
quotas that it turns out when carrying capacity declines
due to competition among whale species. The decrease inabundance caused by the competition is misinterpreted by
the current RMP as an over-exploitation so that catches are

Fig. 1. Stock structure hypothesis of the Antarctic minke whale based on genetic and non-genetic data from JARPA (IWC, 2008). The upper figure shows theencounter rates of whales in 10° squares of latitude and longitude in waters 0°-30°S in October (Kasamatsu, 1995).

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report of the expert workshop to review jarpa ii

set unnecessarily low. That part needs also to be improvedby the use of more realistic multi-species models.
Important information for the application of the current
RMP process is stock structure, abundance and productivity(MSYR). JARPA research already provided some of theinformation.
For example the analyses of genetic and morphometricdata obtained under the JARPA suggested the occurrence ofat least two stocks in the JARPA research area, which forpractical reasons are called ‘Eastern Indian Ocean Stock’(I-Stock) and ‘Western South Pacific Stock’ (P-Stock). The
data suggested an area of transition in the region around 150°165°E (Fig. 1) across which there is an as yet undetermined
level and range of mixing (IWC, 2008).
Abundance estimates in Areas IV and V were conductedbased on the JARPA sighting surveys (Hakamada et al.,2013). ADAPT-VPA analyses based on age and abundance
data from the JARPA suggested MSYR1+ values in the 4-6%
range for Antarctic minke whale (Mori, 2006).
All this information, which is important for effective
management under the RMP, is refined under the JARPA
II in the context of the current RMP process as well in thecontext of the future improvement of the RMP.
Objective 3 of JARPA II is the ‘Elucidation of temporaland spatial changes in stock structure’: (i) Antarctic
minke whale; and (ii) other baleen whale species.
Stock structure information is a basic requirement for
the IWC conservation and management measures. It isessential for the implementation of the RMP. JARPA had
indicated that the Small Areas used in the trial application of
RMP to Antarctic minke whale in 1993 were not consistent
with the stock structure of this species in the research area.
Elucidation of the stock structure would also contribute to
Objective 4.
Some specific scientific questions to be addressed under
this objective are the following.
The aim of the JARPA II under Objective 3 is first to
update the analyses on stock structure of Antarctic minke
whale in the context of the suggested hypothesis, by using
new genetics information obtained in the JARPA II, and thento estimate the mixing proportion of the I and P Stocks inthe transition region, including possible yearly fluctuationin distribution and mixing proportion. The geographicaloverlap between the I and P Stocks under the JARPA II is
examined in a wider sector than that suggested in the IWC
JARPA Final Review (IWC, 2008): 130°-175°E. A different
aim is to investigate the stock structure of fin, humpback andsouthern right whales in the feeding grounds.
• Are the additional genetics analyses based on JARPA II
data consistent with the current stock structure hypothesis
for Antarctic minke whale?
• How the distribution and mixing of the I and P Stocks
changes annually?
• Are the fin, humpback and southern right whales
genetically structured in the Antarctic feeding grounds?
• What is the biological validity of the IWC Management
Areas for baleen whale species?
Objective 4 of JARPA II is ‘Improving the management
procedure for Antarctic minke whale stocks’.
The goals under Objective 4 will be addressed with progressof the work under the other three Objectives. This objective isdivided into two parts: (a) contribution to the implementationof the current RMP through the attainment of key informationsuch as abundance, stock structure and MSYR (contribution
from Objectives 1 and 3); and (b) contribution to the future
improvement of the RMP through the incorporation of a more

realistic (and possibly species-specific) MSYR range into the
RMP algorithm, and the incorporation of the effects arisingfrom the inter-species relationships among whale species(contributions from Objectives 1 and 2).
For (a) above, abundance and MSYR (through SCAA
analyses) will be obtained under Objective 1 while theredefinition of Management Areas and mixing will follow
results on stock structure under the Objective 3. For (b)above, MSYR (through VPA and SCAA analysis) will beobtained under Objective 1 and the incorporation of theeffects arising from the inter-species relationships amongwhales species will be examined after the development of
the ecosystem model has been completed under Objective 2.
Fig. 2 shows the interaction among the four Objectivesof the JARPA II.
Overview of results
Objective 1 ‘Monitoring the Antarctic ecosystem’
Abundance and distribution

Demographic parameters trends of Antarctic minke whale in1940s-1970s were consistent with the pattern expected under
the krill surplus hypothesis; the changes in the Antarctic
ecosystem suggested under the JARPA research (e.g. whale
species composition and distribution) was confirmed.
• SCAA analysis showed that Antarctic minke whale
abundance increased from 1930 until the mid-1970s (SC/F14/O02).
• SCAA analyses showed that abundance trend in recent
decades has been relatively flat in the I Stock and perhaps
declining slowly in the P Stock (SC/F14/O02).
• The total abundance of Antarctic minke whales in Areas
IIIE-VIW (Line Transect Method) has been roughly
stable during the JARPA+JARPA II research period (SC/F14/J03).
• The abundance trend estimates of humpback whale
Stocks D in Area IV (13.6% with 95% CI: 8.4%-18.7%)
and E in Area V (14.5% with 95% CI: 7.6%-21.5%) weresimilar, and they showed a significant increasing trend
during the JARPA+JARPA II period (SC/F14/J04).
• Spatial distribution of humpback whales in Area IV was
expanded in the period 1989-2006 while the distributionof Antarctic minke whales remained stable in the same
period (SC/F14/J18).
• The abundance of blue whales in Areas IIIE, IV, V andVIW combined showed a significant increasing trend
during the JARPA+JARPA II period (8.2% with 95% CI:
3.9%-12.5%) (SC/F14/J05).
• The abundance of fin whales in Areas V and VIW
combined showed a significant increasing trend during
the JARPA+JARPA II period (12.0% with 95% CI:
2.6%-21.5%). The increasing trend was not statisticallysignificant in Areas IIIE+IV (SC/F14/J05).
• The abundance of southern right whale in Area IV
showed an increasing trend, which was not statisticallysignificant (SC/F14/J05).
Biological parameters

Demographic parameters trends of Antarctic minke whale in1940s-70s were consistent with the pattern expected under
the krill surplus hypothesis; the change in the Antarctic
ecosystem suggested under the JARPA research (e.g.
whale species composition and distribution, biology of the
Antarctic minke whale) was confirmed.
• Age at sexual maturity of Antarctic minke whale
decreased between approximately 1940s and 1970s
cohorts.

j. cetacean res. manage. 16 (suppl.), 2015• The age at sexual maturity of Antarctic minke whale
Stocks I and P has remained at between 7-8 years oldfrom the 1970s cohorts. It showed a slight but statisticallysignificant increasing trend from the 1970s cohorts (SC/F14/J08).
• The recruitment rate of Antarctic minke whale Stocks I
and P increased between 1930s and 1970s and in recent
decades it maintained a stable trend at a low level (incomparison with the level observed in the 1960s-1970s
cohorts) (SC/F14/O02).
• The apparent pregnancy rate of Antarctic minke whale
Stocks I and P maintained a stable trend at a high level
during the JARPA+JARPA II period (SC/F14/J09).
• There is the possibility that fin whales are reaching sexual
maturity at a younger age as compared with the period ofcommercial whaling (SC/F14/J10).
Feeding ecology/energetic

The shift in the Antarctic ecosystem suggested under the
JARPA research was confirmed. Deterioration in nutritional
condition were found for the Antarctic minke whale. Less
krill availability for Antarctic minke whale could explain thedeterioration in nutritional conditions for this species. Less
availability could be due to: (i) decrease in krill biomass
because of global warming; or (ii) increase in the abundanceof competing krill predators including baleen whales. The
latter is better supported by the findings.
• The blubber thickness, girth and fat weight of Antarctic
minke whales showed a significant decreasing trend
throughout the JARPA period. There was no clear trend
for the JARPA II period (Konishi et al., 2008); SC/F14/
J13). For both data sets combined the decreasing trend
was less marked in comparison with that found for the
JARPA period alone.
• The stomach content weight of Antarctic minke whales
showed a significant decreasing trend throughout
the JARPA+JARPA II period in offshore areas. No

373

significant trend was found in the Ross Sea for females,
where humpback whales are not distributed (Konishi and
Butterworth, 2013); SC/F14/J14).
• The daily prey consumptions of Antarctic minke
whales per capita during the feeding season based on
two methods were 95.1-127.0 and 182.6-250.3kg for
immature and mature male, and 125.8-138.7 and 268.1325.5kg for immature and mature female, respectively.
These values are equivalent to 2.65-4.02% of the bodyweight (SC/F14/J15).
• The total prey consumption by Antarctic minke whalesin the research area was estimated in 3.51 and 3.98
million tons by each method, respectively. These values
correspond to 7.6% and 8.6% of the estimated krill
biomass in the research area (SC/F14/J15).
• The daily prey consumption per capita of Antarctic
minke whales decreased between the JARPA and JARPA
II periods for all sexual classes, based on the results ofdiurnal change in stomach content mass (SC/F14/J15).
• The daily prey consumption of fin whales per capita
during the feeding season based on three methods ranged
between 276 and 2,136kg. These values were equivalent
to 0.50 and 3.84% of the body weight (SC/F14/J16).
• Antarctic minke and fin whales fed on krill of similar
body size implying that fin whales and the Antarctic
minke whales have similar feeding habit, and no preysize selectivity (SC/F14/J16).
• Body weight of fin whales in the JARPA II period isheavier than those previously reported for the Antarctic
in the 1950s suggesting improved nutritional conditions
in recent decades (SC/F14/J10).
Pollutants• Total Hg levels in krill sampled from the stomachs of
Antarctic minke whales were similar between Areas IV(0.006-0.026 ppm dry wt) and V (0.003-0.052), and noyearly trend was observed for the period 2005-11 (SC/F14/J23).

Fig. 2. Schema summarising the objectives of the JARPA II prepared by the proponents. Several parameters considered ‘indicators’ of change in the ecosystem
are studied and monitored under Objective 1. The modelling exercise under Objective 2 aims to provide interpretations to the pattern of changes observed
under Objective 1: can these changes be explained only by the prey-predators interaction without the need for recourse to environmental change hypotheses?
Objective 4 depends on the results obtained under the other objectives: stock structure of Antarctic minke whale under Objective 3; abundance, MSYR andnatural mortality of Antarctic minke whale under Objective 1.

374

report of the expert workshop to review jarpa ii• Total Hg levels in liver of Antarctic minke whale in Areas
IV and V were 0.003-0.130 ppm wet wt and <0.0010.250, respectively. These levels are considerably lower
than those in other ocean basins. Hepatic Hg levels of
all age groups of Antarctic minke whale in Area IV
decreased significantly, and the age group 15-26 years
old in Area V increased significantly during the JARPA
and JARPA II period (SC/F14/J23).
• Mean concentration of organochlorine in 21-25 years
old minke whale in Area V in 2010/11 were HCB (140ng/g fat wt.), DDTs (100 ng/g fat wt.), PCBs (28 ng/g
fat wt.), CHLs (25 ng/g fat wt.), and HCHs (0.8 ng/g fatwt.). Levels of DDTs, HCHs, HCB and CHLs in Area V
decreased significantly with year, while the yearly trend of
PCBs did not change significantly. The levels of HCHs in
the Antarctic Ocean have varied from slightly decreasing
to a steady state in the middle 1990s (SC/F14/J24).
• Mean concentrations of total Hg in livers and musclesof fin whale were 0.052 and 0.021 (ppm wet wt.),
respectively. Mean concentrations of PCBs, DDTs,
HCHs, HCB and CHLs were 6.5, 13, 0.65, 39 and 4.5(ng/g fat wt.), respectively. Hg levels in liver and muscle
were one order of magnitude lower than those of fin
whales from the western North Atlantic Ocean. OCslevels were extremely lower than those in fin whales
from the middle latitude of the northern hemisphere, andthose in killer whales (top predator) from the Antarctic
Ocean (SC/F14/J25).
Oceanography/marine debris• The effect of the global warming was not detected in the
research area (SC/F14/J20).
• Annual fluctuation on the locality of the southernboundary (SB) was observed in the research area during
the JARPA and JARPA II periods. Southward shiftof the SB generally coincided with a rapid increase of
humpback whales in 1995/96 and 2001/02 suggestingthat oceanographic conditions affect the distribution ofwhales (SC/F14/J21).
• A low incidence of marine debris and entanglements wasobserved in the JARPA/JARPA II period (SC/F14/J22).
Objective 2 ‘Modelling competition among whale species’
While developing ecosystem models requires a substantial
data collection and analytical effort, initial progress wasmade toward this objective in JARPA II.
• Several input data for the models were collected by
JARPA and JARPA II. Progress was made toward thedevelopment of two types of ecosystem models: multispecies production model and Ecopath with Ecosim.
Several additional data and analyses were identified toprogress further this work (SC/F14/J26).
Objective 3 ‘Elucidation of temporal and spatial changes
in stock structure’
The stock structure hypothesis of Antarctic minke whales
derived from JARPA research was confirmed and refinedby JARPA II. Important stock structure information forassessment was also obtained for other large whale species
in the Antarctic feeding ground.
Antarctic minke whale• Results of additional genetics (mtDNA sequencing;
microsatellite DNA) analyses based on the JARPA II
data were consistent with the previous stock structure
hypothesis of at least two stocks in the JARPA II researcharea, which occupy the most western and eastern part of
the JARPA II research area, respectively (I and P Stocks)(SC/F14/J28)• Genetic analyses based on microsatellite showedsignificant yearly changes mainly within the stock inthe most western side of the research area (I Stock).
In general, results of the new microsatellite analyses
suggested stronger fidelity to migration destination infemales than in males (SC/F14/J28).
• The transition area where two stocks mix with each otheris longitudinally wider than originally suggested under
the JARPA. The spatial distribution of the two stocks
has a soft boundary in Areas IVE and VW, which varyannually (SC/F14/J29).
• There are possible sex differences along the soft boundary(SC/F14/J29).
Humpback whale• Genetic pattern of variation based on mtDNA wasconsistent with the geographical locations of breeding
and feeding grounds of Stocks D, E and F. Western
Australia whales are strongly related to Area IV, Eastern
Australia whales are related to Area VW and whales
from New Caledonia and Tonga are related with feeding
grounds east of 160°E. Whales from Cook Island andFrench Polynesia are not related genetically to whalesin the feeding grounds covered by the JARPA II surveys(SC/F14/J30).
• Mixing proportion estimates of Stocks D, E and F in thefeeding grounds were provided (SC/F14/J30).
• Results of microsatellite analyses showed that Areas IIIE,
IV, V and VIW are occupied by different stocks, which isconsistent with the mtDNA results (SC/F14/J31).
Fin whale• Results of mtDNA and microsatellite analyses suggested
genetic structuring in the JARPA II research area (SC/F14/J32).
Southern right whale• Genetic analyses based on microsatellites suggested
site fidelity of whales to Area IV. Results of the mtDNA
analysis suggested longer distance feeding migration
than previously thought, and the possibility that different
stocks migrate into Area IV in summer (SC/F14/J33).
Photo-identification• 3,108 photo-ID images of the blue, humpback andsouthern right whales were digitised and incorporated
into the ICR photo-ID catalogue (SC/F14/J34).
• Some ‘matching’ were found within the feeding groundand between feeding and breeding grounds, which areconsistent with the results of the genetic analyses (SC/F14/J34).
IWC management Areas• Results of the stock structure studies confirmed that thecurrent Management Areas of the IWC are not supported
by biological evidence in the case of the Antarctic minke
whale.
• The IWC boundaries of the Areas III, IV, V and VI match
better with the genetic results found for the humpbackand fin whales.
Objective 4 ‘Improving the management procedure for
Antarctic minke whale stocks’• New information on stock structure and mixing and on
abundance will be important for future RMP application
to Antarctic minke whale, as such information isessential for the RMP’s pre-Implementation assessment
and Implementation Simulation Trials. JARPA II has
shown that the previous trial specifications of the RMP
to Antarctic minke whale in 1993 is no longer valid.

j. cetacean res. manage. 16 (suppl.), 2015• The new information, particularly the stock structure
information, is to render the RMP more efficient, i.e.
providing extra catch without increasing conservation
risk. For example the 10 degree longitude slices usedpreviously as Small Areas are no longer supported by the
JARPA II data.
• SCAA analyses based on JARPA II data showed that
Antarctic minke whale can show growth rates that areappreciably larger than the lower bound for MSYR usedin RMP trials.
Other results• Alternative method for age determination in the Antarctic
minke whale by way of aspartic acid enantiomers wasdeveloped (SC/F14/J12).
• New genetic method based on paternity analyses of
foetuses to estimate the size of the mature male populationof Antarctic minke whale was developed (SC/F14/J07).
• Novel information on physiology and reproductive
biology of the Antarctic minke whale was provided (SC/F14/J35-J37 [published information]).
4. REVIEW OF RESULTS: Monitoring
cetacean habitat
4.1 Proponents’ summary of results
The comprehensive JARPA/JARPA II oceanographic survey
has derived in a very unique set of temperature and salinity
data profiles for almost half of the Antarctic area obtained
systematically in more than 20 years of research. This data
set enabled the oceanographic structure and dynamics inthe research area to be investigated. Southward/northward
movement of the Southern Boundary (SB) of the Antarctic
Circumpolar Current (ACC) could play an important role inthe distribution of whales in the research area. This should befurther investigated using the comprehensive oceanographic
and whale sighting data sets from JARPA and JARPA II.
Additional results, important for the interpretation of other
JARPA II results, is that the surveys provided no evidence of
global warming in the research area unlike the case reported
for the Antarctic Peninsula (Trivelpiece et al., 2011).
JARPA/JARPA II surveys made possible the firstcomprehensive observations of marine debris on sea surface,stomachs of whales, and of entanglements of whales in the
Antarctic. Given the low indices, the effect of marine debris
on whales in the Antarctic is expected to be limited at thepresent time. Continued monitoring is desirable.
SC/F14/J20 analysed oceanographic observation dataobtained by the JARPA and the JARPA II to clarify physicaloceanographic conditions in the JARPA/JARPA II area as abasis for understanding the habitat environment of whales.
About 2,500 profiles were obtained by XBT, XCTD and
CTD observations from 1990 to 2009. Profile data wereconverted to the same format and utilised to describe theoceanographic features of the JARPA/JARPA II area. Basedon this data set the averaged feature of the oceanographic
structure in the research area for two decades was described.
The Southern Boundary (SB) of the Antarctic Circumpolar
Current (ACC), which is detected by a 0°C temperature
contour line on the 27.6 isopycnal surface, characterised
the research area. The position of the SB is controlled bymajor features of bottom topography such as the Kerguelen
Plateau and the Pacific-Antarctic ridge. In the area east of
the Kerguelen Plateau, the position of SB changes on a
decadal timescale. The southward shift of SB in the region
is observed in the early 2000s, and a northward shift isobserved in the later 2000s. Although global warming is an

375

important factor for the Southern Ocean, unlike the Antarctic
Peninsula region, the JARPA and JARPA II temperature datashows no statistically significant warming in the JARPA/
JARPA II research area for the two decades examined.
SC/F14/J21 examined information such as: (a)Kerguelen-Davis Oscillation Index (KDOI); (b) the extent
of sea ice; (c) abundance estimates of humpback andAntarctic minke whales in Area IV from the 1989/90 to
2007/08 seasons collected by the JARPA and the JARPA II;
and (d) zonal average of Southern Boundary of Antarctic
Circumpolar Current (SB) in the Indian Ocean sector. Timeseries of these data showed trends and relations as follows:First, KDOI indicated: (1) the effective data of Kerguelen
had a limited number of years since 2006 while the Davis’s
data had 56 years since 1957, therefore, the analytical periodof KDOI was limited to seven and half years at this stage;and (2) KDOI during 2006 and 2013 showed the variabilityof an upward trend with approximately 2-year periodicity.
Second, sea ice extent in austral summer from 1979 to 2011
showed an upward trend with yearly variability suggestingsome relationship with KDOI. In addition, the lower
abundance estimate of Antarctic minke whale coincidedwith the larger sea ice extent such as 1997/98 and 2007/08,
probably due to navigational limited surveys caused by sea
ice. Third, the rapid southward shift of SB coincided witha rapid increase of humpback whales such as in 1995/96
and 2001/02, probably due to Antarctic Circumpolar Water
southward shift.
SC/F14/J22 summarised the information on marine debris
collected during the surveys of the JARPA and JARPA II for
the period 1987/88 to 2010/11. Three kinds of observations
were considered: marine debris on the sea surface, marine
debris in the stomachs of whales caught (Antarctic minke,
dwarf minke and fin whales), and entanglements. In the
Antarctic research area, south of 60ºS, marine debris
included metals (drum, can), petrochemical products (buoy,ball, bottle, container, fender, net, rope, styrofoma). Buoys
were the most abundant debris (69% of all marine debris
recorded). The highest density index (DI: number of marine
debris observed per 100 n. miles) was recorded in Area V(DI: 0.15), followed by Area IV (DI: 0.12). DI of buoys in
Area IV and V suddenly increased after the 2005/06 australsummer season. The increase of buoy debris coincides withan increase of long-line fishery operations in this area. The
stomachs of a total of 10,041 Antarctic minke whales, 16
dwarf minke whales and 16 fin whales were examined fordebris. A total of 70 marine debris and objects other than prey
were found in the stomachs of the three species, including
feather, stone, wood, plastic and others. The number ofoccurrence of marine debris and objects other than prey inthe fore and main stomachs per 100 Antarctic minke whales
examined was estimated at 0.35. Four cases of entanglement
in a total of 10,041 Antarctic minke whales examined were
found. Those involved fishing hook, monofilament fishing
line, rope and packing band.
4.2 Panel’s conclusions and recommendations
Monitoring the physical and biological habitat is an integral
part of monitoring a whale’s ecosystem, i.e. Objective 1 of
JARPA II. The survey methods of the original proposal stated:‘Oceanographic and meteorological observations will be
carried out while monitoring the environment, including sea
ice, surface temperature, sea surface height and chlorophyll
α concentration over the entire research area, using satellite
data. JARPA II will investigate the relationships between
oceanographic data and species distribution, including

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report of the expert workshop to review jarpa ii

cetaceans, by real-time or time series analysis. JARPA II also
intends to actively cooperate with international organisations
and projects on oceanographic surveys.’
The proposal was implemented during the first four
years of JARPA II by collecting data via XBTs, CTDs, anElectric Particle Counting and Sizing System (EPCS), andan echo sounder at the same times and areas as the visual
whale abundance survey was being conducted. Collectionof such simultaneous data is an important component ofbeing able to relate the physical and biological habitat withwhale distribution and density and ultimately to assist in
developing an understanding of the role of whales in theecosystem.
Unfortunately, this was not done for the last two years(2009/10 and 2010/11) when only TDR data were collected.
SC/F14/J20 and SC/F14/J21 provided a good initialinvestigation into the broad spatial and temporal scale
physical oceanographic characteristics within the JARPA
study area. The authors conducted a variety of data checks
using climatological worldwide data sources and thus recalibrated some data. As stated in SC/F14/J20 this level ofcalibrations is sufficient to study broad scale features in theSouthern Ocean. However, to investigate finer spatial andtemporal characteristics, which is ideal when investigating
the relationships between oceanographic data and species
distributions, it is important to collect in situ water samples
to calibrate and when necessary correct the instrument
readings; as a minimum instruments should be calibrated at
the factory once a year.
The Panel noted that in addition to the data collected by
the JARPA and JARPA II programmes, there are a number ofother sources of similar data (such as those found in world or
Japanese databases) and also associated with other available
habitat characteristics (such as bottom depth, sea ice extent,
distance from sea ice, SST, chlorophyll, and sea surface
height). Many of these databases are available online.
The Panel recognises the contribution of the longterm time series represented by the JARPA and JARPA II
programmes, particularly in the context of simultaneouscollection of whale and environmental data. However, thePanel is concerned that the necessary analytical work to
fulfil the objectives of the programme has not received theattention and resources that it deserves. If the programme isto meet its objectives in the medium to long-term, the Panelmakes the following recommendations:(1) the collection of the full suite of oceanographic data
be resumed in the coming season and the necessary
calibration work be undertaken;(2) the proponents investigate the availability and utility ofother oceanographic and related data, including data on
sea ice, archived in other Japanese or world databases
and incorporate these with the data collected from the
JARPA and JARPA II programmes to form a morecomprehensive dataset (a useful catalogue of available
data with links to many national Antarctic data centres
is available from the Antarctic Master Directory at
www.gcmd.nasa.gov/portals/amd/);(3) the proponents consider make their oceanographic data
available to other international programmes;(4) the existing TDR and EPCS data are analysed to
fully describe the datasets contents and review theirusefulness in future habitat and whale analyses; and(5) concerted efforts begin to analyse the oceanographic
and other environmental data in the light of the obtained
cetacean sightings survey and biological data (moredetails are provided under the relevant agenda items
below).

Item (1) can begin immediately. Work on Items (2)-(5)should also begin as soon as possible and the initial resultspresented within the next two-three years. Items (2) and (3)will also fulfil the aim in the original proposal to actively
cooperate with international organisations and projects
on oceanographic surveys which to date has not beenachieved. In these times of limited funds and resources, suchcooperation is important to maximise efficiency and benefit.
The Panel commends the collection of debris data, bothin the environment and in the stomachs of the whales thatcan provide valuable baseline data. Such information should
regularly (the need to do this annually or less frequently
should be evaluated) be analysed and presented to the IWC.
5. REVIEW OF RESULTS: Elucidate
temporal and spatial changes in stock
structure (SC/F14/J27-J34)
5.1 Proponents’ summary of results
Significance of the stock structure analysis under the past
JARPA is the occurrence of at least two stocks of Antarctic
minke whales in the JARPA research area, which were called‘Eastern Indian Stock’ (I-Stock) and ‘Western Pacific Stock’(P-Stock), and the mixing of the two stocks in the region
around 150°-165°E. In addition it was demonstrated that thecurrent IWC Management Areas are not applicable to thisspecies. Questions remaining were the detailed descriptionof the mixing of the two Antarctic minke whale stocks in thetransition area and the stock structure of other large whale
species, such as humpback, fin and southern right whales in
the research area.
The specific objectives of the stock structure studies
presented are to: (1) examine the stock structure hypothesis
for Antarctic minke whales proposed based on the JARPA
data using only the JARPA II data (SC/F14/J28); (2) describe
temporal and spatial mixing pattern of the I and P Stocks of
Antarctic minke whales in the transition region (SC/F14/
J29); and (3) understand genetic characteristics and stock
structure of other large whale species, such as humpback(SC/F14/J30, SC/F14/J31), fin (SC/F14/J32), and southern
right whales (SC/F14/J33) in the feeding ground.
SC/F14/J27 presented a brief summary of the genetic
analysis protocols, including sampling, used at the Instituteof Cetacean Research for the stock structure studies under
the Objective 3 of the JARPA II. In general the protocols
follow the IWC SC guidelines for DNA data quality control
for genetic studies relevant to IWC management advice(IWC, 2009).
SC/F14/J28 tested the hypothesis on stock structure
derived from the JARPA research by applying two different
kinds of the genetic markers, 338bp of the mtDNA control
region sequences (2,278 individuals) and 12 microsatellite
DNA (mtDNA) markers (2,551 individuals) only to the
JARPA II samples, making this study as an independent test.
The areal and temporal pattern of the stock mixing, which
is one of the objectives of the JARPA II, was examined indetail in a separate document (SC/F14/J29). The statisticalanalysis of heterogeneity followed a step-wise fashion:
first, for each annual survey Areas IIIE, IVW and IVE werecompared in the western sector, and Areas VE and VIW werecompared in the eastern sector. Next the yearly variation in
the western and eastern sectors was investigated. Finallya comparison between western and eastern sectors of the
JARPA II research area was conducted. This was done forfemales only, males only, and both combined. While the
mtDNA analysis detected statistically significant differences

j. cetacean res. manage. 16 (suppl.), 2015

between the western and eastern sectors for females, males,
and total samples, the microsatellite analysis did for females
and total samples. Therefore genetic results based only on
the JARPA II samples were consistent with the previous
hypothesis of at least two stocks in the research area, one
in the most western part (I Stock) and the other in the mosteastern part (P Stock). Furthermore microsatellite analyses
observed substantial yearly variation mainly within the
I Stock, especially for females. These yearly differences
could be explained either by the mixing of the I and P
Stocks in different proportions in different years in partof the western sector (i.e. SC/F14/J29), or by the sporadic
intrusion of an unknown third stock occurring in the westernpart of Area III. Unfortunately, despite the doubled numberof the microsatellite loci from the previous JARPA study as
recommended by the 2006 JARPA final review Workshop,
the level of the stock differentiation was still too low (FST
less than 0.001) to conduct genetic analysis at the individuallevel, that prevented us from conducting an analysis that
could have distinguished the above two possibilities.
SC/F14/J29 took a mathematical approach thatsimultaneously incorporated two different sources of
information, genetic and morphometric data, for estimating
longitudinal segregation of two stocks. This study was inresponse to a recommendation made during the JARPA final
review Workshop, and the model was originally presented to
the IWC SC in 2012 using only the JARPA data (Kitakado
et al., 2012). This time the model was applied to the
JARPA and JARPA II data collected between 1989/90 and2010/11 in Areas IIIE-VIW. A soft boundary was allowed
to vary by year and sex with assuming baseline stocks. A
joint likelihood function derived from the two sources
is defined for the estimation of mixing proportions. The
mixing proportion was modelled by a linear logistic modelwith stock-specific, i.e. putative I-Stock in the western areas(Areas IIIE and IVW) and P-Stock in the eastern areas(Areas VE and VIW), parameters for the two sets of data. It
was observed that the morphometric data were dominant to
the genetic data and helped convergence in the optimisation.
However, the inclusion of the morphometric data altered
the estimation results and tended to give softer, i.e. more
fluctuated, boundaries. On the whole, the result indicatedthat the spatial distribution of the two populations changed
year by year in Areas IVE and VW, which was wider andmore western than we previously proposed. It also suggested
possible sex differences in the dynamics of the mixing
pattern along the boundary.
SC/F14/J30 analysed mtDNA data from 575 humpback
whales obtained in the Antarctic during surveys of the
JARPA/JARPA II and the International Decade for Cetacean
Research/Southern Ocean Whale and Ecosystem Research(IDCR/SOWER), and from 1,057 whales from low latitude
localities of the South Pacific and eastern Indian Ocean todescribe the distribution and mixing of breeding stocks D,
E and F in the Antarctic feeding grounds. This study was
conducted to respond to a recommendation from the JARPA
review Workshop and to recommendations from the IWC
SC in 2012 (IWC, 2013a). Such information is important tointerpret abundance estimates as well as to allocate historicalcatches to stocks. This was one of the few studies that utilised
samples from both low latitude and Antarctic areas. Afterthe alignment of the sequences of the mtDNA control region
from the two data sets, the consensus 329 bp was used. The
low latitude samples were used as baselines representing‘pure’ stocks to examine distribution and mixing of these‘pure’ stocks in the Antarctic feeding area. The results were377

roughly consistent with north-south migration of Stocks D,
E and F. Western Australia whales (D) are strongly related toArea IV, while Eastern Australia whales (E1) are related toArea VW and New Caledonia (E2) and Tonga (E3) whales
were related to the feeding grounds east of 160°E. In contrast,
Cook Island (F1) and French Polynesia (F2) whales were
not related to the feeding grounds covered by the JARPA II
surveys. This paper was presented and discussed at the IWC
SC in 2013 (Pastene et al., 2013).
SC/F14/J31 analysed biopsy samples from humpback
whales obtained during surveys of the JARPA and JARPA
II up to the 2010/11 season as well as IDCR/SOWER using
14 microsatellite DNA loci in order to describe their stock
structure in the Antarctic feeding ground. The samples weredivided into four groups based on the IWC Management
Areas: III (n=93), IV (n=218), V (n=153) and VI (n=64). In
three of 37 cases of duplicate sampling, the second samples
were collected at least a day apart (1 day, 9 years, and 11
years). Heterogeneity tests were conducted for females only,
males only, and both combined. Although a few cases ofsmall temporal genetic differences were detected within theareas, major genetic differences were among the samples
from the different Areas. In addition, stronger differentiation
was seen in females than in males. Despite the increase in
the number of the loci from six in the previous paper for
the JARPA review Workshop to 14, the level of the stock
differentiation (FST=0.003) was still too low to conduct
genetic analysis at the individual level, so that further
analysis would require using samples from their breedingareas to better understand the stock structure in the feeding
grounds. Nevertheless, this microsatellite study confirmedthat humpback whales from different stocks occupied theAreas with higher differentiation in females than in males.
SC/F14/J32 analysed samples of fin whales (biopsies andresearch takes) obtained during JARPA and JARPA II using
mtDNA control region sequencing and microsatellite DNA
to describe stock structure of this species in their feeding
grounds. A total of 55 samples obtained from Area IIIE(n=6), IV (n=23), V (n=24) and VIW (n=2) was genotyped
at 16 microsatellite DNA loci and was sequenced for 479
bp of the mtDNA control region. Although the haplotype
distributions appeared to be quite different among theareas, almost all of the individuals had different haplotypes,
reducing the statistical power. Statistical comparison was
thus made by grouping the haplotypes into two categories:
those specific to each of the Areas and those shared by morethan one Area. Results of the mtDNA heterogeneity test
showed statistically significant differences among Areas
IIIE, IV and V. The microsatellite analysis also showedstatistically significant differences between Areas IV and
V. Results of the genetic analyses therefore suggested thepossibility of genetic structuring of fin whale stocks in
the JARPA II research area that corresponded to the IWC
Management Areas. Their detailed mixing patterns should
be explored with a larger sample size in the future.
SC/F14/J33 analysed biopsy samples from southern rightwhales obtained during the JARPA and JARPA II surveys up
to the 2009/10 season using both microsatellite DNA and
mtDNA markers in order to describe genetic characteristicsof the species in the Antarctic feeding ground. A total of 70
samples obtained from Areas III to V was genotyped at 14
microsatellite DNA loci and was sequenced for 430bp of
the mtDNA control region. In two cases of three duplicate
sampling, the second samples were collected in a later year (4and 8 years, respectively) in the same Area (IV), suggestingsite fidelity to feeding ground. The levels of genetic diversity

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report of the expert workshop to review jarpa ii

observed from both markers were comparable to thosereported from other southern right whale genetic studies. A
total of eight haplotypes was generated from 21 segregationsites and these haplotypes were phylogenetically separated
into two clades. These eight haplotypes were then comparedto the 37 haplotypes (273bp) reported in Patenaude et al.
(2007) that were observed in the samples obtained from the
Indo-Atlantic (Argentina and South Africa) and the IndoPacific (southern Australia and New Zealand) basins. The
comparison revealed that our two clades were the same astheir A and W clades, each of the eight haplotypes matchedto one of their haplotypes, and three were same as theirIndo-Atlantic specific haplotypes. These results suggest thatsome southern right whales undertake much longer-distanceseasonal migration between their feeding and breeding
grounds than we had previously thought and thus whales
from multiple stocks migrate to our research area in the
Antarctic feeding ground.
SC/F14/J34 summarises the information on photoidentification of blue, southern right and humpback whales
obtained during the JARPA/JARPA II surveys in Antarctic
Areas IIIE, IV, V and VIW in the austral summer seasons.
A total of 3,108 photo-identification pictures selected fromall of the pictures obtained between 1989/90 and 2010/11
were deposited in the photo-identification catalogue
administrated by the Institute of Cetacean Research. All of
the deposited pictures were digitised to facilitate archiving.
The JARPA pictures of blue whales were submitted to
the IWC Secretariat to facilitate comparisons with othercatalogues. Similarly, the JARPA pictures of humpback andsouthern right whales were provided to the IWC’s Antarctic
Humpback Whale Catalogue and to other international
research organisations, respectively, under co-operation
studies. A preliminary examination of the pictures within
the feeding grounds as well as between the feeding grounds
and lower latitude localities resulted in several matches.
The JARPA and JARPA II photo-identification data havethe potential to contribute to the better understanding of
the movement, distribution and abundance, and in turn tothe assessment and conservation of blue, southern right andhumpback whales.
In summary, these studies demonstrated that thecomprehensive large-scale JARPA II (as well as JARPA)
survey effectively used a combination of lethal and nonlethal techniques to acquire valuable information for
effective management of large whales in the Antarctic.
The studies implied that different species require different
management strategies on an area by area basis even within
the same feeding grounds for their long-term persistence.
The stock structure results are one of the key information
for the RMP process. The JARPA results were refined under
the JARPA II and this information can be used in the context
of the current RMP process as well as in the context of the
future improvement of the RMP. We believe that our studies
properly addressed and answered all of the objectives weraised and the recommendations from previous reviews that
required a response.
5.2 Panel’s conclusions and recommendations
This section deals with Objective 3 of JARPA II: Elucidationof temporal and spatial changes in stock structure. In addition
to a general paper describing genetic laboratory methodology(SC/F14/J27) and a summary overview paper (Pastene, 2006),
the Proponents presented two genetic papers on minke whales(SC/F14/J28, SC/F14/J29, with the latter also including
morphometric data), two genetic papers on humpback

whales (SC/F14/J30, SC/F14/J31), one genetic paper eachon fin whales (SC/F14/J32) and southern right whales (SC/F14/J33), and a summary of photo-ID information for blue,
southern right, and humpback whales (SC/F14/J34).
The Panel noted that knowledge of stock structure isfundamentally important not only to this objective of the
programme but to all of the objectives. The Panel commends
the considerable field and laboratory effort that had gone intodeveloping a comprehensive long-term dataset, particularly
for the Antarctic minke whales. It also commends the efforts
to develop analytical approaches to incorporate genetic andother markers in an integrated analysis (SC/F14/J29).
5.2.1 Scope of the studies
The Panel agrees that information resulting from the JARPA
and JARPA II programmes has considerably increased ourunderstanding of stock structure within the research area,
which directly addresses Objective 3. However, the Panelnoted that Antarctic minke whales and other baleen whale
species are more-or-less continuously distributed aroundthe Antarctic continent; in contrast, the JARPA II researcharea represents just under half of the circumpolar area.
Given the stated objective, the lack of information provided
for areas outside the programmes’ research area presents
some inherent difficulties in fully meeting the objective to
elucidate spatial and temporal variations in stock structure,even though the information developed under JARPA II isprobably sufficient for the purposes of developing trials toevaluate RMP variants within the area of sampling. Thisissue is discussed in more detail below with respect toAntarctic minke whales.
5.2.2 Genetic data
The Panel welcomes the fact that the authors of SC/F14/J27
referenced the IWC guidelines for DNA data quality control(IWC, 2009) as had also been recommended at the JARPN
II review (IWC, 2010). However, the Panel recommends
that a revised paper be submitted that explains in moredetail how far the guidelines were able to be followed (thepresent paper did note why the proponents had not beenable to follow one recommendation regarding sequencing of
microsatellite loci).
5.2.3 Antarctic minke whales
SC/F14/J28 described new mtDNA and microsatellite
genetic analyses of minke whales. The Panel welcomes thefact that, in response to comments from the JARPA review
Workshop (IWC, 2008), the authors had doubled the numberof microsatellite markers, sequenced a portion of the mtDNA
molecule, and performed additional analyses, including useof the clustering program STRUCTURE (Pritchard et al.,2000) to evaluate evidence for more than one gene pool.
This paper excluded samples from the putative mixing
area, which were considered in SC/F14/J29. The Panelacknowledges the extensive laboratory and analytical work
behind the paper. However, there are a number of places
where additional information is required to fully interpretthe results. For example, if the authors’ hypothesis iscorrect that different proportions of stocks I and P mix inpart of Area IV, then the FIS values for that area should on
average be positive, reflecting a deficiency of heterozygotes.
Furthermore, the loci showing the most positive FIS values
should be the loci having the highest FST values among
the two putative stocks (Waples, 2011). If the two-Stockswith-mixing hypothesis is true, then the analysis suggested
above should also produce a positive correlation between
FIS and FST for samples taken from the area of putative

j. cetacean res. manage. 16 (suppl.), 2015

mixing that were not considered in this analysis. Similarly,
the magnitude of linkage disequilibrium at pairs of locishould be proportional to the product of FST at the two lociinvolved (Waples, 2011). The mean FST estimated between
the two stocks in this case is quite small (<0.001), so these
correlations might be hard to detect. If these analyses are
conducted, it would be reasonable to start by evaluating only
females, which show the strongest signal of differentiation.
The Panel makes the following detailed recommendations concerning SC/F14/J28:(1) samples from the putative pure I and P Stocks havestatistically significant differences in allele frequencies– a revised paper should provide the locus-specific FSTor theta values for these comparisons to identify whichloci are responsible for the differences;(2) Table 4 presents evidence for overall deviations from
Hardy-Weinberg (HW) expectations in both putative
stocks – a revised paper should include FIS values foreach locus in each stock to provide information about
the direction and magnitude of HW deviations and thus
the biological importance of this result; and(3) the discussion section should be expanded to consider
the results of the analyses suggested above.
SC/F14/J29 provides an innovative approach to integrate
multiple sources of data to distinguish stock structure andidentify time-varying distribution patterns by maturity class.
As noted below, the method could be extended to make useof other data sources such as stable isotope ratios of C, N
and O, and hence provide a way to synthesise much of theinformation on stock structure collected during JARPA/
JARPA II. Stable isotope ratios of these elements have beenused to assess population structure and degree of mixing
in other cetacean species with reasonable success (Born etal., 2003; Giménez et al., 2013; Vighi et al., 2014) and, in
addition, this method has the potential to assess the valueof data collected using lethal methods (in this case, themorphometric data).
The Panel welcomes this effort to integrate genetic
and non-genetic data, noting that this is in accord with the
Scientific Committee’s view that stock structure can best be
addressed using a suite of techniques and data types. The
Committee has recognised that developing a quantitative
integrative approach is complex. In terms of developinga revised paper in time for the forthcoming ScientificCommittee meeting, the Panel makes the following
recommendations:(1) notation in the paper should be improved to avoid using
the same index for individual and stock;(2) the estimates of the parameters and their standard errors
should be reported – this information could be used to
identify which of the morphometric measurements ismost informative regarding stock mixing;(3) diagnostic statistics for the fits to the morphometric data
should be presented;(4) the meaning of the statement: ‘The variables G and M
are further assumed independent within an individual,
and also between individuals’ should be clarified – it is
not clear whether this means that matrix G is assumed
independent of matrix M, or that the different elements of
M are assumed to be independent (i.e. is the Σ diagonal?);(5) the parameters for each model should be documented toclarify how many of the parameters of the mean function
and variance-covariance matrix (morphometric data) as
well as the baseline allele frequencies (genetic data) are
estimated parameters; and379(6) the discussion should explore the fact that one of themain results of this study - that morphometric data had
a stronger influence on results than the genetic data
- differs from that found in many other such studies,and the authors should suggest a possible biologicalexplanation for this result.
In the longer-term, the Panel recommends that:(1) the model be formulated as a random effects model(Bayesian or maximum likelihood). This may eliminate
some of the problems associated with lack of convergencefor some of the more complicated models – this mightalso reduce some of the large inter-annual fluctuations
in mixing proportions, which a plot produced during theWorkshop (Fig. 3) shows are generally very imprecise;(2) more flexible functions for the relationship between
longitude and proportion should be considered; and(3) the benefits of applying the integrated model to all data
(i.e. data aggregated by sex and maturity state) should
be re-evaluated since, for example, there are clear
between-sex differences in some of the morphometricmeasurements such as V7 (fig. 3 of SC/F14/J29) a model with sex-specific values for the mean andvariance functions for the morphometric data, but
sex-independent distribution proportions should be
explored.
Management procedure considerations on stock structure
focus on developing plausible interpretations of available
data not simply the single ‘best’ interpretation when
examining uncertainty. In this spirit, the Panel recommends
that during the coming year, the authors of SC/F14/J28 and
SC/F14/J29 consider the merits of an alternative to the twoStocks-with-mixing hypothesis: a single stock that exhibits
one-dimensional isolation by distance along a longitudinal
gradient. This alternative is suggested by visual inspectionof fig. 3 in SC/F14/J29, which shows how morphometricscores for individual whales vary by longitude. Morphology
appears to vary more or less linearly along the zone ofsampling, rather than being constant at the eastern andwestern extremes, with an area of mixing in between. Under
isolation-by-distance, statistical comparisons of samples
from the extreme eastern and western sampling regions
could produce the types of results described in SC/F14/J28(Schwartz and McKelvey, 2009).
The genetic analyses of isolation-by-distance could use
the statistic (a) for comparing genotypes of individuals
suggested by Rousset (2000). With isolation-by-distance, alinear relationship is expected between a and ln(distance),
where distance is measured by longitude. It should bepossible to modify the genetic/morphometric model in SC/
F14/J29 to evaluate a scenario in which morphology varies
along a cline. Conceptually, the study system could beconsidered to correspond to a narrow ribbon, either joined
end to end or not. If samples were available from 360
degrees of longitude rather than just under half that, it would
be possible to determine whether the opposite ends of theribbon are joined (hence forming a type of flattened torus) orremain separate. Under the latter scenario, Antarctic minke
whales would form a type of ring species (Irwin et al.,2005), with extreme forms being reproductively isolated,but connected by gene flow along a gradient. These two
scenarios might not imply that different RMP variants would
be selected for the portion of the range that has been sampled,but they would have profoundly different implications forunderstanding of the biology of the species, and that seems
to be broadly consistent with JARPA II Objective 3.

380report of the expert workshop to review jarpa ii

Fig. 3. Point estimate and approximate 90% confidence interval for the estimation of ‘area-wise mixing proportion’based on ‘genetic and morphometric data’.

Stock-structure Archetype III in IWC (2006) illustrates
an isolation-by-distance scenario of the kind suggested
here. In the present case, application and interpretation
will be complicated by the fact that minke whales are
migratory, and the genetic data are collected on the feeding
grounds rather than the breeding grounds. Furthermore, themagnitude of genetic differentiation is small. Nevertheless,the Panel agrees that evaluating whether the empiricalgenetic and morphometric data are consistent with anisolation-by-distance model in the area of mixing would
provide valuable information. One approach that might beable to distinguish between the two-Stocks-with-mixing
and isolation-by-distance hypotheses is to evaluate how
the inbreeding index (FIS) changes with longitude. Under
two-Stocks-with-mixing, FIS should be zero (on average)in the pure-Stock areas (indicating agreement with HardyWeinberg equilibrium within stocks) and positive in thearea of mixing (indicating a deficiency of heterozygotes
due to the Wahlund effect). Under the isolation-by-distancehypothesis, the value of FIS should be slightly positive andshould not change with longitude.
Pastene (2006 provided a useful historical perspective
and summary of hypotheses and results of studies regarding
stock structure of Antarctic minke whales. This summary wasparticularly valuable as it included discussion of a numberof potential stock-structure indicators in addition to genetics,
including tagging data, pollution studies, ecological markers,
morphometrics, size at physical maturity, and sighting andcatch distributions. However, some types of informationcollected by JARPA II (such as oceanographic data) were

not integrated into the analysis. The Panel recommends that
the Proponents attempt to collect/incorporate other types of
information that were not discussed but which potentiallycould be important to elucidate temporal and geographicalchanges in minke whale stock structure, including building
upon recent developments in satellite tags (the Panel
recognises the practical difficulties but believes that thegains could be important for a number of topics under the
programme in addition to stock structure), stable isotopes ofC, N and O, and fatty acids (IWC, 2014a).
5.2.4 Humpback whales
SC/F14/J30 examined mtDNA in samples (primarily frombiopsies from JARPA, JARPA II, and SOWER surveys) of
humpback whales from Antarctica and lower latitudes. This
study was conducted to respond to a recommendation fromthe JARPA review (IWC, 2008) and to recommendations
from the Scientific Committee to study distribution andcalculate mixing proportions of breeding stocks D, E and F
in the feeding grounds (IWC, 2013a). The Panel welcomes
these results, first presented as Pastene et al. (2013) at the2013 meeting of the Scientific Committee, which provide
valuable information about distribution of humpback whaleson the Antarctic feeding grounds.
SC/F14/J31 reported results for microsatellite DNA
analysis of the Antarctic samples (but not the lower latitude
samples) of humpback whales analysed for mtNDA in SC/
F14/J30. The authors increased the number of loci from 6 to
14 in response to the recommendation of the JARPA review
Workshop (IWC, 2008). Table 3 of SC/F14/J31 shows that381

j. cetacean res. manage. 16 (suppl.), 2015

each sample was in agreement with overall Hardy-Weinberg(HW) equilibrium, but no information was given in the text
about the results of these tests or how they varied amongloci. Other tables show p-values for heterogeneity tests
comparing samples from different areas or times. The Panelrecommends that an updated paper be submitted in which
the tables also include the effect size (e.g. FST values).
5.2.5 Fin whales
SC/F14/J32 reported results of genetic analysis of fin whales
using mtDNA and microsatellites. The total number ofsamples available across all years and sectors (n=52) wasmuch fewer than planned, and this limits any conclusions
about stock structure. The study incorporated biopsy
samples from the JARPA programmes as well as captured
animals. Standard tests of heterogeneity found no evidence
for significant spatial differences in mtDNA, but the authors
did find significant differences when they conducted an
additional analysis that re-sorted the haplotypes into groups
that are shared among regions and those that are not.
While welcoming the new information on fin whale
stock structure - a subject that has not been examined by
the Scientific Committee for many years (Donovan, 1991)- the Panel was concerned at some aspects of the analyses
and interpretation. It therefore recommends that an updated
paper be provided that:(1) omits the analysis using post-hoc groupings unless theauthors can provide a robust statistical justification forthis approach;(2) discusses in more detail the fact that three microsatellite
loci showed significant departures from Hardy-Weinberg(HW) expectations after accounting for multiple testing
before the data are used in other analyses;(3) provides FIS values for each locus in table 7 of SC/F14/
J32, which would indicate whether the HW departures
represent an excess or deficiency of heterozygotes;(that information is essential to understand the possible
causes for the deviations);(4) shows how the samples were arranged into Management
Areas in fig.1; and(5) recognises that conclusions about possible differences
between Areas IV and V are provisional until thenature, cause, and influence of these HW deviations areresolved.
In the longer term, in order to increase sample sizes the
proponents should examine the availability of historic tissues
that may be used for genetic studies (e.g. from commercial
whaling or biopsy samples from other research cruises).
5.2.6 Southern right whales
SC/F14/J33 reported results of genetic analysis of biopsy
samples from 70 southern right whales, with most (n=63)
being taken in Area IV reflecting their distribution within
the research area as searching effort was made throughout
the research area; mtDNA sequences were compared withthose reported in previous studies to allow comparisonsacross studies. As expected, most haplotypes in the present
study matched to those previously reported from the IndoPacific, but three haplotypes appear to be of Indo-Atlantic
origin, which suggests that at least some southern rightwhales might make longer migrations than previously
thought. Although the sampling design provided limited
power to detect geographic variation throughout the wholeof Antarctica, the Panel welcomes this study that represents
the first attempt to genetically characterise southern rightwhales feeding in Antarctica.

5.2.7 Photo-identification data
SC/F14/J34 summarised photo-ID information containedin over 3,000 images of humpback, blue, and southern
right whales taken during JARPA and JARPA II cruises.
The photos were taken opportunistically and SC/F14/J34
reported some preliminary matches.
The Panel welcomes the effort that has been put into thecollection of photographs for photo-ID. It also welcomes
the submission of photographs to relevant international
catalogues. It recommends that further effort is put into
analysing these data in conjunction with these other
Southern Hemisphere catalogues, recognising their potential
to provide information on stock structure, mixing andmovements. It expects that such work could be presented to
the Scientific Committee within the next two years.
6. REVIEW OF RESULTS: Monitoring whale
abundance trends (SC/F14/J03-J07, SC/F14/
J17-J18; SC/F15/O01-O02)
6.1 Proponents’ summary of results
Based on sighting data by JARPA and JARPA II, abundancesof Antarctic minke, humpback, fin, blue, southern right,
sperm, southern bottlenose and killer whales were estimated.
There are good agreements between IDCR/SOWER
abundance estimates and JARPA/JARPA II abundance
estimates for Antarctic minke and humpback whales.
Abundance trends obtained from JARPA and JARPA II arealso generally consistent with biological and demographic
information. This reconfirms the scientific value of thesighting data during JARPA and JARPA II.
6.1.1 Antarctic minke whales
SC/F14/J03 estimated abundance and abundance trend of
Antarctic minke whale based on sighting data collected by
JARPA and JARPA II (1989/90-2008/09 seasons). These
analyses addressed most of the previous recommendations
from the JARPA review Workshop in 2006 (see Annex 4of SC/F14/J01). Abundance estimates were based on thestandard line transect method using the program DISTANCE
under the assumption of g(0)=1. The annual rates of increase
were estimated using log-linear models. Model error was[Text
1]
taken table
into considerationin the estimates. Results:
Average

Minimum

Maximum

Area

Estimate

Estimate

CV

Estimate

CV

IIIE
IV
V
VIW

18,75932,714
101,106
15,486

4,478
15,08867,661
8,434

0.911
0.645
0.308
0.601

48,54063,794
151,072
27,790

0.711
0.509
0.326
0.507

Estimates of annual rates of increase were 1.1% (95%
CI: -2.3, 4.5%) for Areas IIIE+IV and 0.6% (95% CI-2.2%, 3.3%) for Areas V+VIW. Adjustments to allow
for g(0) being less than 1 were made by the applicationof a regression model developed from the results of the
Okamura-Kitakado (OK) method estimate of Antarctic
minke whale abundance from IDCR-SOWER surveys,
which provides estimates of g(0) from the statistics of theminke whale school size distribution in a stratum. With thisadjustment, abundance estimates increased by an average of
88.0% for Area IV and 109.0% for Area V. The annual rates
of increase changed to 2.5% (95%CI-1.3, 6.3%) for Area
IIIE+IV and -0.6% (95%CI-3.9, 2.6%) for Area V+VIW,
which are not substantially different from the base case. It382

report of the expert workshop to review jarpa ii

is suggested that there is good agreement between IDCRSOWER abundance estimates and the abundance estimateswith the g(0) adjustment in Areas IV and V.
6.1.2 Humpback whales
SC/F14/J04 estimated abundance and abundance trend of
humpback whale based on sighting data collected by the
JARPA and the JARPA II (1989/90-2008-09 seasons). These
analyses addressed most of the previous recommendations
from the JARPA final review Workshop in 2006 (see Annex
4 of SC/F14/J01). Abundance estimates were based on thestandard line transect method using the program DISTANCE
under the assumption of g(0)=1. The annual rates of increase[Text table 2]
were estimated using log-linear models. Results:
First year

Last year

Area

Year

Estimate

CV

Year

Estimate

CV

IIIE
IV
V
VIW

1995/96
1989/90
1990/91
1996/97

1,378
5,325
602
1,493

0.190
0.302
0.343
0.185

2007/08
2007/08
2008/09
2008/09

11,90429,067
13,894
3,609

0.261
0.255
0.338
0.322

Estimates of annual rates of increase in Areas IIIE, IV, V and
VIW are 20.8% (95% CI:4.1, 37.6%), 13.6% (95% CI8.4,
18.7%), 14.5% (95% CI7.6, 21.5%) and 6.2% (95% CI-0.9,
13.4%), respectively. Except for Area VIW all the trend
estimates were statistically significant. The JARPA+JARPA
II results thus indicate that humpback whales in Area IV
and Area V are definitely increasing rapidly. There is good
agreement abundance estimates based on JARPA+JARPA II
and those based on IDCR-SOWER in Areas IV and V.
6.1.3 Blue, fin and southern right whales
SC/F14/J05 estimated abundance and abundance trendsof blue, fin and southern right whales based on sighting
data collected by JARPA+JARPA II (1989/90-2008/09).
The estimates are calculated by standard line transect
analysis methods using the program DISTANCE under theassumptionthat g(0)=1. The annual rates of increase were
[Text table 3]
estimated using regression model. Results:
Start –Start season
latestAbund.
CV
seasons

Latest season
CV

Abundance
trend

95% 95%
CILL CIUL

0.345

0.082

0.039 0.125

Fin whale (Indian Ocean stock), Areas IIIE-IV
1995/96- 3,087 0.191 2,610 0.285
0.089
2007/08-0.145 0.324

Fin whale (western S Pacific stock), Areas V+VIW
1996/97- 1,879 0.226 14,981 0.298
0.120
2008/09

0.026 0.215

Southern right whale, Area IV
1989/9042
1.305 1,557
2007/08-0.164 0.281

Abund.

Blue whale, Areas IIIE+IV+V+VIW
1995/962008/09

300

0.308

1,223

0.303

0.059

The abundance trend estimate for blue whales was
statistically significant. The abundance trend estimate forfin whales in Areas V+VIW was statistically significant.
Changes in whale species composition were also confirmed
in Area IV.
6.1.4 Odontocetes
SC/F14/J06 estimated abundance and abundance trendsof sperm, southern bottlenose and killer whales based onsighting data collected by JARPA+JARPA II (1989/902008/09). The estimates are calculated by standard linetransect analysis methods using the program DISTANCE
under
the 4]
assumption that g(0)=1. The annual rates of[Text table
increase were estimated using regression model. Results:
Minimum

Maximum

Abundance trend

Area Estimate

CV Estimate

CV

Estimate 95%CILL 95%CIUL

Sperm whale
IIIE
1,181
IV
1,358
V
597
VIW
101

0.197 2,309
0.192 4,106
0.509 2,185
1.008
736

0.199
0.255
0.254
0.214

0.021
-0.024-0.0390.066-0.041-0.062-0.082-0.090

0.082
0.013
0.005
0.223

Southern bottlenose
IIIE
866 0.569 7,364
IV
1,082 0.266 11,828V
893 0.447 6,729
VIW
908 0.396 2,681

0.383
0.266
0.269
0.233

-0.018
0.104
0.077
0.009

-0.2250.030
0.013-0.099

0.190
0.179
0.142
0.117

Killer whale
IIIE
524
IV
2,268
V
7,184
VIW
943

1.157
0.185
0.277
0.350

0.084
-0.021-0.011
0.043-0.096-0.087-0.048-0.120

0.264
0.044
0.026
0.206

1.036 6,300
0.489 14,374
0.356 18,867
0.593 4,066

Abundance trend are not statistically significant except
those of southern bottlenose in Area IV and V. Abundance
estimates for sperm, southern bottlenose whales are considered
to be underestimated due to assumption that g(0)=1.
6.1.5 Paternity analysis
SC/F14/J07 used a mark-recapture method with paternity
analysis based on microsatellite DNA data from the JARPA
and JARPA II samples of Antarctic minke whales toestimate their stock size and movement pattern. Throughout
the paternity analysis, genotypic data of maximum 12
microsatellite DNA loci examined from 137 foetuses thatwere collected from the pregnant females captured during the
2003/04 JARPA were used to look for their potential fathers
among 1,779 males collected from the 2001/02 JARPA to2010/11 JARPA II. One case of matching among the foetusmother-father trio was found. The assigned father’s bodylength was 8.66m and the estimated age was 12 years old,which did not necessarily, deny the match. This matching
was used to conduct preliminary estimate the mature maleabundance of the I Stock using Petersen mark-recapture
method modified by Chapman.
6.1.6 Distribution
SC/F14/J17 examined the distribution pattern of the Density
Index (DIW: number of whales sighted/100 n.miles) ofblue, fin, sei, Antarctic minke, humpback, southern right,
sperm, southern bottlenose, Ziphiidae and killer whalesin the Antarctic based on the JARPA (1987/88-2004/05)
and JARPA II (2005/06-2008/09) sighting data. A total of353,134 n.miles was surveyed in Areas IIIE, IV, V and VIW,
mainlysouth5]of 60°S. The following table summarises the[Text tablesighting information:
All Areas (IIIE, IV, V and VIW; south of 60°S,
35°E-145°W
Species

Sch.

Ind.

Calf

Mss

DIS

Order
of
DIW DIWBlue whale
286
495
11
1.73
0.081 0.140
8
Fin whale
1,268 5,209 20
4.11
0.359 1.475
5
Sei whale
36
59
0
1.64
0.010 0.017 10Ant. minke
25,507 69,076 1
2.71
7.223 19.561 1
Humpback
10,036 18,770 137 1.87
2.842 5.315
3Southern right
235
298
6
1.27
0.067 0.084
9Sperm whale
3,810 3,926
0
1.03
0.079 1.112
6
S bottlenose
1,666 3,045
31.83
0.472 0.862
7Unid. beaked
3,175 5,457
3
1.72
0.899 1.545
4
H:\SkyDrive\Documents\AC
Supplement
JARPA\Rep02
Killer whale
1,472 20,569
59 16\Rep02
13.97 -0.417
5.825 Tabs.docx
2

15 Januaj. cetacean res. manage. 16 (suppl.), 2015

Among 10 whale species, Antarctic minke whales
were the most frequently sighted species, followed by thekiller, humpback, unidentified beaked, fin, sperm, southernbottlenose, blue, southern right and sei whales in the DIW
order. Maps of the DIW by 1° x 1° square are provided.
These large amounts of data collected by JARPA and JARPA
II including the current maps will contribute to investigating
habitat utilisation of each species and provide more useful
information for whale management in the Antarctic andAntarctic marine ecosystem.
SC/F14/J18 revealed that the spatial distribution of
humpback whales was expanded in Area IV in the periodof JARPA and JARPA II (1989/90-2005/06) while that of
Antarctic minke whale remained stable. The results indicatedthat competition between humpback and Antarctic minke
whales for habitat in Area IV during the period of JARPA
and JARPA II was intensified as abundance of humpback
whales increased (SC/F14/J04). The competition could bereflected in the decline of the energy storage and stomach
contents weight of Antarctic minke whales (SC/F14/J13; SC/
F14/J14). Data obtained in JARPA and JARPA II were usedin SC/F14/J18 and were divided into three periods: early(1989/90, 1991/92 and 1993/94), middle (1995/96, 1997/98
and 1999/2000) and late (2001/02, 2003/04 and 2005/06).
Spatial distribution was estimated using generalised additivemodels (GAM). The presence or absence of whales wasused as a response variable while seafloor depth, distancefrom shelf break and longitude were used as explanatory
variables.
Mean probabilities of the occurrence of Antarctic
minke whales in the survey area in early, middle and lateperiods were 0.41, 0.46 and 0.41 while those of humpback
whales were 0.14, 0.35 and 0.46. Occupied area indices(probabilities of the occurrence of Antarctic minke whales
minus probabilities of the occurrence of humpback whales)
were also calculated. If the index was 1, only Antarctic minke
whales were present in a grid cell while only humpback
whales were present if the index was -1. If the index was0, probabilities of the presence of Antarctic minke whales
and humpback whales in a grid cell were identical. Mean
occupied area indices in early, middle and late periods were0.28, 0.11 and -0.07, respectively.
6.1.7 Summary
In summary, the abundance trend for the Antarctic minke
whales was broadly stable in both Areas IIIE+IV and
V+VIW in the JARPA/JARPA II period. There was a good
agreement between IDCR-SOWER and JARPA/JARPA II
abundance estimates in Areas IV and V. Abundance of blue,
humpback and fin whales are increasing in the JARPA/
JARPA II period. The abundance of southern right whales isincreasing but the trend is not significant. There was a good
agreement between IDCR-SOWER and JARPA/JARPA II
abundance estimates for humpback whales in Areas IV and
V. A significant increasing trend was found for southernbottlenose whales in Areas IV and V.
It should be noted that the increasing trend of blue whaleand fin (Areas V+VIW) whales was not significant by using
JARPA data only. They became significant with the additionof new data from JARPA II.
Whale species composition was changing gradually,
especially in Area IV. A temporal shift in species compositionand spatial distribution of humpback whales was observed in
Area IV, which was not observed in Area V (Area V includes
the Ross Sea where humpback whales do not distribute).

383

6.2 Author’s summary of the SCAA analysis
Punt has been undertaking work on statistical catch-at-age
analysis (SCAA) on behalf of the Scientific Committee formany years as described in SC/F14/O01, using age datafrom commercial catches, JARPA and JARPA II as well as
abundance estimates from IDCR/SOWER and the JARPA
programmes.
SC/F14/O02 applied statistical catch-at-age analysis
to data for Antarctic minke whales. The SCAA model isspatially-structured, can model multiple stocks of minke
whales, and can utilise several data types for parameter
estimation. The application to Antarctic minke whales
considers two stocks (I and P) in five areas which cover
Antarctic Areas III-E to IV-W. The parameters of the model(annual deviations about the stock-recruitment relationship,changes over time in carrying capacity, density-dependence
parameters (productivity and carrying capacity), and theparameters which determine growth by stock, age-specificnatural mortality by stock, and vulnerability by area and‘fleet’) are estimated by fitting the model to data on catches,
catch-at-length, conditional age-at-length, and estimates ofabsolute and relative abundance. A reference case analysis
is selected, and sensitivity explored using retrospective
analyses and by varying the assumptions on which thereference case analysis is based. The reference case analysis
is able to mimic all of the data sources adequately. Most of theanalyses (reference and sensitivity) indicate that Antarctic
minke whales in the assessed area increased from 1930 until
the mid-1970s and have declined thereafter, with the extent
of the decline greater for minke whales in Antarctic Areas
III-E to V-W than for those further east. Natural mortality
is consistently estimated to be higher for younger and older
individuals. Estimates of MSYR1+ are presented, but are
unreliable owing to the lack of contrast.
6.3 Panel’s conclusions and recommendations
6.3.1 Abundance and trends from surveys
The Panel welcomes the presentation of results fromthe JARPA and JARPA II programmes. This represents
a considerable expenditure of research time and a large
dataset for long-term monitoring. It complements the workof the IDCR/SOWER programme that has now finished.
The importance of monitoring trends in abundance forcetacean species is of general conservation and management
interest as well as providing information relevant to the
JARPA II programme objectives and especially as input toecosystem models. It is particularly important in the context
of documenting the recovery of species/populations that had
been extensively depleted by commercial whaling, as wellas investigating variation in species mix compared to theperiod prior to exploitation.
The JARPA review meeting (IWC, 2008) had
examined the survey information available at that timeand had identified a number of items that required further
investigation before acceptable abundance estimates could
be obtained. The Panel agrees that the papers on Antarctic
minke whales (SC/F14/J03) and humpback whales (SC/F14/J04) had adequately addressed most of these concernsand had in some cases identified further work that would beundertaken - see the discussion in Matsuoka et al. (2011)
and Hakamada et al. (2013). It requested clarification of thetreatment of additional variance in the estimation of rate ofincrease and further consideration as to the approach used.
Given the importance of g(0) estimation to the Antarctic
minke whale abundance estimation (IWC, 2013a) and thenecessarily ad hoc approach used in SC/F14/J03 to account384

report of the expert workshop to review jarpa ii

for this in the JARPA and JARPA II programmes where theindependent observer approach had not been implemented,
the Panel recommends that consideration be given to using
an independent observer approach in future surveys.
The papers on the other baleen whale species (SC/F14/J06) had been unable to follow the same analyticalprocedure for some aspects of the abundance estimation due
to smaller sample sizes than were available for the minke
and humpback whale cases, particularly in that they wereunable to obtain separate estimates for sightings vessels only
(as opposed to combined with sightings and sampling vesseldata) only for part of the period.
It is important to remember that the estimates ofabundance and trends presented in the papers to theWorkshop represent numbers (and trends) of animals within
particular geographic areas at particular times of the year.
In some cases, such as Antarctic minke whales, humpback
whales and blue whales, this probably covers the peak of
their distribution in the austral summer. However, for otherssuch as fin whales, this is not the case based on their past
distributions. In perhaps the most extreme case of sperm
whales, only large males penetrate south of 60°S. While
the authors note these issues, the Panel reiterates thatconsiderable care should be given to their interpretation.
In all cases, if the results are intended to provide inferencesof population trends, then careful consideration needs tobe given to the questions of stock structure; this can beparticularly complex for cases/areas where separate breeding
stocks overlap on the feeding grounds. While this is covered
briefly in the papers presented, these matters need to beconsidered more thoroughly by the authors in the discussionsections of their papers.
Recognising the value of these datasets and welcoming
the analyses presented, the Panel recommends that a revised
stand-alone paper for the ‘other’ baleen whale species (i.e.
not the Antarctic minke and humpback whales – these havealready been published) is developed (and submitted for
publication) that:(1) presents a more thorough description of methods andassumptions used (bringing in some of the information
presented in other papers such as SC/F14/O02):(2) highlights the issues related to small sample size and therecommendations made at the JARPA review meeting,including those relating to use of the SSV data and thetreatment of non-surveyed areas);(3) identifies more clearly the differences between ideal
and realised tracklines by year including percentage
achieved coverage;(4) explains more fully the treatment of additional variancecomponents in the estimation of rates of increase;(5) includes more extensive discussion than at present of
the results including interpretation of the results withrespect to stock structure and proportion of range
covered, the implications of the sensitivity results on
the confidence intervals obtained within the ‘base case’,the merits of the inclusion or exclusion of abundance
estimates that rely on considerable extrapolation within
the trend estimation process; and(6) includes an updated power analysis of the effectsof survey interval and estimation of trend to informconsideration of levels of effort and survey design inthe future.
With respect to the toothed whale estimates, there aresome additional issues that arise in the context of g(0),which is considerably more difficult to estimate for sperm

and bottlenosed whales given their diving behaviour. The
authors need to consider, given these issues and the lack ofknowledge of the full distribution of these species, whether
the production of absolute abundance estimates is feasible
or valuable. The Panel also recommends the collection ofdata on the ecotype of killer whales to try to allow estimatesof abundance to be developed for each. This information isof importance to ecosystem modelling given their different
feeding habits and in particular to evaluate the consequencesof predation on minke whales.
6.3.2 SCAA analyses
The Panel welcomes the updated SCAA analysis presented.
It agrees that the SCAA model is both the best currently
available model for examining stock dynamics for the
Antarctic minke whales in the JARPA II area, and that themodel performs well in this regard. There are however anumber of recommendations for improvements and further
work, which are discussed below. The Panel also noted thatcertain results from the SCAA model may not be consistent
with inferences developed from other components of JARPA
II or may suggest potential revisions to the design of JARPA
II itself. These points are also outlined below, and should beconsidered by the proponents.
Abundance estimates

The model is currently tuned to the abundance estimatesfrom JARPA/JARPA II that were available in late 2013.
These should be replaced by the g(0)-corrected estimatesin SC/F14/J03. Given the comparison of estimates from
JARPA and IDCR in SC/F14/J03, model configurations inwhich the JARPA/JARPA II abundance estimates are treatedas absolute measures of abundance should be exploredfurther.
Random effects model

The model uses a random effects structure implemented as
errors-in-variables to estimate year effects in recruitment anda number of other parameters (vulnerabilities, proportion ofeach stock in each area, growth rate and carrying capacity,
SC/F14/O02, appendix E). The functional forms andstandard deviations of these random effects are set externallyto the model based on expert judgement. If these standard
deviations are too small, there is a risk of overly smoothing
the deviations between years, and hence underestimating
the confidence limits. Following a request from the Panel, atest was run to re-estimate recruitments with an essentiallyunconstrained distribution (standard deviation of log
recruitment deviations of 10). The recruitment estimates
resulting from this run were plotted against those estimatedwith the standard deviation used in the base case (seeFig. 3). The linear nature of the scatterplot indicates thatthe estimation was not affected by the choice of standard
deviation. The Panel recommends that similar tests shouldbe performed for the other parameters with annual deviations
to check that the specified standard deviations are not overly
constraining.
MSYR

As the author noted, the variability in MSYR in the sensitivity
tests (SC/F14/O02, table 8) indicates that the model is notable to reliably estimate MSYR. It appears that there is little
contrast in the data required to estimate MSYR. Previous
versions of the model had given MSYR estimates. However
this may have been an artefact of the relatively inflexible
nature of model structure. The Panel recommends that theability to estimate MSYR should be revisited in the future inlight of the availability of new data.

j. cetacean res. manage. 16 (suppl.), 2015

385

Natural mortality model

The piecewise model characterising the relationshipbetween natural mortality and age is, in some respects,
sub-optimal. The need to externally define the breakpoints
raises the risk of introducing model mis-specification. Tests
with the autoregressive function produce a similar mortality
function, and it seems possible that the Siler model would
also give similar results. If so, the Panel recommends
using the Siler function as this will reduce the risk ofmodel mis-specification and be more parsimonious than the
autoregressive function. If the Siler function does not matchthe autoregressive function, then it would be preferable toadjust the breakpoints in the piecewise model so that themodel gives a better (but still parsimonious) approximation
to the autoregressive solution.
The Panel also recommends that sensitivity tests
are conducted to examine the response of the modelled
population to changes in the mean M for both stocks whenthese are assumed to be the same.
Stock Structure

The Panel noted that the model is currently configured
with two non-overlapping stocks. However, the model has
the capability to incorporate a wide variety of different
possible stock structures, and recent results from the JARPA
programmes (see Item 5) suggest possible different stock
structures. For example, two stocks may mix across Areas
IVE and V-W with the proportional contribution of eachstock to the total abundance in this mixing area varying overtime. It would be possible to extend the SCAA model by
fitting to a time series of mixing proportions (such as the
proportions predicted by the model described in SC/F14/
J29) or by fitting directly to genetic and morphometricdata that may discriminate among the two stocks (e.g. byimplementing an approach similar to that described in SC/
F14/J29 directly in the SCAA model). Either approach gives
the possibility that the model can, in the future, evaluate theimpact of different stock-structure hypotheses on the stock
dynamics. The Panel also noted that if further analysis ofgenetic and morphometric variations in minke whales fromthe JARPA II study area supports the alternative hypothesisof a single stock in which individuals are isolated by distance(see Item 5.2) rather than two stocks which mix, substantial
revision to the SCAA model would be required.
CHANGES in CARRYING CAPACITY through time

The base case model estimates that carrying capacity
increased from the mid-1940s to the early 1960s and thendecreased. The Panel agrees that care should be taken
when interpreting trends in carrying capacity that may be
estimated using the SCAA model. Estimated trends incarrying capacity would not necessarily index trends in theavailability of krill. The SCAA model is parameterised suchthat carrying capacity may also index the annual survival of
age-0 minke whales, which are born, nursed, and weaned
at low latitudes. Thus, estimated trends in carrying capacity
may index trends in conditions that have occurred outside
the Antarctic, including trends in predation on minke whale
calves.
Changes in growth rates through time

The model produces estimates of rapid growth in the earlytime period and slower growth in more recent years, witha transition between the early 1980s and around 1990 (SC/F14/O02, fig. 5). The estimated growth rates are, however,largely determined by data on the smallest individuals, andfew small individuals were caught during the commercial
hunt. Therefore, estimated growth rates in the earliest period

Fig. 4. Scatterplot of recruitment at age 1 and different SD assumptions(see text).

may be unreliable. This issue should be considered further
because, at present, it is difficult to reconcile the age-atmaturity time series presented in SC/F14/J08 with thegrowth rates estimated by the SCAA model. The time series
presented in SC/F14/J08 suggests that growth rates did not
decline during the 1980s and were relatively constant for all
cohorts born since about 1965. The Panel recommends thatthe proponents and developers of the SCAA work to resolve
this apparent inconsistency.
Effects of JARPA/JARPA II data

Results were presented indicating that removing JARPA/
JARPA II abundance data (while retaining JARPA/JARPA
II age-length data and the IDCR/SOWER abundance data)
had little effect on the model results. This was interpreted
to indicate that the abundance data are consistent with otherdata (including the age-length JARPA/JARPA II data) fitted
in the model. To further evaluate the effect of the JARPA/
JARPA II data, the Panel recommends that sensitivity tests
removing all JARPA/JARPA II data except catches from themodel (including removing the age-length data), or removing
just the age-length data while retaining the abundance data.
In either case, an assumption about selectivity pattern for the
JARPA/JARPA II period would be needed.
The Panel agrees that sensitivity tests such as thoseundertaken to examine the effects of including (excluding)the JARPA/JARPA II abundance data from the SCAA are
useful for considering the future design of JARPA II. For
example, if the annual JARPA II abundance data have little
effect on results from the SCAA, the proponents might bepresented with an opportunity to spend less time conducting
sightings surveys while spending more time addressing otherissues. For example, if a sighting survey was not conducted
one year, the time at sea could be used to conduct a process
study focussed on field work that examines the possibility ofcompetition between whale species (see Item 6.3.3 above).

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6.3.3 Alternative approaches to abundance estimation
SC/F14/J07 reported results of a preliminary evaluation ofuse of paternity analysis to estimate abundance of Antarctic
minke whales. This exercise genotyped 137 femalefoetus pairs collected during 2003/04 JARPA cruises andsearched for matches among the >1,700 males in the genetic
dataset. One likely match to a male parent was found. The
analytical procedures used and the caveats provided weregenerally appropriate. The Panel welcomes this generalapproach because it has considerable potential to provide
useful information and had the following comments on thispreliminary evaluation:(1) a point estimate of abundance based on a single parentoffspring match is of questionable value, since: (a) the
confidence intervals are so wide - Bravington et al.
(2014) shows how to calculate the number of matches
required to produce an estimate with a specified
coefficient of variation; and (b) the point estimate is
unstable in that the addition of even a single recapture
dramatically reduces the point estimate; and(2) although a brief mention is made of the importanceof random sampling, this crucial topic merits a fuller
treatment, given the fact that all samples are restricted
to an area that represents less than half of the summer
distribution of the species.
6.3.4 Distribution
This section addresses part of Objective 1 of JARPA
II: Monitoring of the Antarctic ecosystem, with specialreference to temporal changes in cetacean distribution
based on sightings data and the specific consequences forthe distribution of minke whales; and part of Objective 2:
Modelling Competition among whale species. Proponents
presented two papers on distribution including: a general
mapping of species occurrence pooling all the observations
from JARPA and JARPA II from 1987 to 2009 and using an
uncorrected index of whale density with 1° x 1° resolution(SC/F14/J17); and a paper (SC/F14/J18) reporting ananalysis of changes in the spatial distribution of minke andhumpback whales in Area VI during the JARPA and JARPA
II period using generalised additive models (GAMs).
The Panel was pleased to see the effort of putting all thesightings data together, providing a retrospective view ofthe distribution of all cetacean species observed, and the useof spatially-explicit modelling of the humpback and minke
whale sightings to address some of the proposed objectives.
The GAM analysis breaks down the JARPA and JARPA
II period from 1987 to 2006 into an early, middle and latephase and reports large scale area occupancy for eachspecies, and a derived area occupancy index to address theextent of overlap in spatial distribution between minke andhumpback whales. The Panel identified two major issues: (1)the analysis of area occupancy should be more informative,
which could be achieved by expressing the results in termsof whale density using line transect-based spatial modelling,as opposed to using simple GAMs, and where possible, theanalysis should be carried out for other species; and (2) theindex to compare occupancy between species in its current
formulation cannot appropriately describe species overlap
and segregation in distribution and their changes over time.
While welcoming the use of more quantitative approaches
to examining distribution, the Panel notes the methods used
are somewhat simplistic. The Panel therefore recommends
that a more rigorous area occupancy analysis should beundertaken that incorporates recent advances in spatialmodelling (Bravington and Hedley, 2012), accounting forwhale detectability and the encounter rate process, including

autocorrelation, and thus reflecting spatially-explicit
differences in density. In addition, the choice of explanatory
variables in spatial models should better reflect theenvironment that describes the distribution of each species,
such as sea-ice, primary production and oceanography,
incorporating a broader dataset than simply using JARPA and
JARPA II data alone (and see Item 4). Recommended remotesensing resources, as used in other analyses (Beekmans etal., 2010) are, inter alia, bathymetry databases (GEBCO orothers) and related indices such as distance to shelf break andother contours; ocean colour for chlorophyll concentration(NASA Sea-viewing Wide Field-of-view Sensor, SeaWiFS) or
related; sea ice concentration and related ice indices, including
distance to the sea ice edge, maximum and minimum seasonalsea ice extent, and sea ice season duration; sea surface
temperature (SST); frontal zone location positions of theSouthern Antarctic Circumpolar Current Front (SACCF) andthe Southern Boundary of the Antarctic Circumpolar Current(SBACC), and sea surface velocities (SSV).
The Panel agrees that the index of area occupancy to
predict presence-absence also needs to be improved by
re-formulating the analysis as a line-transect based spatialmodel. This will increase the accuracy of fine-scale prediction
and will improve the analysis of segregation in distribution
range between minke and humpback whales. In addition, theindex used to investigate the extent of overlap in distribution(zi = px,i – py,i where i is a small scale prediction grid cell, px,iis the probability of presence of minke whales, and py,i ofhumpback whales) is not valid in its present form, as it isbased on models and predictions independently derived foreach species. Moreover, many different combinations of px,iand py,i yield identical zi values, although the interpretationof those values could be very different. A proposed solution
to this problem is modelling together the spatially-explicit
probabilities of absence of both species (p00,i), presence ofminke whales (px0,i), presence of humpback whales (p0y,i),and presence of both species (pxy,i) in response to spatial
covariates. The observed presence of each species in agrid cell can be treated as a sample from a multinomial
distribution, and estimation in a GAM analysis context canbe achieved as in Beare et al. (2003) using a binomial or
Poisson regression approach, or alternative solutions.
As part of their discussion of distribution, the proponents
note that a major motivation of the work presented is in
response to Objective 2, and in particular inter-specificcompetition. Their main emphasis is on Antarctic minke
and humpback whales, and the ecological consequences of
their competition. However, the Panel agrees that despite
the apparent changes in distribution and relative abundanceof these two species, any firm conclusion is premature2
and is but one of several possible hypotheses to explain
existing data and for use in modelling. The Panel notesthat at this stage the proponents have failed to develop a
proposed underlying hypothesis for this competition, or theformulation of a conceptual model to develop an analyticalframework. Studies on whale competition are rare, andmostly investigate the partition of resources utilised bydifferent species on the feeding grounds (Friedlaender et al.,2009; Santora et al., 2010). This requires a characterisation of
the spatial distribution of each whale species in relation withenvironmental variables, and the distribution, abundanceand behaviour of their common prey.
SC/F14/J18 stated that ‘The results indicated that competition between
humpback and Antarctic minke whales for habitat in Area IV during theperiod of JARPA and JARPA II was intensified as abundance of humpback
whales increased’.

2

j. cetacean res. manage. 16 (suppl.), 2015

To this aim, the Panel recommends that the focus of
future effort should include work that will enable specificationof a hypothesis as to how the proposed competition in
the Antarctic ecosystem is taking place as well as otheralternative hypotheses. The Proponents should show thatthe attribution of correlational results to competition is notconfounded by the consequences of oceanographic shifts andchanges in carrying capacity leading to changes in prey andwhale distribution and abundance. This requires conclusive
evidence of foraging preferences, shared use of the same
prey field by different whale species and information that the
prey field is somehow unable to satisfy the demands of all
predators that are sharing the resource. For this, the Panelrecommends the use of dedicated methods such as TDRs
linked to satellite telemetry to characterise species-specificdiving ranges in relation with the size and distribution of krill
aggregations; and use of indirect methods to compare prey
preferences. The Panel refers to the full discussion of related
matters in the context of the SOWER 2000 programme(IWC, 2000b). It stresses the importance of swarm size aswell as average density with respect to studies of energetics
and successful feeding of baleen whales. In order to develop
plausible hypotheses for use in ecosystem modelling as partof JARPA II, the Panel recommends that the proponents
invest effort in some and preferably all of the following:(1) analysis of proximity of whale species in the sighting
survey tracklines – this can be done with existing data andinitial analyses should be possible within the next year;(2) focused studies of prey swarming behaviour and density
in relation with local whale distribution and abundance– this may require a change of fieldwork focus for one
or more seasons and integrated data from focal follows,
telemetry (radio and/or satellite), krill abundance anddensity estimates and biopsy sampling (for inter alia
stable isotope analysis);(3) comparison of prey in stomach contents in areas where
both species overlap in distribution, and in areas where
they segregate, including investigation of whether
whale species and krill length-maturity classes exhibit
distinct spatial segregation in their distribution patterns– this can begin with existing samples and it should bepossible to present a progress report within the next two
years; and(4) analysis of stable isotope ratios of C, N and O from tissue
samples – initial examination of the discriminatory
power of such analyses can occur with existing samples
preserved dry or frozen.
7. REVIEW OF RESULTS: Monitoring krill
abundance and feeding ecology ofwhale stocks (SC/F14/J13-J16, SC/F14/J19, SC/
F14/O03, SC/F14/O05-O07)
7.1 Proponents’ summary of results
JARPA II includes a research component of krill biomass
estimation based on quantitative echo sounder data. Alongwith this data, feeding ecology (e.g. energy storage, stomach
content, and feeding habit) of Antarctic minke and fin whales
was investigated. Analysis of those data would contributeto answering such scientific questions as; whether the
total availability of krill to whales in the Antarctic marine
ecosystem has changed or not, and whether the shift in thewhale species composition has affected prey availability
to different species of whales. This analysis would also
contribute to building ecosystem models for the Antarctic
marine ecosystem.

387

SC/F14/J15 investigated the feeding habits of Antarctic
minke whales based on samples obtained during the surveysof the JARPA and JARPA II for the period 1987/88-2010/11.
Some of the analyses considered recommendations offeredduring the 2006 JARPA final review meeting, e.g. thoserelated to the duration of the feeding period, digestion rateand examination at smaller scales. The Antarctic minke
whales fed mostly on Antarctic krill in offshore area, and on
ice krill in coastal (shallow) area on the continental shelf of
the Ross Sea and Prydz Bay. The whales fed mainly before
05:00am suggesting that feeding activity decreases early inthe day. The daily prey consumption was estimated using
two independent methods, one based on theoretical energy
requirements and the other on diurnal changes of stomach
contents mass. The daily prey consumptions per capita
during the feeding season based on the two methods were
95.1kg, 127.0kg and 182.6kg, 250.3kg for immature andmature males, and 125.8kg, 138.7kg and 268.1kg, 325.5kg
for immature and mature females, respectively. This isequivalent to 2.65-4.02% of the body weight. The daily
prey consumption per capita decreased between the JARPA
and JARPA II periods based on the results of the method
on diurnal change in stomach content mass, for all sexualclasses. The seasonal prey consumption by all Antarctic
minke whales in the research area was 3.51-3.98 million
tons, and this figure was equivalent to 7.6-8.6% of the krill
biomass estimated by acoustic survey in the total researcharea. Further research on feeding ecology will focus on the‘uncertainties in the prey consumption estimates of whales’.
SC/F14/J16 examined the feeding habits of fin
whales sampled during the surveys of the JARPA II for
the period 2005/06-2010/11 (n=16). The fin whales fed
mostly on Antarctic krill in the research area. The daily
prey consumptions of fin whales per capita estimated bythree methods ranged from 276 to 2,136kg. These values
were equivalent to 0.50 and 3.84% of the body weight.
The seasonal prey consumptions for all fin whales in the
total research area based on three methods were 0.54-0.78
million tons, 3.38-4.51 million tons and 2.19-2.93 million
tons, respectively. There was coincidence in the frequency
of body length of the Antarctic krill, E. superba consumed
by fin and Antarctic minke whales. It seems that fin andAntarctic minke whales have similar feeding habits with noprey size selectivity. SC/F14/J10 showed evidences that fin
whales taken by JARPA II are heavier than those taken bycommercial whaling in the 1950s.
SC/F14/J19 presented results of krill biomass estimations
based on quantitative echo sounder data. Estimations weremade based on data obtained in Areas IIIE/IV in the 2007/08
JARPA II survey and in Areas V/VIW in the 2008/09 JARPA
II survey. The biomass estimates obtained in Areas IIIE
and IV were 6.6 and 12.5 million tons, respectively while
those in Areas V and VIW were 24.0 and 3.4 million tons,
respectively. The information on krill biomass for Area IV
was used as input parameters in the ecosystem modellingwork. It should be noted here that comprehensive data forkrill biomass abundance were obtained only for two JARPA
II seasons due to security problems.
In summary SC/F14/J13 and SC/F14/J14 showed
consistent results on deteriorating nutritional conditionsof Antarctic minke whales in the research period. SC/F14/
J15 showed that the daily prey consumption per capita of
Antarctic minke whales decreased between the JARPA and
JARPA II periods for all sexual classes, providing additional
support to the results of SC/F14/J13 and J14 (see Item 8
below). These results suggested that less krill is available

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report of the expert workshop to review jarpa ii

for Antarctic minke whales in recent years (JARPA+JARPA
II period). Direct estimation of krill biomass, however, wasobtained only for two JARPA II surveys due to security
problems.
7.2 Panel’s conclusions and recommendations
7.2.1 Krill abundance
The estimation of krill abundance represents a fundamental
component of JARPA II and is central to its ability to meet
its ecosystem-related objectives. As described in Item
10.2.1.1, a reliable time series of krill abundance is requiredin order to quantify possible competition between whale
species. SC/F14/J19 presented the results of two directedsurveys for krill. While welcoming the work to estimate krill
abundance the Panel had concerns over the interpretation of
results in the paper. In particular, the size compositions used
for classifying acoustic targets as Antarctic krill, E. superba,
appear to be contaminated by data on other euphausiid
species. For example, fig. 3a of that document indicatesthat no Antarctic krill were included in the size-frequency
distribution of animals collected from stratum IV-NW
during 2007/08. The Panel recommends that the biomass
estimates reported in SC/F14/J19 be recomputed using size
compositions that, in northern strata, are representative of
E. superba only since this is the primary species of krill thatminke whales consume in these strata. For strata IV-SW and
V-SE, which respectively include Prydz Bay and the Ross
Sea, the Panel recommends that separate size-frequency
distributions and biomass estimates be made for E. superba
and E. crystallorophias3.
The Panel recommends that the recomputed biomass
estimates reported in SC/F14/J19 be compared to biomass
estimates made from acoustic data collected during JARPA
as soon as possible. As the authors themselves have noted,
to make this comparison, the acoustic data from both JARPA
and JARPA II must be processed in the same way. Suchcomparisons have the potential to provide the first direct
evidence as to whether there have been changes in thebiomass of krill in the JARPA II study area. However, thePanel also highlighted the fact that the data from the JARPA
and JARPA II programmes alone are limited. It therefore
recommends that any such comparisons also include
biomass estimates from the BROKE (Nicol et al., 2000)
and BROKE-WEST (Nicol, 2010) surveys. It is important
to note that during the BROKE survey, acoustic data werecollected at only two frequencies. To compare JARPA and
JARPA II acoustic biomass estimates to that from BROKE,
it might be necessary to reprocess the existing JARPA and
JARPA II data using only the frequencies used in BROKE.
The Panel stresses that limiting processing of acoustic datacollected during future surveys to two frequencies is notrecommended.
The understanding of krill biomass (and any trends in it)
is fundamental to future work under JARPA II if it is to meetits objectives. Whilst recognising issues associated withoutside disturbance, the Panel is concerned that not enougheffort has thus far been made in JARPA II to address thisissue. The Panel notes that the IKMT net used during JARPA
In cases where data from the IKMT tows do not provide sufficient data tocharacterise the size compositions of E. superba (or E. crystallorophias), it
may be possible to use data on the size compositions of these species fromthe fresh (F) component of the diets of whales sampled in any given stratum
if the spatial distribution of sampled whales was random with respect to the
distributions of these two euphausiids and there is some confidence that thewhales did not select for certain sizes of prey.

3

II is appropriate for the objective of observing the species
and size compositions of euphausiids in the JARPA II study
area, but agrees that an insufficient number of IKMT tows
were made during the two krill surveys reported in SC/F14/
J19. The relatively few tows that were made during surveys
conducted in 2007/08 and 2008/09 were not likely to haveprovided representative data on the species composition of
euphausiids nor on the size composition of E. superba in the
study area. For example, only two tows were made in the
IV NW stratum during 2007/08, and E. superba were notcaught in either tow.
The Panel welcomes the fact that a substantial numberof random krill samples (about 200g per sample) werecollected from the stomachs of whales sampled during
JARPA and JARPA II and archived at the ICR (Table 1).
Some of the krill samples have already been processed toprovide size-frequency distributions (using about 50-150
krill per sample). A smaller number of samples have also
been processed to provide maturity-stage distributions of thekrill. All krill samples archived from collections made prior
to 1996/97 have been processed, but a substantial number ofthe samples archived from collections made since 1996/97
have yet to be processed. Size and maturity distributions
are extremely valuable for understanding the dynamics ofthe krill stock in the JARPA II study area and the Panelrecommends that the archived krill samples be processed tocharacterise size-frequency and maturity-stage distributions
as a matter of priority. The Panel also recommends that the
proponents collaborate with members of SC-CCAMLR touse data collected from the archived samples in an effort tostudy the population dynamics of krill.
The Panel recommends that future krill surveys
conducted during JARPA II employ survey design standards,
including the spacing between net-tow stations along theacoustic transect lines, similar to those developed andimplemented for the CCAMLR 2000 (Trathan et al., 2001),
BROKE (Nicol et al., 2000) and BROKE-WEST surveys(Nicol, 2010). It would also be useful for the proponents
to discuss survey design issues with scientists outside the
JARPA II programme that have experience in conducting
krill surveys.
The Panel notes that the backscattering strength threshold(log Sv ≤ -80dB) used thus far during JARPA II is potentiallybiasing low the acoustic krill biomass estimates. The
proponents indicated that the threshold values were selectedto accommodate the excessive noise of the research vessel,
recognising that this would affect the detection of small krill.
The depths over which backscattering data were integrated
to estimate krill biomass (15-150m) were also limited byvessel noise (more typically, backscattering data on krill
are integrated down to 250m). The Panel recommends that
future krill surveys be conducted using a more appropriate
vessel since krill biomass estimates are required to meetthe objectives of JARPA II. Noise standards for conducting
acoustic surveys have been discussed by ICES (1995).
Future krill surveys should be frequent because thedensity of krill in any given stratum may vary significantly
from year to year, and the objectives of JARPA II require atime-series view of how the prey field is changing over time.
The JARPA II study area is very large, and it is probably
not possible to survey the entire study area every year witha single survey vessel. Thus, the Panel recommends either
using multiple survey vessels to synoptically cover the
JARPA II study area every 1-3 years or using one vessel
to survey alternating halves of the study area every year
(similar to the approach taken for sighting surveys).

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j. cetacean res. manage. 16 (suppl.), 2015

7.2.2 Feeding ecology, consumption rates and inputs toecosystem modelling
The Panel considered papers SC/F14/J15, SC/F14/O03
and SC/F14/R01 under this item. The Panel welcomes theconsiderable amount of field and laboratory work undertakenon this subject and analysed in SC/F14/J15. However,it agrees that a major shortcoming of the work originallypresented in SC/F14/J15 was that the results presented
did not quantify uncertainties in the various estimates ofconsumption. This shortcoming was identified in SC/F14/
O03; in SC/F14/R01, the proponents proposed the use of aMonte Carlo approach to address this shortcoming.
The Panel agreed that the approach proposed in SC/
F14/R01 is both a positive development and a useful way
forward. However this work is at its beginnings only;
without the results of a thorough analysis, the Panel is unable
to determine whether JARPA II has provided or will provide
consumption estimates within a sufficiently narrow range.
The Panel recommends that work proposed in SC/F14/R1
be further developed and allocated high priority. Ideally a
new paper should be submitted to the next meeting of the
Scientific Committee. As a minimum, this should advancethe outlined work plan by including in the Monte Carlosimulations, uncertainty in:(1) r (the ratio of low/high feeding intake) and the length of
the feeding season for Method 1; and(2) the extent of night feeding for Method 2.
The Panel also recommends that the work is extended bycomputing a time series of Monte Carlo results for the total
potential consumption of krill using abundance estimates ofAntarctic minke whales (preferably those estimated by the
SCAA model) and the uncertainties around these estimates.
These analyses are an essential consideration as to theextent to which the JARPA II programme in its present
form is able to meet its objectives. The Panel therefore
recommends that the results of the Monte Carlo work be used
to re-evaluate the future research and sampling in JARPA
II. This should include an exploration of the sensitivity of
the Monte Carlo distributions of consumption estimates (forboth individuals and for the total stock(s) of whales) to a
reduction in the uncertainty of each parameter and to the
choice of allometric relationship between body mass andmetabolic rate. Future research should be targeted to reduce
uncertainties in the parameters to which the Monte Carlo
distributions of consumption estimates are most sensitive.
This approach could also be used both to aid determinationof future sample sizes by estimating how many samples
would be needed to reduce uncertainties in consumption
estimates by a desired level and to assess whether lethal or
non-lethal sampling or a combined approach represents best
way of reducing such uncertainties.
The Panel agrees that results presented in SC/F14/J16 (figs 4-6) indicate substantial overlap in the sizes ofkrill that are consumed by Antarctic minke whales and fin
whales when sampled near the same locations and times,
noting also that the sample size for fin whales is small,
restricted to smaller individuals and outside the species main
distribution area. As elsewhere in this report, however, thePanel cautions the proponents over any over-interpretationthat this proves that there is ‘inter specific interaction’ orcompetition between these species in any limiting sense(e.g. see SC/F14/J18), rather than it being one of several
potential hypotheses. SC/F14/J16 suggests that differencesin the size composition of krill consumed by both species
will be determined by the size-specific distribution of krill.

Table 1
Numbers of krill samples that were collected from whale stomachs and havebeen archived at the ICR, including the numbers of these archived krill
samples that have already been processed to characterise size-frequency
distributions of krill eaten by whales.
JARPA

Sample no.

BL data

JARPA II

Sample no. BL data

1989/90
1990/91
1991/92
1992/93
1993/94
1994/95
1995/96
1996/97
1997/98
1998/99
1999/2000
2000/01
2001/02
2002/03
2003/04
2004/05
Total

55
33
45
140
164
161
201
243
264
180
288
151
107
94
113
116
2,355

55
33
45
140
164
161
201
27
32
22
23
24
0
0
0
35
962

2005/06
2006/07
2007/08
2008/09
2009/102010/11

62
80
46
73
49
22

8
4
0
1
2
2

Total

332

17

The Panel notes that while the distributions of minke andfin whales overlap, minke whales are most abundant farther
south and fin whales are most abundant farther north; thesize compositions of krill in the areas where these whales
are most abundant may differ, and, if so, that would tend
to lessen the strength of interactions between them. The
Panel also noted that when the best available point estimateof krill biomass in Area IV during 2007/08 (12.5 million
tonnes, SC/F14/J19) is compared with the best available
point estimates of krill consumption by minke whales (0.37
million tonnes using Method 1 and 0.33 million tonnes using
Method 2, SC/F14/J15) and fin whales (0.05-0.42 million
tonne using three different allometric relationships between
body mass and metabolic rate, SC/F14/J16), the hypothesisthat minke whales and fin whales are competing for krill ina limiting sense does not seem well supported. These point
estimates suggest that together both species of whales might
have consumed about 6% annually of the standing biomassof krill and thus that krill biomass was not limiting.
The Panel’s conclusions and recommendations regarding
the nutritive condition of Antarctic minke whales (blubber
thickness and energy storage – SC/F14/J13; and stomach
contents – SC/F14/J14) are further described in Item 8.3.2.
8. REVIEW OF RESULTS: Monitoring whale
biological parameters (SC/F14/J08-J12, SC/
F14/J35-J37)
8.1 Proponents’ summary of results
The general objective of this component of the programme
was to monitor the changes over years in Antarctic ecosystemfrom viewpoint of biological parameters. Recruitment,
pregnancy rate, age at sexual maturity and other biologicalparameters were selected as target parameters. Biologicalparameters of Antarctic minke whales were estimated byusing JARPA and JARPA II samples and yearly trend was
examined for I-Stock and P-Stock.
8.1.1 Age at sexual maturity
SC/F14/J08 examined yearly trend of age at sexual maturity
in the Antarctic minke whale based on Transition Phase (TP)in earplugs collected by the JARPA (1987/88-2004/05) and
H:\SkyDrive\Documents\AC
Supplement 16\Rep02- JARPA\Rep02
Tabs.docx
the
JARPA II (2005/06-2010/11).
Analysis
was conducted
for both sexes and for two stocks (I and P) separated at
165°E. TP of the JARPA samples were read by two readers

15 Janua

390

report of the expert workshop to review jarpa ii(reader-K and reader-Z) and by a new reader (reader-B) in thecase of the JARPA II samples. Truncation bias was correctedby standard procedures. Yearly trend of mean age at sexual
maturity derived from two reader groups were consistent.
The results confirmed that the age at sexual maturity of bothstocks declined from the mid 1940s cohorts at around 10-12
years old to the early 1970s cohorts at around 7-8 years old,presumably as response to improved nutritional conditions
at that time. Age at sexual maturity remained constant at
7-8 years old till the 1990s cohorts. Statistical analysis toinvestigate trend from the 1970s cohorts showed slight but
statistically significant increasing trend for the I Stock (bothsexes) and P Stock (females). Although slight trend was
detected, the level of current changes is much smaller thanthe decreasing trend level observed from the 1940s cohorts
in response to improved nutritional conditions at that time.
8.1.2 Ageing errors
During the JARPA final review Workshop a recommendation
was made that the comparability of commercial and
JARPA age data be investigated by re-reading a sub-setof the commercial samples in an appropriately designed
blind test. Based on the results of the studies responding
the recommendation, the IWC SC agreed in 2010 that nofurther experiments or analyses on age reading errors wereneeded to resolve ageing related problems raised at the
JARPA review Workshop. SC/F14/J11 presented a statisticalmethod for quantifying age-reading error, i.e. the extent ofbias and inter-reader variability among readers, which use
the data produced in the blind test. The method in SC/F14/
J11 assumes the availability of an independent control reader
who produces reference ages for ageing structures which arealso read by the subject readers. This reader is assumed toprovide unbiased age estimates. Linear structures in bias andvariance are incorporated in a conditional probability matrix
representing the stochastic nature of age-determination foreach reader. A joint likelihood function for the parameters
related to ageing bias, variance and nuisance parameters isdefined based on observed ageing outcomes from both thecontrol and subject readers. The method is applied to data for
Antarctic minke whales taken during Japanese commercial
whaling (1971/72-1985/86) and JARPA (1986/87-2004/05).
A total of 250 earplugs selected according to a predetermined
protocol were used in the analyses to estimate the inter-reader
variation for four Japanese readers. One of the authors acted
the control reader. The Japanese readers and the control
reader differed in terms of both the expected age given thetrue age, and variance in age-estimates. The expected age
and random uncertainty in age-estimates differed among
the Japanese readers, although the two readers in charge of
age-reading for samples taken during JARPA and JARPA
II provided quite similar ageing outcomes. These results
contribute to analyses using catch-at-age data for thisspecies. It should also be noted that the model and approachin this paper can be applied to populations other than the
Antarctic minke whales, if a control reader is available,
even retrospectively. An original version of this paper waspresented to the IWC SC meeting in 2013 as Kitakado etal. (2013). After discussion, the IWC SC agreed that theapproach and results of the study provide useable inputs data
for SCAA analysis.
8.1.3 Alternative ageing techniques
SC/F14/J12 measured the eye lenses of 18 Antarctic minke
whales and of 20 foetuses to examine the approach of age
estimation in Antarctic minke whales by ratio of aspartic
acid enantiomers. In the age estimation equation, thespecific coefficient of hydrolysis effect in preparation andthe constant conversion of aspartic enantiomers (Kasp) were
estimated. The study found that the D/L ratio of aspartic acidin the lenses of minke whales had increased by 0.0102 in
the hydrolysis process (7hr at 108°C), and estimated that
the Kasp was 1.96×10-3 (/year) calculated from the earplug
age and the age index with the D/L ratio of aspartic acid inminke whales. Consequently, formulae Age (year) = 511×Ln((1+D/L)/(1-D/L)) -13.2 was obtained. In the future, thismethod would be applied to other species and individuals
with unreadable earplugs.
8.1.4 Pregnancy rate
SC/F14/J09 examined yearly trend of the proportion ofpregnant whales in the matured female population (PPF)
of Antarctic minke whales based on samples collected by
the JARPA (1987/88-2010/11) and the JARPA II (2005/062010/11) surveys. Linear and logistic regression analyses
were conducted to estimate yearly trend in this biologicalparameter for I-Stock and P-Stock, separated at 165°E. The
PPF of both stocks appeared stable at around 0.9 and whendata from all years were combined, the PPF was 0.932 for
I-Stock and 0.904 for P-Stock. No significant yearly trend
was detected in either stock during the JARPA II period. A
significant increasing trend was detected in the JARPA and
JARPA+JARPA II period analyses for the P Stock by thelogistic regression analysis. The latter result was influenced
by just two lower values in the 1990/91 and 1994/95 seasons.
Although continuation of the high PPF was observed during
the JARPA II period, monitoring of the PPF is important
to detect possible change in nutritional condition andreproduction which affect sustainable management of thiswhale stock.
8.1.5 Nutritive condition
SC/F14/J13 showed the results of the annual trend inenergy storage in sexually mature Antarctic minke whales
based on catch data from all 24 surveys of the JARPA and
JARPA II. These analyses addressed most of the previous
recommendations from the JARPA final review Workshop
in 2006 (see recommendations Marine Ecosystem 4-6 inAnnex 4), as well as more recent recommendations fromthe IWC SC, e.g. analyses included survey track line inrandom effects as recommended in IWC (2013a). Fivevariables which are, or could be, indices of storage ofenergy, were examined: total fat weight in the whale body,
blubber thickness at two lateral measurement points andgirth measured at two specified positions. Three of thesevariables were available from almost all whales sampled,but girth at axilla was measured over 20 years and fat weight
was only measured over 17 years of the JARPA programme,
and only for the first whale sampled on each day. A numberof covariates were also recorded. A large number of linear
mixed-effects statistical models were investigated for eachof the dependent variables, and the Bayesian Information
Criterion (BIC) was used to select the best model. All models
examined had ‘year’ as a possible explanatory variable.
The results show that all five measures of energy storage
declined substantially over time during the JARPA period,
with a more than 10% decline in total fat weight. For all fivedependent variables the values for energy stores are higher
for females than for males. The values increase during thefeeding season and are higher for higher body coverage of
diatoms. This is assumed to be a measure of how long theanimal has spent in Antarctic waters. The results are similar
when each sex is analysed separately, but the decrease inenergy stores was somewhat larger in females than in males.

j. cetacean res. manage. 16 (suppl.), 2015

The results from the JARPA II period were very different
from the JARPA period. There is no clear trend towards
increase, or further decrease, in any of the four measures ofenergy storage. The results suggest that fundamental changes
have taken place in the eastern part of the Antarctic marine
ecosystem during the 1990s. These changes have resulted inless optimal feeding conditions for Antarctic minke whales.
SC/F14/J14 reported the results of an analysis of
temporal trend in stomach content weight in the Antarctic
minke whale, one of the major krill predators in Antarctic
waters. These analyses addressed most of the previous
recommendations from the JARPA final review Workshop
in 2006 (see recommendation Marine Ecosystem 3 inAnnex 4), as well as more recent recommendations from the
IWC SC e.g. analyses are now based on survey transects
as recommended by IWC (2013a). A reported decline inenergy storage over almost two decades indicates that foodavailability for the whales may also have declined until
recently. To test this hypothesis, catch data from 21 survey
years in the JARPA and JARPA II (1990/91-2010/11) wasused to investigate whether there was any annual trend in
the weight of stomach contents of Antarctic minke whales.
Linear mixed-effects analysis showed a decreasing trend
in the weight of stomach contents over the 21 years since
1990/91, which was statistically significant. A similar patternof decrease was found in both males and females, except in
the case of females sampled at higher latitude in the Ross
Sea. These results suggest a decrease in the availability ofkrill for Antarctic minke whales in the lower latitudinalrange of the research area. The results are consistent withthe previously reported decline in energy storage (SC/F14/J13). The decrease in krill availability could be due toenvironmental changes or to an increase in the abundance ofother krill-feeding predators. The latter appears more likely,
given the recent rapid recovery of the humpback whale.
Furthermore, humpback whales are not found in the Ross
Sea, where both Antarctic krill and ice krill are available,
and where no change in prey availability has been suggested
for Antarctic minke whales.
8.1.6 Summary
In summary the analysis of biological parameters of Antarctic
minke whale showed a small but significant increase in theage at sexual maturity after the 1970s cohorts suggesting thatnutritional conditions could be deteriorating for this species.
However the age at sexual maturity has remained low at
between 7-8 years old. Furthermore a constant and highpregnancy rate was observed during the JARPA/JARPA II
period. Therefore in this period high reproductive rates canbe postulated however the recruitment rate has not increasedin the same period. On the other hand there is the possibility
that the age at sexual maturity of fin whale (see below)
could have decreased suggesting improved nutritional forthis species. The changes in age at sexual maturity in the
Antarctic and fin whales are consistent with the change inthe ecosystem suggested by JARPA results.
With respect to nutritive condition, SC/F14/J13 andSC/F14/J14 showed consistent results on deteriorating
nutritional conditions of Antarctic minke whales in the
research period.
8.1.7 Fin whales
SC/F14/J10 presented new biological information of 16
Antarctic fin whales collected from JARPA II biologicalresearch. New information on body proportion, reproductive
status, age/length relationship, body length/body weight
relationship as well ecological markers (external parasites),

391

is presented. For age/body length relationship and age/maturity parameters were compared with data of thecommercial whaling period in the Antarctic which stopped
some 36 years ago. The main results were that body weightof whales in the JARPA II period is heavier than thosepreviously reported for the Antarctic in the 1950s. Further,results suggested the possibility that whales are reachingsexual maturity at younger ages.
8.1.8 Other information on the biology of Antarctic minke
whalesSC/F14/J35 (Nagai et al., 2007) used follicular oocytes,
follicular fluid, and umbilical serum from seven female
Antarctic minke whales in JARPA II to investigate whale
reproductive physiology, especially for improvement of invitro oocyte maturation and related technologies for whales.
Main results were the following: (i) the mean diameter ofthe oocytes from the large follicles was significantly greaterthan those from the small and medium follicles; (ii) theosmolarity of the follicular fluid from the small follicles was
significantly lower than that of the medium follicles; (iii)the concentrations of the total-protein, glucose, albumin andchlorine concentrations of the follicular fluid from the threegroups of follicles were significantly lower than that of theumbilical serum; (iv) for steroid hormones, the progesterone
concentrations were not significantly different among thefluid from the three groups of follicles and the umbilicalserum, but the estradiol 17-β concentrations of the follicular
fluid increased with the size of the follicle (14.3 and 34.6 ng/
ml for small and large follicles, respectively).
SC/F14/J36 (Ono et al., 2009) used heart samples ofAntarctic minke whale collected during the 2005/06 JARPA
II surveys for comparison of Purkinje fibres in different
mammalian species by light and electron microscopy.
From light microscopy of silver impregnated tissues in thecytoarchitecture of a network of Purkinje fibers revealed
the reticular fibres ensheathing individual Purkinje strands
consisting of 2-8 cells in both the ungulates (i.e. sheep
and goats) and cetaceans (whales and dolphins) while theyencircled each Purkinje cell in the primates (humans andmonkeys), carnivores (dogs and seals), and rodents (rats).
By scanning electron microscopy of NaOH digested tissues
findings, the ungulates were found to have a Purkinje fiber
network composed of Purkinje strands; the cells in thestrands were oval and made side-to-side and/or end-to-end
connections while in primates and carnivores it was delicate
and complicated; the Purkinje cells were usually cylindricaland connected end-to-end, the exception being theirpolygonal or stellate shapes at the bifurcations. Purkinjecells in the rodents resembled ventricular cardiac myocytes
in cytoarchitecture. From the comparative observations,this study showed that the whales and seals respectively
had Purkinje cells similar to that of the ungulates, and theprimates and carnivores. These findings indicated that thestructural variety of the Purkinje fiber network may reflect
the conducting function and be related to the phylogeny ofthe mammalian species.
SC/F14/J37 (Sasaki et al., 2013) studied the morphological and immuno-histochemical nature of the structureand function of placenta in the Antarctic minke whale using
the samples of the placenta and non-pregnant uterus obtained
during the 2001/02, 2002/03 and 2004/05 seasons of the
JARPA as well during the 2005/06 JARPA II season. Fromthe hematoxylin and eosin stain and immuno-histochemicallystained samples using the avidin-biotin peroxidase complex(ABC) method or scanning electron microscopy findings,
they revealed morphological changes of placenta through392

report of the expert workshop to review jarpa ii

the gestation period and the localisation of steroidogenic
enzyme in the Antarctic minke whale placenta. From thefindings the placentation and morphological function typicalof cetaceans may become apparent.
8.2 Observer summary of SC/F14/O06-O08
The authors of SC/F14/O06 and O07 summarised theconclusions of their analyses of minke whale body condition
that used the JARPA/JARPA II data. The biology of the
Antarctic minke whale and the highly variable Antarctic
environment are likely to generate within season spatial andtemporal heterogeneity in the distribution of individuals bysex, age, maturity and condition. Consequently attempting tomonitor subtle trends in biological parameters of Antarctic
minke whales such as blubber thickness and stomach
contents requires a carefully considered and consistentsampling strategy. They showed that JARPA and JARPA II
have not followed such a strategy and consequently the data
are confounded and biased. Even after complex statisticalmodelling, any long-term trends inferred from these data
are likely to be imprecise and unreliable. Standard linear
and linear mixed effects models were applied to attempt tocorrect for heterogeneity in the manner in which the data
were collected and these confirmed that interaction terms
were significant. The results of these analyses indicated thatspatial and temporal heterogeneity requires complicatedstatistical modelling and that statistical inferences fromthese models are weak and ambiguous. While some models
suggest statistically significant trends in some body condition
parameters others do not, with the conclusion that the datacollected during JARPA and JARPA II are insufficient for
determining whether there have been trends in Antarctic
minke whale body condition or otherwise.
The authors of SC/F14/O08 also noted that the methods
used to calculate sample sizes had assumed that sampling was
the only source of variability and consequently had failed totake into account spatial and temporal variability. A bootstrap
analysis demonstrated that the original method of calculation
substantially over-stated the ability of JARPA II to detectchanges in body condition and were therefore invalid.
The authors pointed out that backwards model selection
using BIC in the nutritive condition analyses leads to theillogical consequence that a valid a priori hypothesis for
spatial and yearly interaction terms is accepted, but would
then subsequently be discarded even though it is highly
statistically significant. Furthermore the models using
interaction terms have the lowest AICs.
8.3 Panel’s conclusions and recommendations
The Panel recognises the considerable field and laboratory
effort that lay behind the papers on biological parameters.
Discussion of biological parameters is part of the proponents’objective of monitoring the ecosystem. The work is also
directly dependent upon and relevant to consideration ofstock structure (Item 5). It is also dependent on the broadconsideration of the representativeness of the sampling
design (see the JARPA review [IWC, 2008] and Item 12).
The Panel notes that the analyses presented on biologicalparameters assume: (a) that the sampling scheme is broadly
representative of the population; and (b) related to the stock
structure hypotheses agreed at the JARPA review rather thanthose that were subsequently presented to this meeting (Item5). With respect to sampling design in JARPA II, the Panelnotes the disruption caused to the proposed sample size anddesign due to outside interference in the Antarctic and refers
to the discussions on sample size and design at the JARPA

review meeting (IWC, 2008) and in SC/F14/O05-O07. The
Panel’s overall comments on sampling design/size within
JARPA II are given under Item 12.1.2.
8.3.1 Biological parameters8.3.1.1 Age at attainment of sexual maturity

SC/J14/J08 examined the changes in the age at attainmentof sexual maturity (tm) as reflected in the identification ofthe ‘transition phase’ in the earplugs of Antarctic minke
whales. The authors identified a decline from the mid-1940s
cohorts (around 10-12 years) to the early 1970s (7-8 years),remaining largely constant since then but with possibly aslight recent increase in the I-Stock and P-Stock (females).
The Panel agrees that the analyses presented in SC/J14/
J08 require further refinement in addition to updating withrespect to stock structure, as the authors’ themselves note. In
particular, it recommends that a revised paper should:(1) use the approach of Thomson et al. (1999) to betteraccount for truncation bias (which is considered in SC/
F14/J08) as well as the fringe effect, which relates to thelow proportion of animals with ages slightly higher than
the age-at-maturity, an issue likely related to the needfor contrast to detect a transition phase in an earplug;(2) incorporate the effects of age reading errors presented
in SC/F14/J11;(3) take into account the problems of the representativenessof samples in recent years due to disruption of the
programme discussed under Item 12.1.2;(4) compare analyses of tm from the transition phase withthose using corpora counts and, if available, histologicalstudies of testes;(5) compare the results of the revised study with those, reanalysed to the extent possible following Thomson etal. (1999) for fin and sei whales from the Antarctic to
examine whether any observed trends are synchronous,
which may provide information on whether there is across-specific (e.g. outside environmental) cause to thetrends;(6) evaluate alternative models (e.g. where trends areshared across sex, stock or both); and(7) include a more detailed description of the models
evaluated and a more detailed discussion of the resultsthat considers all plausible explanations for observed
trends.
The Panel recognises that incorporating all of thesesuggestions will probably not be possible for the next
Scientific Committee meeting. However, it believes that an
updated paper that at least includes an improved discussion
and a workplan and timetable should be presented.
The Panel noted that SC/F14/J11 had already been
discussed and approved by the Scientific Committee (IWC,
2013a) and is ‘in press’. It endorses the positive conclusionsof the Scientific Committee and welcomes this work.
8.3.1.2 Pregnancy rates

SC/F14/J09 examined pregnancy rate data from JARPA and
JARPA II. The Panel notes that ‘pregnancy rates’ identified in
the paper are more appropriately called ‘apparent pregnancy
rates’. The Panel stresses that pregnancy rate (or apparent
pregnancy rate) data should not be over-interpreted. In termsof effects at the population level the ultimate parameters ofinterest are successful births and calf survival rather thanpregnancy rates per se. The Panel agrees that the analyses
presented in SC/J14/J09 require further refinement inaddition to updating with respect to stock structure, as theauthors’ themselves note. It recommends a number of
improvements that should be incorporated into a revised
version of the paper:j. cetacean res. manage. 16 (suppl.), 2015(1) further discussion of the limitations of the criteria used to
identify a pregnant female – in particular, better accountshould be taken of the fact that small foetuses may be
missed by scientists and that foetuses may be lost due toabortion4 during the hunt or loss during towing leading
to an underestimate of true pregnancy rate; and(2) to assist with (1), the authors should consider examination of other criteria for pregnancy (in some cases thismay be possible from existing data or tissue samples)
including diameter of the largest uterine horn, histologyof the uterine mucosa, examination of the mammary
glands and examination of hormone levels in blubber.
With respect to (2), the Panel recognises that identificationof lactating females is more difficult in Antarctic minke
whales because births occur in winter and lactation lasts
only about three months, while harvesting takes place inthe summer i.e. some months after weaning. The Panelrecommends that the feasibility of using histologicalexamination of mammary glands to assess recent lactation
is examined. If this is possible, the proportions of recently
lactated to [apparently] pregnant females and to mature
females would be a more appropriate index of reproductive
success that would likely be more sensitive to environmental
fluctuations than crude pregnancy rates because perinatal
mortality or abandonments of newborns by females in low
nutritive condition is known to occur in many mammal
species. The possibility of undertaking a feasibility study
using existing samples should be examined and the results
reported to the Scientific Committee.
The Panel notes that while age at attainment of sexual
maturity is the result of conditions affecting the individual
over a period of several years, pregnancy/lactation is more
directly linked to the conditions prevailing in the season
when the whale had been caught or in the immediately
previous one. Given the proponents’ primary objectiveof studying biological parameters in order to monitorchanges in the ecosystem, the Panel recommends thatthe relationship between reproductive parameters and theinformation available on body condition, krill consumption
and environmental covariates such sea surface temperature(SST), location, and other factors be examined on an annual
basis. This may help to explain anomalous years likethe 1990-91 or the 1994-95 seasons, when low apparent
pregnancy rate values were observed.
The Panel also recommends that:(1) existing sampling protocols be examined to ensure thatuterine horn and mammary gland measurements arerecorded and mammary gland tissue and uterine mucosa
are collected for histological analysis; and(2) the programme takes the opportunity to examine theefficacy of approaches to examine pregnancy using
blubber analyses and faecal samples found for someother mysticetes (Kellar et al., 2013; Mansour et al.,2002) – and see Item 12.4.
8.3.1.3 Age estimation

393

and a correction for ‘foetus age’ in the DLo estimate. The
authors were able to measure differences in the DL ratios ofdeveloping foetuses (n=20), which suggests good precisionin their DL measurements. One of the usual challengesof this method is obtaining a sufficient sample of ‘age 0’
animals which is not a problem for the present study.
The Panel welcomes this work and makes the following
recommendations for an updated paper:(1) improve the estimates of SE which were artificially
optimistic in that: (a) they did not incorporate all sourcesof error, such as error in Kasp (the time-temperaturechange constant of aspartic acid), the DLo measurement,
and the DL measurements for each individual; and (b)were applied incorrectly to calculate the SE of the age
estimates (in fact they estimate the mean SE of theregression as a function of age);(2) with respect to (1) a more appropriate method is thebootstrapping as in Rosa et al. (2012) or the Deltamethod as in George et al. (1999);(3) better specification of how the correction for a hydrolysis
effect on the DL measurements was developed andapplied;(4) a fuller explanation in the methods sections as to how
the lens nucleus was dissected without contaminating
it with ‘modern’ tissue from the exterior of the lens orwith blood or fluids;(5) present the earplug estimates side-by-side with the
AAR estimates;(6) for ovaries collected from mature females, include thetotal number of corpora for both ovaries in the analysis
to determine ovulation rates and age relationships(George et al., 2011); corpora data should be collected
as part of future age studies; and(7) correct the labels in fig. 3 and carefully check the use of
Kasp vs 2 Kasp in the manuscript.
In the longer term, the sample size should be increased.

8.3.2 Nutritive condition
The factors considered in the models on which the analyses
in SC/F14/J13 and SC/F14/J14 are based are not derived
from biological hypotheses, but rather arise primarily fromdiscussions within the Scientific Committee. The Panel
recognises that this is a consequence of the nature of the
discussions within the Scientific Committee rather thana failure by the authors. The Panel recommends that theauthors of SC/F14/J13 and SC/F14/J14 first develop a
conceptual model of the system under consideration and usethat to identify a set of covariates to consider in the modelling.
Model selection should always be guided by underlying
knowledge of the system. It is therefore inappropriate toautomatically select the ‘best model’ because such a model
can lead to covariates being selected for which there is no
reason that there are related to response variable. Following
the selection of which factors to consider in the modelling,
the following steps should be undertaken5:

The use of the Aspartic Acid Racemisation (AAR) age
estimation method, adapted for mammals by Bada et al.
(1980), has recently regained interest for use with cetaceans.
It is useful for species where standard methods are notapplicable (e.g. species lacking teeth or readable earplugs)
as well as comparison with standard methods. SC/J14/J15
offers some new approaches to the AAR technique including
a correction for the effect of acid-hydrolysis extractions(1) identify whether any of the covariates are highly
correlated and either: (a) exclude a subset of the covariates
so that the remaining covariates are uncorrelated; or (b)develop new covariates which represent independent
aspects of the current covariates (using for example
PCA);(2) select a ‘full model’ (this may be difficult if the data set
is unbalanced) and base selection of which factors andOf course, it is also possible that abortion may have occurred naturally at amore advanced stage if the whale had not been caught.

5

4These steps are provided for the analysis of environmental and geographic
factors on measures of condition but pertain to analysis of any data.

394

report of the expert workshop to review jarpa ii

their interactions to treat as random effects - the models
should be fitted using REML and a model selection
approach such AIC, BIC or standard hypothesis testing
approach applied; and(3) select the fixed effects structure given the random
effects structure selected at step (2), where the models
are fitted using maximum likelihood; and(4) use REML to fit the best model identified in (3) above.
SC/F14/J13 and SC/F14/J14 do not report many fit
diagnostics. The Panel recommends that any revised papers
provide at least plots of the residuals versus the predictor
variables (including year and stratum), histograms ofresiduals and random effects, plots of residuals spatially, and
Q-Q plots for the ‘best model’ (although there may be more
appropriate diagnostic statistics for these analyses given theunbalanced nature of the data set).
The Panel notes too that the original estimates of thepower to detect changes in blubber thickness only accounted
for sampling error. It recommends as a more appropriate
way to assess power, use of the results of existing data
analyses to characterise both sampling error and othersources of error and to use all error when simulating data as
the basis for power calculations.
The Panel also recommends that future analyses of thedata on the condition of Antarctic minke whales include:(a) consideration of a model in which year is a categoricalvariable and is treated as a random effect if a plot of residuals
against year show there are residual patterns by year; and (b)examination of how robust the results in SC/F14/J14 are tobasing model selection on BIC rather than AIC.
Given the discussion in a number of papers presented byboth proponents and observers, the Panel offers the following
summary of the merits or otherwise of AIC and BIC, bothcommonly used metrics for model selection. The two
metrics, while mathematically similar (twice the negative
log-likelihood plus a penalty term), arise from different
underlying arguments (Kass and Raftery, 1995). BIC willtend to select simpler models than AIC if the number ofdata points exceeds 7, all things being equal (because thepenalty term for additional parameters is larger than for
AIC). Simulations have shown that AIC often selects amore complex (wrong) model over the simpler (and correct)
model (Kass and Raftery, 1995). Whether this necessarilymeans that BIC is always better than AIC is not clear
because AIC attempts to find the best approximating model
rather than true model (which would be rarely in the set ofcandidate models). BIC is, however, generally preferred to
AIC for ‘large’ data sets. However, what constitutes ‘large’in any particular case depends inter alia on the set of models
under consideration. Thus, for any one problem the selection
between AIC and BIC is seldom definitive. Many practicing
statisticians consequently often apply both AIC and BIC,
examine the sensitivity to the different models selected andapply expert judgement.
Despite the complexity of the analyses and the protracted
discussion of appropriate statistical techniques, the Panel
expresses the opinion that the ‘weight of evidence’ that thedifferent measures consistently indicated that there was an
overall decline in body condition of minke whales throughthe JARPA period, as well as the implications of such a result,
warrants careful consideration and further investigation interms of possible causes.
The Panel notes that the recommendation from theprevious review of the JARPA programme (IWC, 2008) toimprove analyses of body condition by using more accurate
proxies, such as the lipid content of the blubber has, for themost part, not been followed by the proponents, although
this may partly be due to the text not being as clear as it
might have been. Although blubber thickness has an overallcorrelation with blubber lipid content, the relation between
the two variables is poor and tends to fit an asymptotic
relationship where lipid contents fall precipitously inindividuals with thin blubber; moreover, blubber would tend
to thicken when body girth decreases (Aguilar et al., 2007,
pp.6-7) and fig.2). As a consequence of both processes,
blubber thickness would tend to underestimate lower body
condition stages. The Panel thanked the proponents forsome preliminary analyses of a small number of samples
during the Workshop but recommends that further studies
should incorporate blubber lipid content analyses for all
samples, and that the collection of current measurements
also continues to ensure comparability with past and futuredata.
The Panel further notes that several cetacean research
programmes include the collection of faeces from wild
populations to assess diet and food habits. The JARPA
II programme is in position to assess the efficacy andaccuracy of this non-lethal approach. In that regard, thePanel recommends that faecal samples (from the colon) becompared with stomach samples for species composition(and see Item 12.4).
8.3.3 Other studies of reproduction not directly relevant to
the objectives of JARPA II
The Panel welcomes papers that make maximum use of
the samples obtained even when not directly relevant to theobjectives of the programme. It made comments on two ofthese.
SC/F15/J35 is a published paper which describes some
new information on aspects of minke whale reproductive
physiology and endocrinology. It appears that it is intended
as a general addition to literature about cetacean reproductive
biology; however, the Panel noted that the in vitro maturationof whale ova a technique that is of uncertain application andthe authors do not explain how or where such technologies
could be used. The Panel had some editorial suggestions
that might have improved the paper but recognise that is nolonger appropriate.
SC/F14/J37 describes the structure and function of theplacenta of the Antarctic minke whale. The Panel notes thatsimilar literature is rare and that this paper is among themost thorough descriptions available. Since morphology is adirect expression of a species’ adaptation to the environment,
it provides a fundamental expression of how the animal
interacts with (and depends upon) that environment toachieve homeostasis. Sudden disruptions of environmental
factors can rapidly have negative consequences on survivalof individuals and on population demographic rates. Sincefunction follows form, ecologically-driven dysfunctions
are best understood once the normal morphology has been
described. In a changing environment, such an understanding
can inform conservation policy and management. For
example, with regard to paper SC/F14/J35, the transmissionof contaminants across the placenta from mother to foetus isdetermined by the type of placentation.
The Panel recommends further anatomical work on the
Antarctic minke whale for structures that have not been fully
investigated in minke or other species.

j. cetacean res. manage. 16 (suppl.), 2015

9. REVIEW OF RESULTS: Monitoring effectsof contaminants on cetaceans
9.1 Proponents’ summary of results
JARPA II has been collecting and analysing data on
contaminants in the body of whales and whale’s prey. This
component provides unique scientific information on theeffects on the contaminants on the health of whales andmovements of the contaminants in the Antarctic marine
ecosystem. The data has been utilised by the scientists
outside of the JARPA/JARPA II programmes as a valuable
source of information to the environmental analysis.
SC/F14/J23 measured total Hg levels in samples from
Antarctic krill, and Antarctic minke whales to examine thefeatures of accumulation and yearly changes of total Hg
in Antarctic Areas IV and V. Total Hg levels in Antarctic
krill from Areas IV and V were in the range 0.006-0.026
and 0.003-0.052ppm dry wt., respectively. There were no
significant differences between Areas. No yearly changes of
total Hg levels in Antarctic krill were observed in samples
from both Areas in the period from 2005 to 2011. Hg levels
in liver of Antarctic minke whales from Areas IV and V
were in the range 0.003-0.13 and <0.001-0.25ppm wet wt.,
respectively. Hepatic Hg levels of minke whales of all age
groups in Area IV decreased significantly with research
years and that of 15-26 year old whales in Area V increased
significantly.
SC/F14/J24 determined the concentrations of persistentorganochlorines in the blubber of five mature male (2125 years old) Antarctic minke whales from Antarctic Area
V sampled by the JARPA II in 2010/11. For comparison,blubber from 40 whales sampled by the JARPA surveys in
the period from 1988/89 to 2004/05, were used. In 2010/11
mean concentrations of HCB (140ng/g fat wt.) were thehighest followed by DDTs (100ng/g fat wt.), PCBs (28ng/g
fat wt.), CHLs (25ng/g fat wt.), and HCHs (0.8ng/g fatwt.). Levels of DDTs, HCHs, HCB and CHLs in Area V
decreased significantly with year, while the yearly trend of
PCBs did not change significantly. HCHs levels in minke
whales in 2010/2011 were similar to those in the JARPA
period from 1996/97 to 2004/05 however they were lower
than those in the JARPA period from 1988/89 to 1996/97.
These results suggested that levels of HCHs in the Antarctic
Ocean have varied from slightly decreasing to a steady statein the middle 1990s.
SC/F14/J25 determined total mercury (Hg) in 10 livers
and 16 muscle samples of fin whales sampled by the JARPA
II in 2005/06-2010/11. In addition PCBs, DDTs, HCHs,
HCB and CHLs were determined in two blubber samplesof fin whales sampled in 2010/11. Mean concentrations of
total Hg in livers and muscles were 0.052 and 0.021 (ppm
wet wt.), respectively. Mean concentrations of PCBs, DDTs,
HCHs, HCB and CHLs were 6.5, 13, 0.65, 39 and 4.5 (ng/g
fat wt.), respectively. Hg levels in liver and muscle of fin
and Antarctic minke whales from the Antarctic Ocean weresimilar, and these were one order of magnitude lower thanthose of fin whales from the western North Atlantic Ocean.
Geographical differences of Hg levels within a species could
be caused by food, habitat and/or age. Levels of OCs, except
for HCB, in fin whales from the Antarctic Ocean were similar
to those in the Antarctic minke whales. These levels were
extremely lower than those in fin whales from the middle
latitude of the northern hemisphere nearby human activity,
and those in killer whales from the Antarctic Ocean. These
results suggest that the OCs levels in whales could mainly beaffected by the position of their food in the food chain andarea. OCs levels in fin and Antarctic minke whales in the395

Antarctic Ocean could be the lowest among whale species in
the world. HCB levels in large whales would be affected bytrophic levels of its preys rather than by spatial differences.
Firstly, it should be noted that the pollutant analyses
summarised above were based on a limited number ofsamples from JARPA II. This was due to the fact that mostof the samples collected for this purpose were lost after thetsunami that affected Japan in 2011. The analysis of theremaining samples however, provided interesting results.
For example they confirmed the Antarctic Ocean as thecleanest area. The data on pollutant accumulation in whale
tissues and prey species in the Antarctic is an important
baseline for comparison with future research in the same ordifferent research areas.
The first examination of pollutants in the fin whalein the Antarctic showed very low levels of mercury andorganochlorines. Again the data set can be used as a baseline
for future comparisons with fin whales from other areas.
9.2 Panel’s conclusions and recommendations
The Panel welcomes the papers presenting the results
on organochlorine and heavy metal analysis in tissuesof Antarctic minke whales and the krill consumed while
recognising that, especially for organochlorine studies, theloss of samples due to the tsunami severely limited the scopeof the studies. It concurs with SC/F14/J23 and SC/F14/J24
that the results indicating very low tissue levels for bothgroups of pollutants pointed towards the lack of effects on
whale populations.
Given the low pollutant levels traditionally observedin Antarctic minke whales based on the previous JARPA
analyses, the Panel recommends that if the programme
continues, pollution studies should be given lower priority
and the resources used to address other higher priority
issues. While sampling and appropriately storing of tissue
and krill samples should continue for reference purposesin case of evidence for a new pollutant in the region or theoccurrence of a shift in trends, the Panel agrees that it would
be sufficient to undertake analyses on a suitably chosensubsample at periodic intervals (say 3-5 years).
The Panel also welcomes the research on marine debris,a type of pollution on which little information is available
for the Antarctic Ocean. It notes that information on thishad also been collected by the now discontinued SOWER
programme. This topic has also been recently identified asa priority by the IWC (2014b). The Panel also notes the
low incidence of both ingested debris and occurrence ofdebris in the sea surface. It recommends continuation ofthese observations and that the data be shared with otherinternational efforts.
10. REVIEW OF RESULTS: Model ofcompetition among whale species
While developing ecosystem models requires a substantial
data collection and analytical effort, initial progress wasmade toward this objective in JARPA II by: (a) providing
several data sets from JARPA/JARPA II as input data; (b)starting developing of two models; and (c) identifying new
data and analyses to progress further this work.
10.1 Proponents’ summary of results
SC/F14/J26 showed the progress made in the developmentof ecosystem modelling for species in Area IV. Two types ofmodelling approaches were employed; one is a multi-species
production model, and the other is the EwE, a comprehensive

396

report of the expert workshop to review jarpa ii(whole-of-ecosystem) model. There are differences in thecomponent species between them, but the baleen whales andkrill play key roles in both. For the statistical estimation in
the multi-species production model, a functional response
function was proposed to link the dynamics of predators(four baleen whale species and crabeater seal) and prey
(Antarctic krill) to both between and within-species
competitions. The model was applied to the data on timeseries of the four baleen whale species, crabeater seal andkrill, and then a likelihood function was defined based on
the data with some additional assumptions on inestimableparameters. However, at this stage the estimated population
trends for the component species did not fit well to the data,
which might be attributed partly to lack of spatial structurein the model and partly to lack of representativeness ofassumed parameters and information on prey abundance.
For the EwE approach, the construction of a mass-balancing
for Ecopath for 27 functional groups was attempted, which
could be the basis for the next stage of analysis for projection
forward using the Ecosim framework. Several suggestions
were made to progress the modelling work.
10.2 Panel’s conclusions and recommendations
10.2.1 Ecosystem modelling
The Panel notes, as did the proponents, that the modelling
work is at a preliminary stage. However, the Panel stresses
that this work is an integral part of assessing whether the
proponents can meet JARPA II Objectives 1, 2 and 4.
Furthermore, the modelling work forms a key platform
for integrating the various aspects of the overall JARPA II
programme into a coherent understanding of the ecologicalrole of studied whale species in the ecosystem. The Panel
therefore recommends that considerably more effort andresources are allocated to this aspect of JARPA II (and seeother recommendations such as those detailed under Items
6.3.4 and 7.2.2). Without this, it is not possible to state
whether the programme can meet its objectives. Such work
will also be of importance in reviewing future priorities forfield work.
10.2.1.1 Krill biomass time series (and see Item 7.2.1)

It is important to calculate the fraction of the krill stock
consumed by each species to model potential competition
between the different whale species. Trends in foodavailability to a given predator are a function of both trends
in competing predators and the available prey biomass; it isdifficult to draw conclusions on possible competition effects
without also knowing trends in prey availability. The Panel
therefore recommends that future high priority be given to
obtaining new estimates to allow an area-based time-seriesof krill biomass estimates, if JARPA II is to meet Objectives
1 and 2. The Panel also agrees with the proponents that thetime series can be extended into the past by reanalysing the
JARPA krill data with revised target strength values to makepast data compatible with more recent observations. This isdiscussed under Item 7.2 but is mentioned again here given
the importance of krill biomass trends as a critical componentof modelling whale consumption and competition.
10.2.1.2 Ecopath model

Further development of the Ecopath model may prove
useful given the limited data and biological knowledge on
the functioning of the Antarctic ecosystem. Parameterisation
should be improved, fitting to the available data where
possible. Development of an Ecopath model has thepotential to not only give an overall picture of ecosystem
links, it may prove useful to form the basis of sensitivity

analyses to highlight where uncertainty in the biologicalunderstanding and data has a high impact on the ecosystemmodels; quantifying uncertainty and examining sensitivity
is valuable in its own right because it can provide input into
defining future research needs, and informing the designof the minimum realistic models. The Panel therefore
recommends that the Monte Carlo approach developed by
Gaichas et al. (2011 to characterise uncertainty in Ecopath
results be applied. The Panel also recommends that, toaccount for observed trends in the abundance of whales in
the JARPA II study area, ‘biomass accumulation’ terms be
included in the development of a revised Ecopath.
10.2.1.3 Minimum realistic multispecies modelling

The multispecies production model described in SC/F14/J26
has the potential to model the interactions among various
whale species. However the model is currently at an earlystage in development. The Panel strongly encourages
continued development of this approach and increased
allocation of effort. The model is rather ambitious for a
first stage in model development, with four whale species
and one seal species. The level of data availability is muchgreater for minke and humpback whales than for the otherspecies, and the Panel suggests that it may be useful to start
the modelling work by constructing a model containing only
these two whale species. Adding additional predator species
would be justified if such a model can be constructed andsuccessfully fitted to data. A stated aim of the modelling is tocompare model results including and excluding competition
among species. However, all versions of the proposed multispecies model will include some degree of ‘competition’because all of the predator species in the models outlinedin SC/F14/J26 are feeding on the same finite krill resource.
Consequently, the current model structure cannot be used totest the hypotheses that competition is impacting Antarctic
minke whales. In principle, this hypothesis can be tested bycomparing the fits of the current model with that of a seriesof single-predator models. However, the Panel notes thatfitting the production model using ADMB may be difficult
for technical reasons. One possible partial solution to thisis to reduce the step length within the model (e.g. from anannual to a monthly time step).
10.2.1.4 general

The Panel recommends that simulation be used to determine
the data needed to reliably distinguish among competing
hypotheses to explain the available data, including the
proponents’ preferred option, competition among species.
Artificial datasets of known properties allow for testing ofmodel behaviour, including identifying whether the model isable to respond in predictable ways to known signals in thedata. This process can also be used to identify the required
level of precision in the data to parameterise the model.
The whale species of interest spend only a fraction of
their lives within the JARPA II area. Whale dynamics will
also be impacted by factors when they are outside thisarea, although in many cases there is unlikely to be data toparameterise such effects. The Panel also notes that there
is a time lag in the biomass dynamics (due to generational
effects), which is poorly modelled in the current production
model. This could be addressed by using a multispecies
version of the SCAA model developed for minke whales, orby introducing the lags explicitly into the production model.
The Panel emphasises that producing ecosystem models
is a long-term exercise, which requires the integration of alarge amount of data as well as ecological and biologicalknowledge of the system. The work conducted to datej. cetacean res. manage. 16 (suppl.), 2015

represents a useful start. However, the Panel recommends
increased collaboration with other researchers from outside
the JARPA II area to improve the modelling. This should
be in the form of incorporating data from adjacent areas
where possible, and by sharing expertise and knowledge
with other ecosystem and multispecies modelling experts.
Furthermore, model development should not only draw ondata from the existing elements of the JARPA II project,
but should also inform the development of the project (andfuture projects) to address data deficiencies identified during
the model development process.
The stated main aim of producing the ecosystemmodels is to provide tools to test competing hypotheses
about ecosystem functions (for example ‘that there is acompetition-driven inverse relationship between minke
and humpback whales’). The models developed must fit toavailable data to have sufficient realism to produce reliable
results. The models must also be sufficiently flexible toincorporate a range of different competing hypotheses andshould do so in a clear and explicit manner.
11. REVIEW OF RESULTS: Improve
management procedure for Antarctic
minke whales
11.1 Proponents’ summary of results
In the short term, the results of the Revised Management
Procedure (RMP) Implementation can be substantially
improved when such information as stock structure,abundance and a range of possible MSYR become available
and more accurate (contributions from Objectives 1 and
3). Knowledge of the Antarctic marine ecosystem and its
dynamics (Objectives 1 and 2) would also improve the
IWC conservation and management system, including theapplication of the ecosystem approach.
New information on stock structure and mixing andon abundance from JARPA II will be important for any
future RMP application to Antarctic minke whales, as suchinformation is essential for the RMP’s pre-Implementationassessment and Implementation Simulation Trials. JARPA II
has shown that the previous trial application of the RMP to
Antarctic minke whale in 1993 is no longer valid.
For example, the new information, particularly the stock
structure information, will render the RMP more efficient,i.e. providing extra catch without increasing conservation
risk. The 10 degree longitude slices used previously as Small
Areas are no longer supported by the JARPA II data. JARPA
II has provided new information on stock structure enablinga better definition of Small Areas.
Furthermore SCAA analyses based on JARPA II data
showed that the Antarctic minke whale can show growth
rates that are appreciably larger than the lower bound for
MSYR used in RMP trials.
The results above are consistent with the view expressedby the IWC Scientific Committee at the 1997 IWC JARPA
Mid-term Review (IWC, 1998b) which was also confirmedby the IWC JARPA Final Review Workshop (IWC, 2008):‘Finally, the results of the JARPA programme while not required
for management under the RMP, have the potential to improve
management of minke whales in the Southern Hemisphere in thefollowing ways: (1) reductions in the current set of plausible scenarios
considered in Implementation Simulation Trials; and (2) identificationof new scenarios to which future Implementation Trials will have tobe developed (e.g. the temporal component of stock structure). The
results of analysis of JARPA data could be used in this way perhaps
to increase the allowed catch of minke whales in the Southern
Hemisphere, without increasing the depletion risk about the levelindicated by the existing Implementation Simulation Trials of the
RMP for those minke whales’ (IWC, 1998b; 2008).

397

11.2 Panel’s conclusions and recommendations
The trials used to select the current Implementation of the
RMP for the Southern Hemisphere minke whales (IWC,1993) were based on the assumption that there are six stocksof minke whales in the Antarctic. The boundary between
half-Areas I-E and II-W (i.e. 60°W) was assumed to be theboundary between the ‘core’ areas for two stocks, with theboundaries of the ‘core’ areas for the remaining stocks set
by selecting five additional core area boundaries at random
from the remaining 11 half-Area boundaries, subject to theconstraint that the ‘core’ areas may not be more than four
half-Areas wide. The trials considered two assumptions
regarding mixing of stocks.
(1) The ‘concentration’ hypothesis in which a stock (theconcentration stock) is selected at random from thesix. This stock occupies its core area only. Each of its
neighbours occupies its own core area plus the adjacenthalf-Area within the first concentration stock. The stock‘opposite’ the first stock is the second concentration
stock. It also occupies its core area only. Each of its
neighbours occupies its own core area plus the adjacenthalf-Area with the second concentration stocks’ corearea.
(2) The ‘overlap’ hypothesis assumes that in each year eachof the six stocks feeds in the nearest half-Area of thetwo core areas immediately adjacent to own core area.
The trials considered MSYR values of 1% and 4%,
and set historical abundance based on estimates fromabundance from IDCR for 1979-90. No Implementation
Reviews have been conducted for the Southern Hemisphereminke whales. Moreover, the trials used to select the current
Implementation were not developed using the protocol
which has been used when conducting the Implementations
for the western North Pacific minke whales and the North
Atlantic fin whales (IWC, 2012). No trial specifications
based on the results from the JARPA/JARPA II were
presented to the Workshop.
The stock structure hypotheses on which the original
trials were based were very broad, with the aim to encompass
most likely actual situations. The information from JARPA
and JARPA II, along with other information obtained since
the original trials were developed, would be very useful if
the current protocol were to be applied, as they should refine
the set of hypotheses on which trials would be based. In
particular, the information on stock structure collected during
JARPA/JARPA II could be used to develop stock structure
hypotheses for Areas III-E to IV-W which are conditioned
on data, while the revised estimates of abundance would
be used for conditioning. The information from the SCAA
and similar analyses could be used to develop hypotheses
related to changes in carrying capacity, natural mortality,and variation in birth rates.
The operating models on which Implementation
Simulation Trials have been based have considered multiple
stocks but have not explicitly allowed for hypotheses related
to competition and ecosystem change6. In principle, thework on ecosystem modelling could be used to develop a setof operating models which allow for competition. However,
the ecosystem models need to be developed with sufficient
resolution (e.g. age- and sex-structure for minke whales).
Single-stock trials and some Implementation Simulation Trials in whichcarrying capacity, natural mortality and MSYR changed over time havebeen conducted to examine the qualitative features of these factors on theperformance of the RMP.

6

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report of the expert workshop to review jarpa ii

12. SUMMARY AND CONCLUSIONS
Before considering the individual components and papers
presented in terms of objectives, the Panel made a numberof general comments that are applicable to all aspects of the
programme.
12.1 General issues
12.1.1 General statement on specification of objectives
The Panel noted several times in its discussions that thegeneral and extremely broad nature of the objectives of theoriginal proposal and its ongoing nature made it difficult tofully review how well the programme could be said to bemeeting its own objectives:(1) ‘monitoring of the Antarctic ecosystem’;(2) ‘
modelling competition among whale species (toinform) future management objectives’;(3) ‘elucidation of temporal and spatial changes in stock
structure’; and(4) ‘improving the management procedure for Antarctic
minke whale stocks’.
The Panel welcomes the clarification of the objectives
and their inter-relationships provided by the proponents in
SC/F14/J01, but notes that within such broad and long-term
overall objectives as (1) and (2), almost any information canbe said to be contributing to them in some way; this madethe task of the Panel to assess how well the programme
was meeting its own stated objectives, somewhat difficult.
It also notes that previous Panels have encountered similardifficulties.
The Panel therefore recommends that for the benefitof any future Panel reviews as well as for the proponents’
own internal review and evaluation process, the proponents
read the guidelines for proposed permits developed by the
Scientific Committee (IWC, 2013b) after JARPA II wasdeveloped, and consider developing refined objectives
and sub-objectives with timelines for progress. This willallow performance to be more easily assessed in future andwill better enable the timing of the next ongoing review(normally, this is every six years).
The Panel also notes that the original special permit
programme also covered fin and humpback whales. It
believes that the proponents should have considered the factthat their original sampling design was not carried out more
carefully and provided information on any implications for
future sampling and the overall multi-species modelling
objective.
12.1.2 Sampling design and protocols
There are separate survey designs for the sampling andthe survey vessels. Several of the analyses such as thestatistical catch-at-age analyses rely on the assumption thatthe sampled animals are reflective of the survey estimatesof abundance. In addition, analyses of data on, for example,
blubber thickness, are more straightforward to conduct andinterpret if the sampling scheme is balanced (and see SC/F14/
O05). The Panel examined in detail the planned and realised
tracklines for some of the surveys (SC/F14/J02). The latter
tracklines often deviate from the planned tracklines due todisruption due to weather, ice conditions and increasingly
to sabotage activities by protestors. The survey protocol
states that the vessel continues along the trackline whensurveying and sampling is disrupted by weather; surveying
and sampling resumes when weather becomes conduciveto scientific study. In contrast, the response to protestors
appears often to have been to restrict the area where
sampling takes place primarily to locations close to the ice-

edge, effectively changing the sampling design and affecting
the representativeness of the sample and thus the value ofthe long-term time series and the ability to meet objectives.
The Panel is concerned that the inability to realisethe designed sampling scheme will severely compromise
the ability of the programme’s objectives to be met (someexamples of this can be seen in SC/F14/O05). While
the Panel recognises that the disruption by protestors is
outside the proponents’ control, it is clear that under these
difficult circumstances, considerable effort must be madeto ensure that the sampling remains representative rather
than only focusing on sample size (the Panel noted that thesize restrictions with respect to fin whales compromise therepresentativeness of that dataset within the research area).
The Panel recommends that an explicit protocol is developed
to specify a priori how the design is modified if disruption
by protestors occurs; simulation studies based on existingdata should assist in this. The aim of this protocol should
be to: (a) maximise the extent to which sampled animals are
representative of the survey estimates of abundance; and (b)ensure that the data remain compatible with the past datasets.
If disruption continues, consideration should be given to notstarting the survey at the same longitude in each study area
so that it is reasonable to expect that all of the study area iscovered over time. Development of protocols to account for
disruptions should be undertaken by analysts as well as bythose who conduct the surveys.
12.1.3 Integration of results from within each component of
the programme and from outside the programme
The Panel recognises that this is the first period of the
JARPA II programme but notes that the programme aroseout of the long JARPA programme and that many of thepapers analysed data from both programmes. It agrees thatthe analyses presented on the different objectives would
have benefitted considerably from better integration of all
aspects of the programme with the modelling work, in orderto have allowed better identification of key parameters andthe degree of accuracy and precision that would allow forrobust conclusions to be drawn. The Panel notes that thiswas recognised in several of the proponents’ presentations.
It recommends that this should be undertaken as a matterof priority and a more complete evaluation of uncertainty
undertaken in terms of model inputs to allow an updated
evaluation of sampling design, size, research methods andpriorities.
The Panel welcomes the considerable work that was
put into field and laboratory work and the developmentof papers. However, as is apparent from a number of its
recommendations, it is essential that further analyticalwork integrating the information available from the various
aspects of the programme is undertaken. In particular, given
the objectives of ‘monitoring changes’ and understanding
the ecosystem, the Panel stresses the importance of trying toobtain as much information as possible on the environment(oceanographic, prey related, etc.) at appropriate geographical and temporal scales, and to integrate the analysesof such data in a coherent manner. Without such efforts,
meeting stated objectives will not be possible. In thiscontext, the Panel notes that the analyses of the extensive
data collected was in many cases overly simplistic andisolated from other aspects of the programme. It strongly
recommends that considerably more effort and resources be
put into the analytical side of the programme, both via morethorough analyses of individual datasets where indicated inthis report and especially through better integration amongst
the datasets. As noted earlier this will also assist greatly inj. cetacean res. manage. 16 (suppl.), 2015

re-examining field priorities and strategy. It is already clearthat the estimation of krill abundance (within or outside
the programme) is essential to meeting ecosystem-related
objectives.
12.1.4 Archiving and use of past samples
The Panel notes the recommendation from the JARPN II
review that if whales are killed for scientific purposes, thenevery effort must be made to maximise the informationfrom them. Whilst the JARPA II programme has undertaken
considerable laboratory work, the Panel has identified anumber of additional studies that will be valuable both forthe programme itself and the wider scientific community (seerecommendations). In addition, an important component ofsuch a programme relates to archiving. It is essential that aproper tissue archiving system is in place that will allow:(1) analytical sample sizes to be increased for studies thatwere carried out on only a subset of the animals killed;
and/or(2) new analyses to be carried out as techniques improve.
Similarly, it is important to compile a relational databasethat links all components and results of analyses for eachanimal with the tissue archive.
The Panel agrees that a number of questions potentiallycould be addressed with material that may have beenpreserved from the historical commercial catch. Examples
include genetics on minke whales from outside the JARPA/
JARPA II study area, genetics on fin and other whale species,
stable isotopes on any species and area, or retrospective
pollution studies on Antarctic minke whales and other species
to better assess occurrence of temporal trends. While reports
from JARPA and JARPA II detail the availability of tissue
samples from these programmes, no similar information isavailable from the historical catch. The Panel recommends
that a detailed list of available historical samples be
produced, specifying details on species, date of sampling,
associated biological data, and preservation method (e.g.
formaldehyde, alcohol, frozen, dry, etc.).
12.1.5 Effect of catches on stocks
The effect of future JARPA II catches on the populationof minke whales in Antarctic Areas III-E to VI-W was
explored using HITTER analyses (SC/F14/J38). SC/F14/J38
suggests that catches of up to 850 animals per annum will
not substantially reduce population sizes. While the Panelagrees broadly with the conclusion that such catches will
not adversely affect Antarctic minke whales in the researcharea, it also notes that the most appropriate way to assess theimpact of future Special Permit catches on stocks is within
the framework of an RMP-type process; that approach
explicitly accounts for uncertainty in a more appropriate
way than a simple HITTER approach with sensitivity tests.
The Panel notes that the IWC Scientific Committee has notundertaken a full Implementation of Antarctic minke whales
under the current Requirements and Guidelines, as a resultof the instruction of the Commission after the establishmentof an IWC Sanctuary in the Southern Ocean.
The actual catches of fin whales were small given theestimated abundance within the research area (which does
not cover the major distribution of fin whales based on past
catches). Given this, and the abundance estimates available
for the research area, the Panel agrees that the catches would
not have affected the stock(s). No humpback whales weretaken in the first phase of the programme. Again it notes thatan RMP-type process is required to fully address questions
concerning the effect of ongoing catches on stocks.

399

12.2 Achievement of objectives
The Panel draws attention to its comments above (see Item
12.1.1) concerning the specification of the objectives andthe difficulties it faced in reaching conclusions on the firstphase of the long-term programme. In particular, it refers
to the need for additional work to assist in future reviews.
In reviewing the components of the programme, the Panelnotes the interactions between objectives and components
of work provided in Fig. 2 above. It has therefore structured
its summary based upon agenda Items 4-11, recognising thatsome of the topics address more than one of the objectives.
12.2.1 Monitoring cetacean habitat (see Item 4.2)
The monitoring of the physical and biological habitat isrelevant to all of the objectives and central to Objective 1.
The Panel notes that thus far, JARPA II has built uponsome of the work undertaken on collecting oceanographic
information but that unfortunately this work has been limitedin the most recent two years.
The Panel recognises the contribution of the longterm time series represented by the JARPA and JARPA II
programmes, particularly in the context of simultaneouscollection of whale and environmental data. However, thePanel is concerned that the necessary analytical work to
fulfil the objectives of the programme has not received theattention and resources that it deserves. If the programme isto meet its own objectives in the medium to long-term, thePanel makes the following recommendations:(1) the collection of the full suite of oceanographic data
be resumed in the coming season and the necessary
calibration work be undertaken;(2) the proponents investigate the availability of otheroceanographic and related data and incorporate thesewith the data collected from the JARPA and JARPA II
programmes to form a more comprehensive dataset;(3) the proponents develop a method to make their data
available to other international programmes;(4) the existing TDR and EPCS data are analysed to fully
describe the datasets contain and review their usefulness
in future habitat and whale analyses; and(5) concerted efforts begin to analyse the oceanographic
and other environmental data in the light of the obtained
cetacean sightings survey and biological data (moredetails are provided under the relevant agenda Items
below).
The Panel welcomes the work being undertaken on
marine debris and encourages its continuation and theregular presentation of the results to the IWC.
12.2.2 Temporal and spatial changes in stock structure (seeItem 5.2)
The work presented relates directly to Objective 3 of the
programme but is fundamental to the other objectives as well.
The Panel commends the considerable field and laboratory
effort that had gone into developing a comprehensive longterm dataset, particularly for the Antarctic minke whales,
as well as the efforts to develop analytical approaches toincorporate genetic and other markers in an integrated
analysis.
The Panel agrees that considerable progress has been
made in understanding stock structure within the researcharea (just under half of the circumpolar region) in accord
with Objective 3. However, it draws attention to the factthat the programme has incorporated little information fromoutside the research area.

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12.2.2.1 Antarctic minke whales (see Item 5.2.3)

The Panel welcomes the work represented in SC/F14/J28 and
SC/F14/J29; the latter representing the innovative integrative
approach, incorporating genetic and non-genetic data. It
made a number of detailed short-term recommendations forrevised papers, highlighting additional analyses that willassist in interpretation of the data.
The Panel made a number of medium-term recommendations to develop further the approach of SC/F14/J29. It also
recommends consideration of other potential hypotheses(e.g. isolation-by-distance along a longitudinal gradient),
recognising that management procedure (Objective 4)
considerations on stock structure focus on developing
plausible interpretations of available data, not simply the‘best’ when examining uncertainty.
Finally, the Panel recommends incorporation of
additional types of data in consideration of stock structure(and any temporal trends).
12.2.2.2 Humpback whales (see item 5.2.4)

The Panel welcomes the examination of humpback
stock structure on the feeding grounds (SC/F14/J30) by
integrating data from biopsy samples collected during
JARPA and JARPA II with other data collected within the
Antarctic and higher latitudes. This work was undertaken in
response to a recommendation from the JARPA review andmakes a valuable contribution to the in-depth assessments ofSouthern Hemisphere humpback whales.
The Panel made some detailed recommendations for arevised paper.
12.2.2.3 Fin whales (see item 5.2.5)

The Panel welcomes the genetic analyses presented in SC/
F14/J32 on fin whales, although the sample size was small.
The study incorporated samples from the catch as well as
biopsy samples collected during JARPA and JARPA II.
However, the Panel has some concerns about some aspectsof the analyses and interpretation. The Panel made somedetailed recommendations for a revised paper. In the longer
term, it suggests increasing the sample size from existingdata by using biopsy samples collected by other programmes(e.g. SOWER) and from earlier commercial whaling.
12.2.2.4 Southern right whales (see item 5.2.6)

The Panel welcomes the genetic analyses presented in SC/
F14/J33 with biopsy samples primarily collected in Area
IV. The study provided the first attempt to geneticallycharacterise southern right whales on the feeding grounds
and provided indications that at least some southern rightwhales may travel further than previously thought. This
information will contribute to future in-depth assessmentsof this species.
12.2.3 Monitoring abundance, trends and distribution (seeItem 6.3)
12.2.3.1 Abundance and trends from survey data
(see Item 6.3.1)

This information is related to Objectives 1, 2 and 4 and is toa considerable degree dependent upon information obtained
under Objective 3.
There were two primary analytical approaches: sightings
data analyses and statistical catch-at-age analyses (SCAA).
The Panel welcomes the presentation of survey resultsfrom the JARPA and JARPA II programmes. It agrees
that this information is contributing significantly to theobjectives of the programme. The survey work represents
a considerable expenditure of research time and a large
dataset for long-term monitoring. It complements the workof the IDCR/SOWER programme that has now finished.

The importance of monitoring trends in abundance incetacean species is of general conservation and management
importance especially in the context of documenting therecovery of species/populations that had been extensively
depleted by commercial whaling, as well as investigating
variation in species mix compared to the period prior to
exploitation.
The JARPA review meeting (IWC, 2008) had
examined the survey information available at that timeand had identified a number of items that required further
investigation before acceptable abundance estimates could
be obtained. The Panel agrees that the papers on Antarctic
minke whales (SC/F14/J03) and humpback whales (SC/F14/J04) had adequately addressed most of these concernsand had in some cases identified further work that would
be undertaken - e.g. see the discussion in Matsuoka et al.
(2011) and Hakamada et al. (2013).
The Panel made a number of detailed recommendations(Item 6.3.1) to develop a revised stand-alone paper for the‘other’ baleen whale papers. In terms of future field work,
the Panel recommends: (1) consideration of the use of
Independent Observer mode in future surveys to address theissue of g(0); and (2) the collection of killer whale ecotype
data.
12.2.3.2 SCAA analyses (see Item 6.3.2)

The Panel notes that the SCAA analyses, whilst using datafrom JARPA and JARPA II, have been directed by the
Scientific Committee and undertaken by non-proponent
scientists (primarily Punt) over many years. The Panelwelcomes the updated SCAA analysis presented. It agrees
that the SCAA model is both the best currently available
model for examining stock dynamics for the minke
whales in the JARPA II area, and that the model performs
well in this regard. The Panel made a number of detailed
recommendations for improvements and further work. The
Panel also notes that certain results from the SCAA model
may not be consistent with inferences developed from othercomponents of JARPA II or may suggest potential revisions
to the design of JARPA II itself. These points concerned interalia MSYR, stock structure and growth rate changes. They
should be considered by the proponents once the revised
SCAA analyses have been undertaken.
12.2.3.3 alternative approaches to estimating
abundance (see item 6.3.3)

The Panel welcomes the preliminary evaluation of the use ofpaternity analysis to estimate abundance of Antarctic minke
whales (SC/F14/J07). It made two suggestions regarding
further work on this matter.
12.2.3.4 Distribution (see Item 6.3.4)

Examining and understanding the distribution of whale
species and the reasons for any changes in distribution iscentral to Objectives 1 and 2. The Panel welcomes the work
undertaken in this regard, primarily based upon the sightingssurvey data and recognises the attempts to quantify changes
in distribution (primarily of minke whales and humpback
whales) over time within the research area. That being said,
the Panel recommends that more robust and comprehensive
analyses of the existing data be undertaken that incorporate
many more potential explanatory variables. In particular, amore rigorous area occupancy analysis should be undertakenthat incorporates recent advances in spatial modelling. The
Panel agrees that much greater emphasis be placed upon thisanalysis in the short-term (1-2 years) in order to determine
the extent to which objectives are being met and to determine
whether improvements are required.

j. cetacean res. manage. 16 (suppl.), 2015

The distributional information has been used by the
proponents to formulate their ‘competition’ hypotheses(Objective 2). The Panel agrees that considerably morework is needed to develop a conceptual model for suchcompetition. This requires additional field effort to thatcurrently being undertaken in order to develop plausible
hypotheses for use in ecosystem modelling that are essential
to meet Objective 2. The Panel recommends that the
proponents invest effort in some and preferably all of thefollowing:(1) analysis of proximity of whale species in the sighting
survey tracklines – this can be done with existing dataand initial analyses should be possible within the nextyear;(2) focused studies of prey swarming behaviour and density
in relation with local whale distribution and abundance– this may require a change of fieldwork focus for one
or more seasons and integrated data from focal follows,
telemetry (radio and/or satellite), krill abundance anddensity estimates and biopsy sampling (for inter alia
stable isotope analysis);(3) comparison of prey in stomach contents in areas where
both species overlap in distribution, and in areas where
they segregate, including investigation of whether
whale species and krill length-maturity classes exhibit
distinct spatial segregation in their distribution patterns– this can begin with existing samples and it should bepossible to present a progress report within the next two
years; and(4) stable isotope analysis from tissue samples – initial
examination of the discriminatory power of suchanalyses can occur with existing samples.
12.2.4 Monitoring krill abundance and feeding ecology

12.2.4.1 Krill abundance (and see items 4.2 and 7.2.1)

Monitoring krill abundance is fundamental to Objectives 1
and 2 of the programme. While welcoming the work that has
been done in JARPA and at the beginning of JARPA II (seeItem 4.2), the Panel is concerned that insufficient recent
effort has been placed upon this component of the work. The
Panel has made a number of recommendations both withrespect to the analysis of existing data and the collection of
future data. In particular, the Panel recommends that futurekrill surveys should be frequent because the density of krill
in any given stratum may vary significantly from year toyear, and the objectives of JARPA II require a time-series
view of how the prey field is changing over time. The JARPA
II study area is very large, and it is probably not possible tosurvey the entire study area every year with a single survey
vessel. Thus, the Panel recommends either using multiple
survey vessels to synoptically cover the JARPA II study area
every 1-3 years or using one vessel to survey alternating
halves of the study area every year (similar to the approachtaken for sighting surveys).
12.2.4.2 feeding ecology (and see item 7.2.2)

Understanding feeding ecology is central to achieving
Objective 2 and is relevant to Objective 1.
The Panel welcomes the considerable amount of field
and laboratory work undertaken on this subject and analysed
in SC/F14/J15. However, it identified a number of important
shortcomings in the work originally presented, in particular
related to quantification of uncertainty. The Panel agrees
that the approach developed by the proponents during theWorkshop to address these shortcomings (SC/F14/R01)
is both a positive development and a useful way forward;
however, without the results of a thorough analysis, the401Panel is unable to determine whether JARPA II has provided
or will provide consumption estimates within a sufficiently
narrow range. The Panel recommends that work proposedin SC/F14/R01 be further developed and allocated highpriority. Ideally a new paper should be submitted to the next
meeting of the Scientific Committee. As a minimum, thisshould advance the outlined work plan by including in theMonte Carlo simulations, uncertainty in:(1) r (the ratio of low/high feeding intake) and the length of
the feeding season for Method 1; and(2) the extent of night feeding for Method 2.
The Panel also recommends that the work is extended bycomputing a time series of Monte Carlo results for the total
potential consumption of krill using abundance estimatesof minke whales (preferably those estimated by the SCAA
model) and the uncertainties around these estimates.
These analyses are an essential consideration as to theextent to which the JARPA II programme in its present
form is able to meet its objectives. The Panel therefore
recommends that the results of the Monte Carlo work
be used to re-evaluate the future research and sampling
in JARPA II. Details are provided under Item 7.2.3. Thisapproach could also be used both to aid determination of
future sample sizes by estimating how many samples would
be needed to reduce uncertainties in consumption estimatesby a desired level and to assess whether lethal or non-lethal
sampling or a combined approach represents best way of
reducing such uncertainties.
The Panel agrees that results presented in SC/F14/J16(figs 4-6) indicate substantial overlap in the sizes of krillthat are consumed by minke whales and fin whales whensampled near the same locations and times, noting alsothat the sample size for fin whales is small, restricted tosmaller animals and outside the species main distribution
area. However, the Panel cautions the proponents over any
over-interpretation that this proves that there is ‘interspecificinteraction’ or competition between these species in any
limiting sense. The information provided in SC/F14/J16
suggests that together, minke and fin whales might haveconsumed about 6% of the standing biomass of krill and thusthat total krill biomass per se was not limiting.
The Panel reiterates that additional field work is needed(see Item 12.2.3.4) to develop plausible hypotheses in order
to meet Objective 2.
12.2.5 Monitoring whale biological parameters (and see
Item 8.3)
The Panel recognises the considerable field and laboratory
effort involved in the papers on biological parameters. This
work is relevant to Objective 1 and to Objective 4 and isdependent upon consideration of stock structure (Item 5). It
is also dependent on the representativeness of the sampling
design (see the JARPA review (IWC, 2008) and Item 12).
The Panel notes that the analyses presented on biologicalparameters assume: (a) that the sampling scheme is broadly
representative of the population; and (b) related to the stock
structure hypotheses agreed at the JARPA review rather thanthose that were subsequently presented to this meeting (Item5). With respect to sampling design in JARPA II, the Panelnotes the disruption caused to the proposed sample size anddesign due to outside interference in the Antarctic and refers
to the discussions on sample size and sampling design at the
JARPA review meeting (IWC, 2008) and in the papers by dela Mare and colleagues at this meeting (see Item 8.2). The
Panel’s overall comments on sampling design/size within
JARPA II are given under Item 12.1.2.

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12.2.5.1 Age at attainment of sexual maturity/
pregnancy rates (and see items 8.3.1.1 and 8.3.1.2)

The Panel welcomes the information provided on theseimportant topics, recognising the amount of field andlaboratory work involved and the analytical work inestimating ageing errors. However, it cautions the proponents
against over-interpretation of the data at this stage. It
recommends additional analytical work to be undertaken
before conclusions can be drawn as detailed under Items
8.3.1.1 and 8.3.1.2.
Recommendations for future field and laboratory work
are made under Item 8.3.1.2.
12.2.5.2 Age estimation (and see item 8.3.1.3)

The Panel welcomes the work presented on the evaluationof the use of aspartic acid racemisation as an alternative
approach to estimating age and made a number ofrecommendations for an updated paper and future work.
12.2.5.3 Nutritive condition (and see item 8.3.2)

The Panel recognises that the factors considered in themodels on which the analyses in SC/F14/J13 and SC/F14/J14 are based are not derived from biological hypotheses, butrather arise primarily from discussions within the ScientificCommittee. The Panel recommends that the authors of SC/
F14/J13 and SC/F14/J14 first develop a conceptual model ofthe system under consideration and use that to identify a setof covariates to consider in the modelling. Model selection
should always be guided by underlying knowledge of thesystem. It is therefore inappropriate to automatically selectthe ‘best model’ because such a model can lead to covariates
being selected for which there is no reason that there are
related to response variable. The Panel made a series ofsuggestions and recommendations as how best to take thiswork forward.
Despite the complexity of the analyses and the protracted
discussion of appropriate statistical techniques, the Panel
expresses the opinion that the ‘weight of evidence’ (i.e. thedifferent measures consistently indicated that there was an
overall decline in body condition of minke whales throughthe JARPA period), as well as the implications of such a
result, warrant careful consideration in terms of cause.
Following the discussion at the JARPA review meeting(IWC, 2008) and the advantages of considering lipid content
of the blubber as well as thickness, the Panel recommends
that any further studies should incorporate blubber lipid
content analyses for all samples, and that the collection ofcurrent measurements also continues to ensure comparability
with past and future data. The Panel also recommends that
faecal samples (from the colon) be compared with stomach
samples for species composition.
12.2.6 Monitoring the effects of contaminants (see Item 9.2)
The Panel welcomes the papers presenting the results
on organochlorine and heavy metal analysis in tissuesof Antarctic minke whales and the krill consumed while
recognising that, especially for organochlorine studies,
the loss of samples due to the tsunami severely limited thescope of the studies. It concurs with SC/F14/J23 and SC/
F14/J24 that the results indicating very low tissue levels forboth groups of pollutants pointed towards the lack of effects
on whale populations. Given the low levels observed, thePanel recommends lower priority for pollutant studies in
the future and agrees that it would be sufficient to undertake
pollutant analyses on a suitably chosen subsample at periodic
intervals (say 3-5 years).
The Panel also welcomes the research on marine debris,a type of pollution on which little information is available

for the Antarctic Ocean. It notes that information on thishad also been collected by the now discontinued SOWER
programme. This topic has also been recently identified asa priority by the IWC (IWC, 2014b). The Panel also notes
the low incidence of both ingested debris and occurrenceof debris in the sea surface. It recommends continuationof these observations and that the data be shared with otherinternational efforts.
12.2.7 Model of competition among whale species (see Item
10.2)
12.2.7.1 Ecosystem modelling (and see Item 10.2.1)

The Panel recognises that the modelling work is at apreliminary stage. The Panel stresses the fundamental
importance of this work to most of the objectives of JARPA
II and an understanding of the role of the studies species
in the ecosystem. The Panel therefore recommends thatconsiderably more effort is allocated to this aspect of JARPA
II (and see other recommendations such as those detailed
under Items 6.3.4 and 7.2.2). Without this, it is not possible
for the Panel to state whether the programme can meet its
objectives. Such work is also of importance in reviewing
future priorities for field work.
12.2.7.2 Krill biomass time series (and see Item
10.2.1.1)

Knowledge of krill biomass (and trends thereof) are central
to the JARPA II programme. Recognising the difficulties
posed due to outside disruption, the Panel recommends
that future high priority be given to obtaining new estimatesto allow an area-based time-series of estimates of krill
biomass, if JARPA II is to meet Objectives 1 and 2. The
Panel also agrees with the proponents that the time series
can be extended into the past by reanalysing the JARPA krill
data with revised target strength values to make past datacompatible with more recent observations (and see Item
7.2).
12.2.7.3 Ecopath model (and see Item 10.2.1.2)

The Panel notes that further development of the Ecopath may
prove useful given the limited data and biological knowledgeof the Antarctic ecosystem and made recommendations toimprove the approach.
12.2.7.4 Minimum realistic multispecies modelling(and see Item 10.2.1.3)

The Panel welcomes the preliminary work undertaken andstrongly recommends further development of the approachwith increased allocation of effort. The Panel made a numberof suggestions to further this approach.
12.2.7.5 General (and see Item 10.2.1.4)

The Panel recommends that simulation be used to determine
the data needed to reliably distinguish among competing
hypotheses to explain the available data, including the
proponents’ preferred option, competition among species.
This may be used to identify: (1) whether models are able
to respond in predictable ways to known signals in thedata; and (2) the required level of precision in the data toparameterise them.
The Panel emphasises that producing ecosystem models
is a long-term exercise, which requires the integration of alarge amount of data as well as ecological and biologicalknowledge of the system. The work conducted to daterepresents a useful start. However, the Panel recommends
increased collaboration with other researchers from outside
the JARPA II area to improve the modelling exercise.
The stated main aim of producing the ecosystemmodels is to provide tools to test competing hypotheses
about ecosystem functions (for example ‘that there is a

j. cetacean res. manage. 16 (suppl.), 2015

competition-driven inverse relationship between minke
and humpback whales’). The models developed must fit toavailable data to have sufficient realism to produce reliable
results. The models must also be sufficiently flexible toincorporate a range of different competing hypotheses andshould do so in a clear and explicit manner.
The Panel made a number of suggestions to further themodelling work.
12.2.8 Improved management procedure for Antarctic
minke whales (see Item 11.2)
The Panel agrees that the information from JARPA and
JARPA II, particularly with respect to stock structure andabundance will greatly improve any future Implementation
Simulation Trials, should these ever be requested by theCommission. Similarly, SCAA and related analyses could
be used to develop hypotheses related to carrying capacity,
natural mortality and variation in birth rates. In principle, thework on ecosystem modelling could be used to develop a setof operating models which allow for competition. However,
the ecosystem models need to be developed with sufficient
resolution (e.g. age- and sex-structure for minke whales).
12.3 The relationship of the research to IWC
Resolutions and discussions
The Panel briefly considered this item. The Commission
has passed a number of Resolutions on matters related toecosystem research and climate change. Resolution 199413 (IWC, 1995) encouraged Contracting Governments andthe Scientific Committee to study environmental changes
and impact on cetaceans. Resolution 1995-10 (IWC, 1996)
encouraged Contracting Governments to study the effects ofpollutants on cetaceans as recommended by the ScientificCommittee’s Workshop on the topic. Resolution 1997-7(IWC, 1998a) encouraged Contracting Governments tocontinue to provide information on environmental changes
and potential effects on cetaceans. Resolution 1999-4 (IWC,
2000a) requested Contracting Governments to provide
the Scientific Committee with data on contaminants incetaceans.
The Panel agrees that a number of aspects of the
JARPA II programme are relevant to these Resolutions.
The submission of the present Workshop papers fulfils therequest that these types of analyses be submitted to the IWC.
Submitting revised or new papers that incorporate the Panelrecommendations will provide better information related to
the Resolutions and discussions.
In addition to the work related to ecosystems andenvironmental change discussed above, the Panel agrees
that the work on stock structure and abundance, in addition
to information from photo-identification studies is directly
relevant to the Scientific Committee’s work on the in-depth
assessments of the Antarctic minke whale, humpback, blue
and right whales.
12.4 Utility of lethal and non-lethal techniques
The Panel stresses that its primary task at this meeting wasto provide an objective scientific review of results of the
JARPA II programme thus far including consideration oflethal and non-lethal methods; its task was not to provide
either general condemnation or approval of lethal sampling
research under scientific permit in general - considerationof this would require discussion of a number of issues
well beyond the purview of a scientific panel - e.g. see theapproach identified in Waples and Plummer (2009).
The Panel noted the more extensive discussion of thisissue at the Workshops to review the JARPN II programme

403(IWC, 2010) and Icelandic programme (IWC, 2014a),
including a table summarising the actual and potential
contributions of various lethal and non-lethal techniques
as applied to North Atlantic minke whales. These reports
provided a good review of the strengths and weaknessesof the then available non-lethal techniques for studies on
the following topics that are also relevant to the JARPA II
Research Programme: feeding ecology; pollutant studies;and stock structure including genetic studies. The Panelalso noted the more recent review undertaken as part of the
SORP (Southern Ocean Research Partnership) programme.
The Panel has not repeated the information from thosereviews again here but took them into account during its
deliberations.
Although a comparison of the utility of lethal andnon-lethal sampling is not an objective of the JARPA II
programme, the samples and data already collected toachieve the objectives can be analysed to investigate thisgeneral research question. This would also be consistent withthe approach of JARPA II to use both lethal and non-lethal
techniques; review of the effectiveness of both lethal andnon-lethal methods in a quantitative manner will certainly
benefit periodic review of the long-term programme.
Therefore, the Panel recommends that the proponents
examine the approaches for comparison used in the Icelandic
programme be conducted and develop an approach toformally and objectively compare the results from different
approaches in the light of the programme’s objectives. Morespecifically, conclusions about diet and nutritional statusbased on analysis of stomach contents could be compared
to conclusions drawn from analysis of fatty acids or stable
isotope ratios based on internal tissues and organs andtissue derived from mimicked biopsies (skin and outermost
blubber). This can then be used by the proponents and future
Panels to better address this issue in a quantitative way.
Under Item 8.3.2, the Panel recommended collecting
faeces to conduct DNA and diet composition analyses.
These samples could also be used in the comparison studyof lethal and non-lethal techniques.
13. ADOPTION OF REPORTThe Panel completed the primary work on its report andrecommendations at the Workshop and subsequently byemail prior to its submission to the proponents on 11 April
2014. Final editing took place in the period following 11
April by email. The final edited report was adopted later in
April 2014.
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Annex A
List of Participants
Panel members
Alex Aguilar
University of Barcelona, Spain
Greg Donovan
IWC Secretariat, Cambridge, UK
Jaume Forcada
British Antarctic Survey, Cambridge,
UK
Daniel Howell
Institute of Marine Research, Bergen,
NorwayDebra Palka
NOAA, USA
André Punt
University of Seattle, USA
Glenn Stauffer
Penn State University, Pennsylvania,
USA
Robin Waples
NOAA, USA
George Watters
NOAA, USA

Proponents
Fisheries Agency of Japan, Japan
H. Moronuki
N. Okazoe
National Research Institute of Far
Seas Fisheries, JapanT. Kishiro,
H. Maeda
T. Miyashita
J. Morishita
H. Murase
National Research Institute ofFisheries Science, JapanT. Watanabe
Tokyo University of Marine
Science and Technology, Japan
H. KatoT. Kitakado
Institute of Cetacean Research,
Tokyo, Japan
T. Bando
D. Goodman
T. Hakamada
H. Kanda

K. Konishi
T. MogoeS. Nishiwaki
L. Pastene
T. Tamura
A. WadaInterpreters
S. Baba
H. Yasokawa
Observers
Australia
W.K. de la Mare
M. Double
NorwayL. Walløe
USA
P. WadeUniversity of Cape Town
D.S. Butterworth

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Annex B
Agenda
1.

2.
3.
4.

5.

6.

7.

Introductory items
1.1. Chair’s opening remarks
1.2
Meeting arrangements and work schedule
1.3
Review of documents and data (SC/F14/J01J38, SC/F14/O01-O08)
Adoption of the Agenda
Proponents’ overview of the research objectives andresults (SC/F14/J01-J02)
Review of results: monitoring cetacean habitat (SC/F14/J20-J22)
4.1
Proponents’ summary of results
4.2
Panel’s conclusions and recommendations
Review of results: elucidate temporal and spatialchanges in stock structure (SC/F14/J27-J34)
5.1
Proponents’ summary of results
5.2
Panel’s conclusions and recommendations
Review of results: monitoring whale abundance trends(SC/F14/J03-J07, SC/F14/J17-J18; SC/F14/O01-O02)
6.1
Proponents’ summary of results
6.2
Panel’s conclusions and recommendations
Review of results: monitoring krill abundance andfeeding ecology of whale stocks (SC/F14/J13-J16, SC/
F14/J19, SC/F14/O03, SC/F14/O05-O07)
7.1
Proponents’ summary of results
7.2
Panel’s conclusions and recommendations

8.

Review of results: monitoring whale biologicalparameters (SC/F14/J08-J12, SC/F14/J35-J37)
8.1
Proponents’ summary of results
8.2
Panel’s conclusions and recommendations
9. Review of results: monitoring effects of contaminantson cetaceans (SC/F14/J23-J25)
9.1
Proponents’ summary of results
9.2
Panel’s conclusions and recommendations
10. Review of results: model of competition among whale
species (SC/F14/J26)
10.1 Proponents’ summary of results
10.2 Panel’s conclusions and recommendations
11. Review of results: improve management procedure for
Antarctic minke whales
11.1 Proponents’ summary of results
11.2 Panel’s conclusions and recommendations
12. Summary and conclusions (SC/F14/J01, SC/F14/J38,
SC/F14/O04, SC/F14/O08)
12.1 Achievement of the objectives
12.2 Other contributions to important research needs
12.3 The relationship of the research to IWC
Resolutions and discussions
12.4 Utility of lethal and non-lethal techniques
13. Adoption of Report

Annex C
List of Documents
Primary documents
SC/F14/J
01. Pastene, L.A., Fujise, Y. and Hatanaka, H. The Japanese
Whale Research Program under Special Permit in the
Antarctic (JARPA II): origin, objectives, research
progress made in the period 2005/06-2010/11, andrelevance for management and conservation of whales
and the ecosystem.
02. Nishiwaki, S., Ishikawa, H., Goto, M., Matsuoka, K.
and Tamura, T. Review of general methodology andsurvey procedure under the JARPA II.
03. Hakamada, T. and Matsuoka, K. Estimates of abundance
and abundance trend of the Antarctic minke whale in
Areas IIIE-VIW, south of 60°S, based on JARPA and
JARPA II sighting data (1989/90-2008/09).
04. Hakamada, T. and Matsuoka, K. Estimates of abundance
and abundance trend of the humpback whale in Areas
IIIE-VIW, south of 60°S, based on JARPA and JARPA
II sighting data (1989/90-2008/09).
05. Matsuoka, K. and Hakamada, T. Estimates of abundance
and abundance trend of the blue, fin and southern rightwhales in Areas IIIE-VIW, south of 60°S, based on
JARPA and JARPA II sighting data (1989/90-2008/09).
06. Hakamada, T. and Matsuoka, K. Estimates of abundance
and abundance trend of the sperm, southern bottlenoseand killer whales in Areas IIIE-VIW, south of 60°S,
based on JARPA and JARPA II sighting data (1989/902008/09).

07. Kanda, N., Goto, M. and Pastene, L.A. Paternity
analysis on Antarctic minke whales using JARPA and
JARPA II samples.
08. Bando, T., Kishiro, T. and Kato, H. Yearly trend in
the age at sexual maturity of Antarctic minke whales
examined by transition phase in earplugs collectedduring JARPA and JARPA II surveys.
09. Bando, T. and Hakamada, T. Yearly trend in the proportionof pregnant animals among mature female Antarctic
minke whales in the JARPA and JARPA II period.
10. Mogoe, T., Bando, T., Maeda, H., Kato, H. and Ohsumi,
S. Biological observations of fin whales sampled by
JARPA II in the Antarctic.
11. Kitakado, T., Lockyer, C. and Punt, A.E. A statisticalmodel for quantifying age-reading errors and its
application to the Antarctic minke whales.
12. Yasunaga, G., Bando, T. and Fujise, Y. Preliminary
estimation of the age of Antarctic minke whales basedon aspartic acid racemization.
13. Konishi, K. and Walloe, L. Time trends in the energy
storage in the Antarctic minke whales during the JARPA
and JARPA II research periods.
14. Konishi, K., Hakamada, T., Kiwada, H., Kitakado, T.
and Walloe, L. Decrease in stomach contents in the
Antarctic minke whale (Balaenoptera bonaerensis) in
the Southern Ocean [this paper cannot be cited except in
the context of IWC meetings until is formally publishedin Polar Biology].

j. cetacean res. manage. 16 (suppl.), 2015

15. Tamura, T. and Konishi, K. Prey composition andconsumption rate by Antarctic minke whales based on
JARPA and JARPA II data.
16. Tamura, T. Preliminary analyses on prey consumption
by fin whales based on JARPA II data.
17. Matsuoka, K. and Hakamada, T. Distribution pattern ofwhale species sighted in the Antarctic based on JARPA
and JARPA II sighting surveys (1987/88-2008/09).
18. Murase, H., Matsuoka, K., Hakamada, T. and Kitakado,
T. Preliminary analysis of changes in spatial distributionof Antarctic minke and humpback whales in Area IV
during the period of JARPA and JARPA II from 1989
to 2006.
19. Wada, A. and Tamura, T. Estimation of krill biomass
based on JARPA II acoustic surveys.
20. Watanabe, T., Okazaki, M. and Matsuoka, K. Results ofoceanographic analyses conducted under JARPA and
JARPA II and possible evidence of environmental changes.
21. Naganobu, M., Matsuoka, K., Murase, H. andKutsuwada, K. Consideration on the Kerguelen-Davis
Oscillation Index (KDOI) influencing variability on
environmental ecosystem in the Prydz Bay Region, east
Antarctic: data exploration.
22. Isoda, T., Tamura, T., Nishiwaki, S and Pastene, L.A.
Observation of marine debris in the Antarctic based on
JARPA and JARPA II data.
23. Yasunaga, G., Bando, T. and Fujise, Y. Pattern of
mercury accumulation in the Antarctic minke whale andits prey based on JARPA II data.
24. Yasunaga, G., Bando, T. and Fujise, Y. Pattern oforganochlorine accumulation in the Antarctic minke
whale based on JARPA II data.
25. Yasunaga, G. and Fujise, Y. A note on mercury andorganochlorine accumulation in the Antarctic fin whalebased on JARPA II data.
26. Kitakado, T., Murase, H., Tamura, T. and Yonezaki, S. An
attempt to ecosystem modelling for species in Area IV
in the Antarctic Ocean using JARPA and JARPA II data.
27. Kanda, N., Goto, M., Oikawa, H. and Pastene, L.A. A
note on sampling and laboratory procedure protocols of
the genetic work at the Institute of Cetacean Research.
28. Pastene, L.A., Goto, M. and Kanda, N. An update of thegenetic study on stock structure of the Antarctic minke
whale based on JARPA II samples.
29. Kitakado, T., Schweder, T., Kanda, N., Pastene, L.A.
and Walloe, L. Dynamic population segregation by
genetics and morphometrics in Antarctic minke whales.
30. Pastene, L.A., Kitakado, T., Goto, M. and Kanda, N.
Mixing rates of humpback whales from Stocks D,
E and F in the Antarctic feeding grounds based on
mitochondrial DNA analyses [SC/65a/SH13; this paper
can be cited only in the context of the IWC meetings].
31. Kanda, N., Goto, M. and Pastene, L.A. Stock structureof humpback whales in the Antarctic feeding grounds asrevealed by microsatellite DNA data.
32. Goto, M., Kanda, N. and Pastene, L.A. Genetic analysis
on stock structure of fin whales in the Antarctic basedon mitochondrial and microsatellite DNA.
33. Kanda, N., Goto, M., Nishiwaki, S. and Pastene, L.A.
Long-distance longitudinal migration of southern rightwhales suspected from mtDNA and microsatellite DNA
analysis on JARPA and JARPA II biopsy samples.
34. Matsuoka, K. and Pastene, L.A. Summary of photoidentificationentification information of blue, southern
right and humpback whales collected by JARPA/
JARPA II.
35. Nagai, H., Mogoe, T., Ishikawa, H., Hochi, S., Ohsumi,
S. and Fukui, Y. 2007. Follicle size-dependent changes

407

in follicular fluid components and oocyte diameter in
Antarctic minke whales (Balaenoptera bonaerensis).
Journal of Reproduction and Development 53(6): 12651272.
36. Ono, N., Yamaguchi, T., Ishikawa, H., Arakawa, M.,
Takahashi, N. Saikawa, T. and Shimada, T. 2009.
Morphological varieties of the Purkinje fiber network
in mammalian hearts, as revealed by light and electron
microscopy. Arch Histol Cytol. 72(3), 139-149.
37. Sasaki, M., Amano, Y., Hayakawa, D. Tsubota, T.,
Ishikawa, H., Mogoe, T., Ohsumi, S., Tetsuka, M.,
Miyamoto, A., Fukui, Y., Budipitojo, T. and Kitamura,
N. Structure and steroidogenesis of the placenta in the
Antarctic minke whale (Balaenoptera bonaerensis).
Journal of Reproduction and Development 59 (2):159167.
38. Hakamada, T. An examination on the effect on thestocks of JARPA II catches.
‘O’ Documents
SC/F14/O
01. Punt, A. A summary history of the application of statisticalcatch-at-age analysis to Antarctic minke whales.
02. Punt, A. Assessment of Antarctic minke whales using
statistical catch-at-age analysis.
03. Leaper, R. Summary of previous Scientific Committee
discussions relating to minke whale consumption rateestimates in SC/F14/J15.
04. Vikingsson, G. and Pampoulie, C. Comments on thescientific outputs of the first phase of the JARPA II
Special Permit program.
05. Wotherspoon, S., Double, M., McKinlay, J., Candy, S.,
Andrews-Goff, V., de la Mare, W.K. JARPA and JARPA
II cannot monitor trends in the Antarctic ecosystem due
to flawed sampling strategies.
06. de la Mare, W.K., Candy, S., McKinlay, J., Wotherspoon,S., Double, M. What can be concluded from the statisticalanalyses of JARPA/JARPA II body condition data?
07. de la Mare, W.K., Candy, S., McKinlay, J., Wotherspoon,S., Double, M. JARPA II sample size calculations ignore
spatial and temporal variability.
08. Double, M., de la Mare, W.K. and Gales, N.G.
Observers’ statement to the JARPA II Special Permit
Expert Panel Review Workshop.
Response documents
SC/F14/R
01. Tamura, T. A response to document SC/F14/O03‘Summary of previous Scientific Committee discussions
relating to minke whale consumption rate estimates in
SC/F14/J15’ by R. Leaper.
02. Konishi, K., Hakamada, T. and Pastene, L. Response to
document SC/F14/O05 ‘JARPA and JARPA II cannot
monitor trends in the Antarctic ecosystem due to flawed
sampling strategies’ by S. Wotherspoon et al.
03. Hakamada, T. and Konishi, K. Response to Document
SC/F14/O07 ‘JARPA II sample size calculations ignore
spatial and temporal variability’ by W. K. de la Mareet al.
04. Pastene, L. A response to Australia’s observers’
statement (SC/F14/O08) to the JARPA II Special Permit
Expert Panel Review Workshop
05. Konishi, K. and Walløe, L. Response to the paper SC/
F14/O06: ‘What can be concluded from the statisticalanalyses of JARPA/JARPA II body condition data?’by William de la Mare, Steven Candy, John McKinlay,
Simon Wotherspoon and Michael Double.

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report of the expert workshop to review jarpa ii

Annex D
AnnexbyD tablesData Produced
JARPA II that were Available for the[Text table 1]
Review Workshop

Abundance estimate several species1
Seasons
1. Angle and distance experiments
2. Ice edge line
3. Effort data
4. Weather data
5. Sighting Antarctic minke whale
6. Sighting fin whale
7. Sighting humpback whale
8. Sighting blue whale
9. Sighting southern right whale
10. Sighting sperm whale
11. Sighting S bottlenose whale
12. Sighting[Texttablekiller2] whale

Sample size

2005/06-2010/11
2,617 tests
2005/06-2010/11
4,234 points
2005/06-2010/11 43,161 activities
2005/06-2010/11 34,694 records
2005/06-2010/11
7,344 sch.
2005/06-2010/11
605 sch.
2005/06-2010/11
4,570 sch.
2005/06-2010/11
146 sch.
2005/06-2010/11
150 sch.
2005/06-2010/11
894 sch.
2005/06-2010/11
310 sch.
2005/06-2010/11
352 sch.

Ecological data (oceanographic, marine debris, krill)
Seasons

Sample size

13. Temperature (XBT)
14. Temp. salin. (XCTD)
15. Temp. salin. (CTD)
16. Temp. salin. (EPCS)
17. Marine debris (stomach)2
18. Marine debris (sea surface)2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11

19. Echo sound (krill abund./dist.)
20. IKMT net
[Text
table
3]krill
21. Body
length2007/08, 2008/09
2007/08, 2008/09
2007/08-2008/09

18 stations
347 stations
361 stations
482 days
3,280 whales
88 cases ofdebris obs.
326 days
68 stations
68 stations

Antarctic minke whale (biological, feeding ecology, pollutants, stock structure data)
Seasons

Sample size

Biological data
22. Catching date
23. Catching location
24. Sex25. Body length
26. Age (earplug)3
27. Age (racemization)4
28. Transition phase5
29. Presence/absence of corpora6
30. Testis weight7
31. Fetus length
32. Fetus weight
33. Fetus number8
34. Fetus sex
35. Lactation condition

2005/06-2010/11 3,264 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 38 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 1,701 whales
2005/06-2010/11 1,563 whales
2005/06-2010/11 1,127 whales
2005/06-2010/11 1,127 whales
2005/06-2010/11 1,701 whales
2005/06-2010/11 1,127 whales
2005/06-2010/11 1,701 whales

Feeding ecology/energetics
36. Blubber thickness (two points)
37. Body weight
38. Freshness stom. contents
39. Main prey
40. Organ weight incl. fat weight
41. Girth (two points)
42. Stom. content (IWS)
43. Stom. content weight
44. Lipid content in blubber

2005/06-2010/11 3,264 whales
2005/06-2010/11 1,597 whales
2005/06-2010/11 1,925 whales
2005/06-2010/11 332 whales
2005/06-2010/11 82 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 2,953 whales
2005/06-2010/11 20 whales

Pollutants/health945. Heavy metals (whale)
46. Organochlorine (whale)
47. Heavy metal (prey)
48. Gross pathological observations ofinternal organs10
Stock structure
49. Body proportion (8 measurements)
50. mtDNA (sequences) (from catches)
51. mtDNA (RFLP) (from catches)
52. Microsatellite DNA (from catches)

2005/06-2010/11 212 whales
2005/06-2010/11 5 whales
2005/06-2010/11
30 prey
2005/06-2010/11 3,264 whales
2005/06-2010/11 3,264 whales
2005/06-2010/11 2,278 whales
2005/06
764 whales
2005/06-2010/11 2,551 whales

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7

j. cetacean res. manage. 16 (suppl.), 2015

409

table
4]
Antarctic fin[Textwhale(biological,
feeding ecology, pollutants, stock structure data)
Seasons

Sample size

Biological data
53. Catching date
54. Catching location
55. Sex
56. Body length
57. Age (earplug)3
58. Transition phase5
59. Presence/absence of corpora6
60. Testis weight7
61. Foetus length
62. Foetus weight
63. Foetus number8
64. Foetus sex
65. Lactation condition

2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11

17 whales
17 whales
17 whales
16 whales
16 whales
0 whales
8 whales
8 whales
3 whales
3 whales
8 whales
3 whales
8 whalesFeeding ecology/energetics
66. Blubber thickness (14 points)
67. Body weight
68. Freshness stom. contents
69. Main prey
70. Organ weight including fat weight
71. Girth (three points)
72. Stomach content (IWS)
73. Stomach content weight
74. Lipid content in blubber

2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11
2005/06-2010/11

16 whales
15 whales
14 whales
15 whales
15 whales
16 whales
16 whales
15 whales
10 whales

2005/06-2010/11
2005/06-2010/11
2005/06-2010/11

16 whales
2 whales
16 whales

Pollutants/health975. Heavy metals (whale)
76. Organochlorine (whale)
77. Gross pathological observations ofinternal organs10

Stock structure
78. External measurements (41)2005/06-2010/11 16 whales
79. mtDNA (sequences) (catches/biopsy) 2005/06-2010/11 C:17; B:13
80. Microsatellite DNA (catches/biopsy) 2005/06-2010/11 C:17; B:13

[Text table 5]

Stock structure other species
Seasons

Sample size

Humpback whale
81. mtDNA (sequences) (biopsy)
82. Microsatellite DNA (biopsy)
83. Photo-identification data

2005/06-2009/10 121 whales
2005/06-2009/10 127 whales
2005/06-2010/11 1,201 pictures

Blue whale
84. mtDNA (sequences) (biopsy)
85. Photo-identification data

2005/06-2010/11
2005/06-2010/11

11 whales
376 pictures

Southern right whale
86. mtDNA (sequences) (biopsy)
87. Microsatellite DNA (biopsy)
88. Photo-identification data

2005/06-2010/11
2005/06-2010/11
2005/06-2010/11

33 whales
33 whales
671 pictures

Annotations8
1
The figure given corresponds to the total females examined.
Standard Line Transect data. It should be noted that in some JARPA II
9
The 2011 earthquake and tsunami affected heavily the samples collectedsurveys some areas could not be covered due to external interferences andfor pollutant studies. This explains the particular smaller samples size forsabotages from anti-whaling groups, and that some kind of extrapolation
this item.
will be necessary.
102This figure corresponds to the total number of whales examined for
The figure given corresponds to the total number of stomachs examined.
3
abnormal tissues or organs in gross pathology.
JARPA II age data of Antarctic minke whale were obtained by a new reader
with expertise and training enough for this kind of work. The figure given
The data listed here do not necessarily match the numbers
here are the total number of earplugs examined. Age information could beobtained for 81.8% of the total samples. An ageing calibration exercise
for those items indicated in the cruise reports of the JARPA
was carried out (Kitakado et al., 2013). In the case of the fin whales age
II for the period 2005/06-2010/11. This is due to the fact that
information could be obtained for 100% of the samples.
a substantial number of samples for some items were lost4This sample size corresponds to the results of a pilot study to investigate
due to the earthquake and tsunami that affected Japan on
the feasibility of the racemization method for ageing purposes. At thisstage these data were not produced for the purpose of biological parameters
11 March 2011 (IWC, 2012). These tables shows the actual
estimates but for examining the feasibility of the technique.
number of data by each item. Data for those items obtained5
The figure given corresponds to the total earplugs examined. Transition
during the JARPA surveys were also available for the review
Phase information in the Antarctic minke whales could be obtained forWorkshop.
approximately 42.1% of the total samples (mature+immature).
6
Ovary samples were lost as an effect of the 2011 earthquake and tsunami so
References
information on the number of corpora is not available. Information on theInternational
Whaling
Commission.
2012. Report of the Scientificpresence/absence of corpora (information necessary for determining sexual
Committee. J. Cetacean Res. Manage. (Suppl.) 13:1-74.
maturity in females) is based on examination of the ovaries conducted at
Kitakado, T., Lockyer, C. and Punt, A.E. 2013. A statistical model forthe field.
7quantifying age-reading errors and its application to the Antarctic minke
While in JARPA both testis weight and histological approaches were usedwhales. Paper SC/65a/IA04 presented to the IWC Scientific Committee,
for determining sexual maturity in males,H:\SkyDrive\Documents\AC
in JARPA II maturity of males
Supplement 16\Rep02 - JARPA\Rep02 Tabs.docx
15 January 201514:18
10
June 2013, Jeju Island, Republic of Korea (unpublished). 18pp. [Paper
was determined only by the testis weight criterion (due to ‘man-power’available from the Office of this Journal].
limitation and economical considerations).