Report of the Expert Panel of the
Final Review on the Western North
Pacific Japanese Special Permit
Programme (JARPN II)

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

529

Report of the Expert Panel of the Final Review on theWestern North Pacific Japanese Special Permit Programme(JARPN II)*12
EXECUTIVE SUMMARY
The Expert Panel Workshop of the final review on the western North Pacific Japanese Special Permit programme (JARPN II)
was held in the Toyomi Center Building, Tokyo from 22-26 February 2016.
The primary objective of the Expert Panel Workshop for this final review was to review the scientific aspects of the
JARPN II programme in the light of the stated programme objectives. In particular, the Panel was to:(1) assess the extent of the programme’s scientific output;(2) assess the degree to which the programme coordinated its activities with related research projects;(3) evaluate other contributions to important research and information needs outside the original set of objectives; and(4) evaluate how well the objectives of the research were met, and the extent to which results have led to demonstrated
improvements in the conservation and management of whales and/or other marine living resources.
The Panel received a total of 55 primary papers, 37 ‘for information’ papers, 1 observer statement and a response by the
proponents to that statement. An important component of the review was to examine progress made by the proponents withresponse to the recommendations of the 2009 Expert Panel. The proponents’ responses to those recommendations and thePanel’s evaluation are discussed in the report and summarised in Annex D.
The Panel noted that this ‘final’ review was somewhat unusual in that the field component of the JARPN II programme
was not expected to finish until 2016. The Panel’s general comments and recommendations on: (a) timing; (b) the natureof final reports; (c) the work being undertaken from 2014-16 to compare lethal and non-lethal techniques; and (d) ways toimprove consideration of progress with recommendations are given under Item 11.1 of this report. Annex G contains somesuggestions for potential guidelines for an integrated final report from a special permit programme.
With respect to JARPN II’s scientific output, the Panel noted that the programme thus far had results in 31 peer-reviewed
papers related to the programmes primary objectives and 30 arising from ancillary studies that contributed to research notrelated to the primary objectives. It had had also produced a large number of IWC papers that had contributed to ScientificCommittee work on the RMP and in-depth assessments. The Panel strongly encouraged the submissions of further analyses
to peer-reviewed journals.
The Panel welcomed much improved collaboration with other research projects compared to 2009 (most of which waswithin Japan). It encouraged additional collaboration with respect to any future analyses of the data.
In terms of evaluating the extent to which the results met the objectives of the programme and have improved conservationand management, the Panel considered this in two stages. The first was to examine how well they had met sub-objectives
developed by the proponents after 2009 that had been finalised in 2014. The Panel’s views are summarised in Table 10 of thereport. The second stage was to review how well the proponents had met their three main objectives (noting the timing issue
raised under Item 11.1) and to consider how the work had contributed to conservation and management. The Panels views are
given below (apart from with respect to the sperm whale component which it agreed had produced little of scientific value).
OBJECTIVE 1: FEEDING ECOLOGY AND ECOSYSTEM STUDIESThe ultimate goal of this objective was to provide multispecies management advice. As noted by the 2009 Panel, this was
an extremely ambitious task and one likely to take many years. The level of field and laboratory work has been impressive
and the examination of uncertainty with respect to the prey consumption and prey preferences has been greatly improved
since 2009 although analytical improvements can still be made. However, the question of the effects of sampling design (seeItem 3.4.2) requires further consideration and, primarily as a result of a lack of allocated resources (despite the 2009 Panelrecommendation), the modelling work remains preliminary.
Even allowing for the complexity of the issue, there are examples of MRM/MICE1 models that that can be parameterisedby fitting to data which are used to provide input to tactical assessment models and there are better developed food web andextended single species models; with additional resources, progress could (and should) have been made in the developmentof intermediate model types. The Panel concludes that at this stage of development, the modelling results are not suitable
for addressing strategic management questions2. At present, at least, the results have not led to improved conservation andmanagement of cetaceans or of other marine living resources or the ecosystem.
OBJECTIVE 2: MONITORING ENVIRONMENTAL POLLUTANTS IN CETACEANS AND THE MARINE ECOSYSTEM

This objective related to monitoring pollutants in the environment and cetaceans including: (a) pattern of accumulation incetaceans; (b) bioaccumulation through the food chain; and (c) the relationship between pollutants and cetacean health. The
Panel notes that the achievement of this objective was hampered considerably by the loss of samples as a result of the tsunami.
It also acknowledges the efforts made to follow the recommendations of the 2009 Panel. The level of field and laboratory work
has been good and understanding of chemical pollutants and cetaceans off Japan has been greatly improved. However, thePresented to the Scientific Committee meeting as SC/66b/Rep06.
Minimum Realistic Models/Models of Intermediate Complexity for Ecosystem Assessment.
2
Ecosystem models such as Ecopath with Ecosim, Atlantis, and other large complex models which are difficult to parameterise by fitting to data are not suitable for tactical management anywhere in the world at present and probably far into the future. Single species models with predation and multispecies (MICE)
models could be used to provide tactical advice in the future.
*
1

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REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

Panel concludes that only partial progress has been made towards addressing the objectives and more effort needs to be put
on improved analyses and interpretation of results (see discussion and recommendations under Item 8.4). This is especiallytrue in terms of the relationship of pollutants and cetacean health, which is most relevant to improved conservation andmanagement of cetaceans. It is not clear from the papers presented if (and if so how) the work undertaken has contributed tothe conservation of other marine resources or the ecosystem.
OBJECTIVE 3: STOCK STRUCTURE OF LARGE WHALESThe broad objectives simply related to the stock structure of large whales (common minke whales, sei whales, Bryde’s whales
and sperm whales), although this was clarified at the 2009 Panel workshop to be primarily related to developing or narrowing
the number of hypotheses to be considered by the IWC Scientific Committee in its work related to the RMP and in-depth
assessments. The level of field, laboratory and analytical work has been impressive, as was the effort put into responding to
the 2009 Panel recommendations. The Panel did make some recommendations for improved analyses, particularly related to
power and the ability to distinguish amongst weakly-differentiated populations. The Panel concludes that the stock structurecomponent of JARPN II has made, and will continue to make, important contributions to the conservation and managementof cetaceans by providing fundamental data and analyses for the RMP Implementation Reviews of common minke whales andBryde’s whales, and the in-depth assessment of sei whales.
In general, the Panel recognised the extensive field and laboratory components of the programme but was concerned that
this was not matched by the analytical effort. To this end it made almost 40 recommendations for improved analyses, of whicharound 15 could be achieved in the short-term (by the 2016 or at the latest 2017 Annual Meeting of the Scientific Committee).
Annex E contains a summary table of all Panel recommendations and expected actions by the proponents with a timeline.
1. INTRODUCTORY ITEMSThe Expert Panel Workshop of the final review on thewestern North Pacific Japanese Special Permit programme(JARPN II) was held in the Toyomi Center Building, Tokyo
from 22-26 February 2016. The list of participants is given
as Annex A.
1.1 Opening remarks
The Scientific Committee Chair welcomed the Panel
Members, Observers and Japanese Proponents to Tokyoand thanked the Fisheries Agency of Japan for hosting theWorkshop on the final review of the western North PacificJapanese Special Permit programme (JARPN II). Morishita(IWC Commissioner for Japan) also welcomed the Paneland all participants.
The meeting was organised following the previous styleof expert workshops. Mornings comprised open sessions
with summary presentations by the proponents and theopportunity for questions and discussion (Panel members,
proponents and observers present), followed by afternoon
closed sessions for the Panel to discuss the morning topics
and begin to outline relevant sections of its report and assign
writing tasks.
1.2 Appointment of Chair and rapporteurs
Fortuna, as Chair of the IWC Scientific Committee, chaired
the Workshop.
Donovan co-ordinated the report writing. All membersof the Panel contributed to the sections of the report.
1.3 Adoption of Agenda
The adopted Agenda is given as Annex B.
1.4 Review of available documents
The list of documents is given as Annex C. A total of 55
primary papers (SC/F16/JR01-55) were available, alongwith 37 for information papers, one Observer’s statement
and one response by Japan (SC/F16/O01-O02) to theObserver’s statement (SC/F16/O01).
1.5 Structure of the report
For most items in the report, the first section is a summaryof the relevant part of the JARPN II by the proponents - thePanel has not edited those sections and they represent theviews of the proponents only; these sections are shown in asmaller font and indented and the table and figure numbers
are preceded by a ‘P’. The final sections of each item
represent the comments and conclusions of the Panel. Item
11 presents the overall conclusions of the Panel alone.
According to Annex P, the report must be available to the
proponents by 11 March 2016. It will be made available on
the IWC website on 22 April 2016.
A summary of all of the recommendations from the2009 Panel report, the response of the proponents to thoserecommendations and the present Panel’s evaluation of the
response to those recommendations is given as Annex D. A
summary of all of the recommendations made by the present
Panel is provided as Annex E.
2. OBJECTIVES OF THE WORKSHOPThe Head of Science provided an introduction to the Annex
P review process. This was revised at the 2015 ScientificCommittee meeting in response to Commission Resolution
2014. The revised Annex P was recommended by consensus
by the Committee to the Commission (IWC, 2016). A simple
schematic of the process is given as Fig. 1.
The overall objective of the Annex P process is to provide
a full, fair, independent, balanced and objective review. The
first component of this is the Expert Panel workshop. The
Panel members, who should not have conflicts of interest,
are selected by the Chair and Vice-Chair of the ScientificCommittee and Head of Science, in consultation with aStanding Steering Group (SSG) comprising the previous
four chairs of the Scientific Committee. The number of panel
members depends on the subject matter of the review and theaim is to include a balance of experts from outside and inside
the Committee, depending on subject. For the present final
review, the selection also took into account membership ofthe mid-term review in 2009 for consistency and experience.
In addition to the Chair, Head of Science and one memberof the SSG, the Panel included three regular members of theCommittee, three former members who have not attended
for some years, and five non-members of the Committee;
one member participated by correspondence only.
The primary objective of the Expert Panel Workshop forthis final review was to review the scientific aspects of the

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

JARPN II programme in the light of the stated programme
objectives. In particular, the Panel was to:(1) assess the extent of the programme’s scientific output,
and whether this was appropriate in light of the statedresearch objectives and the time elapsed;(2) assess the degree to which the programme coordinatedits activities with related research projects; this included
assessment of whether the degree of coordination was
sufficient to ensure that the field and analytical methods
were appropriate and best practice to achieve the statedobjectives and whether the degree of coordination was
sufficient to avoid unnecessary duplication;(3) evaluate other contributions to important research andinformation needs that were not part of the original setof objectives of the research programme;(4) consider any other relevant matters as decided by the
Scientific Committee; and(5) evaluate how well the initial, or revised, objectivesof the research were met, and the extent to which
results have led to demonstrated improvements in theconservation and management of whales, for broad
categories of objectives 1 (‘improve the conservationand management of whale stocks’) and 2 (‘improve theconservation and management of other living marine
resources or the ecosystem of which the whale stocks
are an integral part’).
For a final review, the Panel is not asked to comment
on matters such as effect of catches, lethal versus non-lethal
techniques or future field studies. However, aspects of thefirst two matters are briefly considered in the context ofprogress with the recommendations of the 2009 ‘mid-term’review (IWC, 2010).
The review of the initial proposal was held by the ScientificCommittee in 2000 (IWC, 2001a, pp.61-64). As noted
above, the mid-term review was held in 2009 (IWC, 2010)
and consideration of progress with the recommendationsof that review was an important component of the Panel’s
evaluation of the final results of the programme.
It was noted that this is an unusual ‘final’ review, in thatthe field component of the JARPN II programme is notformally intended to be completed until 2016 – originally
this review was expected to be a review of an ongoingprogramme and it was only in 2015 that it was changed to a‘final’ review; these are normally expected to occur within
three years of the close of a programme to allow consolidated
and integrated analyses of the full datasets. The Panel makessome general comments on how this has affected the present
review and recommendations on the implications of this531

for the future in its conclusions under Item 11. Given the‘adjustments’ made to JARPN II from 2014 (see SC/F16/
JR54) in the light of the deliberations at the International
Court of Justice (ICJ) with respect to Japan’s special permit
programme in the Antarctic, the Panel’s primary focus wason the period up to 2013.
3. SHORT GENERAL OVERVIEW OF JARPN II
3.1 Proponents’ overview of general objectives and anychanges over time

The three main objectives of JARPN II were the following:
Objective 1: Feeding ecology and ecosystem studies,including: (i) prey consumption by cetaceans; (ii) prey
preference of cetaceans; and (iii) ecosystem modelling.
Objective 2: Monitoring environmental pollutants incetaceans and the marine ecosystem, including: (i) pattern ofaccumulation of pollutants in cetaceans; (ii) bioaccumulation
process of pollutants through the food chain; and (iii)
relationship between chemical pollutants and cetacean health.
Objective 3: Stock structure of large whales, including: (i)common minke whale; (ii) Bryde’s whale; (iii) sei whale; and(iv) sperm whale.
These main research objectives have not been changed
since the start of the program. However and following 2009JARPN II review workshop recommendations, several subobjectives within each main objective were identified. The
background of each objective and sub-objective is explained in
more details under Agenda Items 4, 5, 6, 7 and 8.
The origin of JARPN II is related with the research outputs
from the previous JARPN program. JARPN provided both
qualitative and quantitative information on prey species ofwhales in the western North Pacific. The analysis of stomach
contents demonstrated that common minke whales fed on
several prey species, all of which were also the target ofcommercial fisheries in Japan. The JARPN Review Workshop
in 2000 (IWC, 2001b) made a series of recommendations tostrengthen the feeding ecology part of the JARPN program.
Following this development, JARPN II was started in 2000
with the primary purpose to study the interactions between
fisheries and cetaceans through ecosystem modelling in a‘hot-spot’ area in the Pacific side of Japan, one of the mostproductive areas in the world. The output of JARPN II canassist in the formulation of effective ecosystem-based fisheries
management policies.

3.2 Summary of general conclusions and recommendations from the 2009 Workshop
A short overview of general recommendations (i.e. relevant
to two or more of the objectives) made at the 2009 Workshop(IWC, 2010) was provided by the Chair. Summaries of
the topic-specific recommendations are provided under
the relevant agenda items. Overall, the 2009 Panel had
recognised that an enormous amount of scientific work
has been undertaken in the field, laboratory and in analysis
during the first six years of the programme. In addition to its
comments under specific objectives of the programme it hadraised several general issues.
3.2.1 Objectives
The 2009 Panel had noted that the objectives for the
programme were general and long-term – this made itdifficult to undertake a full mid-term review. The 2009 Panelrecommended that well specified and quantified objectives
and sub-objectives should be developed as soon as possible,
with reference to timelines.

Fig. 1. Schematic of the Annex P process from the start of an Expert
Workshop (see text).

3.2.2 Sampling design, sample size and areas
The 2009 Panel had recommended that for each objective it
is necessary to specify the quantities of interest that need to
be determined to achieve the objective, and for each quantity

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REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

of interest, all the sources of uncertainty in its estimation
should be identified and quantified, and in particular it should
be determined which of these are functions of sample size.
The results of this analysis should then be used to determine:(a) how much the research has contributed, in quantitative terms, to achieving the objectives;(b) what further quantitative progress towards theobjectives can be expected from completing the
programme;(c) the extent to which increasing/decreasing sample
size would enhance/reduce the rate of progress
towards achieving the objectives; and(d) the extent to which the sample design is the mostappropriate for achieving the objective and inparticular for maximising the information gained
from the chosen sample size.
3.2.3 Data analyses and modelling
The 2009 Panel noted that for many of the analyses,
associated uncertainty had either not been addressed or
only partly addressed. Insufficient use had been made ofexisting datasets (e.g. JARPN and early JARPN II data) forcomparative and sensitivity analyses, cross-validation ofmodels and assessment of uncertainty.
Particular attention had been drawn to the fact that mostprogramme emphasis to date had been on data collection
rather than full analyses and model development. It had
recommended that more resources must be dedicated tothis ‘if there is to be a reasonable chance of meeting the
programme’s objectives in a reasonable time-frame’. It had
also noted the value of even simple or preliminary ecosystemmodels to assist with and refine sampling design and sample
size.
The 2009 Panel had also noted that several papers
presented had overstated the conclusions that could be made.
The present Panel’s general view of how the proponents
have responded to the 2009 review is given under Item
3.4. Specific recommendations are dealt with objective by
objective under the relevant agenda items. The full list of
2009 recommendations is tabulated in Annex D, along witha summary of the view of the proponents on whether andhow they had been addressed and the conclusions of the
Panel.
3.2.4 Other matters not the focus of a final review

LETHAL AND NON-LETHAL RESEARCH TECHNIQUESThe 2009 Panel had recommended that the proponents add
a sub-objective to their programme to allow a quantitative
comparison on all aspects of lethal and non-lethal techniquesin the context of the programme’s other objectives and subobjectives, including suggestions as to how such a subobjective could be realised, including analyses of precisionand uncertainty, evaluation of practicalities and logistics
of field and laboratory techniques. Aspects of this are
considered under Item 3.4.
EFFECTS OF CATCHES UPON THE STOCKSThe 2009 Panel had noted that Scientific Committee
guidance as to how to evaluate the effect of catches uponstocks may be valuable. It noted that this might include: (a)providing projections where the permit catch is equal to zero
as well as for the permit catches alone; (b) using a rangeof MSYR values; and (c) using Implementation Simulation
Trials (ISTs) as the basis of an evaluation where these exist(not the same as using the RMP/CLA to provide the advice
itself). With respect to common minke whales it had notedthat the approach presented by the proponents at the time

was not sufficient and had noted the need to use updated
ISTs when these had been developed as part of an RMP
Implementation Review. The 2009 Panel had accepted thatthe catches of Bryde’s and sperm whales would not pose a
problem to the stocks and had asked that additional analyses
be undertaken for sei whales before it could provide advice.
Aspects of this are discussed under Item 11.
FURTHER REVIEWThe 2009 Panel had noted that given its recommendations
for additional work regarding sample size and the effects ofcatches on two of the stocks, it could not complete its review
and asked the Scientific Committee to consider the mostappropriate way to ensure that it was completed. Aspects ofthis are discussed under Item 11.
3.3 Proponents’ overview of broad sampling scheme
including sample size, areas and strategy includingchanges over time and response to the 2009 Workshop
recommendations

SC/F16/JR02, SC/F16/JR03, and SC/F16/JR04 described themethodology and procedures of the JARPN II field surveys. In
general, survey procedures were the same to those presented to
the 2009 Workshop, which reviewed data and results obtained
between 2002 and 2007. The JARPN II involved two survey
components, an ‘offshore’ component which was covered by the
research base vessel Nisshin Maru and sighting and sampling
vessels, and a ‘coastal’ component which was covered bysmall-type whaling vessels. The offshore component covered
the broad offshore areas in the western North Pacific from subarea 7 to sub-area 9, and the coastal component was conductedin the two coastal areas, Sanriku in spring and Kushiro inautumn.
In principle, whale sampling was made along predetermined track lines, and all of the sighted whales weretargeted for sampling excluding mother and calf pair (SC/F16/JR03, SC/F16/JR04). Annual sample size for the offshore
component was started as 100 common minke, 50 Bryde’s and
10 sperm whales in 2000-2001 feasibility studies. A total of 50
sei whales was added in the full-scale offshore survey started
in 2002, and the sample size was changed to 100 in 2004. The
coastal component was started in 2002 targeting 50 minke
whales as a feasibility study, and the full-scale coastal survey
was started in 2004 with sample size of 60 minke whales foreach coastal area per year.
As same as in the previous 2000-07 surveys, dedicatedsighting surveys and prey species surveys were also carried out
as part of the multidisciplinary research program. Dedicatedsighting surveys were conducted to estimate the numberof whales migrating to the research area (SC/F16/JR02).
Methodology of the sighting surveys followed the IWC survey
guidelines. Non-lethal experiments, such as photo-ID andbiopsy were conducted from the sighting vessels. Co-operativeand concurrent surveys of whale sampling and acoustic/
trawl preys and oceanographic survey were conducted by the
dedicated prey survey vessels (SC/F16/JR04).
SC/F16/JR54 describes the adjustments made to JARPN
II from 2014. Following the Judgment of the International
Court of Justice (ICJ) in March 2014, the Government of Japan
voluntarily reviewed the JARPN II program in response to the
Judgment. The voluntary review resulted in the reprioritisationof research objectives as well as recalculation of sample sizes.
Sampling of sperm and common minke whales in the offshore
component was suspended. Comparative studies for verifying
the feasibility and practicability of non-lethal methods,
including biopsy sampling and faeces collection, are being
carried out from 2014 to 2016.
At the request of the Panel, more information is provided
in Annex F.

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533

Fig. 2. Overview of the survey design under JARPN II. There are several survey components under JARPN II: (a) dedicated sighting surveys; (b) whale
sampling surveys (coastal and offshore); (c) concurrent whale/prey surveys in offshore areas; and (d) prey surveys in coastal areas. In the example figure here,
pre-determined track lines for whale sampling surveys in the offshore component are shown in orange; on effort track lines of dedicated sighting (single line)
and whale sampling (parallel lines) surveys in green. Concurrent whale/prey surveys are shown by coloured squares. Only the research areas are shown for
Sanriku and Kushiro in this figure. Kindly produced by the proponents.
Table 1
Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to objectives (see text).
Brief summary of 2009 recommendations/suggestions

Panel evaluation

Develop refined, more quantified sub-objectives and short-term objectives

Partly met but no timeline

3.4 Panel comments and conclusions on general issues
and sampling areas. It had noted that this should be done
3.4.1 Objectives
Table 2 for the quantities of interest (parameters) relevant to the
The Panel welcomedfact evaluationthat the ofproponents
hadquantified with
sub-objectives
by: (1) undertaking
Summarytheof Panel’s
the proponents’
response torevised
the 2009 andrecommendations
respect to samplingdeveloped sub-objectives for the JARPN II programme
design andas
sample size(see text). analyses on the existing datasets, including
exploratory
recommended
by the 2009 Panel. However, it notes thatpower and sensitivity analyses, foreachof the components
Brief summary of 2009 recommendations/suggestions
Panelevaluationwhilst work to develop sub-objectives had begun in 2010 theyof the research; and (2) examining the effects of sample sizefor eachinterest
foraftereach objective
sources of uncertainty and relationship with sample
had not beenQuantify
finaliseduntilquantity
2014, offive
years
the review.
on the ability and time to meet Not
the addressed
sub-objectives (IWC,
size and use to evaluate progress on meeting objectives, effect of changing sample size on meeting
In addition,objectives
the sub-objectives
had not been accompanied
2010, p.432).
and suitability of sampling design.
by shorter-term objectives (i.e. targets and timeline) (Table
The Panel agrees that this recommendation had not been
1). Whilst development of a timeline became moot in 2015
addressed. However, it recognised that given that this is a final
when Japan asked that the present review be considered as
review, the emphasis now is rather on whether the samplinga final review, in 2014, it was expected that the 2016 review
designand3 sample sizes were adequate to meet the objectives.
Table
would represent
a periodic review.
is considered further
under
specific design
agendaSummary of Panel’s evaluation of the proponents’ response to the This
2009 recommendations
with respect
to sampling
anditems
samplebelow.
size (see text).
The Panel notes that Annex P requires that new proposals
WithPanel
respect to sampling design and areas, the Panel
do contain sub-objectives and a timeline and agrees that thisrequested
additional
information from the proponents
Brief summary of 2009 recommendations/suggestions
Panel comments
evaluation
should be an essential component of any long-term researchduring the Workshop, and specifically, maps showing theLargely
Kanda et al. (2014b) (a revised version of Kanda et al., 2014a)
genetic assessments
shouldinclude a a
brief
description ofprogramme, Theirrespectiveof whether
it contains
lethalsurveyaddressed
design bypresented
year andseason (earlyor late),and
along
with protocols
a summaryof the samplinglaboratory
procedures to ensure data quality (with reference to IWC guidelines
sampling component.
realised tracks andsampling
These
are given
employed
for the locations.
genetic studies
under Objective
3 ofinthe JARPN
for DNA data quality).
II conductedat the Instituteof Cetacean Research.
Annex F. The sampling
schemeis complicated(see Fig.The2 individual
3.4.2 Sampling design and sample size
stock structure-related documents also confirm that the IWC
and Annex F), withchanges over time in where sampling for3.4.2.1 SAMPLING DESIGN AND AREAS
guidelines for DNA data quality (Tiedemann et al., 2012) haveprey occurred (thesampling areas) owing in
The 2009 Panel had noted that for several of the objectives
beenco-operative
adhered to.
part
to
the
need
to
coordinate
withfishery surveys.
there was insufficientjustificationofsamplesizes,
design
Revised papers should include estimates of genetic divergence (along
Largely
FST estimates were provided
when appropriate. Levels ofwith levels of uncertainty) in addition to probabilities of homogeneity.

addressed

uncertainty associated with divergence estimates were, however,not reported.

P values (and divergence estimates) should be reported for all locicombined rather than for each locus separately.

Addressed-

Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to objectives (see text).

534

Brief summary of 2009 recommendations/suggestions

Panel evaluation

Develop refined, more quantified sub-objectives and short-term objectives

Partly met but no timeline

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II
Table 2
Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to sampling
design and sample size (see text).
Brief summary of 2009 recommendations/suggestions

Panel evaluation

Quantify for each quantity of interest for each objective sources of uncertainty and relationship with sample
size and use to evaluate progress on meeting objectives, effect of changing sample size on meeting
objectives and suitability of sampling design.

Not addressed(see Table 2). The only objective for which an attempt has
With respect to the offshore component, the design forbeen made to fully quantify uncertainty is estimation of total
the sightings surveys seems reasonable and the fulfilled
Table 3
consumption(e.g. see SC/F16/JR15) where a target CV
tracklines are adequate. However, with respect to the0.2
was chosen.with
Therespectvalueto of
0.2 wasapparently
sampling, there
areof worrying
differences
betweenthe to the ofSummary
Panel’s evaluationof the proponents’
response
2009
recommendations
sampling
designand sample based
size (see text).
upon Winship and Trites (2003) work on Stellar sea lions.
proposed sampling tracklines (which it appears may be
Panel
However,thosePanelauthors’attempts to address objectives
broadly adequate
althoughof quite
variable in terms of blocks
Brief summary
2009 recommendations/suggestionscomments
evaluation
related to the use of consumption estimates for estimating
and direction from year to year - the rationale for the design
LargelyKanda
et al. (2014b)(a revised versionof Kandacall
et al., 2014a)
The genetic assessments should include a brief description preyof preference
and ecosystemmodelling,
if anything,
is not presented)
and the actual sampling. The Panel waspresented a summary of the sampling and laboratory protocols
procedures to ensure data quality (with reference to IWC guidelines addressed
intoquestionthechoiceofasingle
value
of
0.2.
As
noted
informed thatforthis
function of decisions that had toemployed for the genetic studies under Objective 3 of the JARPN
DNAwasdata aquality).
by the 2009 Panel,a moreatappropriate
to examine
be taken in the face of poor weather but an examination of
II conducted
the Institute of method
Cetacean Research.
The individual
this is to use evensimpleecosystemdocuments
models for
JARPN
stock
structure-related
alsotheconfirmthat the IWC
the maps (Annex F) shows that in many years, the samples
guidelines
for DNA data quality(Tiedemannet al.,of2012) have
II area to evaluate
the consequencesof different
levelswere not taken along the pre-determined tracks and in somebeen adheredto.
precision of estimatesof consumption(and other parameterscases (e.g. 2012 in the ‘late’ season), many of the catches
Revisedpapersshouldincludeestimatesofgenetic
divergence(alongLargelyF
estimates
appropriate.
of interest) and hence
samplewere
sizes.provided
Aspectswhenof thisissue areLevels ofST
lay along new lines with no apparent rationale. This raises
with levels of uncertainty) in addition to probabilities of homogeneity.
addressed
uncertainty
associatedwith divergence estimates were, however,discussed
further
under
Item
6.4.
important concerns as to whether such samples can be
not reported.
The Panel agrees
to make some brief comments intaken as representativeof the animals within the surveyed
P values (and divergence estimates) should be reported for all loci
Addressed
responseto
SC/F16/JR54,
although this is formally outside
areas. As a minimum,
the proponents
shouldhave addressed
combined rather
than for each locus
separately.
the scope of this review whose focus is on the period up
this issue forMultiple
those analyses
for
which
this
is
an
importanttesting issues: (a) apply False Discovery Rates; (b) exercise
Addressed
to 2013prior to -the Government of Japan’s changes to theassumption (e.g.
manyfeeding
ecologystudies).
discretionin ofthe thenumberof pairwise
comparisonsevaluated.
programme in the context of the ICJ ruling (related to a
The Panel
recommends
thaton a thenew
document
based
Providemore details
analyses
involving
the program Addressed reprioritisation of the objectives, the addition of an objective
upon Annex FSTRUCTURE.
is developed by the proponents and submitted
and a change in sample size):to the 2016 Annual
Meetingofthe
ScientificCommittee.
PartlySampling procedures were adequately described, but no
Include a brief discussion of experimental design with respect to samp(1) the
only parameterused
address
samplesize towas
In addition tolingthe
information
ondesign
sightings,
the document
addressed
explanationsof thetosampling
rationalein relation
resolving the(explaining
how thespecificallyaddresses uncertainties
consumption
andthis
was inrelationtouncertainties
related tostock
structure
wereprovided.
relatedto stocke.g. whether the spatial and temporal related to total
should record,for eachyearstructure,and season:coverage of samples of minke whales has been sufficient to test a target CV of 0.2 – however the rationale for this was(a) the predeterminedtracklines
forsampling
andtheadequately the alternative stock structure hypotheses).
unclear as two explanations were given, one related torationale for those lines; andRedo the Boundary Rank analyses (Taylor and Martien, 2002) with WinshipNo longer and- Trites (2003) in SC/F16/JR54 and another(b) the actual
thosetracklines and the
new data coveragefor common ofminke
whales.
relevantinwhich additional information presented to the meeting
rationale for any decisions taken to deviate from theIntegrate Korean by-catch samples into the new datasets to look at related
Addressed- approach used by the Government ofto thepredetermined
including
rationale for any
heterogeneitylines
for common
minkethewhales.
Norwayinits
special permit programme in the 1990s
new Undertakelines developed.
PartlyAssessments of statistical power were performed with regards tothe assessments of power to simulate data to evaluate power (NMMRP,
1992);The papertoshould
issue
whether
actualaddressed
homogeneity
under different
detect a address
specified thefractionof aofputative
stockthe(e.g.
the hypothetical(2) with
respectrejecting
to commonminke
whales migration
this ledrates.
tominke whales)
overall sample
using
sampling canWbestock
saidoftocommon
be representativeof intheananimals
inlowersamplesizes
intheinshorecomponentandthethe surveyed simulatedarea anddata.
discuss the extent to which this may
suspension of sampling of minke whales in the offshore
Undertaketests for population genetic(drift-mutation-migration)
Partlywas tested for long-term demographic popaffect those objectives/parameters/analyses
for whichthis iscomponent–Non-equilibrium
however the implicationsof this in terms ofequilibrium.
addressed
ulation trends via mismatch distribution analysis of mitoor may be important (e.g. feeding ecology studies). Finally,meeting the chondrial
originalDNA.
objectives
were
notfullyof explored;
However, assessments
short-term deviations
the Panel also notes that the JARPN II sampling area does(3) with respectfrom
to mutation-drift
sei and Bryde’s
whales,present
equilibrium(Piry theet al.,
1999) were not
not cover the spatial extent of any of the stocks (e.g. JE,
undertaken.
However, theseassessments
may but
be infeasible.
sample sizes
were estimatedto be
too smallno
O, OW, and OE)
postulated
Theof mutation- changes
weremade – again the implications with
Employapproaches tothatbedo foundnot relyeast
upon ofthe Japan.
assumption
Notpaper shoulddrift-migration
also comment
upon when
the implicationsof thisequilibrium
estimating population
divergence.
addressed
respect
to meeting the original objectives were not fully
for any analyses
that
are
or
might
be
dependent
upon
an
explored;Attempt the detection of related pairs of individuals
Partly andSpatially-explicit relatedness analyses are underway but only
assumption that the samples are representative of biologicaladdressed
results wereshown at theWorkshopitself.
(4) theadditionpreliminaryof an objective
to compare
lethaland nonpopulations (e.g.
with multivariate
respect toanalyses
age data).
The Panel agrees
techniquesin linerecommendation
Undertakeof morphological
data with respect to lethal
Largely
Resultswas
presented
forwithsomethecases,
mainly for O and J stock
stockofstructure.
addressed
commonminkethat a document
this nature (i.e. with respect to proposedfromthe 2009
Panel,
butwhales.
the explanation of this withand actual samplingandeffort) shouldbe partof integrative respect
sizes
was confusing
and thePollutant
protocols
Use of pastandsighting
present contaminantdata shouldbe an
Not to sample
There areinsufficient
Persistent Organic(POP) data forstudy of or
stock
structure.
applicable
this to be were
productive.
forthe comparison
not provided – the Panel refers
any future periodic
final
reviews (see Item 11).
the adviceProof
providedas howto do
suchbya taggingcomparison
The sampling
for the toinshore
Initiate scheme
satellite tagging
narrow component
the range of (theplausible stock to Partly
of principle
has beenachieveda single common
addressed
and two Bryde’s whales, but many more must be
given
in theminke
2009 whale
report.
Kushiro and structure
Sanrikuhypotheses.
sampling areas and their 30 and 50n.
tagged if the results are to be used to inform discussions.
miles radii sub-blocks) appears to be based upon logistics
The Panel recommends that a single document be
and no consideration of representativeness, either within theprovided
1 to the 2016 Annual Meeting that:sampling areas or for the populations. The implications of(1) provides a clearer rationale for the changes in sample
this for the analyses based upon such assumptions must be
sizes and any implications for meeting the original
considered by the proponents in a more quantitative manner.
objectives of the programme (e.g. related to time series
The Panel recommends that this be fully addressed inof data, analyses and sample size); andrevised papers submitted to the 2016 Annual Meeting.
(2) provides the field and analytical protocols for the
3.4.2.2 SAMPLE SIZE
comparison of using lethal and non-lethal techniques
The Panel agrees that the 2009 recommendation related tofor each key parameter taking into account the advice
sample size and meeting objectives had not been addressed
provided in 2009.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017535

Fig. P1. Schematic representation of research components under Objective 3 of JARPN II.

4. STOCK STRUCTURE
Although this is not the first primary objective of the
programme, it is fundamental to understanding many of theother aspects of the programme (e.g. distribution, abundanceand status) and thus was discussed first.
4.1 Proponents’ summary of objectives including
modifications, if any, since the start of the programme

The main objective 3 of the JARPN II research plan was toinvestigate the stock structure of large whales such as thecommon minke, Bryde’s, sei and sperm whales in the North
Pacific.
Stock structure information is important for the development of RMP ISTs in the case of the common minke andBryde’s whales, and for the in-depth assessments (IA) of otherlarge whale species such as the sei, sperm and right whales.
Fig. P1 shows a schematic representation of the data, analyses
and outputs within this main objective.
Common minke whale
The IWC SC completed the Implementation Review (IR) of
the western North Pacific common minke whale at its 2012
and 2013 Annual Meetings (IWC, 2013; 2014). Most of theinformation on stock structure presented and discussed during
the IR was based on data and information from JARPN and
JARPN II. A total of 22 sub-areas were set for the aim of the
IR (Fig. P2a), and three stock structure hypotheses were used
(Fig. P2b-d).
The plausibility of the three stock structure hypotheses was
discussed at the 2012 IWC SC Annual meeting. A group of
five geneticists summarised their interpretation of the relative
support for and against the five hypothesised stocks (JE, JW,
OE, OW and Y) involved in the different hypotheses (IWC,
2013, p.135). It should be noted that their evaluation was madebased on the available genetic information only despite plentyof non-genetic information was available for the IR.
Despite this effort by geneticists, it was not possible for
the IWC SC to agree on plausibility of the three stock structure
hypothesis. As a consequence, all three main stock structure
hypotheses were ‘no agreement’ and were therefore treated asif they had been assigned ‘Medium’ plausibility in the trials(IWC, 2013, p.126).
There are, however, some interesting points in theevaluation of different stocks by the five geneticists. Fromtheir summary table (IWC, 2013, p.135), it is clear that theplausibility of additional structure in the J stock is low, andconsequently further analyses under the JARPN II were notfocused to investigate subdivision within this stock. The J stock
distributes mainly in the Sea of Japan but mixes with the O
stock in the coast of northern Hokkaido and in the Pacific sideof Japan. What it is important is the study of the dynamics of
the J stock around the Japanese coast, and the interaction with
the O stock, mainly in the Pacific side of Japan. This is the basis
for the first sub-objective under Objective 3 of JARPN II:

Objective 3, Sub-objective 1
Monitoring of the spatial and temporal distribution of J
stock on both west and east coasts of Japan using genetics
and non-genetics approaches, and all sources of samples
available e.g. JARPN, JARPN II and by-catches (Relevant
documents: SC/F16/JR38, SC/F16/JR39).
With regard to the O stock, the genetic evidence summarisedin the summary table (IWC, 2013, p.135) cannot discard
definitively the possibility of additional structure within the O
stock. Therefore the second sub-objective within Objective 1is the following:
Objective 3, Sub-objective 2
Using genetic and non-genetic data from JARPN and
JARPN II, investigate whether or not the sub-division of the
O stock into OW and OE is plausible. The genetic analysis
should include those approaches mentioned in Table 1 asproviding support for the existence of the OW (e.g. PCA
analyses) (Relevant documents: SC/F16/JR40-41; SC/F16/
JR43; Kishiro and Miyashita (2011) [SC/F16/JR42]).
Bryde’s whale
The RMP Implementation for western North Pacific Bryde’s
whale was completed by the IWC SC in 2007 (IWC, 2008,p.9). During the Implementation two sub-areas (Fig. P3a; IWC(2009, p.7), and four stock structure hypotheses (Fig. P3b;
IWC (2007a, p.8), were considered.
The IWC SC examined the plausibility of the four
hypotheses based on the genetics and non-genetics information
available in 2006. That information was summarised in IWC(2007b). Based on such information, the IWC SC assigned
the following plausibility to the four hypotheses: Hypothesis
1=High; Hypothesis 2=High; Hypothesis 3=High; Hypothesis
4=Medium.
It should be noted that a substantial number of genetic
samples have been accumulated since the Implementation of
Bryde’s whale in 2007. For example larger sample sizes are
now available for Sub-areas 1W and 1E, from JARPN II,
dedicated sighting and IWC POWER surveys. This allowed
new analyses to be conducted to evaluate the plausibility of
Hypothesis 4, which propose two stocks within sub-area 1. In
addition, genetic samples are now available for Sub-area 2 fromthe IWC POWER surveys. Genetic analyses of such samples
would allow the evaluation of plausibility of Hypotheses 2 and3, which propose two stocks in those sub-areas.
Therefore the following sub-objective related to
Bryde’s whale, which is relevant in the context of the RMP
Implementation, was defined:
Objective 3, Sub-objective 3
To investigate the plausibility of: (i) stock sub-division
within Sub-area 1 as proposed under Hypothesis 4; and (ii)
sub-division between Sub-areas 1 and 2 as proposed under
Hypotheses 2 and 3, using all genetic samples available
from different sources till 2014, different genetic markers
and satellite tracking (Relevant documents: SC/F16/JR44,
Murase et al. (2016) [SC/F16/JR45]).

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REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

Fig. P2.
(a)
(b)
(c)
(d)

The 22 sub-areas used for the Implementation Simulation Trials
for North Pacific common minke whale.
Schematic diagram of Hypothesis I.
Schematic diagram of Hypothesis II.
Schematic diagram of Hypothesis III.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

537

Fig. P3. (a) Sub-areas used for the RMP Implementation of western North Pacific Bryde’s whale. (b) Stock structure hypotheses selected for the RMP
Implementation of western North Pacific Bryde’s whale.

Sei whale
The in-depth assessment of North Pacific sei whale started atthe 2015 IWC SC meeting. The IWC SC agreed to proceedwith two initial alternative stock structure hypotheses: (i) asingle stock in the entire North Pacific as proposed by Kanda
et al. (2015), based on several pieces of evidences included thegenetics; and (ii) a five-stock hypothesis proposed in Mizroch etal. (2015), based mainly on the interpretation of mark-recapture
data: Japan coastal; North Pacific pelagic; Aleutian Islands andGulf of Alaska; eastern North Pacific migratory; and Southern
North American coastal stock (coastal California) (IWC,
2015b). The IWC SC agreed that discriminating between these
two hypotheses is difficult in the absence of genetic data fromthe potentially extirpated stocks, and thus both hypotheses areplausible (IWC, 2015a).
The sub-objective related to sei whale is the following:

Objective 3, Sub-objective 4
To investigate the plausibility of a single stock of sei whalein the pelagic regions of the North Pacific (‘North Pacificpelagic’), using all genetic samples available from different
sources till 2014, and different genetic markers (Relevant
documents: SC/F16/JR46-47; Kanda et al. (2015) [SC/F16/JR48]).
Other species
There are a number of genetic samples collected under JARPN
II from sperm whale (catches) and North Pacific right whales(biopsy). The number of samples is small and then, detailed
analyses on stock structure were not possible. However, genetic
studies based on microsatellite and mtDNA were conducted onthese two species to investigate levels of genetic diversity andthe utility of such genetic markers for future studies on stock
structure, in addition to a new worldwide phylogenetic study inthe case of the right whale (see documents SC/F16/JR49-51).

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REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

4.2 Overview of conclusions and recommendations fromthe 2009 Workshop
A short overview of the conclusions and recommendations
made at the 2009 Workshop (IWC, 2010) on the stock
structure component of JARPN II was provided by the Headof Science.
The Panel had noted the considerable work undertakenby the proponents that had led to a uniquely large genetic
dataset for common minke, Bryde’s and sei whales in
the region. The analyses that were presented had been
generally sound and provided useful information for RMP
Implementation Reviews (common minke and Bryde’s
whales) and the future in-depth assessment of sei whales,
although it was not possible at that time to conclude thatthe number of plausible hypotheses had been reduced. It had
noted that the incorporation of non-genetic information was
valuable. However, it noted that the programme had madeno meaningful contribution with respect to sperm whales
and was unlikely to do so in the future.
In terms of the main problems identified, these wereidentified as being related to either: (a) poor specificationof the original objectives; and/or (b) generic difficulties (i.e.
not associated with this particular programme) related to
weakly-differentiated populations.
Some general limitations identified by the 2009 Panelthat should be addressed related to the need to:(a) assess demographic linkage (there was an over relianceon hypothesis testing in the analyses presented);(b) examine the statistical power of results suggesting
no differentiation;(c) consider non-genetic-equilibrium scenarios; and(d) examine the implications of the use of the Bonferroni correction.
In order to address these issues the Panel had made anumber of recommendations and suggestions.
In terms of ‘simple issues’, the Panel recommended thatin the revision of papers presented and in future work the
proponents should:(a) explain data quality procedures;(b) estimate divergence and uncertainty not justhomogeneity;(c) report p-values for all loci combined;(d) consider the use of the ‘False Discovery Rate’ (FDR)
rather than the Bonferroni correction approach;(e) improve the presentation of results for analyses
using the ‘STRUCTURE’ program; and(f) better explain the influence of sampling design
related to specific stock structure hypotheses.
With respect to more extensive issues, the Panel had
recommended that the proponents should:
i. integrate data on common minke sampled by the
Republic of Korea into analyses;
ii. improve the assessment of the power for various
approaches (e.g. using simulation studies);
iii. test for population genetic equilibrium;iv. estimate divergence using non-equilibrium
approaches;
v. examine the spatial distribution of relatives;
vi. consider additional analyses with respect tomorphology and pollutants; andvii. develop a telemetry programme.
The Panel had also stressed that the small sample size forsperm whales precluded any useful information (it had madean overall recommendation for the sperm whale componentof JARPN II to be removed).

4.3 Proponents’ summary of the results (incl. response
to 2009 Workshop)

Studies on stock structure of large whales under JARPN
II were based on a large and comprehensive data/sample
set, and involved the use of both genetics (mtDNA control
region sequences and microsatellite DNA) and non-genetic(morphometric, morphologic, satellite tracking).
Regarding to genetics, the combination of large sample
sizes and powerful genetic markers was an appropriate tool to
examine stock structure in weakly differentiated populations asshown below.

Objective 3, Sub-objective 1
Monitoring of the spatial and temporal distribution of J stock
common minke whale on both west and east coasts of Japan
using genetics and non-genetics approaches, and all sourcesof samples available e.g. JARPN, JARPN II and by-catches.
SC/F16/JR38 examined a total of 4,275 western North
Pacific common minke whales with a set of 16 microsatellite
DNA loci and the program STRUCTURE to assign individual
to either J or O stocks. Samples were available from JARPN/
JARPN II (1994-2014; n=2,637), and by-catches (2001-14;
n=1,638), from different management sub-areas (SA) around
Japan. Results of the Bayesian clustering analysis confirmedthat the whales came from two genetically differentiatedstocks, J and O stocks. The number of unassigned individuals(‘unknown’) decreased with the increase in the number ofmicrosatellite loci used, and they were widely distributed.
By using 16 loci, more than 90% of the individual whales
were assigned to either stocks. Almost all of the individuals
collected from the Sea of Japan side (SA6 and SA10E)
belonged to the J stock, whereas almost all of the individuals
from the offshore North Pacific (east of SA7WR) belonged
to the O stock. Intermediate areas (SA7CN, 7CS and SA11)
contained individuals from both stocks. The SA2 was mainly
occupied by the J stock. In SA2 the J stock was predominant(around 80% in proportion) around the year. In SA7CS andSA7CN the proportion of the J stock increase in autumn/winterand decrease in spring/summer. A phylogenetic tree of mtDNA
haplotypes showed several clades but none supported by highbootstrap values. There was no stock-specific clade althoughmost of the individuals assigned to the J stock shared a sameclade. Most of the individuals assigned to the O stocks shareclades where the J stock individuals were less frequent. The
unknown samples were widely distributed through the clades.
SC/F16/JR39 focused on the unique white patch on the
flipper of the common minke whale to differentiate between J
and O stocks. Animals collected from JARPN II research during
2012 and 2013 were used; assignment of individual whales to
the O and J stocks was based on microsatellite analysis (n=220).
The morphological differences in the size and pattern of thewhite patch on the flipper of each whale was examined. The
length of the white patch along the anterior (ventral) margin of
the flipper tends to be proportionally larger in O stock animals.
The pattern of the boundary area of the white patch named asthe ‘Grayish Accessary Layer (GAL)’ was remarkably different
between stocks. Of the total animals with ‘no GAL’ type,
94% were J stock. Conversely, of the total animals with GAL
expanding over the half the flipper width, 98% were O stock.
It is concluded that there were clear morphological differencesin the body color pattern of the flipper between J and O stocks.
In conclusion, JARPN II was able to monitor the O and J
stock common minke whales around Japanese waters. Almost
all animals from the Sea of Japan belonged to the J stock while
all animals in the Pacific side east of SA7WR belonged to the
O stock. Intermediate sub-areas (7CN, 7CS, and 11) contained
animals from both stocks.
In SA2 the J stock animals were predominant through theyear while in SA7CS and SA7CN the proportion of J stock
animals increased in autumn/winter and decreased in spring/summer. The O stock had a reverse trend.
The number of ‘unknown’ individuals in the STRUCTURE
analysis decreases as the number of loci increases. They are

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

widely distributed in the research area and through the mtDNA
phylogenetic clades. Therefore whales assigned to J and O stocks
by the STRUCTURE analysis can be subjected to examinationof sub-structure using different analytical approaches, and there
is no problem of ‘circularity’ in such analyses.
Objective 3, Sub-objective 2
Using genetic and non-genetic data from JARPN and JARPN
II, investigate whether or not the sub-division of the O stock
common minke whale into OW and OE is plausible. The
genetic analysis should include those approaches mentioned
as providing support for the existence of the OW (e.g. PCA
analyses).
SC/F16/JR40 examined the genetic population structure of‘O’ stock common minke whale in the western North Pacificbased on mitochondrial DNA control region sequencing(487bp) and microsatellite DNA (16 loci). Samples used in thetests of homogeneity were obtained during the surveys of the
JARPN and JARPN II in sub-areas of the Pacific side of Japanbetween 1994 and 2014 (n=2,071 for microsatellite; n=2,070
for mtDNA). Whales were assigned to the ‘O’ stock by the
analysis of STRUCTURE presented in SC/F16/JR39. Tests
based on both genetic markers and different grouping of thesamples showed no evidence of sub-structuring in the ‘O’ stock
common minke whale in the Pacific side of Japan. A simulation
exercise showed that the statistical power of the homogeneity
test was high. In addition, a Discriminant Analysis of Principal
Components (DAPC) based on the total samples used in SC/
F16/JR38 showed clear differentiation between J and O stock
whales but no evidence of sub-structuring within the O stock
samples. Consequently the results of this study suggested a low
plausibility for the hypothesis of sub-division of the O stock
common minke whale into OW and OE.
SC/F16/JR41 examined stock structure of western
North Pacific common minke whales by using externalmeasurement data collected during 1994 and 2014 JARPN and
JARPN II surveys. Most of the analyses conducted followed
recommendations from the 2009 JARPN II review workshop.
External measurements of mature males were first compared
between O and J stock animals assigned by the microsatellite
DNA analysis. Then, only O stock animals were comparedamong sub-areas. The analytical procedures used were theAnalysis of Covariance (ANCOVA) and DAPC. Significant
differences were detected between O and J stock whales. J stock
animals had longer head region compared to O stock animals.
No significant differences were detected in O stock animals
among sub-areas. The results of the present study provided
no evidences for sub-structuring of the O stock into OW andOE as proposed in one of the hypotheses used in the RMP
Implementation, as common minke whales from coastal andoffshore sub-areas did not differ in morphometric characters.
Kishiro and Miyashita (2011) [SC/F16/JR42] described anexperiment on satellite tracking conducted on common minke
whales in the Pacific side of Japan, specifically in coastal wateroff Hokkaido in autumn 2010. Using a handy air gun, a satellitetag (Argos transmitter) was attached on one common minke
whale on 13 September (estimated body size: 7.8m). The
movement of the whale was tracked for a period of 27 days. The
whale stayed in the coastal waters off Kushiro, for at least fourweeks in the autumn season. The JARPN II review workshop
in 2009 had recommended satellite tracking experiments on thewhale species studied, as a long-term task. This study started
such experiments in the common minke whale. Further satellite
tracking experiments have the potential to elucidate whether or
not a coastal-resident stock of common minke whale occur inwaters off Hokkaido (OW) as proposed by one of the current
stock structure hypotheses.
In SC/F16/JR43 the catch-at-age data for minke whales in
the western North Pacific provided by the JARPN/JARPN II
were used to refine existing RMP ISTs in a simple way, so as
to investigate the relative plausibility of the single- and two
stock hypotheses for the O stock whales in the Pacific sideof Japan. While the single stock scenario seems consistent539

with these age data, it is difficult to reconcile the two stock
hypothesis (OW and OE) with them because of the relative
absence of particularly younger whales in a supposedly
separate discrete OE stock. The analysis demonstrated theimportance for management purposes of obtaining age data
for the common minke whales in the western North Pacific,
which in turn necessitates lethal sampling. Such age data need
to be incorporated in the conditioning of revised RMP ISTs forcommon minke whales in this region.
During the Power Point presentation the proponents
informed on the ongoing kinship analysis in common minke
whale. A total of 22 cases of potential parent-offspring cases
were identified.
In conclusion, the most parsimonious interpretation ofdifferent analyses conducted on J-purged O stock animals(hypothesis testing including an evaluation of the power,
discriminant analysis of principal components, morphometrics,
and catch-at age data) is that a single O stock occurs from theJapanese coast till at least approximately 170°E. Based on the
updated scientific evidences presented at this workshop, theplausibility of an OW stock is ‘Low’ or ‘Very low’.
An important implication of the research conducted under
this sub-objectives is that in future RMP ISTs, the conditioning
needs to take age-structure data into account to better reflect
the underlying dynamics of this population.
As explained in Section 4.1, the next sub-objective involves
Bryde’s whales. The RMP Implementation for western North
Pacific Bryde’s whales completed by the IWC SC in 2007
involved two sub-areas and four stock structure hypotheses.
The following sub-objective addresses the plausibility of threeof the stock structure hypotheses.
Objective 3, Sub-objective 3
To investigate the plausibility of: (i) stock sub-division
within Sub-area 1 as proposed under Hypothesis 4; and (ii)
sub-division between Sub-areas 1 and 2 as proposed under
Hypotheses 2 and 3, using all genetic samples available fromdifferent sources till 2014, different genetic markers andsatellite tracking.
SC/F16/JR44 examined a total of 1,019 and 1,026 samplesof North Pacific Bryde’s whales with microsatellite DNA (17loci) and mitochondrial DNA sequencing (299bp), respectively,
to examine the plausibility of four stock structure hypotheses
used by the IWC SC during the 2007 RMP Implementation.
Samples were from different sources: JARPN II (catches),
Japanese dedicated sighting surveys (biopsy); IWC/POWER
surveys (biopsy) and past commercial whaling (catches).
No significant genetic heterogeneity was found between theWestern and Eastern Sectors of sub-area 1, a result supported
by high statistical power. However both genetic markers
showed significant differences (for males, females and sexes
combined) between sub-areas 1 and 2. Phylogenetic analysis
of mtDNA haplotypes revealed no subarea-specific clades. It is
proposed that a longitudinal sector around 180° could represent
a hard boundary or a transition area where the two stocks mix.
Based on these results, it is suggested that the plausibility of thestock structure hypotheses for western North Pacific Bryde’s
whale used in the 2007 Implementation whale should be reexamined. The results of this study suggest that the two-stock
hypotheses (Hypotheses 2 and 3) could be more plausible thanthe one-stock hypothesis (Hypothesis 1) and the three-stock
hypothesis (Hypothesis 4).
Murase et al. (2016) [SC/F16/JR45] reported the movementof two individual Bryde’s whales using satellite-monitored
radio tags in offshore waters of the western North Pacific (subarea 1). One whale was recorded for 13 days 4 hours 57 minutes
from 13 to 26 July 2006. The other whale was recorded for 20
days 5 hours 5 minutes from 24 July to 13 August 2008. It has
been documented that the subarctic-subtropical transition area
(around 40°S) is one of the feeding areas of Bryde’s whales insummer. However, the results of this study revealed that some
Bryde’s whales move from the subarctic-subtropical transitionarea to the sub-tropical area even in summer. This study

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REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

provided the first information on continuous movement ofBryde’s whales in the offshore western North Pacific in summer.
The JARPN II review workshop in 2009 had recommended
satellite tracking experiments on the whale species studied, as
a long-term task. This study started such experiments in the
Bryde’s whale. Further satellite tracking experiments have thepotential to complement the genetic studies on stock structure
in sub-areas 1 and 2.
In conclusion the results of the genetics, which included anevaluation of the statistical power, mark-recapture and satellite
tracking analyses suggested a single stock in sub-area 1.
Genetic analyses suggested the possibility of additional stock
structure in sub-area 2.
Based on these results, it may be necessary to re-consider
the plausibility of the four hypotheses on stock structureagreed during the RMP Implementation in 2007. Hypotheses
1 (single stock) and 4 (three stocks) should be assigned lower
plausibility.
Objective 3, Sub-objective 4
To investigate the plausibility of a single stock of sei whalein the pelagic regions of the North Pacific (‘North Pacificpelagic’), using all genetic samples available from different
sources till 2014, and different genetic markers.
SC/F16/JR46 examined genetically a total of 1,554 sei
whales with mtDNA control region sequencing (487bp) andmicrosatellite DNA (17 loci) to investigate population genetic
structure of this species in the North Pacific. Samples wereavailable from different sources, JARPN II (catches) (200214), POWER (biopsy) (2010-12) and past commercial (catches)(1972-73). For the heterogeneity test two longitudinal sectors
were defined in the North Pacific: Western and Eastern at 180°,
which covered this ocean basin widely from approximately
145°E to 135°W. No significant spatial genetic heterogeneity
was found by the two genetic markers. A phylogenetic treeof 82 mtDNA haplotypes showed several clusters, but none
was supported by high bootstrap values. Whales from bothWestern and Eastern sectors were widely distributed throughthe clusters. Taken as a whole, the genetic information in thisstudy is consistent with the view that the oceanic regions of theNorth Pacific are occupied by a single stock of sei whale.
SC/F16/JR47 used microsatellite DNA markers to analyse
samples of sei whales collected widely from the North Pacificat the same time of the year in order to test spatial genetic
heterogeneity in this ocean basin. Although we have beenreporting results of the genetic studies on the North Pacificsei whales to previous IWC/SC meetings, this study is thefirst to utilise temporally similar (collected at the same year),yet geographically very different, samples (covered west-end
to east-end of the North Pacific). This study used samples
collected from the northwestern (JARPN II), northcentral(POWER), and northeastern (POWER) areas of the North
Pacific in the same summer seasons in 2010, 2011 and 2012.
No evidences of significant genetic differences between thesamples from JARPN II and POWER in each of the three
years were found. Each yearly sample was then combined
as JARPN II as well as POWER samples, respectively. No
significant genetic differences were detected between these
two samples. We used genotypic profiles of each whale in the
POWER biopsy samples to find any cases of matching to theindividuals in the JARPN II samples, no matching was found atall. In conclusion, this study failed to demonstrate evidence ofmultiple stocks of sei whales in the North Pacific.
Kanda et al. (2015) [SC/F16/JR48] presented a review ofpast studies on sei whales in order to describe stock structure
hypotheses for the species in the North Pacific. Evidence
obtained from different kinds of the analyses using markrecapture, sighting, catch history, and genetic data shed lighton patterns of distribution and migration of the sei whales,facilitating the hypothesis development. The mark-recapture
data indicated that whales from the same breeding area
distribute widely in the feeding area over almost the entire
North Pacific. Although historical catch data from commercial

whaling era had shown heterogeneous distribution of the sei
whales, genetic evidence indicated no temporal and spatial
genetic differences among the whales obtained from the entire
North Pacific. The heterogeneous catch distribution appeared
to reflect non-random operations of the commercial whaling aswell as patchy distribution of their prey species. Overall, basedon the series of the available evidence we propose a single
stock hypothesis for sei whales in the North Pacific.
In conclusion, results of the genetic analysis confirmed
the view of a single stock of sei whale in pelagic regions of
the North Pacific, as agreed previously by the IWC SC. This
information is useful for the IWC SC’s in depth assessment of
this species in the North Pacific.
Other species and analyses
SC/F16/JR49 examined genetic variation at 15 microsatellite
DNA loci and mitochondrial DNA (mtDNA) control region
sequences (338bp) in sperm whales collected during JARPN
II from 2000 to 2013 in order to examine the effectivenessof these genetic markers for studies of stock structure in thisspecies. Analyses of mtDNA and microsatellite markers in a
total of 56 sperm whales (16 males; 40 females) confirmed thatthese genetic markers were variable enough to explore stock
structure of sperm whales. The overall heterozygosity over 15loci was 0.730 while the nucleotide and haplotype diversity
were 0.0038 and 0.7188, respectively. Statistical tests foundno evidence of deviation from the expected Hardy-Weinberg
genotypic proportion at all of the 15 microsatellite loci. At
this point, no signal of multiple stocks of sperm whale in thewestern North Pacific off Japan was detected.
SC/F16/JR50 examined genetic variation at 14
microsatellite DNA loci and mitochondrial DNA (mtDNA)
control region sequences (275bp) in right whales from thewestern North Pacific and Antarctic Area IV. Genetic analyses
were based on biopsy samples collected during the surveysof the JARPN II in 2011 and 2012 (n=15), and JARPA II in
1993/94-2009/10 (n=67). The overall heterozygosity was
0.630 and 0.650 for North Pacific and southern right whales,
respectively, while the nucleotide diversity/haplotype diversity
were 0.0222/0.9048 and 0.0234/0.7743, respectively. Statisticaltests found no evidence of deviation from the expected HardyWeinberg genotypic proportion in each of the oceanic basins.
The Kimura’s two parameter net interpopulational distance
was 0.0358 (mtDNA) while the Nei’s genetic distance (Da)was 0.7582 (microsatellite DNA), between North Pacific andsouthern right whale. A phylogenetic tree separated mtDNA
haplotypes of the North Pacific, North Atlantic and southern
right whales.
SC/F16/JR51 examined the distribution of the number ofnucleotide substitutions between all pairs of individuals withinwestern North Pacific O and J stock common minke, Bryde’s,
sei and right whales, to investigate whether the pattern of
distribution is indicative of exponential population growth (inevolutionary terms), and thus of non-equilibrium. According
to Slatkin and Hudson (1991), unimodality of the frequency
distribution is indicative of exponential population growth,and this pattern was found in the O stock common minke andsei whales. In contrast multimode pattern in the frequency
distribution was found in the J stock common, Bryde’s andright whales, which is inconsistent with exponential population
expansion.
In conclusion, the genetic markers used are useful for
future studies on stock structure in the sperm and North Pacificright whales. Right whales from different ocean basins wereconfirmed as phylogenetically distinct. Preliminary analyses
suggested that the O stock common minke, sei and possibly
Bryde’s whales are under exponential population growth.
Respond to recommendations from the 2009 JARPN II Review
Workshop
Most of the simple, more extensive and long term mattersrecommended in 2009 were addressed in the analyses on stock
structure presented to the workshop (see proponents’ response
column in Annex D).

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

4.4 Panel review, conclusions and recommendations
4.4.1 Overview
The Panel notes the substantial amount of field, laboratory
and analytical work undertaken under the JARPN II
programme. The proponents presented information andanalyses on stock structure related issues in 25 papers (SC/F16/JR38-41; Kishiro and Miyashita (2011) [SC/F16/JR42];
SC/F16/JR43-44; Murase et al. (2016) [SC/F16/JR45]; SC/
F16/JR46-47; Kanda et al. (2015) [SC/F16/JR48]; SC/
F16/JR49-51; and 11 of the ‘ForInfo’ papers), with a focus
on elucidating stock structure hypotheses for common
minke whales, Bryde’s whales and sei whales, as well as
providing information on sperm whales and right whalesin the North Pacific and Sea of Japan. The data presented
included genetic data (mainly microsatellite genotypes andmitochondrial control region DNA sequences) as well as
age data, morphological data (e.g. length and flipper patch
variation) and telemetry data.
The Panel notes that most documents on stock identity
relied heavily upon reference to previous IWC ScientificCommittee unpublished papers, IWC reports and other SC/
F16/JR documents, which made it difficult to assess theinformation and employed protocols. The Panel refers to
its over-arching recommendation relating to the need for asingle consolidated and integrated paper for each objective(see Items 10 and 11).
The Panel agrees that most of the recommendations andsuggestions proposed by the 2009 Panel have been addressed,
for example those related to data quality procedures,
estimates of genetic divergence, p-values, applying FDR
and assessment of statistical power (and see Item 4.4.3). It
noted earlier recommendations suggesting the application of
non-equilibrium methods to the analysis of stock structure.
However, the Panel recognises that such approaches do notwork well at very low levels of genetic divergence (i.e. FST<0.01), which may imply that they cannot be successfully
applied to these data.
A key and fundamental issue raised by the 2009 Panel,that is applicable to studies within and outside the JARPN
II programme, is the need to assess demographic linkage,i.e. what is a ‘stock’ within the specific context in which it isbeing used and hence whether different stocks, should theyoccur, are detectable using the applied analytical approaches.
This is discussed further below.
ASSESSMENT OF GENETIC DATA TO IDENTIFY DIFFERENT‘STOCKS’The genetic data sets generated are uniquely comprehensive
for microsatellite-based studies, typically consisting of 1417 microsatellite loci typed in 100s (e.g. right whales) toseveral thousand samples (e.g. common minke whale), a
non-trivial effort which the Panel commends.
The focus of most analyses was testing the current RMPdefined stock hypotheses, i.e. the objective was to provide
information relevant to management and conservation.
The population genetic assessments undertaken weresimilar across most species, including the estimation ofstandard indices of genetic diversity, the estimation of thedegree of genetic divergence among sample partitions andhomogeneity tests. In addition, clustering algorithms (e.g.
STRUCTURE and DAPC) were applied to some data sets.
In response to the 2009 Panel, an assessment of the power toreject homogeneity at different migration rates was estimated(using EASYPOP).
With the exception of Bryde’s whale (see below)
statistically significant deviations from homogeneity were
only detected between samples from different hemispheres

541(i.e. Southern Hemisphere and western North Pacific rightwhales) or between the O and J stocks of common minke
whales. In all other homogeneity tests, FST estimates wereclose to zero and homogeneity was not rejected. The
assessment of statistical power suggested high power toreject panmixia at migration rates below 100 (~FST <0.001,
p>0.8).
The proponents inferred the low levels of spatiotemporal genetic divergence and general inability to reject
homogeneity as support for a single stock in the case ofcommon minke whale O stock, sei whales and sperm whales.
The Panel notes that although the presented analyses
show evidence of spatial genetic homogeneity in most
species, the biological interpretation (and hence management
implications) of low genetic divergence is unclear in general,
including for JARPN II.
The Panel also notes that in the absence of deriving thequalitative and quantitative expectations at different levelsof genetic divergence among stocks, it is impossible todetermine if genetic homogeneity implies strong support fora single stock. Similarly, statistical power cannot be assessed
properly if the target effect size is unknown. Accordingly,
the Panel proposes that a quantitative working definitionof what constitutes a ‘stock’ in the targeted species/area
is developed by the proponents for consideration by the
Scientific Committee (see Item 4.4.3.1).
The Panel notes that several assessments were of aqualitative, rather than quantitative, nature. One such
example is SC/F16/JR38 where the possibility of a sub-O
stock origin of ‘unassigned’ common minke whale samples(i.e. samples with assignment probabilities below 90% toeither O or J stock) was rejected based upon the apparent
randomness in phylogenetic placement of mitochondria
control region haplotypes and the absence of a restricted
geographic location of such unassigned samples.
In general, the genetic data analyses continue to rely
heavily on homogeneity tests as well as moment-based
estimators of genetic divergence, such as FST. This point
was made by the 2009 Panel as well, which suggested that
non-equilibrium approaches (i.e. for divergence) be applied
to the data, which could explain the low levels of genetic
divergence; a suggestion that the Panel reiterates (see Item
4.4.3.1).
Although some assessments employed non-spatial
approaches (e.g., STRUCTURE and DAPC for the caseof O stock common minke whales, SC/F16/JR40) mostassessments relied upon testing for spatial heterogeneity
and/or estimating spatial divergence, which harbours a riskof failure to detect the presence of two ‘stocks’ within asingle spatial stratum (see Item 4.4.2.1).
While the sample sizes in most species are exceptional,
the Panel notes that increasing the number of genotyped
loci per sample, as opposed to the number of individuals, islikely to add substantially to the precision and accuracy ofthe estimates of population genetic parameters (i.e. θ, Nm andpopulation divergence time).
The 2009 Panel recommended that data quality
procedures be explained. In terms of the genetic data
generated for this review, most primary papers referred to
Kanda et al. (2014b) with regards to data generation andquality control of the presented genetic data. Kanda et al.
(2014b) adheres to the IWC guidelines for DNA data quality
standards. However, the Panel notes that no proceduresor estimates of genotyping and DNA sequence error rates
were reported in Kanda et al. (2014b) or in the SC/F16/JR
documents.

Quantify for each quantity of interest for each objective sources of uncertainty and relationship with sample
size and use to evaluate progress on meeting objectives, effect of changing sample size on meeting
objectives and suitability of sampling design.

542

Not addressed

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II
Table 3

Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to sampling design and sample size (see text).
Brief summary of 2009 recommendations/suggestions

Panelevaluation

Panel comments

The genetic assessments should include a brief description ofprocedures to ensure data quality (with reference to IWC guidelines
for DNA data quality).

Largely
addressed

Kanda et al. (2014b) (a revised version of Kanda et al., 2014a)
presented a summary of the sampling and laboratory protocols
employed for the genetic studies under Objective 3 of the JARPN
II conducted at the Institute of Cetacean Research. The individual
stock structure-related documents also confirm that the IWC
guidelines for DNA data quality (Tiedemann et al., 2012) havebeen adhered to.

Revised papers should include estimates of genetic divergence (alongwith levels of uncertainty) in addition to probabilities of homogeneity.

Largely
addressed

FST estimates were provided when appropriate. Levels of
uncertainty associated with divergence estimates were, however,not reported.

P values (and divergence estimates) should be reported for all locicombined rather than for each locus separately.

Addressed-

Multiple testing issues: (a) apply False Discovery Rates; (b) exercise
discretion in the number of pairwise comparisons evaluated.

Addressed-

Provide more details on the analyses involving the program
STRUCTURE.

Addressed-

Include a brief discussion of experimental design with respect to sampling (explaining how the design specifically addresses uncertainties
related to stock structure, e.g. whether the spatial and temporal
coverage of samples of minke whales has been sufficient to test
adequately the alternative stock structure hypotheses).

Partlyaddressed

Sampling procedures were adequately described, but no
explanations of the sampling rationale in relation to resolving theuncertainties related to stock structure were provided.

Redo the Boundary Rank analyses (Taylor and Martien, 2002) withnew data for common minke whales.

No longer
relevant-

Integrate Korean by-catch samples into the new datasets to look atheterogeneity for common minke whales.

Addressed-

Undertake the assessments of power to simulate data to evaluate power
to detect a specified fraction of a putative stock (e.g. the hypothetical
W stock of common minke whales) in an overall sample using
simulated data.

Partlyaddressed

Assessments of statistical power were performed with regards torejecting homogeneity under different migration rates.

Undertake tests for population genetic (drift-mutation-migration)equilibrium.

Partlyaddressed

Non-equilibrium was tested for long-term demographic population trends via mismatch distribution analysis of mitochondrial DNA. However, assessments of short-term deviations
from mutation-drift equilibrium (Piry et al., 1999) were notundertaken. However, these assessments may be infeasible.

Employ approaches that do not rely upon the assumption of mutationdrift-migration equilibrium when estimating population divergence.

Not
addressed-

Attempt the detection of related pairs of individuals

Partlyaddressed

Spatially-explicit relatedness analyses are underway but only
preliminary results were shown at the Workshop itself.

Undertake multivariate analyses of morphological data with respect tostock structure.

Largely
addressed

Results presented for some cases, mainly for O and J stock
common minke whales.

Use of past and present contaminant data should be an integrative
study of stock structure.

Notapplicable

There are insufficient Persistent Organic Pollutant (POP) data forthis to be productive.

Initiate satellite tagging to narrow the range of plausible stock
structure hypotheses.

Partlyaddressed

Proof of principle has been achieved by tagging a single common
minke whale and two Bryde’s whales, but many more must be
tagged if the results are to be used to inform discussions.

1
SC/F16/JR43 outlines a statistical catch-at-age model
for common minke whales east of Japan (the areas where
the putative OW- and OE-stocks are found). It is fitted toestimates of abundance and catch age-composition data
under two stock structure hypotheses (mimicking the A andC stock structure hypotheses of the current Implementation
Simulation Trials). The model is noted to be a first step
towards using the age data for conditioning and to selectamong stock structure hypotheses. Age data have been usedpreviously when conditioning operating models to evaluatevariants of the RMP (e.g. for the North Atlantic fin whales)
and have informed other implementations (e.g. for western
North Pacific Bryde’s whales). Thus, if the Implementation
Simulation Trials for the western North Pacific minke
whales are to be revised in the future, the age data should
be included in the conditioning process. The specificassumptions underlying SC/F16/JR43 will need to be refined
before the age data are included in the conditioning process.

In particular, consideration should be given to: (a) using
alternative selectivity patterns; (b) using a spatial structurethat better matches that underlying the trials; and (c) fitting
to the actual abundance estimates rather than the output
from trials. In terms of selectivity, it would be expected thatthe selectivity pattern would be dome-shaped for the inshore
areas, at least for the case in which there is a single O-stock.
Ageing of commercial data could be used to inform thepre-1987 selection pattern, which is currently pre-specified
based on values estimated for North Atlantic minke whales.
See Item 9.1.1 for additional discussion of the age data.
Concerning the specific sub-objectives 1- 4 (Objective
3, above) the Panel agrees that substantial progress wasmade with regards to sub-objective 1 in terms of the spatiotemporal distribution of J stock common minke whales in
the waters off either side of Japan. The Panel notes that anassessment of putative OW and OE common minke whale
stocks was undertaken following the recommendations by

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017the 2009 Panel. However, the Panel realises that further
progress on this sub-objective relies (due to the absence ofgenetic heterogeneity) on identifying the defining dispersal
rates between the putative OW and OE ‘stocks’ and thegenetic expectations at these dispersal rates.
The Panel agrees that the same issue pertains to identifying
stocks in North Pacific sei whales (sub-objective 4) whereequally low levels of spatial heterogeneity were observed.
The Panel agrees that progress was made in terms of
reducing the number of RMP stock hypotheses of North
Pacific Bryde’s whales (sub-objective 2) although additional
data analyses with existing data could potentially assess
further sub-structuring within Sub-areas 1 and 2.
4.4.2 Evaluation of progress with the 2009 recommendations (see Annex D for a complete summary table)
The 2009 Panel had made a large number of recommendations
to further the work on stock structure. All but two of these
had been fully or partly addressed as shown in Table 3.
4.4.3 Recommendations

4.4.3.1 MEDIUM-LONG TERM RECOMMENDATIONSThe Panel has developed the following recommendations
for the medium-term, i.e. normally to be completed 2-3
years after the 2016 Annual Meeting.
(1) In order to facilitate a more definitive discrimination
between single and multiple stock hypotheses, work
should be undertaken to determine the demographic
dispersal rates among areas at which whales indifferent areas can be managed as a single stock. Suchan assessment of ‘critical’ dispersal rates by specificcase (i.e. species, area[s] and management objective)
and the corresponding levels of genetic divergence,
should enable the rejection of 2+ stock hypotheses andhence confirmation of a single stock where applicable.
This general and difficult issue had been raised by the2009 Panel, but without specific recommendations as
to how it might be achieved. However, recently anillustrative example of how this might be achieved has
been presented in Van der Zee and Punt (2014) and thisapproach is recommended to the proponents.
(2) As recommended by the 2009 Panel, analytical
approaches should be applied that do not assume
mutation-drift-migration equilibrium, such as the IM
methods (Hey, 2010).
(3) Serious consideration should be given to using genomewide SNP genotyping approaches, such as RAD
sequencing and GBS (Elshire et al., 2011; Miller et al.,2007). This will increase the data per sample thereby
improving the accuracy and precision of genetic
parameter estimates and facilitate additional analyses(Hey and Machado, 2003; Robinson et al., 2014).
(4) A focussed satellite tagging programme should be
developed to greatly increase sample size to assess
individual migration in the context of stock structure
hypotheses more thoroughly.
4.4.3.2 SHORT TERM RECOMMENDATIONSThe Panel has developed a number of recommendations
for the short-term, i.e. to be completed ideally by the 2016
Annual Meeting but certainly by the 2017 Annual Meeting.
(1) All inferences regarding ‘randomness’ of observations(e.g. unassigned common minke whales) should besubstantiated by a statistical assessment of the presumed
randomness.
(2) The presence of multiple stocks within sample partitions
should be assessed (employing, e.g., STRUCTURE andDAPC) for Bryde’s and sei whales.

543(3) More explicit information on quality checks be provided
in each study as well as study-specific estimates or
genotyping and DNA sequencing error rates.
In addition, in Table 4 the Panel makes a numberof specific comments on individual papers to assist the
proponents in their revisions, ideally for the 2016 Annual
Meeting. These comments are made in the context of theover-riding recommendation for a consolidation of papers
found under Items 10 and 11.
5. FEEDING ECOLOGY AND ECOSYSTEMSTUDIES: PART 1. GENERAL INFORMATIONTO INFORM ECOSYSTEM MODELLING ON
OCEANOGRAPHY, DISTRIBUTION, ABUNDANCEAND STATUS OF WHALE STOCKS
5.1 Proponents’ summary of objectives including
modifications, if any, since the start of the programme

The following three sub-objectives under main Objective 1
were considered under this agenda item.
Sub-objective 1: To investigate the oceanographic
conditions that are relevant for the understanding of preyspecies distribution and abundance in the research area.
Sub-objective 2: To investigate the distribution pattern ofbaleen whales in the research area and the possible factors
affecting such pattern.
Sub-objective 3: To estimate abundance of baleen andsperm whales using JARPN II sighting data and standard
IWC SC methodology.
Initially, in-situ oceanographic observations (e.g. CTD)
were planned to obtain the data (Government of Japan, 2000).
However, analyses presented to this workshop relied on thedata from an ocean model and satellites as the techniques
and technologies are developed substantially in the period of
JARPN II. In the abundance estimation using standard method(i.e. design based method) presented to this workshop, Beaufort
Sea state was used as a covariate of detection function. These
two are major modifications since the start of the programme.

5.2 Overview of conclusions and recommendations fromthe 2009 Workshop
A short overview on the 2009 conclusions and recommendations on the component of JARPN II on oceanographic
features related to whale distribution, distribution andabundance of whales was provided by the Chair. On these
aspects, the 2009 Panel had noted that the programme
was progressing towards addressing its objectives andhad recommended that this work continues. In termsof relationships between oceanographic features andwhale distribution the 2009 Panel had made a number ofrecommendations in relation to data analyses:(a) to incorporate into the index of density, the sightability of detected groups (e.g. effective strip half
widths that include appropriate covariates such as
weather conditions);(b) to test whether any chosen model is an improvement
over a null, uninformative model and to validate
model results (e.g. by comparing the modelledresults, not only with index of densities from the
JARPN II, but also with JARPN and/or othersurvey data and by exploring cross-validation type
techniques);(c) to conduct more spatial modelling analyses (e.g.
using other appropriate modelling techniques suchas GAMs or logistic regressions); and(d) to incorporate shipboard oceanographic data infuture models, together with potential additional
oceanographic/biological features, including modelling the satellite or in situ measurements ofchlorophyll to estimate primary productivity.

544

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II
Table 4
Comments on individual papers presented related to stock structure to assist with their revision.

SC/F16/ Comments
JR38

The Panel noted the somewhat arbitrary choice of a 90% assignment probability as criterion for assigning individuals to a specific cluster. The
Panel acknowledged that the assignments of samples to the O and J stock ‘clusters’ likely did capture general spatio-temporal trends in thesamples. The observation that number of successful assignments (e.g. at >90% assignment probability) is correlated with the number of loci iswell-known. However, this observation does not prove the absence of further stock structure as suggested in SC/F16/JR38 without further
assessment. Such an assessment would entail simulations with increasing numbers of loci including a third ‘cluster’ at a level of divergence that
it would go ‘undetected’ by a STRUCTURE estimation of the number of clusters. The inferred randomness in haplotype affinities of specimens
classified ‘unknown’ should be tested for statistically. In addition, representation of mtDNA data in un-rooted haplotype networks (integrating
haplotype frequency, relatedness and geographical occurrence) might be superior to the NJ trees presented in SC/F16/JR38.

JR39

The Panel noted that the sample size for the ‘no GAL’ flipper type was low (n=16). Overall the results in SC/F16/JR39 confirmed the oft
observed distinction of the J and O stock in the common minke whale, but do not allow for unambiguous stock assignment of individuals, assome morphotypes are found both in J and O stock individuals.

JR40

The Panel noted the very low degree of genetic divergence among areas, as has been reported earlier. In light of the low degree of divergenceamong cluster O stock assigned samples it is unsurprising that clustering algorithms, such as STRUCTURE and DAPC fail to detect any
additional sub-structuring. This result is further exacerbated by the ‘purging’ of assumed non-O stock samples. The Panel agreed to theassessment of statistical power by simulation. However, the biological/management interpretation of the failure to detect additional clusters
depends upon what dispersal (and hence Nm) rates are considered critical in terms of delineating stocks in this specific case. As a rough illustrationconsider 133 migrants per generation. If the generation time is at 10 years this migration rate (not necessarily the same as the dispersal rate)would correspond to ~13 migrants/year. In the case of an effective population size at 6,660 individuals, 13 migrants correspond to 0.2% which
may be little migration from a demographic perspective, but sufficient to maintain the observed genetic cohesion. This example underlines theneed to define what dispersal rates are relevant for the specific management objectives for the North Pacific common minke whale.

JR41

The Panel generally agreed with the analysis which provides an indication that the divergence among J and O stock is reflected not only ingenetics, but also in morphometry. However, it notes that PCA plots of individuals might aid visualisation of potential outliers/further
substructure. It also notes that the objective for using PCA was to reduce 11 morphometric measurements to fewer principal components (PC)to use in discriminant analyses, e.g. to discriminate between J stock and O stock minke whales. The sample size of J stock whales was smaller
than that of O stock whales: 24 compared to 760. O stock whales were sampled in all 20 years included in the analysis, but J stock whales in
only 9; 376 O stock whales were sampled in those 9 years. All 760 O stock whales were used in the discriminant analysis comparing O and J
stocks, but only the years with J stock whales should have been used.

JR43

The use of age data is discussed under Item 9.

JR44

The Panel noted that the homogeneity tests conducted in SC/F16/JR44 were aimed only at assessing spatial heterogeneity within sub-area 1 as
well as between sub-areas 1 and 2. However, no additional analyses (e.g., STRUCTURE or DAPC) were undertaken to assess the possible
presence of multiple clusters (i.e., populations) in each sub-area (or both sub-areas) making difficult to make inferences about the number ofstocks in the sub-areas. Therefore, the conducted analysis is only able to reject stock structure hypothesis 1. It cannot distinguish between
hypotheses 2 and 3 and did not test hypothesis 4.

JR46

The Panel noted that the homogeneity tests performed tested for spatial heterogeneity only and not for the presence of multiple clusters (e.g.
using STRUCTURE or DAPC). Consequently, the conclusion of a single stock is premature. It would be helpful to be more explicit in how thecovered area relates to the stock hypotheses forwarded by Mizroch et al. (2015).

JR47

The Panel noted that the sample sizes in the POWER data set are small (n=13, 29 and 35) which introduces potential issues of statistical power.
For example, the comparison between the JARPN II (2010) and POWER (2010) samples resulted in a FST at 0.011. In other assessments at
similar degrees of divergence with larger sample sizes (as well as simulations aimed at testing statistical power, see SC/F16/JR40) such a degreeof divergence resulted in rejection of panmixia.

JR48*

The Panel agreed this review of available data is relevant to sei whale stock structure in the North Pacific. The presented information favours asingle-stock hypothesis, but did not explicitly conduct a statistical evaluation of alternative hypotheses. It is unclear whether the presented markrecapture data contain (or are identical to) those presented by Masaki (1976), then interpreted as being indicative of three stocks. Also, no
consideration of the stock hypotheses forwarded by Mizroch et al. (2015) is provided.

JR49

The Panel notes that any inference of stock structure on sperm whale is preliminary, given the small sample size and incomplete geographicalcoverage. Nonetheless, at this stage, a hypothesis-free analysis (e.g. STRUCTURE, DAPC) could be informative. The analyses should take into
account the possibility that multiple individuals may have been sampled from the same matrilineal pod, which may bias estimations of population
divergence based upon mitochondrial (and potentially nuclear) data (Bogstad et al., 2015).

JR51

The Panel understood that the analysis was performed to address previous recommendations aimed at assessing the presence and effects ofdeviations from the mutation-migration-drift equilibrium that is an underlying assumption to many population genetic inferences and parameter
estimations. However, mismatch distributions in mitochondrial DNA generally capture (if Ne is large) longer time scale expansions in a single
population. In terms of stock structure, the aspect of concern is whether a low divergence between populations represent recent divergence (butlittle gene flow) or high gene flow; cases with potentially very different management consequences. An example of approaches to estimate Nm
at mutation-migration-drift disequilibrium is the so-called IM (isolation with migration) methods.

Kanda
et al.
(2009)The Panel notes that this paper principally covers the same subject as SC/F16/JR38 and SC/F16/JR40. Here, STRUCTURE outputs are provided
for k=2 and k=3. This analysis favours the two stock (i.e. J and O) hypothesis. Generally, it would be desirable to adjoin the information of thispaper and the papers SC/F16/JR38 and SC/F16/JR40 in order to arrive at a synopsis statement on common minke whale stock structure.

Kanda
et al.
(2010)The validity of the inferred divergence within the Korean samples as well as between some Korean and Japanese samples is, however, difficult
to assess, as p-values for pairwise FST estimates are not provided and apparently no correction for multiple pairwise comparisons was performed.

Kanda The Panel generally welcomes this description of procedures. With regard to error assessments, however, it is necessary to provide quantitative
et al. information with regard to the number of samples re-typed. In addition, the encountered error rates should be reported.
(2014b)Kanda
et al.
(2006)The Panel notes that this paper covers a similar topic as SC/F16/JR46, albeit with much fewer samples. No hypothesis-free analysis (e.g.
STRUCTURE, DAPC) was performed on the data, such that the inference of a single stock appears premature. Further comments made to
SC/F16/JR46 also apply here.

*Kanda*Kanda et al.
(2015).et al. (2015).

2

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

In terms of fieldwork, the 2009 Panel had alsosuggested that the proponents consider conducting future
oceanographic surveys over an area larger than at present,
not only to further investigate oceanographic relationships,
but also to improve abundance estimates for a variety of
species.
With respect to abundance data, the 2009 Panel hademphasised that this is of the great importance for estimating
consumption, biomass, energy requirements, exchange rates
and, therefore, ecosystem modelling, and had recommended
that the proponents should: (a) investigate whether datacollected over the 1994-2007 period for a variety of large
whales can be used to provide information on trends; (b)work up the photo-identification data and compare withcatalogues elsewhere in the North Pacific; and (c) improve
the precision of the abundance estimates that are used to
extrapolate to population-level rates, for both the coastal(possibly with regular well-designed aerial surveys) and theoffshore regions (with a full synoptic survey).
5.3 Proponents summary (incl. response to 2009
Workshop)
5.3.1 Oceanography

5.3.1.1 PROPONENTS SUMMARY OF THE RESULTS(INCL. RESPONSE TO 2009 WORKSHOP)

SC/F16/JR05 examined oceanographic conditions in the
JARPN II survey area on a broad scale while SC/F16/JR06
examined oceanographic conditions on a local scale (i.e. off
Kushiro). Data from the ocean forecast system, FRA-ROMS,
were used in the analyses. FRA-ROMS is an ocean forecast
system developed by Fisheries Research Agency (FRA) basedon Regional Ocean Modeling System (ROMS).
In the previous workshop held in 2009, the panel suggested
that the salinity CTD data must be corrected/calibrated using
the water samples that were simultaneously collected with the
CTD data. The suggestion was not applicable directly as theanalyses presented to this workshop relied on the data from an
ocean model and satellites. Nevertheless, calibrated CTD data
were used as one of the inputs to the model.
Changes in area (km2) of four water types (Oyashio, cold,
warm and Kuroshio) by each month (April-September) in thewhole JARPN II survey area from 2000 to 2013 was investigatedin SC/F16/JR05. There was no statistically significant trend forthe area except the cold water in September. Negative valuesof annual mean of the Pacific Decadal Oscillation (PDO) index
were dominant in the period from 2000 to 2013. Generally, sea
surface temperature in the western North Pacific is high in thenegative phase of the PDO. The results of this paper indicatedthat overall oceanographic conditions in the whole JARPN II
survey area from 2000 to 2013 were relatively stable although
year to year variations and spatial heterogeneity of distributionof water types were observed. It was reported that the climate
regime in the North Pacific changed in 1998. The regime of theperiod of JARPN II cannot be determined until next regime
shift is detected although the PDO indicated that the regime
might be shifted after 2013.
Oceanographic conditions in the survey area of JARPN II
coastal component off Kushiro were investigated in SC/F16/
JR06. Oyashio was dominant water type in subsurface in thesurvey area. Mean water temperature at 10m water depth in
the survey in September was generally decreased from 2000
to 2004 then increased to 2006. It decreased again to 2009. It
was relatively stable from 2009 to 2013. Spatial distributionof water temperature at 10m depth was highly variable fromyear to year.
In conclusion, JARPN II from 2000 to 2013 were conducted
in relatively stable oceanographic conditions on a broader
scale while oceanographic conditions on a local scale (i.e.
off Kushiro) in the same period were highly variable. Further
monitoring of oceanographic conditions in the survey area isrequired to determine the regime of the period of JARPN II.

545

5.3.2 Distribution of large whales

5.3.2.1 PROPONENTS SUMMARY OF THE RESULTS(INCL. RESPONSE TO 2009 WORKSHOP)

A total of four papers including two peer reviewed papers were
presented to the workshop (SC/F16/JR07; Murase et al. (2014)[SC/F16/JR08]; SC/F16/JR09; Sasaki et al. (2013) [SC/F16/
JR10]. In addition, two papers related to this agenda item werealso presented (SC/F16/JR16 and SC/F16/JR25).
Meso and micro scales linking feeding ecology with
distribution were considered in the papers. The following
definition of the scales was described in the IWC/SC report(IWC, 2003, p.67). At meso scale, whales move over days andweeks in search of preferred local abundance of food while
they dive and search for food within localised area at micro
scale. Meso scale distribution was considered in SC/F16/JR07;
Murase et al. (2014) [SC/F16/JR08]; SC/F16/JR09; and Sasaki
et al. (2013) [SC/F16/JR10] while micro scale distribution wasconsidered in SC/F16/JR16 and SC/F16/JR25. The results fromthese papers were integrated to obtain a synthesis on spatial
distribution of whales in relation to feeding ecology.
Spatial distribution and relative abundance of common
minke, sei and Bryde’s whales in JARPN II (2003-13) were
estimated using GAMs in SC/F16/JR07. All species shifted
their distribution area toward the north of the survey area from
May to September but the extents of shift different among
species. Relative abundance of common minke whales washigh in coastal area of Japan. Relative abundance of sei whales
was high in the offshore area of the survey area where SST
was moderate within the area. Relative abundance of Bryde’s
whales was high in the southern part of the survey area where
SST was high. The results suggested that spatial distributionsof three baleen whale species were segregated in the JARPN II
survey area although some overlaps occurred. Extent of directcompetition (e.g. competitive exclusion of feeding area) could
be minimal among the species but indirect competition for preymight occur as they share same prey species.
Preliminary estimates of prey consumption of sei whales
in JARPN II (2003-13) obtained in SC/F16/JR16 (see also
Item 6.3.1). The paper was an extension of SC/F16/JR07. Twolevels of models are constructed to achieve the goal. Firstly,
relative abundance of sei whales in relation with oceanographic
conditions was estimated by using a GAM (SC/F16/JR07).
Secondary, amount of prey consumed by a sei whale in relation
with oceanographic conditions was also estimated by using
GAM. Finally, prey consumption of sei whales in the JARPN
II survey area is calculated as the product of these two models.
SST was selected as environmental covariate in the first andsecond models. However, the shape of functional form forthe first level model (prey consumption) was relatively flat incomparison with the second level model (abundance).
Diving behaviour of sei whales and vertical distribution of
their prey were studied in SC/F16/JR25 (see also Item 6.3.2).
Small acoustic time depth transmitters (pingers) were attached
to two sei whales and their behaviours were recorded for10.2 and 32.0 hours, respectively. The results illustrated theircomplex feeding behaviour at micro scale.
In conclusion, the integrative approach conducted under
JARPN II revealed new insight on spatial distribution ofwhales in relation to feeding ecology. Spatial distributionof sei whales at meso scale could be largely determined byoceanographic conditions such as SST (SC/F16/JR07). Sei
whales could then search for their prey under the optimal
oceanographic conditions at micro scale (SC/F16/JR16 andSC/F16/JR25). Relationship between meso and micro scale
should be investigated further to clarify mechanism of feeding
ecology of whales with distribution. Such study will contributeto improve analyses on abundance and ecology of whales.
The relationship between the distribution of sei whales andoceanographic fronts was investigated using GAM in Murase etal. (2014) [SC/F16/JR08]. Sei whales were concentrated northand south of the Subarctic Front (SAF) and the areas from 250
to 300km north and from 100 to 200km south of the SAF were
estimated as high-density areas of sei whales. The entire inter-

546

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

frontal zone between the polar front (PF) and SAF featured anelevated concentration of sei whales, and the area south of the
PF and along the SAF was identified as an important feeding
ground of sei whales in July from 2000 to 2007.
The monthly distribution patterns of blue, fin, humpback
and North Pacific right whales from May to September inthe western North Pacific were investigated in SC/F16/JR09.
Such information has not been available since the cessation ofcommercial whaling. Further continuation of the systematicsighting surveys including in foreign EEZ areas is required to
improve information on seasonal distribution of baleen whales.
The habitat differentiation between sei and Bryde’s whalesin the western North Pacific was investigated in Sasaki et al.
(2013) [SC/F16/JR10]. Data obtained from May to August
2004 and 2005 in JARPN II were used. This study examined
the relationship between oceanographic features derived fromsatellite data and the distribution of sei and Bryde’s whales
using basic statistics. We investigated oceanographic features
including sea surface temperature (SST), sea surface chlorophylla (Chl-a), sea surface height anomalies (SSHA), and depth of
the habitat. These two whale species used habitats with different
SST, Chl-a, and SSHA ranges. The 0.25 mg/m3 Chl-a contour(similar to the definition of the Transition Zone Chlorophyll
Front) was a good indicator that separated the habitats of sei and
Bryde’s whales. Then generalised linear models were used tomodel the probabilities that the whale species would be present
in a habitat and to estimate their habitat distribution throughout
the study area as a function of environmental variables. The
potential habitats of the two species were clearly divided, andthe boundary moved north with seasonal progression.
In conclusion, these three papers (Murase et al., 2014;
Sasaki et al., 2013) [SC/F16/JR09-10] and SC/F16/JR09)
provided new information of spatial distribution of baleen
whales in the western North Pacific. The results could be
used as baseline information for in-depth assessment of these
species such as sei whales.
5.3.3 Abundance and status of stocks

5.3.3.1 PROPONENTS SUMMARY OF THE RESULTS(INCL. RESPONSE TO 2009 WORKSHOP)

SC/F16/JR11 examined the number of western North Pacificcommon minke whales distributed in JARPN II coastal survey
areas. In order to examine an impact of common minke whales
on Japanese fisheries in Kushiro and Sanriku regions throughestimating the amount of prey consumed by minke whales
or using an ecosystem model, it was required to estimate thenumber of common minke whales distributed in each of thesurvey areas during the JARPN II survey periods. Because it
was considered that the impact of minke whales was importantin the operation area of the coastal fishery, the numbers ofcommon minke whales there were estimated .The estimatednumber off Kushiro were 461 and 433 in early (May-June) andlate season (September) in 2012, respectively. The estimatednumber off Sanriku in the early season was 124 in 2012. Notethat these numbers are not abundance estimates of the minke
whale stock in the areas because the sighting data we used forthe estimation covered only a part of the stock distribution.
SC/F16/JR12 examined the number of western North
Pacific common minke, Bryde’s and sei whales in the JARPN
II offshore survey area. In order to examine an impact oflarge whales, such as common minke, Bryde’s and sei whales
on Japanese fisheries through estimating the amount of prey
consumed by these whales or using ecosystem models, it was
required to estimate the number of these whales in the JARPN
II survey area (subareas 7, 8 and 9 excluding foreign EEZ).
Considering the migration pattern of these whales in the areasuggested by previous analysis, the number of the whales
needed to be estimated separately for the early and late seasonsfor each whale species. The estimates were 3,629 (in 2009) and
2,122 (in 2011 and 2012) in the early and 3,080 (in 2008) in thelate season for the common minke whale assuming g(0)=0.789;
2,957 (in 2009) and 1,851 (in 2011 and 2012) in the early and13,306 (in 2008) in the late season for the Bryde’s whales;

4,734 (in 2009) and 2,988 (in 2011 and 2012) in the early and5,086 (in 2008) in the late season for the sei whales, assumingg(0)=1. It is important to note that these estimates should not
be used for assessment because the estimated figures represent
only a part of the population considered.
SC/F16/JR13 examined the number of blue, fin, humpback
and North Pacific right whales in the JARPN II offshore survey
area based on 2008-14 JARPN II surveys. The numbers areto be used for prey consumption estimation and ecosystemmodelling in the western North Pacific. Given that the area is a
migration corridor of the whales, the numbers were estimatedfor early season (May-June) and late season (July-September).
The estimates were 38 (in 2009) and 161 (in 2011 and 2012) inthe early and 958 (in 2008) in the late season for blue whales;
413 (in 2009) and 1,369 (in 2011 and 2012) in the early and3,958 (in 2008) in the late season for the fin whales; 1,136(in 2009) and 1,921 (in 2011 and 2012) in the early and 392(in 2008) in the late season for the humpback whales; 1,147(in 2011 and 2012) in early season and 416 (in 2008) in lateseason for the North Pacific right whales. It is important to notethat these estimates should not be used for assessment because
the estimated figures represent only a part of the populationconsidered.
SC/F16/JR14 examined the number of sperm whales in
the JARPN II offshore survey area based on 2008-14 JARPN
II surveys. The numbers are to be used for prey consumption
estimation and ecosystem modelling in the western North
Pacific. Given that the area is a migration corridor of the whales,
the numbers were estimated for early season (May-June) andlate season (July-September). The estimates were 11,459 (in2009) and 11,652 (in 2011 and 2012) in the early and 10,843(in 2008) in the late season for the sperm whales, assuming thatg(0)=1. It is important to note that these estimates should not
be used for assessment because the estimated figures represent
only a part of the population considered.
In conclusion, the number of whales distributed in the
research area was estimated using the Line Transect method
and IWC SC standard methodology. This was done for common
minke, Bryde’s, sei, blue, fin, humpback, North Pacific rightand sperm whales. Numbers obtained are not representativeof the abundance of each stock but they represent the numberof animals distributed in the research area at specific research
periods. The numbers obtained were used for the estimation of
prey consumption in common minke, Bryde’s and sei whales,
and as input for the ecosystem modelling (all species except for
the North Pacific right whale).
Respond to recommendations from the 2009 JARPN II Review
Workshop
Most of the matters recommended in 2009 related to items
5.3.1.1, 5.3.2.1 and 5.3.3.1 were addressed in the analyses
summarised above (and see proponents’ comments in Annex D).

5.4 Panel review conclusions and recommendations
5.4.1 Overview
5.4.1.1 OCEANOGRAPHYThe Panel agrees that the proponents have begun toincorporate oceanographic information into their analyses;
the selection of appropriate oceanographic parameters is
crucial to detecting important changes in oceanographic
conditions that are relevant for understanding prey and whale
distribution and abundance. In trying to address this, SC/
F16/JR05 examined changes in areas by four water types
by month within the entire survey area and concluded that
oceanographic conditions had been relatively stable during
2000-13. However, the parameters used were based onmean temperature at deeper depths (100m and 200m), i.e.
less variable parameters than at shallower depths. It is notclear whether these parameters are sensitive enough to detect
important oceanographic changes; their behaviour should be
examined in relation to recent regime shifts in 1988 and 1998.

547

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017
Table 5
Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to oceanography, distribution and abundance(and see text).
Brief summary of 2009 recommendations and suggestions
Pool or compare results with other datasets to increase sample size and increase the possibility of data covering periods ofchanging relationships (e.g. previous regime changes), thus allowing patterns to be detected.
Consider conducting future oceanographic surveys over an area larger than at present.
In the long term, integrate into analyses oceanographic and other data collected on the cruises (bottom depth, water column
temperature, salinity, and density) with satellite derived data, such as SST, chlorophyll, and sea surface height.
Correct/calibrate CTD data using simultaneously collected water samples.
Conduct additional analyses (including using techniques such as GAMs or logistic regressions).
Incorporate the sightability of detected groups into the index of density and test whether the chosen model is an improvement
over a null, uninformative model and try to validate the model results using cross-validation techniques and use of outside data.
Investigate whether sightings data from 1994-2007 can be used to provide information on trends.
Work up photo-identification data and compare with other North Pacific catalogues.
Increased effort to obtain better estimates should be a high priority.

Panel evaluation
Addressed.
Addressed.
Partly addressed.
No longer applicable.
Addressed.
Partly addressed.
Not addressed.
Partly addressed.
Partly addressed.

Another paper, SC/F16/JR06 examined mean water
explicit prey consumption) over time. As the proponents
note, this is exacerbated by inconsistency between survey
temperature at 10m depth in September off Kushiro andTable P1
areas and coverage during the JARPN II surveys, but given
indicated that the temperature was highly variable from 2009
Estimates of daily prey consumption (kg) - offshore.
its fundamental importance to the modelling work, increased
to 2013. However, the Panel noted that this area has beenSpecies
Immature male(kg) (seaMaturemale (kg)
Immature female (kg)Maturefemaleeffortsmustbe(kg)
put into examining this (see Item 5.4.3.2).
characterisedby significant
SST
surfacetemperature)1
warmingduringthesummer(July-September)
from
2010
.
Common minke whale
86-94
129-141
83-94
158-166
5.4.1.3 ABUNDANCE
Bryde’swhalenoted that that419-434577-637 prey species
417-428642-707The
Panel
the dominant pelagic
The
Panel
notes that abundance estimates are important for
Sei whale
397-421524-539610-647forminke whales changed
from saury
and anchovy436-468
toa number of the programme’s objectives, including inter alia
sardine and mackerel from 2012 in this area (SC/F16/JR24).
the assessment of total consumption, energy requirements
Summer warming off Kushiro may have contributed to aand their use in ecosystem modelling; as well as in thedecrease in saury (cold-waterTablefish)
assessment of exchange rates and stock structure. They areP2 migration and an increasein Estimatessardineof andmackerel(warm-waterfish)
migration
intoalso important for the work of the IWC Scientific Committee
seasonal
prey consumption(thousand tons)
by common
minke whale,
Bryde’s whaleand sei whale– offshore.
this area during summer,
althoughthis
may also have beenon conservation and management via Implementation
relatedpotentially
Reviews and in-depth assessments.
Period to natural fluctuations
Prey of the stocks. This
Consumption
important change in oceanographic condition and pelagic
The proponents presented abundance estimates for
2000-07
Japanese anchovy
724
prey composition was Pacificnot discussedinthepaper.
baleenand sperm whales for the JARPN II research area. In
saury
56
Mackerels
43
addition to estimates presented at the 2009 review, data were
5.4.1.2 DISTRIBUTION
Total
1,117
collected and analysed for dedicated surveys conducted2008-13anchovy
674
The
Panel notes the Japaneseclose relationshipbetween distribution
from 2008-2014 using procedures in accordance with IWC
Pacific saury48
and abundance, in partMackerels
given that much of the information
70
Scientific Committee guidelines; g(0) was estimated fromcomes from the same surveys.
this report,
Total For the purposes of
1,226
independent observer (IO) data for minke whales (Okamura
especially with respect to progress with recommendations, it
et al., 2010), and was assumed to be 1.0 for larger baleen
has followed the same allocation of topics amongst the twoand sperm whales. The 2009 Panel noted that confidence
as provided in the 2009 report.
Table P3intervals of existing estimates were wide, which required
The Panel welcomes the papers presented on distribution
increasing
survey effort as a priority to improve the previousof daily prey consumption(kg) – coastal.
of large whales and the considerable effort Estimates
undertakentoestimates. The 2009 Panel also noted that extrapolations
Sanriku
produce new analyses of whale distribution
since the midwere not suitable to examine Kushiroeffects on stocks. Estimatestermreview. The Panelalso welcomesCVthe effort95%
to combine
Maturity
Consumption
CI LL
95% CIof
ULabundanceConsumption
CV minke, 95%
LL Bryde’s
95% CIwhales
ULof commonsei CIand
oceanographic, prey distribution and whale survey data in a
(SC/F16/JR12),
blue,
fin,
humpback
andNorth
Pacific
Immature male
98
0.25
56
149
82
0.19
53
115 right
spatialmodellingframeworkto achieve
Immature
female1060.26 a more integrated
60
162whales (SC/F16/JR13)81
0.19 sperm whales
53 (SC/F16/JR14)
112
and
in
Mature166
0.26habitat and 91
249
116
0.22
69
168
view
ofmalethe whale’s distribution,
theirfeeding
the
JARPN
II
offshoresurveyarea
bystratum
areavailable
Mature female
223
0.30
112
356
155
0.23
91
225
ecology.
for years 2008, 2009, 2011 and 2012.
Considerable progress has been made in the
The Panel welcomes the new estimates and especially
improvement of the use of spatial modelling to examine
the increased effort directed at improved precision whilst
whale distribution, and in a first attempt at combiningTable P4accommodating heterogeneity in detectability and othermultiple data sources to achievea more integrated viewof uncertainty.
New numbers of North PacificEstimates of seasonal prey consumption (tons) – sources
coastal. n=estimatednumber of whales.
of the whale ecology in the JARPN II survey areas. This
common minke whales in coastal
areas by stratum are now
Sanriku
Kushirohas increased the understanding of whale movement andavailable for early (May-June) and late (July-September)YearnConsumption
CV
CI LL
95% CI2012
UL
nConsumptionWhile
CVthe new
95% estimates
CI LL
95%
CI UL upsegregation
betweenspecies in the areas,
and 95%
contributed(SC/F16/JR11).
provideto a 2002
first attempt at mapping food consumption by species
to date 551
numbers,3,469
a number 0.17
of concerns
2,747remain. Abundance
5,436
2003
888 are provided for all areas and species within thegeographicallyand by season. The common use of satellite
estimates
2004
2,182
0.23
1,436
3,446
records
to input
oceanographic
variablesin spatialmodels 5,767research338
the0.23geographic
was not
2005
401
4,234
0.16
3,066
290area, although
2,601
1,569 coverage3,757
is also
improvement
since
review. 2,588consistent
2006 a welcome
216
1,822
0.18 the 2009
1,293
221 among
1,596
2,370
areas and0.19
between 1,128
years. This hasneither
2007
130
782
0.25
568
1,515
However,
there are a number of methodological issues
improved
thecomparability
betweensurveyyears,or the2008
whichneed to be addressed to improve the work presented
precision of estimates across years. This, combined with
2009
(see 2010
Item 5.4.3.2). This is particularly true with respect tothe fact that the surveys do not cover the full summer range
2011
combiningsurveydata
across
years
to
try
to
addresschangesof the putative
stocks,makes
in either
2012
124
850
0.20
599
1,325
433
3,264
0.22 estimation
2,408 of trends5,838
in Average
distribution247and abundance(and0.18
potentially 1,653
spatially- 3,227abundance
difficult.
Thisalso affects3,727
the work
2,302
407 or distribution
2,316
0.21
1,643
on the programmes’ objectives that use abundance and trend
1
Japanese Meteorological Agency http://www.jma.go.jp/jma/indexe.html. 3
information.

548

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

5.4.2 Evaluation of progress with the 2009 recommendations
The Panel welcomes the fact that the proponents had
addressed or partly addressed all but one of the 2009
recommendations as summarised in Table 5 (with the
exception of one recommendation which the Panel considers
no longer relevant).
5.4.2.1 OCEANOGRAPHYThe Panel agrees that use of the data from an ocean model andsatellites is more appropriate for monitoring oceanographic
condition of the survey areas than using sparse shipboard
observations (CTD data) made during the sightings survey.
This renders the 2009 recommendation regarding calibrationof CTDs no longer applicable.
5.4.2.2 DISTRIBUTION

The Panel agrees that the proponents have addressed or
partly addressed most of the 2009 recommendations withrespect to distribution. As noted earlier, the main weaknesses
relate to problems with survey coverage and areas by yearthat render examination of changes in distribution (andlinking that to other factors such as prey distribution andabundance) difficult. This problem is also relevant to theabundance estimation, and is important with respect tofeeding ecology and ecosystem modelling.
5.4.2.3 ABUNDANCEThe Panel agrees that the proponents have only partly
addressed two of the three recommendations from the 2009
review whilst the other, despite its importance (examining
trends) has not been addressed other than by providing an
explanation of the difficulties.
The 2009 Panel noted that abundance estimates are ofgreat importance for improving consumption estimation
and reducing uncertainty, and thus recommended theimprovement of the precision of the abundance estimatesthat are used to extrapolate to population-level rates, for boththe coastal and the offshore regions. In particular, to focus on
the sources or causes of variability in order to understand themechanistic linkages involved. Additionally, for the coastal
region, it considered the possibility of regular well-designed
aerial surveys, and a full synoptic survey for the offshore
region. The present Panel appreciates the effort to use
improved methods to estimate abundance, and in particular
the effort to identify sources and causes of variability
using multi-covariate analysis of the detection function.
In particular, the proponents incorporated consideration ofschool size, year and Beaufort as potential covariates of thedetection function. However, as the proponents acknowledge,
the estimates are not representative of the entire stocks
concerned because of limitations in area coverage; they
thus cannot be extrapolated to population level. Thus theincreased allocation of survey effort in the recent JARPN II
surveys, whilst improving the precision of area abundance
estimates, has not lead to improved precision in populationlevel abundance estimates.
The Panel notes that although some progress has been
made with respect to collating the photo-identification datacollected during the JARPN II programme, work to compare
the data with other catalogues is only slowly progressing.
5.4.3 Recommendations

5.4.3.1 OCEANOGRAPHY

Further efforts should be made by the proponents todetermine appropriate oceanographic parameters formonitoring in the context of prey and whale species. The
offshore survey area and that off Kushiro correspond to highprimary productive regions during summer, with significant
interannual variations (Shiozaki et al., 2004). In the short-term, the Panel recommends that at least chl-a concentration
should be examined as a potential proxy for the foodenvironment for whales; chl-a concentration is considered
to reflect zooplankton production as food for prey species.
In the medium-long term, the Panel recommends further
oceanographic monitoring to compare with prey species
distribution and abundance in the new regime.
5.4.3.2 DISTRIBUTION

In the medium-term, the Panel recommends that considerable
effort be put into the methodological improvement of thespatial modelling in the various analysis related with theobjectives on distribution of large whales and oceanography.
A particular focus must be on the combination of survey datafrom the different years to make them more comparable interms of distribution (and abundance) over time; use of datafrom other sources (e.g. the IWC POWER programme).
This work is not only valuable in itself but is essential for abetter parameterisation of ecosystem models.
The Panel also recommends that additional effort beplaced on fulfilling the 2009 recommendation with respect
to the photo-identification data. This will contribute to
the understanding of large scale movements and whale
distribution within and outside the JARPN II survey area forseveral species.
Before the 2016 Annual Meeting, the Panel recommendswith respect to the papers that used spatial modelling (SC/F16/JR07; Murase et al. (2014) [SC/F16/JR08]; SC/F16/
JR09; Sasaki et al. (2013) [SC/F16/JR10] and SC/F16/
JR16) that revised versions are developed that:(1) include statistical summaries on model fit (R2 and %deviance explained) and model comparison and spatial
covariate selection (e.g. AIC, BIC, GCV scores), which
are currently lacking;(2) avoid extrapolation of the regression models to datapoor areas or areas lacking coverage (especially whencombining food consumption with sightings data); and(3) include variance plots of the fitted prediction surfaces inorder to address precision and data sparseness.
5.4.3.3 ABUNDANCE

With respect to abundance, the Panel recommends:(1) exploration of methods to account for sampling
differences between areas and years to obtain measuresof short- and long-term variation and trends andestimates the extent of additional variance due tochanges over time in spatial distribution; inter alia
this will be essential for modelling efforts (e.g. foodconsumption models and ecosystem models); and(2) comparison of results from the design-based estimatesof abundance with those of model-based estimates,
which can be directly obtained from spatial models of
distribution (e.g. SC/F16/JR16) and using additional
data (e.g. from IWC POWER and other North Pacificsurveys) - model-based estimates could potentiallyaddress problems of unequal sampling coverage between
surveys and with further development could also accountfor additional sources or causes of variability.
6. FEEDING ECOLOGY AND ECOSYSTEMSTUDIES: PART 2, FIELD AND LABORATORYSTUDIES
6.1 Proponents’ summary of objectives including
modifications, if any, since the start of the programme

As explained under section 3.1, within the main Objective
1 there are three main objectives: (i) prey consumption by

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017549

Fig. P4. Schematic representation of research components under Objective 1 of JARPN II.

cetaceans; (ii) prey preference of cetaceans; and (iii) ecosystemmodelling, and within them there are several sub-objectives.
The following sub-objectives were identified:
Sub-objective 4: To estimate the prey consumption bybaleen whales using JARPN II data and samples, and taking
into account the uncertainties identified at the 2009 JARPN II
review.
Sub-objective 5: To evaluate the feeding impact by whales
on fisheries resources using JARPN II data and samples, andinformation from commercial fisheries and other researchsources in coastal areas.
Sub-objective 6: To estimate prey abundance using JARPN
II data, complemented with information available from othersources.
Sub-objective 7: To investigate the prey preference ofwhales in offshore areas, using JARPN II data and samples.
Sub-objective 8: To investigate feeding habits of baleen
and toothed whale species in the research area, and theenvironmental factors involved in determining such habits.
Sub-objective 9: To investigate the yearly trend in body
condition of baleen whales using JARPN II data and samples.

6.2 Overview of conclusions and recommendations fromthe 2009 Workshop
A short overview on the 2009 conclusions andrecommendations for the JARPN II components on prey
consumption, (including biomass estimation of preyspecies), prey preference (including whales’ feeding habits)
and body condition was provided by the Chair.
The 2009 Panel had highlighted a few critical aspects of
the work undertaken, including that insufficient work had
been undertaken to address uncertainty and that the rationale
for choice of sampling areas was unclear. It also concluded
that the presented consumption rates were not reliable
until further analyses have been undertaken. Therefore,
it recommended that the proponents should: (a) provide aclear rationale for sampling areas; (b) provide the scientificrationale used for the modelling formulations and proposed
ranges; (c) clearly explain methods used to extrapolate
from daily to annual rates and amounts; (d) incorporate
information from other studies (e.g. JARPN I, PICES,ESSAS); and (e) present the estimates of consumption bywhales in terms of fisheries and prey biomass.
In terms of treatment of uncertainty, the 2009 Panel
had indicated that the proponents should: (a) incorporate

several reasonable models for estimating daily consumption
as a function of body mass and include in their reports therange of possible results; (b) use that range in subsequent
analyses (including any ecosystem modelling) that employ
these daily/annual consumption estimates; and (c) undertake
sensitivity analyses for the range of parameter values
used in the consumption equations. The 2009 Panel had
also provided specific instructions on how to characterise
uncertainty and for which parameters. All details can befound in Annex F of the 2009 Panel report (IWC, 2010).
The 2009 Panel also recommended that in medium- tolong-term approaches for predictive models, the proponents
should: (a) combine oceanographic data, prey distributions
and sighting survey data statistically to investigate how prey
and whale distributions are associated with oceanographic
conditions, and how whale distributions are related to
distributions of prey; (b) combine data on prey distributions
as observed in the area where the whales were caught
with the diet of the whales (referred to as the micro scale)statistically to evaluate how well the whale’s diet reflects preyavailability in the area where it was caught; and (c) compare
results from (a) and (b) with the results on selectivity alreadyproduced and presented at the 2009 Workshop.
6.3 Proponents summary of the coastal and offshore
components (including response to 2009 Workshop)

Most of the recommendations from the 2009 Workshop related
to Item 6.3 were addressed in the analyses summarised below(see details in Annex D).

6.3.1 Prey consumption, including biomass estimation of
prey species

SUB-OBJECTIVE 4: TO ESTIMATE THE PREYCONSUMPTION BY BALEEN WHALES USING JARPN IIDATA AND SAMPLES, AND TAKING INTO ACCOUNT THE
UNCERTAINTIES IDENTIFIED AT THE 2009 JARPN IIREVIEW
SUB-OBJECTIVE 5: TO EVALUATE THE FEEDING IMPACT
BY WHALES ON FISHERIES RESOURCES USING JARPNII DATA AND SAMPLES, AND INFORMATION FROMCOMMERCIAL FISHERIES AND OTHER RESEARCH
SOURCES IN COASTAL AREAS

SC/F16/JR15 estimated the prey consumption by common
minke, Bryde’s and sei whales in the western North Pacific.
Prey species of whales were identified by examining theirstomach contents, and the amount of prey consumed in the

550

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II

research area was estimated by using information on prey
In Sanriku the estimated sandlance consumption
corresponded to 30-40% of the fisheries catch in two years
consumption per capita and the numbers of whales distributed.
before the tsunami. In Kushiro the estimated seasonal
There were seasonal and geographical changes in the preys
consumption of Pacific saury and walleye Pollock from 2002
consumed in each whale species. The extent of differences of
to 2012 corresponded to approximately 2-3% of the fisheries
estimates of consumptions among several models was 2.4-3.6
catch on these resources.
times. Based on the results obtained by three equations combined
In conclusion the estimates of prey consumption of whales
and Monte Carlo simulations, the daily prey consumptions per
in coastal and offshore waters were made with an improved
capita (kg) were estimated as shown in Table P1.
level of precision. These estimates on prey consumption
The CVs of the seasonal prey consumption by whales wereare useful as input data in ecosystem models. The spatialin the range 0.3-0.4. These values were equivalent to 22-48%,
distribution of sei whales at mesoscale was largely determined
2-7% and 5-66% of the biomass of these fish resources in thewestern North Pacific, respectively. The total line includes allby oceanographic conditions such as SST.
prey species, not just these three.
SUB-OBJECTIVE 6: TO ESTIMATE PREY ABUNDANCE
SC/F16/JR16 examined the preliminary attempt of spatial
USING JARPN II DATA, COMPLEMENTED WITH
estimation of prey consumption by sei whales in the JARPNINFORMATION AVAILABLE FROM OTHER SOURCES
II survey area using data obtained from 2002 to 2013. Two
Murase et al. (2012) [SC/F16/JR18] examined basin-scalelevels of models are constructed to achieve the goal. First,
distribution pattern and biomass estimation of Japaneserelative abundance of sei whales in relation with oceanographic
anchovy in the western North Pacific using a quantitative
conditions is estimated by using a generalised additive model
echosounder. This was the first attempt at such a study in this(GAM). Secondary, amount of prey consumed by a sei whale
region. Data were collected in summer from 2004 to 2007. The
in relation with oceanographic conditions is also estimated bybiomass was estimated using data collected at 38kHz. Species
using GAM. Finally, prey consumption of sei whales in thecompositions in the backscatterings from pelagic fish were
JARPN II survey area is calculated as the product of theseassigned based on the results of trawl hauls taking accounttwo models. Data obtained from 2002 to 2013 are used inof sea surface temperature (SST). Japanese anchovy tended
the analysis. Spatial distribution of prey consumption shiftedto be high density to the west of 153°E and were distributed
toward north as the season progress. Estimated amount ofin an SST range of 9-24°C. Although the temporal and spatial
prey consumption by sei whales using the spatial model wascoverage of the survey differed each year, at least 1.5-3.4
comparable to estimates based on traditional methods (SC/F16/
million tons of Japanese anchovy were present in the surveyJR15). SST was selected as environmental covariates in thearea between 2004 and 2007. To take account of the spatial
first and second models. However, the shape of functional form
coverage of the survey each year, the most reliable biomass
for the first level model (prey consumption) was relatively flat
estimate for this region in the time period was 3.4 million tons
in comparison with the second level model (abundance). The(coefficient of variation 0.22).
Table 5
results indicated that spatial distribution of sei whales at mesoSC/F16/JR19
examined estimation of prey species biomass
Tableto5 the 2009 recommendations with respect to oceanography, distrib
SummaryofPanel’s
evaluationoftheproponents’
response
scale were largely determined by oceanographic conditions
based on 2008 and 2009 JARPN II acoustic surveys around the(and see text).respect to oceanography, distribution and abund
Summary
Panel’s
evaluationof theproponents’
response
to the 2009
recommendations
such as SST. Sei whales
couldofthen
searchfor their
preywithSanrikuregion.
The survey withwas conducted concurrently with(andseetext).
the optimal oceanographic conditions
as indicatedbyrecommendations
feeding
sampling survey of common minke whales. Five stratified
Brief summaryof 2009
andasuggestions
behaviour study (SC/F16/JR25).
Futurestudyresultson feedingwere
surveyed.
lines were
set coveringin the Panel
Brief summary Poolof 2009
and othersuggestions
or recommendations
comparewithdatasets blocks
to increasesamplesize andZigzag
increasetrack
the possibilityof dataperiodsevaluatofecology of baleen Pool
whalesshouldpursuitsuchanintegrated
blocks.
trawlertype
RV,
Takuyo-maru,
the survey.
changing(e.g. previous
regimechanges),thus
allowing
patterns
to be detected.
Addressedor compare
results relationships
with other datasets
to increasesamplesize Aandincreasethepossibilityof data conducted
covering periodsofapproach further. changing relationships
Acoustic
data
wererecorded
continuously along track lines by
Considerconducting
over
an
area
than at present.
(e.g.
previous future
regimeoceanographic
changes), thussurveys
allowing
patterns
to larger
be detected.
SC/F16/JR17 estimatedthe preyby common
In theconsumption
long term,
integrate
into analyses
andatechootherdata collectedon the cruises
depth,
water Addressedcolumna quantitative
sounder.
Samplings
using a(bottom
midwatertrawl
Consider conducting
futureoceanographic
surveysover anoceanographic
arealarger thanpresent.
temperature,salinity,
anddensity)derivedsuch as
SST,
chlorophyll,
andandsea sizesurfaceheight.
minke whales off Sanriku
and term,
Kushiroregions.
Prey
speciesof with satellite
wereconducted
identify
speciescompositions
In the longintegrate
into
analyses
oceanographic
and netother
datadata,
collected
onto
thecruises(bottomdepth,
water
columnPartly addres
Correct/calibrate
using
simultaneously
collected
watersamples.
whales were identified
by examining
theircontents,
andderived
temperature,salinity,
andstomach
density)CTD
withdata
satellite
data, ofsuch
as SST, backscatterings.
chlorophyll,and seasurfaceoceanographic
height.
acoustic
Verticalconditions
Conduct
additional
analyses(including
using
techniquessuch as GAMs
or logistic
regressions).
Correct/calibrate
CTD
data
using
simultaneously
collected
watersamples.
the amount of prey consumed
in theresearcharea wasestimated
wererecordedby usinga CTD.
Surface oceanographicNo longer appli
Incorporate
sightability
detected groups
the index
of recorded
density
andcontinuously
test whether the
chosenmodel
is an
improvement
Conduct
additional
analysesthe(including
techniques
suchconditions
asintoGAMs
orwerelogistic
regressions).
Addressedby using information
on prey
consumption
percapitausing
andofthe
along
track
lines.
Theovera null, uninformativemodelinto
andthetryindex
to validate
the model
results
usingthecross-validation
techniques
and use of outside data.
theBasedsightabilityof detectedgroupsof density
and testwhether
an improvement
numbers of whalesIncorporate
distributed.
on theresults obtained
totalbiomass
estimationof chosensand model
lance isadult
were 8,076t, Partly addres
Investigate model
whetherdata from1994-2007
canusing
be usedto provide informationontrends.
over a null, anduninformative
andsightings
try to validate
modelsandresults
cross-validation
techniques
anduse of outside
data.wereby three equations combinedMonteCarlo simulations,the thecomparelancejuvenile
1,237t, Japanese
anchovy
Worksightings
up photo-identification
data andother North
Pacificwere
Investigate whether
data from 1994-2007
can be used withto provide
information
oncatalogues.
trends.
Not addresse
daily prey consumptions (in kg) Increased
per capita
ofcommon
minke
0.18t
in
block
B
and
C
in
2008.
The
total
biomass
estimation
effortto obtain
better estimatesa highcatalogues.
priority.
Work up photo-identificationdata
and comparewith othershould
North bePacific
Partly addres
whales in Sanriku Increased
and Kushiro
are
given
in
Table
P3.
The
of sand lance adult were 2,512t, sand lance juvenile were Partly addres
effort to obtain better estimates should be a high priority.
estimated seasonal consumption (in tons) in the two areas is
315t, Japanese anchovy were 0.64t in block B and C in 2009,
given in Table P4 and variation by prey species in Table P5.
respectively. It was not possible to evaluate any trend in sandTable P1
Table P1
Estimatesof daily prey consumption (kg) - offshore.
Species

Estimates of daily prey consumption (kg) - offshore.
Immature male (kg)Mature male (kg)Species

Immature male (kg)Common minke whale
Bryde’s whale 86-94
Common minke whale
Bryde’s whale Sei whale419-434
Sei whale

397-421

Mature male (kg)
86-94
419-434
129-141
397-421577-637524-539

Immature female (kg)

Immature female (kg)129-141577-63783-94
524-539417-428436-468

610-647

Table P2
Table
Estimates of seasonal
preyP2consumption (thousand tons) by common minke whale,
whale
andbysei
whale –minke
offshore.
Estimates of seasonal prey consumptionBryde’s(thousandtons)
commonwhale,
Bryde’s whale and sei whale – offshore.
PeriodPrey

Period2000-072008-13

Consumption

PreyConsumption
2000-07
Japanese anchovy
Pacific
sauryJapanese anchovy
724
Mackerels
Pacific saury
56
Total
Mackerels
43
2008-13
Japaneseanchovy
Total
1,117
Pacific
sauryJapanese anchovy
674
Mackerels
Pacific saury
48
Total
Mackerels
70
Total

Mature female (kg)Mature female (kg)
83-94
417-428158-166
436-468642-707

724
56
43
1,117
674
48
70
1,226

1,226

Table P3

158-166
642-707610-647

TotalJapanese anchovy
Pacific saury
Mackerels
Total

2008-13

1,117
674
48
70
1,226

551

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017
Table P3
Estimates of daily prey consumption (kg) – coastal.
Sanriku
Maturity
Immature male
Immature female
Mature male
Mature female

KushiroConsumption

CV

95% CI LL

95% CI ULConsumption

CV

95% CI LL

95% CI UL

98
106
166
223

0.25
0.26
0.26
0.30

56
60
91
112

149
162
249
356

82
81
116
155

0.19
0.19
0.22
0.23

53
53
69
91

115
112
168
225

Table P4
Estimates of seasonal prey consumption (tons) – coastal. n=estimated number of whales.
Sanriku
Year
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
AveragenConsumption

KushiroCV

95% CI LL

95% CI UL

401
216

4,234
1,822

0.16
0.18

3,066
1,293

5,767
2,588

124
247

850
2,302

0.20
0.18

599
1,653

1,325
3,227

n

Consumption

CV

95% CI LL

95% CI UL

551
888
338
290
221
130

3,469

0.17

2,747

5,436

2,182
2,601
1,596
782

0.23
0.23
0.19
0.25

1,436
1,569
1,128
568

3,446
3,757
2,370
1,515

433
407

3,264
2,316

0.22
0.21

2,408
1,643

5,838
3,727

3
Table P5
Estimates of seasonal prey consumption (tons) by species – coastal.
Sanriku
Year

Krill

Sandlance

KushiroAnchovy

Krill

Anchovy

SardineSaury

Mackerels

Pollock

Squid

76

488
49
627
11
41
409

665
1,204
220
971
338
2

0
0
0
0
0
724

460
843
18
198
170
0

0
0
0
0
0
154

791
85
1,546
264
233
1,421

1,066
0
190
153
0
554

199

271

567

121

282

26

723

327

2002
2003
2004
2005
2006
2007
2012

302
-

3,709
1,522

223
300

118

656

Average

210

1,962

whales were found in several regions of the transition zone
lance and Japanese anchovy abundance with just two surveys
between the subarctic boundary and the Kuroshio front. This
conducted in small areas. However sand lance and Japanesewhale species preferred Japanese anchovy as prey, for whichanchovy estimates were used to compare sand lance andthe distribution density was significantly higher in the main
Japanese anchovy consumption by large whales and as input
distribution area of the whale than in the adjacent areas. These
data for the development of ecosystem models for this area.
results indicated that the summer distributions of Pacific saury
In conclusion, Japanese anchovy was the main prey item Table 6
and Japanese
anchovygreatly
influence
distributions
for thebaleenin offshore
andproponents’occasionallyin coastal
Summaryof whales
Panel’s evaluationof theresponse
to the 2009 recommendations
with respect
to feeding
ecologystudiesthe(see text).
of these whale species, suggesting that the whales’ habitat
areas, while sandlance is important for common minke whales
Briefof 2009
recommendations
Panel evaluation
selection is closely related to their prey 2016selection.
in summary
the Sanriku
area.
Prey biomass estimates obtained were
SC/F16/JR21(Murase
et
al.,
2007)
conducted
a studyused fuller
as input
dataforin sampling
the ecosystem
Providerationaleareas. model developed in both
Not addressed.
ofcommonminke
andBryde’swhalesinthewestern
offshoreandSanrikuareas.
Characterise uncertainty following advice provided for key parameters.
Partly addressed.North
Pacific in the
andof2001
summerData analyses should: (a) incorporate several reasonable models for estimating daily consumption
as a 2000
functionbody massand seasons to estimate preyPartly addressed.
6.3.2
Prey
preference,
including
habitscetaceansandas(c)thisis an importantparameter inincludethe rangeof results;(b) use thatwhales’
range in feeding
subsequent
analyses (including anyselection
ecosystemof
modelling);
undertake
SUB-OBJECTIVE
7: TO
INVESTIGATE
ecosystem models. Whale sighting and sampling surveys andsensitivity analyses forthe rangeof parameter THE
valuesPREY
used in the consumption equations.
PREFERENCE
OF WHALESOFFSHORE
AREAS,
Undertake additional
analyses to IN
identify
the greatest
sources USING
of uncertainty and determine
appropriate
sampling
and analyticalprey surveys
using
quantitativeechosounderand mid-water
Not addressed.
JARPN
II DATA AND SAMPLES.
strategies.
trawl were carried out concurrently in the study. Biomasses
SC/F16/JR20(Watanabe
et modellingal., 2012)
presented
first ranges.
Providescientific rationale
for used
formulations
andtheproposed
Partlyaddressed.
of Japanese anchovy, walleye pollock andkrill,
which were
Explainmethods analysis
used to extrapolate
from daily to annual
rates
and amounts.
quantitativeof the characteristicsof thedistribution
major prey species of common minke andAddressed.
Bryde’s whales,
Incorporate
from otherofstudies.
areas andinformation
stomach contentscommon minke, sei and Bryde’s
were estimated using an echosounder. The Addressed.
results suggested
Present
estimatesof consumption
by whales inandtermspreyof fisheries
and prey biomass.
Addressed.
whalesin relation
to oceanographic
environmentsthat common minke whales showed prey selection
for JapaneseCombinethe oceanographic
data, prey
distributions
sightingminke
survey data statistically to investigate how prey and whalein mid-summer
in the western
North
Pacific.andCommonAddressed.
anchovywhile
they
seemed
to
avoid
krill
in
both
the offshore
distributions are associated with oceanographic conditions, and how whale distributions are related to distributions of prey.
whales were distributed within subarctic regions and theandwithcoastalregions
walleyepollock
continental shelf
Combine data on prey distributions as observed in the area where the whales were caught
the dietof theandwhales(referred
to in theAddressed.
northernmostof thecoinciding
region.inSelection
for shoaling
pelagic fish was similar to thatas themicroscale) region
statisticallyto transitional
evaluate howdomain,
well the whale’s
diet withreflects prey availability
the area where
it was caught.
the mainhabitatpreferred
prey,
Pacific
Sei onwhalesthe eastern
North Atlantic.
Bryde’s whale
selection
Compareresults
fromofthetheirapproaches
listedabove
with saury.
the results
selectivity alreadyinproduced
and presented
at the Workshop.
Notshowed
addressed.
were mainly found in the northernmost part of the transition
for Japanese anchovy in August 2000 and July 2001, while itzone and showed prey preference for Japanese anchovy, whichshowed prey selection for krill in May and June in 2001.
was significantly more abundant in the main distribution area
SC/F16/JR22 examined prey preferences of commonof the whale than in its adjacent areas. ‘Hot spots’ of Bryde’s
minke, Bryde’s and sei whales in the offshore component ofTable 7
Summary of sources of uncertainty and the treatment of these in the analyses presented to the Workshop.

552

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN IIJARPN II from 2002 to 2007 using data from the concurrent
surveys of cetacean sampling and prey of cetaceans. The
surveys were conducted as a part of the offshore componentof JARPN II from 2002 to 2007. A prey preference index,
Manly’s α, was used in the analysis. The sum of Manly’s α
for all prey species is 1 and a large value of Manly’s α for aprey species indicates a preference for it. Common minke
whales showed preference toward pelagic fishes as previously
reported. Bryde’s whales showed preference for anchovy. Sei
whales showed preference for copepods. Although the prey ofthree baleen whale species overlapped, Manly’s α suggested
their trophic niches were different from each other. Common
minke and sei whales coexisted in the same survey blocks, but
their prey utilisation patterns were different.
In conclusion, differences in prey preferences among
the three baleen whale species in the offshore area could be
explained by different feeding strategies of sei and common
minke whales. Results for the coastal waters off Kushiro
suggested that prey preference differed according to maturity
stages of the common minke whales. The feeding strategy ofthis species in this coastal locality might change in response tochanges in the environment.
SUB-OBJECTIVE 8: TO INVESTIGATE FEEDING HABITS
OF BALEEN AND TOOTHED WHALE SPECIES IN THE
RESEARCH AREA, AND THE ENVIRONMENTAL FACTORSINVOLVED IN DETERMINING SUCH HABITS

SC/F16/JR23 examined decadal change in food compositionof common minke, sei and Bryde’s whales in the western
North Pacific. Stomach content data obtained in May-October
during 2000-13 off the Pacific coast of Japan, were examined.
The three species were highly dependent on small pelagic
fishes, i.e. Japanese anchovy, Pacific saury and mackerels in
addition to copepods and euphausiids. The yearly trend offood compositions differed among the three whale species.
Sei whales showed a drastic change from Japanese anchovyin early 2000s to mackerels and Japanese sardine after 2010.
This synchronised with the catch record of Japanese fishery in
the western North Pacific. Copepods and euphausiids are also
important prey species for sei whales and were steadily available
in the blooming period. Bryde’s whale had a more simple preycomposition with Japanese anchovy and euphausiids as main
prey species, and the composition of the two main prey species
were highly variable among years and no remarkable change
since 2000-13. Food composition in common minke whale inoffshore waters (east of 150°E) showed the Japanese anchovyand saury as the major species, but the composition differedamong years. Among the three whale species, sei whale
distributed most widely in latitude through Kuroshio extension
to north of the subarctic front feeding at a variety of preyspecies in the JARPN II study area where abundant pelagic fish
carried by Kuroshio-current and Neocalanus copepods in the
blooming season, were available.
SC/F16/JR24 examined the relationship between maturity
and feeding habit of common minke whales in the coastal region
off Kushiro. A total of seven dominant preys, including one
species of krill (Pacific krill), one of squids (Japanese common
squid) and five of fishes (Japanese anchovy, Japanese sardine,chub mackerel, Pacific saury and walleye Pollock) wereidentified in 589 stomachs of common minke whales. Feeding
habits of common minke whales off Kushiro in autumn differed
between immature and mature whales. These results suggested
that prey preference of common minke whales in the coastal
waters off Kushiro in autumn differed with their maturity stage.
Feeding strategy of common minke whales might change toadapt to local environments. Differences can be explained by
the trade-offs of cost of foraging activity for prey and/or energy
demands between immature and mature whales.
SC/F16/JR25 examined the feeding behaviour of sei
whale. Diving behaviour of sei whales and vertical distributionof their prey were recorded simultaneously in 2013 JARPN II
survey to study their feeding behaviour at micro scale. Small
acoustic time depth transmitters (pingers) were attached
to two sei whales and their behaviours were recorded for10.2 and 32.0 hours, respectively. Vertical distributions anddensities (volume backscattering strength, SV) of their prey
were recorded by an echosounder following swimming pathof the individuals. The diving behaviour deeper than 10m wasclassified into two shapes (U-shape, V-shape). It was assumed
that U-shape was related to feeding behaviour, especially lunge
feeding, while V-shape was related to other behaviour. It wassuggested that sei whales actively fed on prey around dusk.
Swimming depth of the whales was shallower than 10m aftersunset while deep scattering layers (presumably myctophids)
migrated from below 60m to around 30m. The results might
indicate that they did not feed on prey in deep scattering layers
at night. However, the possibility that sei whales feed on preynear surface at night cannot be discarded. The results of thisstudy revealed that sei whales changed their diving behaviour
in response to availability of their prey in daytime.
SC/F16/JR26 examined the feeding habits of sperm whalesin the western North Pacific in spring and summer based on
analysis of stomach contents of 56 animals examined from
May to September in the years 2000-2013. A total of 49
undigested and half-digested prey items were found, including
28 species of cephalopods and six species of fish. The Index
of Relative Importance (IRI) showed that Belonella borealisand Histioteuthis spp. were the dominant prey in the Subarctic
Region, while B. borealis and Galiteuthis phyllura were thedominant prey in the Transitional Domain. B. borealis andTaningia danae were the dominant prey in the Northern partof the Transition Zone, while T. danae and Histioteuthis spp.
were the dominant prey in the Southern part of the Transition
Zone. In the Kuroshio Zone, T. danae and Octopoteuthis spp.
were the dominant preys. The composition of prey items
changed in relation to transitional change between north andsouth. Canonical Correspondence Analysis (CCA) indicatedthat environmental and biological factors significantly
contributed to the prey composition of sperm whales. This
study demonstrated that sperm whales moved along waters
with different oceanographic conditions, feeding on a variety
of prey species. Larger whales tend to feed at offshore waters.
This flexibility and different size distribution in sperm whale
seems to be important to maintain large body size and large
abundance in the western North Pacific. The commercially
important Neon flying squid O. bartrami was not an important
prey of the sperm whales sampled in spring and summer, only
one of which was a mature male.
In conclusion, the yearly trend of prey compositions in
Bryde’s and sei whales was different in the offshore waters.
Bryde’s whales showed no trend, feeding every year on krill
and Japanese anchovy. On the other hand drastic yearly changes
were observed in the prey species of sei whales, shifting fromJapanese anchovy to mackerels and Japanese sardine after
2010. Prey availability (biomass) is likely to determine preycomposition for large baleen whales. In coastal waters off
Sanriku yearly changes in the prey species of common minke
whale, were not observed. On the other hand, in coastal waters
off Kushiro, the dominant prey species shifted from Japanese
anchovy and Pacific saury to Japanese sardine and mackerels
after 2011. It is suggested that the feeding strategy of common
minke whales might change to adapt the local environments.
The results of diving behaviour of sei whales by acoustic
devices indicated that sei whales did not feed on prey in deep
scattering layers at night. Such information could validate
some of the assumptions made for the estimation of prey
consumption in future. Sperm whale moved along different
oceanographic waters in the research area, feeding on several
mesopelagic squids.

6.3.3 Body condition of whales

SUB-OBJECTIVE 9: TO INVESTIGATE THE YEARLY TRENDIN BODY CONDITION OF BALEEN WHALES USING JARPNII DATA AND SAMPLES

SC/F16/JR27 examined the annual trend in energy storage insei, Bryde’s and common minke whales during the JARPN II
period. Regression analyses showed that blubber thickness in

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017sei whales has been increasing during the JARPN II period.
The increase per year was estimated at approximately 0.1cm
for mid-lateral blubber thickness. ‘Body length’ and ‘Date’
were included in the best model at the 5% level, while noyear effects were included in the best model for other blubber
thickness and girth measurements. The blubber thickness for
mid-lateral probably is the most sensitive for detecting energy
storage. In Bryde’s and common minke whales, no significant
trends were observed in the regression analyses. The increasein the trend of body condition indicators and the recent changeof prey composition suggest that food availability has changed
favourably for sei whale in the study area.
In the review meeting, the author of SC/F16/JR27 conducted
additional analyses based on the reviewer’s comment that atechnical error had been made. Following this correction, linear
models with minimal covariates, i.e. body length, date, latitude,
longitude and sex were first tested for four response variables,
and then an interaction of the maturity stage with year wasadded separately. The year effects of the models differed among
species and indices of nutritional condition, showing either noor negative effects for sei, a negative effect for Bryde’s andboth negative and positive effects for common minke whales.

6.4 Panel review, conclusions and recommendations
6.4.1 Overview
The data and information from JARPN II studies on whale
food habits and prey preferences is substantial. The sampling
programme was generally well-coordinated across a range
of vessels and platforms, and there is a substantial amount ofconcurrently collected data. However, as noted under Item
3.4.2, there are concerns about how representative the whales
sampled are with respect to: (a) the study areas; and (b) the
populations. The Panel reiterates the importance of the
proponents providing papers to the 2016 Annual ScientificCommittee Meeting evaluating the potential impact of therealised sampling and sample sizes on the feeding ecology
studies, as recommended under Item 3.4.2.
The Panel notes that the available datasets are pertinent
to a broad range of topics, with wider relevance than within
the JARPN II programme objectives. The Panel agrees
that diet data can be used to: (i) determine whale prey
preferences; (ii) develop functional response curves whensampling is accompanied by simultaneous assessments ofprey abundance; and (iii) estimate the impacts of whaleson their prey (when used in conjunction with estimates of
total consumption and whale abundance). The diet data fromJARPN II are compatible with these goals and, in addition,
may serve as indicators of the potential prey selectionof these whale species in other regions of the sub-Arctic
oceans. These studies form a critical input into ecosystemmodelling and can be used directly to understand species
interactions (Bogstad et al., 2015).
6.4.2 Evaluation of progress made on recommendations in2009
As summarised in Table 6, the Panel notes that the proponents
had addressed or partly addressed all but three of the 2009
recommendations (related to sampling areas, sampling andanalytical strategies to reduce uncertainty and a comparisonof the 2009 results with those from different approaches).
More detailed consideration of these topics is given below.
The estimation of food consumption is based on models
that account for per capita daily consumption, the weight ofanimals of different sexes and life stages, the caloric value of
prey species, the assimilation efficiency, and the number ofanimals by sex and stage. The uncertainty of the estimates ofconsumption was quantified using a Monte Carlo approachin which the model parameters were assigned triangular553

distributions. Table 7 summarises the sources of uncertainty
identified by the 2009 Panel and assesses which sourceswere considered in the Monte Carlo procedure. Most, but
not all, of the sources of uncertainty were accounted for.
The Panel recommends that all sources of uncertainty bequantified and an evaluation of which parameters contributethe most to uncertainty be conducted.
Triangular distributions were used to express uncertainty.
However, the reason for selecting this distribution is unclear.
The method selected to account for uncertainty should be
improved to better reflect the statistical distributions ofparameter uncertainty (e.g. log-normal for abundance).
Mass was introduced into the equations as a triangular
distribution with the mean, maximum and minimum mass
samples for each maturity and sex class. The Panel agrees
that the triangle distribution placed too much weight in thetails of the distribution, and that an approach such as a simple
bootstrap of the data would have been more appropriate.
Captive studies are impractical for whales and thus
alternative approaches are needed to estimate consumption.
The proponents presented several allometric-consumption
equations which provided a range of energy/consumption
estimates, meant to bracket upper and lower limits ofpossible levels of consumption. The work that examinedmeal size using fresh stomach content material and foraging
behaviour suggested that some allometric relationships ofconsumption may be too low (equation 2: SC/F16/JR15).
The Panel recommends that this aspect of the analyses bedeveloped further to refine the range of suitable allometricenergy intake/consumption relationships that would haveprovided a more useful range of consumption estimates.
The proponents also statistically examined relationships
between oceanographic conditions, oceanographic data,
prey distributions and sighting survey data to investigate
how prey and whale distributions are associated withoceanographic conditions, and how whale distributions are
related to distributions of prey. They statistically combined
data on prey distributions as observed in the area where thewhales were caught with the diet of the whales (referred toas the micro scale) to evaluate how well the whale’s dietreflects prey availability in the area where it was caught.
Diet composition information is a key component of the
JARPN II programme. In JARPN II, diet was reconstructedby focusing on the relative contribution of the fresh material,
and weighting this by the number of digested prey identified
to species. The fresh samples represent the most recent
ingestion of prey. The impact of using the most recently
ingested prey to weight the reconstruction is not clear,but may be important if the species composition changes
over the period that prey are retained within the first andsecond stomach. In many diet composition studies, thediet is reconstructed using all material recovered from thestomach, not just by using recovery of fresh samples. This
reduces the need to understand or place as much importanceon understanding the mean residence time of each main preygroup in the forestomach. For example, if fish or zooplankton
differ in their rates of digestion, using only fresh material may
overestimate the contribution of the group that is digested
more slowly. Normally, the total number and mass of fishconsumed are estimated through the number of otoliths or
beaks, each scaled up to estimated original prey mass andsummed to obtain a total mass for the meal. The relative
contribution of each prey species to the stomach contents
is determined based upon its estimated mass contribution to
the total meal. The proponents collected information on preysize using the fresh stomach content material, but this couldAverage

554

210

1,962

199

271

567

121

282

26

723

327

Table 6REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II
Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to feeding ecology studies (see text).

Brief summary of 2009 recommendations

2016 Panel evaluation

Table 6of Panel’s
evaluationof the proponents’ response to the 2009 recommendations with respect to feeding ecology studies
text).
ProvideSummary
fuller rationale
for samplingareas.
Not(seeaddressed.
Characterise uncertainty following advice provided for key parameters.
Partly addressed.
Briefsummaryshould:of 2009(a)recommendations
Data analyses
incorporate several reasonable models for estimating daily consumption as a function of body mass and 2016 Panel evaluation
Partly addressed.
include
the
range
of
results;(b) use that
range in subsequent analyses (including any ecosystem modelling); and (c) undertake
Provide fuller rationale for samplingareas.
Not addressed.
sensitivity
analyses
for
the
range
of
parameter
valuesusedintheconsumption
equations.
Characterise uncertainty following advice provided for key parameters.
Partly addressed.
Undertakeadditional
to identify
thereasonable
greatest sources
uncertainty
determine
appropriate
sampling
and analyticalData
analysesshould: analyses(a) incorporate
severalmodelsoffor
estimatingand
daily
consumption
as a functionof body
mass andNot addressed.
Partlyaddressed.
strategies.
includethe range of results; (b) use that range in subsequent analyses (including any ecosystem modelling); and (c) undertake
Providescientificrationaleforusedmodelling
formulations
andproposedranges.
Partlyaddressed.
sensitivity analyses for the range of parameter values used in the consumption equations.
Explain methods
usedanalyses
to extrapolate
from thedaily
to annualratesofandamounts. and determine appropriate sampling and analytical
Addressed.
Undertakeadditional
to identify
greatest
sources
uncertainty
Notaddressed.
Incorporate information from other studies.
Addressed.
strategies.
Present
estimatesofconsumptionbywhalesinterms
offisheries
andpreybiomass.
Addressed.
Provide scientific rationale for used modelling formulations and proposed ranges.
Partly addressed.
Combinemethods
the oceanographic
data, prey
distributions
and ratessighting
data statistically to investigate how prey and whale
Explainused to extrapolate
fromdaily to annual
and survey
amounts.
Addressed.
Addressed.
distributionsinformation
are associatedwithoceanographic
conditions, and how whale distributions are related to distributions of prey.
Incorporate
from
other
studies.
Addressed.
Combine
data
on
prey
distributions
asobserved
in
the
area
where
the
whaleswerecaught
with
the
diet
of
the
whales(referredtoAddressed.
Present estimates of consumption by whales in terms of fisheries and prey biomass.
Addressed.
as the microscale)statisticallydata,
to evaluate
how well theand
whale’s
dietsurvey
reflectsdata
preystatisticallyavailabilitytoininvestigate
the area where
was andcaught.
Combinethe oceanographic
prey distributions
sighting
howitprey
whale
Addressed.
Compare results
the approaches
listed above
with the results
on selectivity
already produced
andtopresented
at theNot
addressed.
distributions
are from
associatedwith oceanographic
conditions,and howwhale distributions
are related
distributionsofWorkshop.
prey.
Combine data on prey distributions as observed in the area where the whales were caught with the diet of the whales (referred toas the microscale) statistically to evaluate how well the whale’s diet reflects prey availability in the area where it was caught.
Compare results from the approaches listed above with the results on selectivity already produced and presented at the Workshop.

Addressed.

Not addressed.

Table 7
Summary of sources of uncertainty and the treatment of these in the analyses presented to the Workshop.
Issue

StatusTable 7sources of uncertainty and the treatment of these in the analyses presented to the Workshop.
Per capita consumptionSummary
in area ofofinterest
Parameter uncertainty in the relationship between energy consumption and body mass (multiple and exponent). Results shown for three relationships.
Issue
StatusResidual variance of species values around the mean curve.
Not considered.
Proportionofannualenergyrequirement
obtainedduringsummerfeedingseasonandlengthofthefeeding
Assumedtobetriangular
distributed.
Per capita consumption in area of interest
season. uncertainty in the relationship between energy consumption and body mass (multiple and exponent). Results shown for three relationships.
Parameter
Variance variancein mean bodymass values(stratified
by sexlife
stage, e.g. mature/immature).
Assumed tobeconsidered.
triangular distributed
Residualof species
aroundthe andmean
curve.
Not(separately
bytriangular
sex and life
stage).
Proportion of annual energy requirement obtained during summer feeding season and length of the feeding Assumed
to be
distributed.
season.
Diet composition
Variance inbodymass (stratified
andlife preystage,groupe.g. mature/immature).
of meanaverageundigested
biomassby
ofsex
each
main
in the forestomach.
Variance of the mean residence time of each main prey group in the forestomach.
Varianceof the average energy content per unit biomass of prey by prey type.
Diet
composition

Assumed to
to be
be triangulartriangular distributed.
distributed
Assumed(separately
sex and life stage).
Notbyconsidered.
Included in the triangular distribution (butsamplesizetois below
for somedistributed.
prey items).
Assumedtriangular
Included inNot
the considered.
triangular distribution.
Not
considered.
Included in thetriangular
distribution (but

Variance of average undigested biomass of each main prey group in the forestomach.
Variance ofof thethe mean
averagebody weightof eachundigested
preygroup
items inbythe
species.
Variance
residencetime ofmain prey
forestomach.
Variance ofof thethe averagerelative frequencies
of eachspeciesby counts
of by
individuals
Variance
energy content
per unit
biomassof prey
prey type.and/or hard parts.
sample size is low for some prey items).
Abundance
Variance of
thepossibly
average bodyweight inof undigested
items by(meanspecies.
(andcovariance)
estimates ofprey
abundance
number of whales present) in survey Included
Assumedintothe
be triangulartriangular distribution.
distributed.
Variance
the relative
frequencies
each species
by countsandof individuals
and/or hard parts.
Not considered.
season byof
sub-area
and time
period, of
includingg(0) variance,
process error.

Abundance
Variance (and possibly covariance) in estimates of abundance (mean number of whales present) in survey Assumed to be triangular distributed.
also
estimatedfromsize allometricseasonbeby sub-area
and time
period,otolith-preyincluding g(0) variance,and process error. individual. The equations should be compared to understandrelationships, which would have increased sample size. Preysize information is also needed for analysing the contributionof whale predation to natural mortality of commercial fishstocks, particularly where different size classes may be 4
targeted by whales or fisheries (e.g. sand lance, walleye
pollock). The size composition of prey was presented insome (e.g. SC/F16/JR17), but not all papers, and considered 4
suggestive of differential selection of particular prey sizes
by each species, but this was not quantified. The energy
content of prey is expected to change seasonally. However,energy density of prey appears to have been examined for
only a limited number of samples for each prey species andit is not clear if seasonal changes in energy density wereconsidered. The Panel agrees that further work is needed(see Item 6.4.3).
When estimating energy requirements, it is importantto account for the costs of energy storage, growth andreproduction. The proponents recognise this (equation 1,
paper SC/F16/JR17), but it is not clear how costs of growthand energy storage have been incorporated into the analyses.
It is possible that the information on condition factor (SC/F16/
JR27) could have contributed to bioenergetics analysis. SC/
F16/JR17 identified several allometric equations that havebeen used in other studies to estimate consumption. These
provide a range of estimates of energy requirements for anwhat each formulation was estimating. For example, Equation
3: SMR=863.6M0.783 (Sigurjónsson and Víkingsson, 1997)
provides an estimate of energy requirements, described asthe ‘near basal energy requirement, times an activity factorof 1.5’, but this does not take into account growth, seasonalchanges in condition or reproduction. If the consumption
models were not modelling energy gain (growth in protein
and fat deposition, as well as reproduction explicitly), thenan alternative could have been to apply a separate multiplier
to take into account some of the additional costs associatedwith growth and reproduction. This would also contributeto model uncertainty, although uncertainty has not been
identified in model development.
A critical factor in estimating consumption is the spatial
distribution of prey and whales throughout the year. The
JARPN II surveys were conducted during May/June andJuly-October. Estimates of consumption have been provided
for the intervening months, but it is not clear how densityand diet consumption have been extrapolated outside of
the areas and months covered during the surveys and dietstudies. The Panel agrees that the proponents clarify this(see Item 6.4.3.2).
Stomach contents provide information on species, size
and relative contribution of prey to the diet. In common withall methods used to determine diet composition, the use of

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

stomach contents has weaknesses. For example stomach
contents only reflect what was consumed during the last 1-2
meals, within the last 24h. The 2009 Panel recommended theuse of other methods to obtain information on diet compositionin addition to stomach data, such as stable isotopes or fatty
acid composition, but these methods have not been used.
Both methods can use material either from sampled whales
or from biopsies. These methods also have limitations suchas information on what specific prey can appear in the diet,a need for a prey library, and an understanding of tissue
turnover rates, as well as uncertainties about where the prey
were ingested. However, the present Panel reiterates theneed to consider other methods because of the potential not
only to assess diet, but also to statistically evaluate overlap
in trophic niche and distribution (Gavrilchuk et al., 2014;
Ryan et al., 2013; Witteveen et al., 2009). The use of thesealternative methods provides insights into prey ingestion
over periods that vary according to the tissue examined:
several days or weeks (skin), several months (muscle), or
long-term (bone) (Browning et al., 2014; Giménez et al.,2016; Yoshida and Miyazaki, 1991). Thus, they may extend
the sampling window to determine diet composition fromthe last 24h to an improved understanding of long-term dietcomposition. Moreover, stable isotope analysis of sequential
layers in baleen, a structure that grows continuously andis composed of metabolically inert tissue that does notexperience isotopic turnover, provides information of
the isotopic body pool composition during a period that
depending on the species involved may extend up to the 2-4
years previous to sample collection (Aguilar et al., 2014),
thus allowing inference on migration route and breeding
ground. The greatest information on diet composition
can therefore be gained from using multiple methods inconjunction.
The ultimate objective of the dietary studies is to convert
individual consumption to population consumption ofdifferent prey. This is a complex challenge because samplingof whales and prey is spatially and temporally limited. It is
not clear how the conversion of the consumption estimatesand prey preference from the level of the individual whaleto the entire population, taking into account the temporal/
spatial changes in diet composition and the spatial/temporalchanges in whale distribution, was undertaken. Although
any approach is likely to be limited, clarification of methods
and how uncertainty was considered is needed.
6.4.3 Recommendations
6.4.3.1 MEDIUM-TERM

The Panel has developed the following recommendations
for the medium-term i.e. normally to be completed 2-3 years
after the 2016 Annual Meeting:(1) all sources of uncertainty should be quantified and anevaluation of which parameters contribute the most to
uncertainty be conducted and taken into account in theanalyses and modelling;(2) the studies on allometric relationships should be
developed further to refine the range of suitable
allometric-energy intake/consumption relationships;(3) the analyses of diet composition should consider theeffect of seasonal changes in energy density of thevarious prey species; and(4) stable isotope analysis of whale tissues and their preyshould be introduced not only into the assessment of diet,but also to statistically evaluate overlap in distribution
and trophic niche between baleen whale species.

555

6.4.3.2 SHORT-TERM

The Panel has developed the following recommendations
for the short-term i.e. to be completed ideally by the 2016
Annual Meeting but certainly by the 2017 Annual Meeting:(1) the sampling distribution for the parameters should be
used in the assessment of the uncertainty associatedwith the estimation of consumption; and(2) clarification should be provided on how density and dietconsumption have been extrapolated outside the areas
and months covered during the surveys and diet studies.
7. FEEDING ECOLOGY AND ECOSYSTEMSTUDIES: PART 3 ECOSYSTEM MODELLING
7.1 Summary of objectives including modifications, ifany, since the start of the programme (by proponents)

With respect to ecosystem modelling, a new sub-objective wasadded:

Sub-objective 10: To develop several ecosystem models,in both coastal and offshore areas, using JARPN II dataand samples as input. Model outputs are likely to provide
information on: (i) the ecosystem structure; (ii) effects of preyavailability and consumption on the population dynamics ofcommon minke and sei whales with consideration of levelsof energy intakes; and (iii) predation impacts of common
minke whales consumption on sandlance stock off Sanriku.

7.2 Overview of conclusions and recommendations fromthe 2009 Workshop
A short overview on the 2009 Workshop conclusions andrecommendations on the ecosystem modelling componentof JARPN II was provided by the Head of Science.
The 2009 Panel had noted that the overall objective
regarding ecosystem management was both highly ambitious
and very general. It had recommended the development of
sub-objectives and a timeframe. It highlighted that it was
unlikely that output sufficient for providing management
advice would be available in a short time frame and it mighttake much longer. The preliminary approaches presented
were a reasonable start but the conclusions were overstatedand certainly not suitable as a basis for management. The
Panel stressed that a variety of modelling approaches are
required. It noted that the data from sperm whales made noworthwhile contribution to the modelling work.
The 2009 Panel made a number of general recommendations including:(a) considerably more resources must be allocated tomodelling work;(b) models should be used to evaluate priorities by
estimating which are the key parameters (and where
effort should be targeted to reduce uncertainty) in
the context of management-related outputs;(c) a wider range of models must be considered;(d) future work should aim towards fitting dynamic
models to time series of data;(e) separate models should be developed for thedifferent ecological regions;(f) considerably more effort should be directed atquantifying uncertainty with respect to all aspectsof the input and model assumptions; and(g) effort should be put into the incorporation of naturalvariability in dynamic processes.
The 2009 Panel had also made a number of specificrecommendations related to:(a) ensuring that all likely significant predators areincluded in the models, not just whales;

556

REPORT OF THE EXPERT PANEL ON THE FINAL REVIEW OF JARPN II(b) not using Type I functional response relationships
as these are unrealistic;(c) the need to improve the Ecopath component withrespect to the EwE (Ecopath with Ecosim) approach
before moving to Ecosim e.g. by:i.
reviewing the species considered;ii. reviewing the parameter values used;iii. rebalancing Ecopath using different app   roaches; andiv. quantifying uncertainty(d) further work on the MRM (minimum realistic
model) approach with an emphasis on fitting to timeseries.

accounts for a certain proportion of the current adult biomass
for the sandlance population although the level of proportion issensitive to the model assumption.
The addendum of SC/F16/JR29 reported that there wasa data-handling error in the analysis above. Due to time
constraints, the authors of SC/F16/JR29 were not able to
finalise all the analyses by the time of the review workshop.
In the addendum, the outline of formulation and one example
of outcomes (without any predation effect), were shown.
The outcome suggested that a percentage of consumption byminke whales accounted for around 10% of total biomass of
sandlance. The number would be higher if expressed in termsof adult natural mortality, as the estimate of natural mortality isless than 1. However, due to preliminary nature of this analysis,
the authors insisted that the result should be taken as at best
broadly indicative only. The authors plan to submit a full paperwith more details on this analysis as well as the analysis withconsideration of predation effects.
In conclusion, the ecosystem models presented tothis workshop were improved significantly based on therecommendations/suggestions raised in the previous workshop
held in 2009. They would serve as baseline models to test
various types of marine ecosystem management options.
It should be noted that the Panel of the workshop in 2009
pointed out that it might take a long time to obtain results ofecosystem modelling that are sufficiently reliable to informmanagement advice. The panel also pointed out that substantial
data collection and analytical efforts are required to accomplish
the goal. The proponents have developed ecosystem models
bearing those suggestions in mind. Although JARPN II was
terminated as a program, continuation of similar research isnecessary to develop ecosystem models for the purpose ofmanagement advice.

7.3 Proponents’ summary of the ecosystem modelling
work (including response to 2009 Workshop)

SC/F16/JR28 presented the results of ecosystem modelling in
the western North Pacific from 1994 to 2013 using Ecopath
with Ecosim (EwE). The recommendations/suggestions raisedby the 2009 JARPN II review workshop were addressed in thepaper except the treatment of uncertainties. Firstly, Ecopath
in 2013 was constructed as available data for the modelling
is relative rich. Ecopath in 1994 was then constructed basedon the model in 2013. Finally, Ecosim is constructed based on
Ecopath in 1994 using available time series data from 1994
to 2013. Regime of the period is relative stable in comparison
with the past. A series of pre-balance diagnostics, ‘PREBAL’(Link, 2010) was conducted for both the 2013 and 1994 models
to evaluate the initial static energy budget of Ecopath. An
ecosystem network analysis indicator, mixed trophic impact(MTI), was used to assess the positive or negative effect ofchanges in the biomass of a species/group on the biomass ofthe other species/groups in the steady state ecosystem. Orderof Trophic level (TL) of baleen whales was as follows (fromhigh to low): common minke (4.1), Bryde’s (3.9), sei (3.7),
humpback (3.5), fin (3.3) and blue (3.2) whales. These species
are in intermediate TL in the ecosystem. MTIs suggested thatchanges in biomass of forage fish impact most of species/groups from low to high trophic levels. Baleen whales
impact forage fish negatively but the magnitude is weak. The
Ecosim model with forced biomass trends of four forage fish
species (Japanese sardine and anchovy, and chub and spottedmackerels) having 10 predator and prey search blocks attain thelowest AIC. Estimated trends of biomasses and total mortality
by using the model are reasonably fitted to input time series
data especially for cetaceans targeted by JARPN II. Overall
results appear to be reasonable but it is still preliminary largely
because of incompleteness of input data. The following arepoints to be improved in further exercises: (1) consistency of
spatial resolution of input data; (2) development of regionalmodels within our EwE area; (3) collection of diet composition
data in regular interval; (4) resolution and quality of data on
non-commercial and lower trophic level species; and (5)evaluation of the sensitivity of Ecopath models to input data.
Uncertainties in the model will be considered along with these
additional works. The developed EwE would be used as a basemodel for broad-scale strategic management consideration ofwhales as well as other species such as small pelagic fish.
SC/F16/JR29 aimed at assessing predation impact onsandlance population by common minke whales off Sanriku
region. A state-space delay-difference model, which is a twostage population dynamics model with a stock-recruitment
relationship, was used for the sandlance population to employ
two independent time series indices for the juvenile and mature
population sizes as well as catch and age-composition data.
Predation impacts on the sandlance were assessed throughcommon minke whales’ consumption expressed as a functional
response. To take into account several stochastic flexibilities
such as process errors, a Bayesian method was used to estimate
the parameters and latent variables in the model. The results
showed that the predation by the common minke whales

7.4 Panel review, conclusions and recommendations
7.4.1 Overview
The Panel reiterates the 2009 Workshop comment thatdeveloping ecosystem models to the level that they could
contribute in the provision of specific management advice
constitutes a major, complex and ambitious undertaking, andthat developing ecosystem models requires substantial datacollection and analytical efforts. Ecosystem modelling is an
ideal tool for integrating a range of different processes anddata sources into a single coherent framework. Moreover,the Panel also reiterates the 2009 Workshop statement that
there are benefits to developing a wide range of models andthat the best ecosystem modelling approach depends on thequestion being addressed.
The Panel agrees that substantial progress has been
made since 2009 on the Ecopath/Ecosim (EwE) model andan extended single species sandlance model with predationby common minke whales was presented for the Sanriku
region. However, the work presented to the Panel remains
preliminary, and the Panel was not able to review the results
for the sandlance model as an error was found in the data
used in the model. It is clear that substantial additional work
remains to be done in order to develop a MRM/MICE1
and before any ecosystem model could be used to provide
managers with strategic advice as envisioned in the original
objective. The Panel does not recommend using ecosystems
models for tactical management.
The Panel agrees that the work presented in SC/F16/
JR28 and SC/F16/JR29 forms a reasonable basis for further
work. However, the work is incomplete in several ways
and the modelling portion of the project was perhaps mostnegatively impacted by the review occurring before the
Minimum Realistic Model/Models of Intermediate Complexity for Ecosystem Assessment.

1

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557

Table 8
Summary of the Panel’s evaluation of the proponents’ response to the 2009 recommendations on ecosystem modelling (see text).
Brief summary of the 2009 recommendations/suggestions
Generic recommendations
Considerably more resources must be allocated to the modelling work –without this, the likelihood that the objective of the programme will bereached in a reasonable timeframe will be minimal.

Panel comments
Partly addressed. Although progress has been made since the 2009
workshop, this is less than would have occurred had sufficient resources
been allocated to this aspect of the programme. This is reflected in the
preliminary nature of the work provided to the Panel.

The models developed should be used to identify the areas of uncertainty
with the greatest impact on model outputs of relevance to management,
and hence to guide the prioritisation of future data collection and theassociated sample size/sampling design.

Not addressed. Given the preliminary nature of the modelling, the work
had not fed through into the prioritisation of data collection during the
programme or to issues of sample size or sampling design

A wider range of models needs to be considered if the objectives of the
programme are to be met. Further work should aim towards fitting dynamic
models to time series of data, especially abundance indices.

Partly addressed. Two models have been developed, although a planned
MRM/MICE has not been developed. Both of the models presented are fit
to the data, but the Ecosim model cannot provide adequate fits to the data
unless the biomass of small pelagic species are pre-specified.

The area covered by JARPN II is not spatially homogeneous, and serious
consideration should be given to developing separate models for three
regions distinguished by the inshore or shelf region, the sub-Arctic oceanic
region of the Oyashio current and the sub-tropical region of the Oyashio
and Kuroshio transition zone.

Partly addressed. It has been addressed for the EwE model, but theextended single-species model has only been applied to one interaction in
one region.

There is a need to take much wider account of uncertainty at all stages ofthe modelling process, including that associated with the prey consumption
rates of whales.

Not addressed. Limited sensitivity analysis has been undertaken for theEwE model, while the extended single-species model is not sufficiently
well developed for sensitivity testing to be undertaken. It is also dependent
on progress with the feeding ecology work (see Item 6.4).

The importance, ultimately, of developing models which incorporate
natural variability in dynamic processes (e.g. recruitment variability forprey species) was emphasised, although it was recognised that this might
not be possible for certain ecosystem modelling packages.

Partly addressed. The extended single-species model includes process error
in recruitment and the population dynamics (but the model currently underdevelopment has not been able to estimate the associated parameter well)Specific recommendations
Include other important sandlance predators in the single species model.

Not addressed.

It is important to concentrate first on improving the Ecopath component of
this EwE analysis before moving on to the next step of extending themodelling effort from a static to a dynamic model such as Ecosim.

Largely addressed. While data limitations remain, the model presented to
the Panel provides a reasonable synthesis of the available information (butsee Item 7.4.3.2).

The species included in the Ecopath analysis should be reviewed giving
attention to Ecopath models developed for other regions; in particular the
inclusion of gelatinous zooplankton should be considered.

Addressed.

The values of the parameters of this Ecopath analysis should be comparedwith values for those others, with attention directed towards any instancesof major discrepancies.

Partly addressed. PREBAL diagnostics were applied, so this was addressedto the extent that PREBAL diagnostics are based on general comparisonsamong models. However, specific comparisons with other North Pacificfood web parameterisations were not included.

Alternative approaches for balancing the Ecopath model should beconsidered e.g. rather than use values for some parameters drawn fromother regions, placing a bound on some relationships (e.g. P/C<0.6).

Partly addressed, but see recommendations (Item 7.4.3.2 (2)).

Analyses must take full account of the uncertainties associated with model
inputs for Ecopath, e.g. using Ecoranger.

Not addressed. Ecoranger is not necessarily recommended, but anothermethod should be considered.

Further work on MRM approaches is encouraged and should focus inparticular on fitting such models to time series of data

Not addressed. Need recognised by proponent scientists. Critical forfurther progress.

originally envisioned end of the programme. The extended
single species model shows promise as step towards a MRM/
MICE to evaluate impacts of whales on fisheries.
The Panel reiterates that the EwE as developed is notsuitable for use in addressing the strategic management
questions outlined by the proponents involving whalefishery interactions. Firstly, reasonable fits were obtained
only when the model was driven with the biomass of small
pelagic species which precludes its use to investigate theimpact of whale predation on these small pelagics. Secondly,
the EwE software imposes constraints on a full evaluation of
the statistical properties of the model and an evaluation of
uncertainty. Previous work (Gaichas et al., 2011) has shownthat a single ‘best fit’ to historical data without accounting
for parameter uncertainty does not provide a sound basis forforecasts.
In conclusion, with respect to the extent to which theproponents have met their Objective 1, Sub-objective 10 5(‘To develop several ecosystem models, in both coastal andoffshore areas, using JARPN II data and samples as input’),
the Panel agrees that it has been only partly met in that whilst
two models have been developed, they are preliminary and aplanned MRM/MICE has not yet been developed.
The model outputs are likely to provide information
on ecosystem structure. However, with respect to theeffects of prey availability and consumption on the population dynamics of common minke and sei whales withconsideration of levels of energy intakes, the Panel agrees
that whilst a link has been demonstrated, the effects havenot been investigated. Finally, with respect to estimating
the predation impacts of common minke whales on thesandlance stock off Sanriku, the Panel agrees that whilst
work is in progress, it has not been completed.
The Panel stresses that the lack of progress with respect
to ecosystem modelling compared to that envisaged in 2009is not a criticism of the scientists involved. Disappointingly,
despite the recommendation from the 2009 Panel and the fact

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that it is central to being able to meet the primary objective,
insufficient resources have been put into this aspect of the
programme.
7.4.2 Evaluation of progress with the 2009 recommendations (see Annex D for a complete summary table)
The 2009 Panel had made a number of generalrecommendations to further the work on ecosystemmodelling. None had been fully addressed but several had
been partly addressed. Two, related to: (a) using models
to identify areas of uncertainty with the greatest impact onmodel outputs to improve sampling; and (b) taking wider
account of uncertainty at all stages of the modelling process,had not been addressed. As noted earlier, the relative lack ofprogress relates to the lack of resources allocated. Detailed
comments are given in Table 8.
The 2009 Panel also made a number of specificrecommendations for ecosystem modelling based on thepapers presented at the time. Two were largely or fully
addressed (related to improving Ecopath), two were partly
addressed (related to parameter values in Ecopath) andthree were not addressed (related to the inclusion of otherpredators in the sandlance model, taking account of theuncertainties in model inputs, and fitting time series of data
within the MRM). Detailed comments are given in Table 8.
7.4.3 Recommendations
The Panel has developed the following recommendations
for the medium-term i.e. normally to be completed 2-3 years
after the 2016 Annual Meeting.
7.4.3.1 GENERAL RECOMMENDATIONS(1) The generic recommendations identified by the 2009
Panel remain, and the Panel endorses them.
(2) Clear objectives on the ultimate use of the models mustbe established to make further progress. One model
may meet a general objective of better understanding
ecosystem linkages, while a very different model
may be more suited to delivering tactical advice forfishery management. For example, EwE modellingdemonstrates linkages between small pelagic species
and whales, both in the static mass balance and in the
dynamic model. However, it is not clear how suchmodelling could directly inform strategic management.
(3) Models can be better used in concert. The results offood web modelling should be used to establish key
predation linkages to include in extended single-species
or multispecies models. This best exploits the strengthsof different types of models. Specifically, Ecopath
should be used to define key relationships for further
study within targeted statistical MRM/MICE models
linked to specific objectives. In such a way the suiteof available modelling tools can be used to integrate
available knowledge.
(4) Stable isotopes provide information on long term
feeding patterns to inform models about trophic
relationships between whales and their prey (see also
Item 6.4).
7.4.3.2 EWE MODEL(1) An evaluation of data quality for each input parameter,the ‘pedigree’ (Gaichas et al., 2015) is needed tocharacterise uncertainty in model inputs.
(2) Further evaluation of PREBAL and other diagnostics
should be conducted. The Panel was concerned byrelatively low Ecotrophic Efficiencies for sardine andsaury, which may indicate that predation on these
species is not fully modelled. This would be worrying
given the focus of the modelling on the relationshipbetween small pelagic fishes and whales.
(3) The estimated vulnerabilities and other fit diagnostics
could be presented more clearly and evaluated further.
Software constraints inherent to Ecosim should not limit
the consideration of uncertainty; sensitivity analysis
using ranges of consumption estimates, for instance,
could be done.
7.4.3.3 EXTENDED SINGLE-SPECIES MODEL(1) The model should be developed to ensure that the
majority of predation mortality is captured. A food
web model (e.g. Ecopath) quantifies mortality from all
predators in the ecosystem, and could be used to informwhich major predators to include in this work.
(2) Additional diagnostics are needed. Specifically, there
is a need to examine the fits to the fishery-independent
survey data, the proportion information, and trends infishing mortality. In additional, posterior predictive
checks can be used to evaluate model fit in Bayesianmodels.
(3) The current spatial boundaries of the model and the useof fishery CPUE as an index of abundance should be
more thoroughly justified.
(4) If CPUE for the dip net fishery is not considered likely
to index abundance, the focus for model fitting shouldbe the fishery independent survey.
(5) There is unlikely to be sufficient information to estimate
the functional form of the feeding relationship andsensitivity to alternative plausible relationships should
be examined.
(6) Some of the posteriors in SC/F16/JR29 appear
implausible (e.g. the posterior for the intrinsic growthrate for the prey larger with most of its mass on values
larger than two and the posterior mode for juvenile
survival close to 1 in the absence of minke whales). Thisis likely a consequence of conflicts between the data
sources or confounding of parameters due to insufficientdata. The causes of the implausible posteriors can be
explored by changing the weights assigned to the data
sources and fitting the model.
8. MONITORING ENVIRONMENTAL
POLLUTANTS IN CETACEANS AND THE MARINEECOSYSTEM
8.1 Summary of objectives including modifications, ifany, since the start of the programme (proponents)

In 1992 the Commission endorsed the plan of the IWC SC topursue studies on environmental changes and their impacts
on cetaceans. In particular, there was concern that pollutants
may have a negative effect on the health of cetaceans resulting
ultimately in a decrease in the abundance of the stocks.
The main Objective 2 of the JARPN II research plan was‘monitoring environmental pollutants in cetaceans and themarine ecosystem’, which is composed of the following threesub-objectives.
Objective 2, Sub-objective 1: Pattern of accumulation ofpollutants in cetaceans (Relevant documents: SC/F16/JR3034).
Objective 2, Sub-objective 2: Bioaccumulation process ofpollutants through the food chain (SC/F16/JR30).
Objective 2, Sub-objective 3: Relationship between
chemical pollutants and cetacean health (SC/F16/JR35; Niimi
et al. (2014) [SC/F16/36]; Shimizu et al. (2013) [SC/F16/37].

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017559

Fig. P5. Schematic representation of the research components under Objective 2 of JARPN II.

A schematic representation of the research components
under Objective 2 of JARPN II is shown in Fig. P5. Air andseawater samples, whale samples and their prey items werecollected in the JARPN II surveys. Levels of pollutants suchas mercury (Hg), organochlorines (OCs) and biomarkers
in the samples were measured in laboratories of the ICR or
the collaborative groups. And these data were analysed forachievement of each sub-objective.

8.2 Overview of conclusions and recommendations fromthe 2009 Workshop (Chair, Head of Science)
A short overview on the 2009 Panel conclusions andrecommendations on the pollutant component of JARPN II
was provided by the Head of Science.
The 2009 Panel had concluded that the JARPN II
pollutant studies represent a valuable contribution topollutant work. It agreed that the programme was addressing
its objectives but it had recommended further work. In
particular it noted that revised papers should include a
general risk assessment for various pollutants, based on
current ecotoxicological knowledge and information fromother wildlife on likely thresholds for adverse health effects
to be detected. For future studies it had recommended thatpolychlorinated biphenyl (PCB) concentrations should bereported on a lipid weight basis and that combined analysesof PCBs and Hg should be undertaken. Furthermore, it notedthat it was important for the proponents to develop a morebalanced, structured study design to allow statistically robust
consideration of hypotheses, with all of the necessary datacollected from each of the targeted individuals and with acontrol or comparison group. It was suggested that bycaught
J-stock animals could be suitable as one comparison group.
Recognising the continuing development of techniques, it
had recommended that tissues should be archived for future
retrospective analyses. It also suggested that priority should
be given to ways to determine absolute age as an additional
covariate for the interpretation of the results.
In terms of additional or more integrated studies, the2009 Panel had recommended that the proponents should:(a) include fatty acid profiles and stable isotope ratios
to help discriminate among reasons for temporalchanges (e.g. dietary changes or exposure variation
with constant diet);(b) place more emphasis on air and water studies; and(c) undertake simple mass balance studies to improveknowledge of the partitioning and offloading ofcontaminants and potential impact of changes in
exposure, noting that this would require additional
analyses of blood, bile, faeces and urine.
Finally, it had recommended that the contaminant results
be linked to prey consumption studies.

8.3 Proponents summary of the results (incl. response to2009 Workshop)

Studies on comprehensive monitoring and assessment ofenvironmental pollutants under the JARPN II were based ona large and comprehensive data/sample set, and involved theuse of pollutant levels (Organochlorines and Hg) in whales andtheir prey items.
The combination of large sample size and detailed
biological data was an appropriate tool to monitor pollutant
levels in whales and their prey items, and assess whale health.
OBJECTIVE 2, SUB-OBJECTIVE 1: PATTERN OF
ACCUMULATION OF POLLUTANTS IN CETACEANS

SC/F16/JR30 examined temporal trends in total Hgconcentrations in muscle of mature male whales using
samples obtained during JARPN and JARPN II (1994-2014).
Common minke whales from sub-areas 7, 8, 9, off Kushiro and
off Sanriku; sei whales from sub-area 9 and Bryde’s whales
from sub-areas 8 and 9 were sampled. Multiple regression
analyses were carried out to determine whether year sampled,longitude, latitude, date, whale body length, blubber thickness
and/or main prey species were significantly correlated with
Hg concentrations in the muscle samples from the whales.
Significant correlations with year and main prey species wereobserved in common minke whales from sub-areas 7 and 9. A
significant time trend was also found for the sei whales but withno significant main prey species effects. However, body length
had a small but significant positive coefficient for sei whales,suggesting that older sei whales might have somewhat higher
Hg levels. Thus temporal trends in total Hg were detected insome but not all species and areas and could be affected bychanges in prey species. SC/F16/JR30 also suggested thatbackground levels of total Hg in the western North Pacific werestable during the 1994-2014 period.
SC/F16/JR31 looked for temporal trends in PCBs forcommon minke whales from sub-areas 7 (period 2002-12),8 (period 2002-09), 9 (period 2002-13), off Kushiro (period
2002-14) and off Sanriku (period 2003-14) from the western
North Pacific. Multiple regression analyses were carried out.
Data included adjustment for years, longitude, latitude, date,whale body length, blubber thickness and main prey species.
No significant correlations with year and food items were
found. It is suggested that background levels of PCB in thewestern North Pacific were stable through the 2002-14 period.
SC/F16/JR32 examined the patterns of PCB congeners,
DDT isomers, HCH isomers, HCB and CHL isomers in the
blubber of five mature males of each of common minke, sei
and Bryde’s whales taken from the western North Pacific in
2012. For comparison, those compounds were also determinedin the blubber of five mature males of Antarctic minke whales
taken from Antarctic Area V in 2010/11. Concentrations of
PCBs were highest among organochlorines in the whales
from the western North Pacific, whereas they were lower than

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concentrations of HCB, DDTs and CHLs in Antarctic minke
whales. Principal Component Analysis showed differencesof trophic level and habitat of PCB congener profiles in thewhales. Over 4 chlorinated chlorobiphenyl (CB) congeners
in the studied whales contributed to the difference of trophic
levels, and the CB-32, 16 and 25 contributed to the geographicaldifference. The main component isomers from pesticide
products originating in DDTs and HCHs were comparatively
lower, and those originating in CHLs were not detected in thewhales from the western North Pacific. These results suggestthat in the western North Pacific, a great deal of time would
have passed from the release of DDTs, HCHs and CHLs into
the environment.
SC/F16/JR33 examined differences of total Hgconcentrations in muscle and liver of J- and O-stocks ofcommon minke whales off Sanriku. Concentrations of total Hg
in muscle and liver of 35 O- and 24 J-stock immature minke
whales taken in the 2012 and 2013 JARPN II surveys were
measured. Multiple linear regression analyses of total Hgconcentrations of the whales were carried out. These included
adjustment for confounders, age index, sex, stock, blubber
thickness and year. Stock had no discernible effect. Thesefindings suggest that there is no stock-dependent difference of
total Hg exposure risk for the minke whales from off Sanriku.
SC/F16/JR34 examined the pollutants status of sperm
whales in the western North Pacific. Total Hg, PCBs, DDTs,
HCHs, HCB and CHLs were determined in samples of sperm
whales in the period 2001-13. Mean concentrations of total Hg
in muscle of sperm whales in the periods 2001-05 and 2011-13
were 1.9 and 1.5 (ppm wet wt.), respectively. No significant
difference was observed in their total Hg levels between thetwo periods. Mean concentrations of PCBs, DDTs, HCHs,
HCB and CHLs in blubber samples in 2012 were 1.9, 0.74,
0.040, 0.077 and 0.65 (μg/g fat wt.), respectively. Levels of
total Hg in muscle samples of sperm whales in the present
study were slightly lower than those from Ayukawa, Japan in
1978 and 1979, and from the southern North Sea in 1994 and1995. Levels of organochlorines, except for CHLs, in sperm
whales from the western North Pacific were similar or lower
than those in sperm whales from the middle latitudes of thenorthern hemisphere nearby human activity. In addition, there
is no evidence that levels of total Hg in muscle of sperm whales
increased in the period of 1970s to 2000s.
In conclusion, year-to-year trends in Hg and PCB levels
in baleen whales from the western North Pacific were notobserved, suggesting that the background levels of thosepollutants were stable during the 2002-2014 research period.
Results of the organochlorines isomer analyses in whale
tissues showed that almost no recent inputs of DDT, HCH and
CHLs have been released into the JARPN II research area.
Initially it was expected that J stock common minke whales
had more exposure to contaminants than O stock whales due totheir coastal distribution. However, no significant differenceswere found in the total Hg exposure between J and O stocks
minke whales off Sanriku. However it should be noted that
future analyses should include older animals to confirm thisconclusion.
OBJECTIVE 2, SUB-OBJECTIVE 2: BIOACCUMULATIONPROCESS OF POLLUTANTS THROUGH THE FOOD
CHAIN

SC/F16/JR30 examined the yearly changes of total Hg ofcommon minke whales from sub-areas 7, 8, 9, off Kushiro
and off Sanriku, sei whales from sub-area 9 and Bryde’s
whales from sub-areas 8 and 9. Multiple regression analyses
were carried out. Data included adjustment for confounders,sampling years, sampling longitude, latitude, sampling date,body length, blubber thickness and main prey species. No
significant correlations between total Hg and sampling years
were observed in almost all whales except for minke whales
from sub-areas 7 and 9 and sei whales from sub-area 9. Total
Hg levels in common minke whales from sub-areas 7 and 9 andsei whales from sub-area 9 were simultaneously correlated withmain food items. These findings suggest that yearly changesof total Hg in common minke whales from the western North
Pacific could be affected by changes of their prey species.
In conclusion, the analysis of the relationship between total
Hg levels in whale tissue and main prey species in whale’s
stomach revealed an effect of prey species on the level ofcontaminant in whales. This suggested that the level of Hg
in whales depended on the trophic level position of the preyspecies.
OBJECTIVE 2, SUB-OBJECTIVE 3: RELATIONSHIPBETWEEN CHEMICAL POLLUTANTS AND CETACEANHEALTH

SC/F16/JR35 examined the health risk of radioisotopes (RIs)
for large whales from the western North Pacific. From 11 March2011 onward, RIs were released to the marine environment
from the Nuclear Power Plant in Fukushima following anearthquake and tsunami. To assess the presence of these RIs in
the large whales, the I131, Cs134 and Cs137 levels in muscle
samples of 53 common minke, 16 Bryde’s, 32 sei and 3 sperm
whales from JARPN II surveys were measured in the period
2011-15. I131 was detected in muscle samples of large whales,
except for two common minke whales off Kushiro in 2012.
Ranges of Cs134 + Cs137 concentrations in common minke,sei, Bryde’s and sperm whales were ND-31, ND-9.8, ND-7.1
and ND-0.59 Bq/kg wet wt., respectively. The radioisotope
levels in all whales examined have been decreasing since2011, and were also extremely lower than the radiation safety
threshold for humans. Therefore, risk of acute toxicity levels
for I131, Cs134 and Cs137 would be extremely low in the large
whales from the western North Pacific.
SC/F16/JR36 (Niimi et al., 2014) determined hepaticconcentrations of persistent organochlorines (OCs) in thecommon minke whale from the North Pacific. To investigate
the effects of OCs on the transcriptome, the study constructeda hepatic oligo array of this species where 985 uniqueoligonucleotides were spotted and further analysed therelationship between the OC levels and gene expression
profiles of liver tissues. The stepwise multiple linear regression
analysis identified 32 genes that correlated with hepatic OC
levels. The mRNA expression levels of seven cytochrome
P450 (CYP) genes, CYP1A1, 1A2, 2C78, 2E1, 3A72, 4A35,
and 4V6 showed no clear correlations with the concentrationof each OC, suggesting that the accumulated OCs in the liver
did not reach levels that could alter CYP expression. Amongthe genes screened by the custom oligo array analysis, hepatic
mRNA expression levels of 16 genes were further measured
using quantitative real-time reverse transcription polymerase
chain reaction. The mRNA levels of vitamin D-binding protein(DBP) were negatively correlated with non-ortho coplanar
polychlorinated biphenyl (PCB) levels. Androgen receptorassociated coregulator 70 (ARA70) expression levels showeda significant positive correlation with concentrations of nonortho coplanar PCB169. These correlations suggest that
coplanar PCB-reduced DBP expression could suppress vitamin
D receptor-mediated signalling cascades in peripheral tissues.
Alternatively, the suppression of vitamin D receptor signallingcascade could be enhanced through competition with theandrogen receptor signalling pathway for ARA70. In addition, anegative correlation between kynureninase and PCB169 levels
was also observed, which suggest an enhanced accumulation ofan endogenous aryl hydrocarbon receptor agonist, kynurenine
in the minke whale population.
SC/F16/JR37 (Shimizu et al., 2013) examined morbillivirus
infection in marine mammals. Mass die-offs caused by thisinfection have repeatedly occurred in bottlenose and striped
dolphins, both of which belong to the family Delphinidae,
but not in other cetaceans. However, it is unknown whether
sensitivity to the virus varies among cetacean species. The
signalling lymphocyte activation molecule (SLAM) is a
receptor on host cells that allows morbillivirus invasion andpropagation. Its immunoglobulin variable domain-like (V)region provides an interface for the virus hemagglutinin (H)561

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017
Table 9
Summary of Panel’s evaluation of the proponents’ response to the 2009 recommendations with respect to pollutants (and see text).
Brief summary of 2009 recommendations

Panel evaluation

Papers should include a risk assessment statement.
Analyses be carried out by age when age data become available.
Examine levels in the liver to facilitate comparison with other studies.
Use GAMs when examining non-linear trends in the Hg levels.
Carry out future studies on a lipid weight basis.
Sample for PCBs and Hg from same individuals to allow combined analyses.
Revise paper SC/J09/JR25 (accumulation of total and methyl mercury and selenium) to highlight important ecotoxicologicalfinding.
Examine T-Hg total body burden estimates, sampling additional organs (e.g. brain, skin, blubber).
Examine T-Hg in brain to compare more coastal bycaught animals to the coastal and offshore JARPN II samples.
Continue comparative molecular phylogenetic research using mRNA isolated from fresh tissues.
Develop a balanced, structured study design and collect all necessary data from each targeted animal; include a control orcomparison group (e.g. bycaught J stock animals).
Archive tissues for future retrospective analyses.
Give priority to have absolute age as an additional covariate for the interpretation of the results.
Include fatty acid profiles and stable isotope ratios in analyses.
Use air and water samples in a ‘fate and behaviour’ study.
Undertake simple mass balance studies.
Eventually link contaminant results to the prey consumption studies.
Undertake power analyses for relationship between sample size and ability to detect changes at various levels.
Evaluate covariates (e.g. age and sex) to determine animals chosen for more extensive sampling.
Examining the same individuals for each of the contaminants is emphasised.

Partly addressed.
Partly addressed.
Addressed.
Not addressed.
Partly addressed.
Partly addressed.
Addressed
Not addressed.
Not addressed.
Addressed.
Partly addressed.
Addressed.
Partly addressed.
Not addressed.
Addressed.
Not addressed.
Partly addressed.
Not addressed.
Partly addressed.
Not addressed*.

*apart from for two sperm whales.

protein. In this study, variations in the amino acid residues of the
V region of 26 cetacean species, covering almost all cetacean
genera, were examined. Three-dimensional (3D) models ofthem were generated in a homology model using the crystal
structure of the marmoset SLAM and measles virus H protein
complex as a template. The 3D models showed 32 amino acidresidues on the interface that possibly bind the morbillivirus.
Among the cetacean species studied, variations were foundat six of the residues. Bottlenose and striped dolphins havesubstitutions at five positions (E68G, I74V, R90H, V126I, andQ130H) compared with those of baleen whales. Three residues(at positions 68, 90 and 130) were found to alternate electriccharges, possibly causing changes in affinity for the virus. This
study shows a new approach based on receptor structure forassessing potential vulnerability to viral infection. This method
may be useful for assessing the risk of morbillivirus infection
in wildlife. In conclusion, I131, Cs134 and Cs137 levels inlarge whales from the western North Pacific were much lower
than safety threshold in humans.
Information of new biomarkers was provided, which
can be used in future studies on adverse effect of OCs andvulnerability to viral infection in whales.

8.4 Panel review, conclusions and recommendations
8.4.1 Overview
The second objective of the JARPN II programme related
to ‘monitoring environmental pollutants in cetaceans andthe marine ecosystem’. This included investigations of:(a) patterns of accumulation of pollutants in cetaceans; (b)the bioaccumulation process of pollutants through the foodchain; and (c) the relationship between chemical pollutants
and cetacean health.
Papers detailing the results of the chemical analyses anddiscussion of the findings in relation to this objective and theirsub-objectives were provided for the panel’s consideration(SC/F16/JR30-35; Shimizu et al. (2013) [SC/F16/JR36];
Niimi et al. (2014) [SC/F16/JR37]). These papers greatly
improved the previous knowledge of chemical pollutants inlarge cetaceans from the North Pacific waters neighbouring
Japan. The studies focused primarily on mercury (Hg) andorganochlorine compounds (OCs), although one study (SC/F16/JR35) examined radioactive caesium (Cs) and iodine 6(I) concentrations in sperm and baleen whales, a subject ofparticular relevance after the Fukushima accident in 2011.
These papers provided new information on concentrationsof these pollutants in common minke whales, sei whales,
Bryde’s whales and sperm whales, their spatial and temporalvariation, and, in the case of Hg, their relationship to those in
prey. The Panel commends the effort undertaken both in thefield and in the laboratory. The results are valuable although,as detailed below, in a number of aspects they require
improvement and adequate ecological interpretation as well
as more in-depth assessment of the risk they represent towhale populations.
8.4.2 Evaluation of progress with the 2009 recommendations
The Panel acknowledges the progress made since the
mid-term review (IWC, 2010) and the substantial work
undertaken to take into account the 2009 recommendations.
Of 20 recommendations or suggestions, 13 had beenaddressed or partly addressed and 5 had not been addressed(see Table 9). However, from a general perspective it
considers that, although substantial effort has been put into
the field and the laboratory work, the statistical analyses aswell as the physiological and ecological interpretation of theresults requires improvement.
For example, the original statistical analyses presented
in some papers were clearly incorrect. The results of thelinear models presented in papers SC/F16/JR27, SC/
F16/JR30 and SC/F16/JR31 indicated that they had been
incorrectly identified and fitted, leading to coefficients thatwere either identical, very large with high standard errors or
equal to zero; also, the categorical variables were in somecases not correctly specified. These analyses were then respecified during the Workshop to include an intercept term
and the models were re-fitted to the data during the meeting.
However, the mid-term review recommendation for the useof generalised additive models to investigate non-linear
patterns in the Hg data has not yet been carried out.
The 2009 Panel had also recommended presenting OC
concentrations calculated on the basis of the extractable
lipid content of the sample. The present Panel was
pleased that the OC concentrations reported in paper
SC/F16/JR32 were given as lipid weight but noted that
others (e.g. SC/F16/JR31) still presented concentrations

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calculated on a fresh tissue basis, making it difficult toreconcile total concentrations and thus to compare thefindings of the temporal trends with the results from thepublished literature. Whilst the proponents suggested thatthe lipid content in the blubber of the sampled animals
was broadly similar and therefore wet weight comparisons
would be appropriate, it has been well demonstrated thatblubber lipid content, even in cetaceans in relatively good
body condition, can vary between individuals and evenminor variations among individuals are likely to influence
the OC results. Similarly, Hg concentrations should be more
appropriately calculated on the basis of the dry weight ofthe sample (dry weight basis) to allow comparison withpublished studies in other regions.
Age is known to be a critical variable when interpretingthe tissue concentrations of most OC and Hg (Aguilar etal., 1999). Following one of the 2009 recommendations,
SC/F16/JR33 examined Hg concentrations through amultivariate analysis that included an index of age as acovariate. However, other papers continued to use bodylength as a covariate. This was because difficulties in ageing
whales are only recently being overcome and thus age was
not available for many of the sampled individuals at the timeof writing the manuscripts. This is a serious impediment for
the understanding of the dynamics of pollutants in whales,
and the Panel stresses the value of including age in all
statistical analyses as soon as this becomes available for anyof the existing studied individuals.
A more general 2009 recommendation had related tothe integration of results. The present Panel reiterates thisand considered that some of the results and analyses could
have been combined to give a more comprehensive, overall
picture. For example, the analysis of Hg levels in muscle
and liver samples were reported in two separate papers. The
findings from the temporal trends paper could have beencombined with the findings from the minke whale O andJ coastal stock comparison study into a single paper. This
would have allowed the latter to be more readily and easilyinterpreted within the context of the former.
The patterns of accumulation of the various OC classes
and the differences among the different cetacean species
were explored as recommended by the 2009 Panel, but thestatistical analyses carried out were limited in scope; morequestions could have been answered and more potential
patterns explored using more in-depth approaches. A
principal component analysis was included in paper SC/
F16/JR33 but the objective for applying this particularly
statistical technique (and with a limited sample size) was
not clearly stated. Linear models were used to determine theimportant factors affecting pollutant concentrations but, as
stated above, the most important confounder of age was inmost cases not included.
The bioaccumulation of pollutants in the cetaceans andtheir food items was also considered in papers SC/F16/
JR30-34, as recommended in 2009 (IWC, 2010), but thePanel recognises that these studies were severely hamperedby the loss of samples caused by the 2011 earthquake.
However, some central aspects of bioaccumulation were
not addressed. For example, neither the effect of trophic level
on OC or Hg patterns of accumulation, nor the calculationof bioconcentration factors, were reported. This could havebeen done through the integration into the analysis of stable
isotope values, but despite the 2009 recommendation, thesewere not determined. Papers SC/F16/JR30 and SC/F16/
JR31 attempted to assess temporal trends, but the statisticalanalyses conducted only investigated linear declines orincreases in contaminants which may fail if the variation
over time was not linear. Generalised additive models(GAMs) would be more adequate to address this issue.
The relationship between chemical pollutants andcetacean health was only marginally addressed. SC/F16/
JR35 did refer to the potential risk to cetacean health (e.g.
an increase in risk of the development of thyroid cancer)
following exposure to radioisotopes, and SC/F16/JR36(Niimi et al., 2014) reported on cytochrome P450 transcript
induction in relation to the concentration of specific PCB
congeners in liver samples from minke whales. However, theexpectation of the 2009 Panel was that an assessment of the
impact on the health of the cetaceans and their populations
would be included in the Hg and OC exposure papers SC/
F16/JR30-34 as well. There are many studies and threshold
estimates for the effects on cetaceans of PCBs in particular,
e.g. ~20ppm in blubber is a good ‘rule of thumb’ based on
Kannan et al. (2000). However, there was little attempt toassess the risk to the populations or to discuss whether thepollutant levels were high or low in comparison to elsewhere
and to the estimated thresholds for effects in cetaceans orlaboratory animal models (Hall et al., 2006; Schwacke et al.,
2002; Wells et al., 2005).
In summary, the Panel concludes that the proponents had
addressed the objectives for the pollution studies but thatmany more questions could be answered using these data
if the better statistical analyses and a more comprehensive
physiological and ecological interpretation were conducted.
Overall the findings (with the exception of the two papers
published in the primary literature) were not discussed indetail or in context, particularly with reference to the manyother published studies relating to the bioaccumulation andimpact of PCBs and Hg on cetaceans at both the individual
and population levels.
8.4.3 Recommendations

8.4.3.1 MEDIUM-TERM

The Panel has developed the following recommendations
for the medium-term i.e. normally to be completed 2-3 years
after the 2016 Annual Scientific Committee Meeting.
(1) Since body length is a poor proxy for age, particularly insexually mature whales, age data should be incorporated
into the multivariate analysis of pollutant concentrations
as soon as they become available.
(2) Stable isotope values should be included in the analyses
to investigate the bioaccumulation process of pollutants
through the food chain.
(3) The risk that these chemical pollutants present to the
populations’ abundance or distribution should be more
widely assessed.
8.4.3.2 SHORT-TERM

The Panel has developed the following recommendations
for the short-term i.e. ideally to be completed in revised
papers to the 2016 Annual Scientific Committee Meeting.
(1) The statistical analyses should be improved, based on
clear, well formulated hypotheses.
(2) The OC concentrations now presented on a fresh weightbasis should be recalculated as values on a lipid weightbasis, and the Hg concentrations now on a fresh weightbasis should be recalculated a those on a dry weightbasis. This can be done straightforwardly throughthe lipid content and the dry weight of the samples,
variables that are always produced during the analyticalprocessing of the tissue.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017(3) Trends in pollutant concentrations should be exploredusing generalised additive models (GAMs) or othernon-linear approaches, in addition to the linear models.
(4) The pollutant concentrations found should be evaluatedin comparison with data from previous studies
conducted in comparable species and available in theliterature.
9. REVIEW OF OTHER CONTRIBUTIONS TO
IMPORTANT RESEARCH NEEDS
9.1 Proponents’ summaries

9.1.1 Age determination
SC/F16/JR52 (Maeda et al., 2013), SC/F16/JR53 and SC/F16/
JR55 presented basic information on earplugs and age readingfor common minke and sei whales sampled by the JARPN
and JARPN II. Age reading using earplugs of common minke
whales are generally believed to be difficult and impracticalbecause of their softness and poor formation of growth layers.
However, under JARPN and JARPN II surveys, all earplugs
were carefully collected and tried to read growth layers. Further,in 2007, new sampling techniques (Gelatinised extraction
method) was developed to prevent damage of earplugs at thecollection stage (Maeda et al., 2013) [SC/F16/JR52]. As aresult, age readability of this species could be improved from
8.7% in the past commercial whaling to 44.1% (45.2% for male,and 41.2% for female) in the JARPN and JARPN II surveys.
Inter reader calibration experiment indicated the age reading
outcomes of two readers appeared similar with no substantial
differences. Based on these results, earplug of common minke
whales in the western North Pacific was revealed as a valid
ageing tool (SC/F16/JR53). Age reading of other species suchas sei whale is ongoing (SC/F16/JR55). Using these age data,
further studies on life history, stocks and population dynamics
of the whales will be conducted in the near future.
9.1.2 Genetics
Sasaki et al. (2005) determined the complete mtDNA sequencesof 10 extant Mysticeti species, inferred their phylogenetic
relationships, and estimated node divergence times. The analysis
concurred with previous molecular studies in the ordering of
the principal branches, with Balaenidae as sister to all othermysticetes base. The analysis also suggested that four lineages
exist within the clade of Eschrichtiidae+Balaenopteridae,
including a sister relationship between the humpback and fin
whales, and a monophyletic group formed by the blue, sei, andBryde’s whales, each of which represents a newly recognised
phylogenetic relationship in Mysticeti.
Nikaido et al. (2005) studied the phylogenetic relationships
of baleen whales comprising 11 extant species based on 36
informative SINE loci. One of the intriguing conclusions wasthat balaenopteridae and eschrichtiids radiated very rapidly
during a very short evolutionary period. During this period,
speciation occurred in balaenopterids and eschrichtiids while
new inserted SINE loci remain polymorphic.
Pastene et al. (2007) tested the hypothesis that elevatedocean-surface temperatures can facilitate allopatry among
pelagic populations and thus promote speciation. Thishypothesis was tested by genetic analyses of populations of
Antarctic and common minke whale. The study suggested
that the two species diverged in the Southern Hemisphere lessthan 5Ma. This estimate places the speciation event during aperiod of extended global warming in the Pliocene. Different
populations of common minke whales likely diverged after the
Pliocene some 1.5Ma when global temperatures had decreased.
Pastene et al. (2010) investigated population structure andpossible migratory links of common minke whales in westernSouth Atlantic and western South Pacific using mtDNA.
Whales from these two oceanic basins were phylogeneticallydistinct, and the genetic distance between them was similar tothat between North Pacific and North Atlantic common minke
whales.

563

Glover et al. (2010) based upon analyses of mtDNA andmicrosatellites, documented the case of a single Antarctic
minke whale in 1996, and a hybrid with maternal contribution
from Antarctic minke whale in 2007 in the Arctic Northeast
Atlantic. This was the first documentation of Antarctic minke
whales north of the tropics, and, the first documentation of
hybridisation between minke whale species.
Glover et al. (2013) investigated the genetic ancestry of apregnant female minke whale captured in the North Atlantic in
2010, and her foetus, using data from mtDNA, microsatellite
and sex determining marker. The analyses demonstrated thatthe mother was a hybrid displaying maternal and paternal
contribution from North Atlantic common and Antarctic minke
whales, respectively. Her female foetus displayed greater
genetic similarity to North Atlantic common minke whales
than herself, strongly suggesting that the hybrid mother had
paired with a North Atlantic minke whale.
9.1.3 Physiology
Birukawa et al. (2008) examined kidneys of baleen and sperm
whales to test the hypothesis that cetaceans have unique actionsof UTs (urea transporters) to maintain fluid homeostasis inmarine habitat. Two protein kinase C consensus sites arepresent in the baleen whale UT-A2s, however, a single protein
kinase C consensus site was identified in the sperm whale UTA2. These different phosphorylation sites of whale UT-A2s
may result in the high concentrations of urinary urea in whales,by reflecting their urea permeability.
9.1.4 Reproductive biology
Bhuiyan et al. (2008) determined the cumulus-oocytecomplexes (COCs) recovery rates with respect to reproductive
status per sei and Bryde’s whales. The study indicated thatin vitro fertilised (IVF) in sei whales is possible to achievecleaved embryos developing to morula stage. This was the firstin vitro embryo attempt in sei and Bryde’s whales.
Hiwasa et al. (2009) investigated effects of threesemen extenders and storage temperatures on post-thaw
characteristics of Bryde’s whale spermatozoa. The study
showed that a synthetic semen extender, AndroMed, could be
used for cryopreservation of whale spermatozoa in addition
to Tris-based extenders containing bovine serum albumin or
egg yolk. Storage of the post-thaw Bryde’s whale spermatozoa
was better at 5°C than room temperature or 35°C. The frozenthawed Bryde’s whale spermatozoa maintained their motility
and viability for at least two days at room temperature and for
four days at 5C.
Lee et al. (2009) examined the feasibility of using subzonal
cell injection with electrofusion for interspecies somatic cell
nuclear transfer (iSCNT) to produce sei whale embryos andto improve their developmental capacity by investigating
the effect of osmolarity and macromolecules in the culture
medium on the in vitro developmental capacity. The results
demonstrated that sei whale-porcine hybrid embryos may be
produced by SCNT using subzonal injection and electrofusion.
Bhuiyan et al. (2010) investigated an effective embryo
reconstruction method and an effective post-activation agent
for in vitro production of sei whale interspecies somatic cell
nuclear transfer (iSCNT) embryos. The study concluded thatbovine oocytes have the ability to support development of sei
whale nuclei up to the 6-cell stage.
Suzuki et al. (2010) investigated whether the equilibration
steps (three or five steps), whale follicular fluid (WFF) addition
and type of sugars (sucrose or trehalose) were effective for theviability and in vitro maturation (IVM) of vitrified immature
oocytes in sei, Bryde’s and common minke whales. The tested
step number of cryoprotectan equilibration, WFF addition
and type of sugars did not improve the maturation rate ofvitrified baleen whale oocytes. However, the study showedthat immature oocytes derived from three baleen whale species
in the western North Pacific could be vitrified and matured invitro at about 30% levels.

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Inoue et al. (2014) examined seasonal changes in thetestis of the common minke whale in the North Pacific from
April to October. Results suggested that the spermatogenetic
activity of the common minke whale has seasonal changes,
namely, it reduce from May to June and is activated in Augustin preparation for the next breeding season.
Kitayama et al. (2015) examined the structure and functionof placentas in common minke, Bryde’s and sei whales with theaim of confirming the structural characteristics of the chorion,
including the presence of the areolar part, and clarifying
steroidogenic activities and foetomaternal interactions in
the placentas of these whales. The study suggested that, in
cetaceans, uteroferrin is used to supply iron to the foetus, andthat trophoblast cells synthesise oestrogen in whale placentas.
The study immunohistochemically revealed the localisationof aromatase and uteroferrin in cetacean placentas during
pregnancy for the first time.
9.1.5 Morphology
Nakamura et al. (2012) investigated the allometric growth
pattern of common minke whales from the North Pacific bycomparing skull length and skull width with body length. The
skull proportion of large Balaenoptera whales (blue and fin
whales) showed positive allometry, but that of the common
minke whale showed negative allometry, despite being a related
species. Such differences in intraspecific growth patterns could
be the result of adaptation driven by feeding strategy.

9.2 Panel conclusions and recommendations
The Panel welcomes the provision of the information on
additional studies undertaken in addition to those envisionedby the original objectives of the programme. This is inaccord with the recommendations of previous Panels thatevery effort be made to maximise the information obtained
from whales that are caught.
The Panel was particularly impressed by the work
undertaken to improve ageing techniques for baleen whales.
In particular it commended the progress in the developmentof the gelatinised extraction method and encourages its
continued development. Age is a critical variable for manyof the aspects of the JARPN II programme. In the case of
the common minke whale, the low readability of earplugs,
which is considered the most reliable source of absolute age
determination in baleen whales at present, was a difficultythat now appears to have been in part overcome. Thus, with
the new technique, the readability of earplugs increased
from a previous 8.7% of individuals sampled to 44.1%. The
Panel recommends investigation into whether there is any
relationship between age or sex and readability that may
affect the representativeness of the earplugs that can be read.
The addition of absolute age data has great potential
to improve analyses relevant to all three of the primaryobjectives. The Panel encourages the proponents to age as
many of the existing samples as possible and to incorporate
age where appropriate in updated analyses (e.g. see therecommendations on pollutant studies, Item 8.4.3).
10. INTEGRATION OF RESULTS
10.1 Proponents’ summary

Previous review workshops of JARPN II and JARPA II haverecommended integration among the research components ofthese whale research programs. Such integration is importantto respond appropriately the scientific questions raised in the
programs as well to identify research needs for the future. Twokinds of integrative analyses were identified in JARPN II: (i)analyses integrating similar data from different sources; and (ii)analyses integrating different study fields/data on a particularresearch topic (these study/data can be from the same or fromdifferent sources).

The proponents identified a total 26 integrative analyses of
the two kinds, with most of them already achieved in JARPN
II. These integrations were important to respond a total of 17
scientific questions related to the main three research objectivesof JARPN II. Examples are shown below.
One example of the first kind (i) was the integration of thesame DNA data from different sources (e.g. DNA from Korean
and Japanese by-catches, JARPN and JARPN II) in the case of
the common minke whale. Such integration allowed the genetic
analyses on stock structure to be conducted in a much large
temporal and geographical scales. One example of the second
kind (ii) was the integration of different analytical approaches(e.g. genetics, morphometric, discriminant analysis of principal
components, age distribution, geographical distribution ofrelatives) to examine stock structure in the case of the common
minke whale. Such integration allowed the postulation of
hypotheses on stock structure with larger plausibility. Bothintegrations, summed to appropriate responses to previous
recommendations, allowed the attainment of sub-objectives
related to common minke whale within main objective 3.
Another example of the second kind (ii) was the ecosystemmodelling exercise which integrated abundance information ofwhales, prey consumption by whales, oceanographic data andbiomass of prey. Such integration, together with appropriate
responses to previous recommendations, allowed a substantial
improvement in the models. They would serve as baselinemodels to test various types of marine ecosystem management
options.

10.2 Panel comments and conclusions
The Panel recognised at least three levels of integration:(1) within sub-objectives;(2) within objectives; and(3) amongst the JARPN II objectives as a whole.
The Panel agreed that the proponents had attempted
to undertake some integration within (1) and (2) but that
there was relatively little integration at stage (3) – either
using JARPN II data alone or using additional data andinformation from other sources. For example, under (1)within the stock structure sub-objectives for common minke
whales, the proponents had undertaken analyses using
genetic data of various kinds (including genetic data from
Korea outside the JARPN II samples and morphological dataand in some cases age data and limited telemetry data) andadditional analyses were proposed. The proponents had also
undertaken spatial modelling using additional exploratory
variables collected outside as well as within JARPN II to
improve the limited analyses from 2009; again further work
has been recommended on this.
Under level (2), the feeding ecology work had begun
to incorporate information from a variety of sources within
and outside the JARPN II programme including abundanceof prey and cetaceans, oceanography in addition to theinformation on prey consumption, selectivity and diet.
However, the pollution component had only integrated
across sub-objectives in a limited way and several of the
Panel’s recommendations are focussed on improving this
aspect of the work.
There has been relatively little work that integrates
amongst the JARPN II objectives and the most obvious
areas might be to incorporate the stock structure hypotheses
into the ecosystem modelling work.
Finally, the Panel agrees that there is a general lack ofintegration amongst the papers addressing the same topic –this made it difficult for the Panel to evaluate many aspectsof the programme including the level of integration. This isconsidered further under Item 11.1.2.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 201711. PANEL CONCLUSIONS WITH RESPECT TO
ANNEX P
11.1 General issues
Before considering the specific items of the review outlined
in Annex P, the Panel draws attention to a number ofimportant general matters that have affected its review.
11.1.1 Timing
As explained in SC/F16/JR54, the closing of the JARPN
II programme reflected a political decision related to theGovernment of Japan’s response to the International Court ofJustice decision regarding JARPA II, rather than a scientificevaluation that the JARPN II programme had attained its
objectives or sub-objectives. In fact, this ‘final review’ of
JARPN II is occurring before the formal completion of the
programme in 2016, although the sample sizes and priorities
for the period 2014-16 were revised by the Government ofJapan (SC/F16/JR54).
Annex P envisions final reviews taking place within
three years of the finish of a programme to reflect the factthat sufficient time needs to be given to the proponents todevelop a comprehensive and integrated final report. It isclear from the discussion and recommendations above that
despite the hard work of the scientists, resulting in a large
number of working papers, that the analyses would havebenefitted from considerably more time. Similarly, more time
would have enabled the scientists to produce an integrated
final report. The Panel recommends that the ScientificCommittee considers including a guideline in Annex P
either relating to the minimum time after completion of aprogramme that a final review can take place or establishinga small review group to determine whether the materials
presented for a final review are in a sufficient state for aworkshop to take place (this may also be worth considering
for new and periodic reviews).
In addition, the fact that: (a) the programme wascompleted early for political rather than scientific reasons;and (b) there were no formal intermediate targets by timeline,
meant that it was difficult for the Panel to properly assess theresults of the programme against the original objectives.
11.1.2 The nature of ‘final reports’Annex P does not provide guidelines for the scope andstructure of final reports. However, the Panel’s experiencein undertaking this review shows that formal guidance isnecessary. The Panel recommends that Annex P should be
revised to include such guidelines and offers the following
comments to assist in that process.
The Panel’s task was made considerably more difficult
because the methods, analyses and conclusions were foundwithin a very large number of documents of varying levelsof completeness and quality. The Panel also noted that somedocuments (e.g. SC/F16/JR54 and part of section 4.3 of SC/
F16/JR01) included information or discussion beyond theterms of reference for this final scientific review. Although
the proponents produced a good brief overall summary
document (SC/F16/JR01), it contained insufficient detail
to allow a proper review and details of sampling design,
strategy, field protocols, analytical methods and conclusions.
For this, the Panel members had not only to examine over 90
working papers and documents, but also references to otherunpublished sources (e.g. IWC papers) over the JARPN II
period. This lack of integration, at least by objective, appears
to be a function of the timing of the review (see Item 11.1.1)
but it is not an efficient way to work and can make it rather
difficult for the Panel (and especially members from outside565

the IWC system) to conduct a thorough review. A suggested
outline for an integrated final report (and associatedmaterials) is provided as Annex G.
11.1.3 Lethal and non-lethal techniques
Although formally outside the scope of this review whose
focus is on the period up to 2013, as discussed under Item
3.4, Japan has modified and reprioritised the JARPN II
programme until it is officially completed in 2016. One
aspect of this relates to the addition of an objective tocompare lethal and non-lethal techniques was in line with therecommendation from the 2009. This topic is central to many
issues raised in Annex P for reviews of new and ongoing
permits and the difficulties in addressing the issue have beenraised by all of the expert Panels thus far. In this light, thePanel highlights the second part of the recommendation
given under Item 3.4.2.2 that the proponents provide a single
document to the 2016 Annual Meeting that provides the field
and analytical protocols for the comparison of using lethal
and non-lethal techniques for each key parameter taking into
account the advice provided in 2009.
11.1.4 Review of progress of recommendations
As illustrated above, one important component of thisreview was an examination of the response of proponents
to the recommendations of the 2009 review. In addition, thePanel also notes that the 2009 Panel had stated that given
the extra work it had requested of the proponents on certain
key matters (including with respect to assessing the effectsof catches on some of the stocks) it had not been able tocomplete its review. The 2009 Panel had requested theScientific Committee to consider ‘the most appropriate waythat this review is completed’.
The Panel recognises that the Scientific Committee
has agreed that it is not necessary to review in detail the
results of ongoing permits every year. However, it believes
that the regular and final reviews (and potentially reviewsof new permit proposals) would be facilitated by a short(just a paragraph or two) biennial update by proponents as
to progress with each of the recommendations after theirinitial response in the Annual Meeting following the review
Workshop; this should also benefit the proponents’ work.
The Panel recommends that the Scientific Committee
should consider a mechanism (e.g. revision to Annex P)
to provide for such a brief annual review of progress withrecommendations. It also reiterates the request of the 2009
Panel that the Scientific Committee develops a mechanism
to allow for the completion of expert Panel reviews if a Panelstates that its review is incomplete until further information/
analyses is provided.
11.2 Assessment of the programme’s scientific output
given the stated objectives and length of the programme
The Panel refers to its earlier comments regarding the timingof the final review and the reasons for the timing of theclose of the programme; this also affects to some extent its
ability to assess the programme’s scientific output given thestated objectives and length of the programme. As shownin Annex H, the JARPN II programme thus far has resulted
in 31 published papers related to the primary objectives ofthe programme and another 30 published papers that arosefrom additional studies not related to the primary objectives.
In addition, the JARPN II programme has resulted in alarge number of papers to the IWC Scientific Committeethat made important contributions to the work on RMP
Implementations and in-depth assessments. It is clear fromthe review that: (a) considerable scientific work has been

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Table 10
Overview of how well the proponents have met their stated sub-objectives within the overall objectives of JARPN II.

Objective/Sub-objective
Objective 1: Feeding ecology and ecosystem studies
Sub-objective 1.1: Investigate the oceanographic conditions that are
relevant for the understanding of prey species’ distribution andabundance in the research area.

Panelevaluation Comments
PartialAlthough some work has been done, additional work is needed to investigate more
appropriate explanatory variables (see Item 5.4).

Sub-objective 1.2: To investigate the distribution pattern of baleen
whales in the research area and the possible factors affecting such
pattern.

Good

Good progress has been made with this sub-objective in what is a developing fieldof spatial and habitat modelling. However, more work is required to try to integrate
the information from different seasons and other surveys within and outside theresearch area (see Item 5.4.2).

Sub-objective 1.3: To estimate abundance of baleen and sperm whales
using JARPN II sighting data and standard IWC SC methodology.

Very
good

Abundance estimates were presented using design-based methods. Effort now
needs to be put into exploring methods for determining trends and comparison withmodel-based estimates.

Sub-objective 1.4: To estimate the prey consumption by baleen whales
using JARPN II data and samples, and taking into account theuncertainties identified at the 2009 JARPN II review.

Good

Good progress was made with incorporating many aspects of the uncertainty
identified in 2009, although some additional sources were identified (see Table 6)
and improved methods to quantify the uncertainty have been recommended (seeItem 6.4.2). The potential impact of sampling design requires evaluation (see Item
3.4).

Sub-objective 1.5: To evaluate the feeding impact by whales on Progress Some progress has been made but the problems with model development (see subfisheries resources using JARPN II data and samples, and information made objective 1.10 in this table) and aspects of uncertainty mean that the proponents arefrom commercial fisheries and other research sources in coastal areas.
not able to identify the feeding impact by whales in a robust way (see Item 6.4.2).
Sub-objective 1.6: To estimate prey abundance using JARPN II data, Sufficient This work has been achieved, at least to inform initial modelling efforts. Additional
complemented with information available from other sources.
work to estimate the uncertainty of extrapolating prey abundance outside thesurveyed blocks/seasons would be useful (see Item 6.4).
Sub-objective 1.7: To investigate the prey preference of whales in Progress Prey preference studies have been undertaken based upon stomach content data andoffshore areas, using JARPN II data and samples.
made prey abundance information but further work is required to address issues ofseasonality, uncertainty and sample design.
Sub-objective 1.8: To investigate feeding habits of baleen and toothed Progress Some work was completed on trends in prey by species and feeding differences by
whale species in the research area, and the environmental factors made habitat but additional analyses are required before firm conclusions can be reached.
involved in determining such habits.
Work began using time depth recorders but sample size is small.
Sub-objective 1.9: To investigate the yearly trend in body condition ofbaleen whales using JARPN II data and samples.

Partial

In addition to the need analyse to further examine power, the question of sampling
design also needs to be addressed.

Sub-objective 1.10: To develop several ecosystem models, in both Progress
coastal and offshore areas, using JARPN II data and samples as input. madeOutput of the models are likely to provide information on: (i) the
ecosystem structure; (ii) effects of prey availability and consumption
on the population dynamics of common minke and sei whales withconsideration of levels of energy intakes; and (iii) predation impacts ofcommon minke whales consumption on sandlance stock off Sanriku.

Although progress has been made in some areas, insufficient resources have been
allocated to this component of the programme. Although two models have beendeveloped they are preliminary and a planned minimum realistic model is notcomplete. As such the modelling efforts are not suitable to provide management
advice or characterise effects of prey on whale dynamics or impacts of whales on
fisheries (see Item 7.4).

Objective 2: Monitoring environmental pollutants in cetaceans and the marine ecosystem
Sub-objective 2.1: To investigate pattern of accumulation of pollutants Partial Aspects of this issue have been addressed and the Panel recognised the difficulties
in cetaceans and their food items.
caused by the loss of samples in the tsunami. However, some central aspects were
not addressed or analyses were incomplete as discussed under Item 8.4.
Sub-objective 2.2: To investigate the bioaccumulation process ofNot
This was not properly addressed and would require inter alia integration with stablepollutants through the food chain.
achieved isotope analyses (see Item 8.4).
Sub-objective 2.3: To investigate the relationship between chemical
pollutants and cetacean health.

PartialSome work was presented (e.g. regarding thyroid cancer and CYP450 induction)
but there was little attempt to use comparative studies and consider possible
population level effects.

Good

This work was thorough and contributed to the RMP Implementation Review.

Sub-objective 3.2: Using genetic and non-genetic data from JARPN
and JARPNII, investigate whether or not the sub-division of the O stock
into OW and OE is plausible. The genetic analysis should include those
approaches mentioned in Table 1 as providing support for the existenceof the OW (e.g. PCA analyses).

Good

This work was thorough and contributed to the RMP Implementation Review.

Sub-objective 3.3: To investigate the plausibility of: (i) stock subdivision within Sub-area 1 as proposed under Hypothesis 4; and (ii)
sub-division between Sub-areas 1 and 2 as proposed under Hypotheses
2 and 3, using all genetic samples available from different source till
2014, and different genetic markers included satellite tracking.

PartialThis work will contribute to the forthcoming RMP Implementation Review but
additional analyses are recommended to assist in understanding the power of theresults obtained and the telemetry programme, whilst showing that it is possible,
has as yet only a very small sample size (2).

Sub-objective 3.4: To investigate the plausibility of a single stock of
sei whale in the pelagic regions of the North Pacific (‘North Pacificpelagic’), using all genetic samples available from different sources till
2014, and different genetic markers.

PartialThis work will contribute to the forthcoming in-depth assessment but additional
analyses are recommended to assist in understanding the power of the results
obtained, although it is recognised that past experience may show that the power is
low.

Objective 3: Stock structure of large whales
Sub-objective 3.1: Monitoring of the spatial and temporal distributionof J stock on both west and east coasts of Japan using genetics and nongenetics approaches, and all sources of samples available e.g. JARPN,
JARPN II and by-catches.

7

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

undertaken and that the output has been accepted in peerreviewed journals and has influenced the work of the IWC
Scientific Committee; but also that (b) a much greater
emphasis should have been put on improved analyses andmodelling – that would increase considerably the value ofthe scientific output of the existing data collected. The Panel
therefore strongly encourages the proponents to follow therecommendations provided in this report and submit further
work to peer-reviewed scientific journals.
11.3 Consideration of the level of co-ordination withother relevant research projects
The Panel welcomes the much-improved collaboration withother research projects compared to that in 2009. It noted thatmost of that co-operation occurred within Japanese institutes(academic and governmental). This is perhaps not surprising
for the coastal components which are within Japanese waters
but it encourages additional co-operation with scientists
from other research projects that address similar issues but
for other regions with respect to any further analyses that are
to be undertaken.
11.4 Evaluation of how well the stated objectives havebeen met and the extent to which the results have
improved conservation and management
11.3.1 Evaluation by the most recent sub-objectives
The Panels’ view of how well the recently developed subobjectives have been met is given in Table 10. The overall
Panel evaluation of the work presented against the original
objectives, and comments on the extent to which the work
has contributed to conservation and management is provided
in the text below (Item 11.3.2) by objective.
11.3.2 Evaluation by original objectives with comments oncontributions to conservation and management
The effect of the conclusion of the programme for politicalreasons on the ability to meet objectives was discussed
under Item 11.1. The 2009 Panel had ‘severely questioned’the scientific value of the sperm whale component of the
programme (IWC, 2010, p.433) for reasons of sample size;
the present Panel agrees that the sperm whale component of
JARPN II has produced little of scientific value as expected.
OBJECTIVE 1: FEEDING ECOLOGY AND ECOSYSTEMSTUDIESThe ultimate goal of this objective was to provide
multispecies management advice. As noted by the 2009
Panel, this was an extremely ambitious task and one likely
to take many years. The level of field and laboratory work
has been impressive and the examination of uncertainty
with respect to the prey consumption and prey preferences
has been greatly improved since 2009 although analyticalimprovements can still be made. However, the question of theeffects of sampling design (see Item 3.4.2) requires further
consideration and, primarily as a result of a lack of allocatedresources (despite the 2009 Panel recommendation), themodelling work remains preliminary.
Even allowing for the complexity of the issue, there
are examples of MRM/MICE1 models that that can beparameterised by fitting to data which are used to provide
input to tactical assessment models and there are better
developed food web and extended single species models;with additional resources, progress could (and should) havebeen made in the development of intermediate model types.
The Panel concludes that at this stage of development, theMinimum Realistic Models/Models of Intermediate Complexity for Ecosystem Assessment.

1

567

modelling results are not suitable for addressing strategic
management questions2. At present, at least, the results
have not led to improved conservation and managementof cetaceans or of other marine living resources or the
ecosystem.
OBJECTIVE 2: MONITORING ENVIRONMENTAL POLLUTANTS IN CETACEANS AND THE MARINE ECOSYSTEM

This objective related to monitoring pollutants in theenvironment and cetaceans including: (a) pattern ofaccumulation in cetaceans; (b) bioaccumulation throughthe food chain; and (c) the relationship between pollutants
and cetacean health. The Panel notes that the achievementof this objective was hampered considerably by the loss ofsamples as a result of the tsunami. It also acknowledges theefforts made to follow the recommendations of the 2009
Panel. The level of field and laboratory work has been goodand understanding of chemical pollutants and cetaceans
off Japan has been greatly improved. However, the Panelconcludes that only partial progress has been made towards
addressing the objectives and more effort needs to be put
on improved analyses and interpretation of results (seediscussion and recommendations under Item 8.4). Thisis especially true in terms of the relationship of pollutants
and cetacean health, which is most relevant to improved
conservation and management of cetaceans. It is not clear
from the papers presented if (and if so how) the work
undertaken has contributed to the conservation of othermarine resources or the ecosystem.
OBJECTIVE 3: STOCK STRUCTURE OF LARGE WHALESThe broad objectives simply related to the stock structure oflarge whales (common minke whales, sei whales, Bryde’s
whales and sperm whales), although this was clarified at the2009 Panel workshop to be primarily related to developingor narrowing the number of hypotheses to be consideredby the IWC Scientific Committee in its work related to the
RMP and in-depth assessments. The level of field, laboratory
and analytical work has been impressive, as was the effort
put into responding to the 2009 Panel recommendations.
The Panel did make some recommendations for improved
analyses, particularly related to power and the ability to
distinguish amongst weakly-differentiated populations.
The Panel concludes that the stock structure component of
JARPN II has made, and will continue to make, important
contributions to the conservation and management ofcetaceans by providing fundamental data and analyses for the
RMP Implementation Reviews of common minke whales andBryde’s whales, and the in-depth assessment of sei whales.
12. ADOPTION OF REPORTThe report was adopted by email on 11 March 2016. The Chair
thanked all members of the Panel for their patience, tirelessdedication, and for having served the Scientific Committeewith their undoubted competence. She was grateful tothem for having donated their time to this activity, which
is not a trivial matter, and for having allowed a scientificand friendly discussion during the entire process (open andclosed sessions, and email exchanges at impossible hours).
The Chair also thanked the Proponents for their kindness
and logistical support.
Ecosystem models such as Ecopath with Ecosim, Atlantis, and other large
complex models which are difficult to parameterise by fitting to data are notsuitable for tactical management anywhere in the world at present and probably far into the future. Single species models with predation and multispecies (MICE) models could be used to provide tactical advice in the future.

2

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Van der Zee, J.P. and Punt, A.E. 2014. Evaluting ciritical dispersal rates forwhale management under the IWCs Revised Management Procedure: anapplication for North Atlantic common minke whales. J. Cetacean Res.
Manage 14: 125-32.
Watanabe, H., Okazaki, M., Tamura, T., Konishi, K., Inagake, D., Bando,
T., Kiwada, H. and Miyashita, T. 2012. Habitat and prey selection ofcommon minke, sei and Bryde’s whales in mesoscale during summer inthe subarctic and transition regions of the western North Pacific. Fish.
Sci. 78: 557-67.
Wells, R.S., Tornero, V., Borrell, A., Aguilar, A., Rowles, T.K., Rhinehart,
H.L., Hofmann, S., Jarman, W.M., Hohn, A.A. and Sweeney, J.C. 2005.
Integrating life history and reproductive success data to examine potential
relationships with organochlorine contaminants for bottlenose dolphins(Tursiops truncatus) in Sarasota Bay, Florida. Sci. Total Env. 349(1-3):
106-19.
Winship, A.J. and Trites, A.W. 2003. Prey consumption of Steller sea lions(Eumetopias jubatus) off Alaska: how much prey do they require? Fish.
Bull. 101: 147-63.
Witteveen, B.H., Worthy, G.A., Wynne, K.M. and Roth, J.D. 2009.
Population structure of North Pacific humpback whales on their feeding
grounds revealed by stable carbon and nitrogen isotope ratios. Mar. Ecol.
Prog. Ser. 379: 299-310.
Yoshida, N. and Miyazaki, N. 1991. Oxygen isotope correlation of cetaceanbone phosphate with environmental water. Journal of GeophysicalResearch: Oceans 96(C1): 815-20.

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Annex A
List of Participants
PANEL
Alex Aguilar
University of Barcelona
Spain
Greg Donovan
IWC Secretariat
UK
Jaume Forcada
British Antarctic Survey
UK
Caterina Fortuna
National Institute for Environmental
Protection and Research
ItalySarah Gaichas
NOAA Fisheries Service
Northeast Fisheries Science Center
USA
Ailsa Hall
Sea Mammal Research Unit
Scottish Oceans Institute
University of St Andrews
UK
Michael Hammill
Canadian Department of Fisheries
and Oceans
CanadaDaniel Howell
Institute of Marine Research
BergenNorwayTaro Ichii
Oceanic Resources Division
National Research Institute of Far
Seas Fisheries
Japan

Jason Link [by correspondence]
NOAA Fisheries
USA
Per Palsbøll
Faculty of Mathematics and Natural
Sciences
GELIFES-Groningen Institute for
Evolutionary Life Sciences
The NetherlandsAndré Punt
School of Aquatic and Fishery
Sciences College of the Environment
University of Washington
USA

PROPONENTS
Fisheries Agency of Japan (FAJ)
Haruya Kumakiri
Hiroyuki Morita
Hideki Moronuki
Naohito Okazoe
Ministry of Foreign Affairs ofJapan (MOFA)
Tatsuo Hirayama
Sayuri Muraki

Ralph Tiedemann
Institute of Biochemistry and Biology
University of Potsdam
Germany

National Research Institute of Far
Seas Fisheries, Japan (NRIFSF)
Toshiya Kishiro
Hikari Maeda
Tomio Miyashita
Joji Morishita
Hiroto Murase
Hideyoshi Yoshida

Judith E. Zeh
Department of Statistics
University of Washington
USA

National Research Institute ofFisheries Science
Masachika Masujima
Makoto Okazaki

INTERPRETERS
Saemi Baba
Hiroko Yasokawa
OBSERVERS
William K. de la Mare
Australia
Lars Walløe
NorwayDoug S. Butterworth
University of Cape Town
South Africa

Tokyo University of Marine
Science and Technology, Japan(TUMSAT)
Toshihide Kitakado
Institute of Cetacean Research,
Tokyo, Japan (ICR)
Takeharu BandoDan Goodman
Mutsuo Goto
Takashi Hakamada
Kenji Konishi
Luis A. Pastene
Tsutomu Tamura
Genta Yasunaga

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017571

Annex B
Agenda
1.

2.
3.

4.

5.

Introductory items
1.1 Opening remarks
1.2 Appointment of Chair and rapporteurs
1.3 Adoption of Agenda
1.4 Review of available documents
1.5 Structure of the report
Objectives of the Workshop
Short general overview of JARPN II
3.1 Proponents’ overview of general objectives andany changes over time
3.2 Summary of general conclusions andrecommendations from the 2009 Workshop
3.2.1 Objectives
3.2.2 Sampling design, sample size and areas
3.2.3 Data analyses and modelling
3.2.4 Other matters not the focus of a final
review
3.3 Proponents’ overview of broad sampling
scheme including sample size, areas and strategy
including changes over time and response to the2009 Workshop recommendations
3.4 Panel comments and conclusions on generalissues
3.4.1 Objectives
3.4.2 Sampling design and sample size
3.4.2.1 Sampling design and areas
3.4.2.2 Sample size
Stock structure
4.1 Proponents’ summary of objectives including
modifications, if any, since the start of the
programme
4.2 Overview of conclusions and recommendations
from the 2009 Workshop
4.3 Proponents’ summary of the results (incl.
response to 2009 Workshop)
4.4 Panel review, conclusions and recommendations
4.4.1 Overview
4.4.2 Evaluation of progress with the 2009
recommendations (see Annex D for acomplete summary table)
4.4.3 Recommendations
4.4.3.1 Medium-longterm
recommendations
4.4.3.2 Short term recommendations
Feeding ecology and ecosystem studies: Part 1.
General information to inform ecosystem modellingon oceanography, distribution, abundance and statusof whale stocks
5.1 Proponents’ summary of objectives including
modifications, if any, since the start of the
programme
5.2 Overview of conclusions and recommendations
from the 2009 Workshop
5.3 Proponents summary of ‘oceanographic
component’ (incl. response to 2009 Workshop)
5.3.1 Oceanography
5.3.1.1 Proponents summary of theresults (incl. response to 2009
Workshop)
5.3.2 Distribution of large whales

6.

7.

5.3.2.1 Proponents summary of theresults (incl. response to 2009
Workshop)
5.3.3 Abundance and status of stocks
5.3.3.1 Proponents summary of theresults (incl. response to 2009
Workshop)
5.4 Panel review, conclusions and recommendations
5.4.1 Overview
5.4.1.1 Oceanography
5.4.1.2 Distribution
5.4.1.3 Abundance
5.4.2 Evaluation of progress with the 2009
recommendations
5.4.2.1 Oceanography
5.4.2.2 Distribution
5.4.2.3 Abundance
5.4.3 Recommendations
5.4.3.1 Oceanography
5.4.3.2 Distribution
5.4.3.3 Abundance
Feeding ecology and ecosystem studies: Part 2, field
and laboratory studies
6.1 Proponents’ summary of objectives including
modifications, if any, since the start of the
programme
6.2 Overview of conclusions and recommendations
from the 2009 Workshop
6.3 Proponents summary of the coastal and offshore
components (including response to 2009
Workshop)
6.3.1 Prey consumption, including biomass
estimation of prey species
6.3.2 Prey preference, including whales’
feeding habits
6.3.3 Body condition of whales
6.4 Panel review, conclusions and recommendations
6.4.1 Overview
6.4.2 Evaluation of progress made on
recommendations in 2009
6.4.3 Recommendations
6.4.3.1 Medium-term
6.4.3.2 Short-term
Feeding ecology and ecosystem studies: Part 3
ecosystem modelling
7.1 Summary of objectives including modifications,
if any, since the start of the programme (byproponents)
7.2 Overview of conclusions and recommendations
from the 2009 Workshop
7.3 Proponents’ summary of the ecosystemmodelling work (incl. response to 2009
Workshop)
7.4 Panel review, conclusions and recommendations
7.4.1 Overview
7.4.2 Evaluation of progress with the 2009
recommendations (see Annex D for acomplete summary table)
7.4.3 Recommendations
7.4.3.1 General recommendations

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8.

9.

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REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

7.4.3.2 EwE model
7.4.3.3 Extended
single-species
model
Monitoring environmental pollutants in cetaceans
and the marine ecosystem
8.1 Summary of objectives including modifications,
if any, since the start of the programme(proponents)
8.2 Overview of conclusions and recommendations
from the 2009 Workshop
8.3 Proponents summary of the results (incl.
response to 2009 Workshop)
8.4 Panel review, conclusions and recommendations
8.4.1 Overview
8.4.2 Evaluation of progress with the 2009
recommendations
8.4.3.1 Medium-term
8.4.3.2 Short-term
Review of other contributions to important research
needs
9.1 Proponents’ summaries
9.2 Panel conclusions and recommendations
Integration of results

11.

12.

10.1 Proponents’ summary
10.2 Panel comments and conclusions
Panel conclusions wrt Annex P
11.1 General issues
11.1.1 Timing
11.1.2 The nature of ‘final reports’
11.1.3 Lethal and non-lethal techniques
11.1.4 Review of progress of recommendations
11.2 Assessment of the programme’s scientificoutput given the stated objectives and length ofthe programme
11.3 Consideration of the level of co-ordination withother relevant research projects
11.4 Evaluation of how well the stated objectives
have been met and the extent to which the results
have improved conservation and management
11.3.1 Evaluation by the most recent subobjectives
11.3.2 Evaluation by original objectives
with comments on contributions toconservation and management
Adoption of Report

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017573

Annex C
List of Documents
PRIMARY DOCUMENTS
SC/F16/JR
1. Tamura, T., Kishiro, T., Bando, T., Yasunaga, G.,
Murase, H., Kitakado, T. and Pastene, L.A. The Japanese
Whale Research Program under Special Permit in thewestern North Pacific Phase-II (JARPN II): results andconclusions in the context of the three main objectives,and scientific considerations for future research.
2. Matsuoka, K., Hakamada, T. and Miyashita, T.
Methodology and procedure of the dedicated sighting
surveys in JARPN II (2008-2013) - Offshore component.
3. Kishiro, T., Yoshida, H., Yasunaga, G., Bando, T.,
Mogoe, T. and Kato, H. Methodology and survey
procedure under the JARPN II - coastal component ofSanriku and Kushiro - during 2008 to 2014, with special
emphasis on whale sampling procedures.
4. Bando, T., Yasunaga, G., Tamura, T., Matsuoka, K.,
Murase, H., Kishiro, T. and Miyashita, T. Methodology
and survey procedure under the JARPN II - offshore
component - during 2008 to 2014, with special emphasison whale sampling procedures.
5. Okazaki, M., Masujima, M., Murase, H. and Morinaga,
K. Oceanographic conditions in the JARPN II survey
area from 2000 tom 2013 using FRA-ROMS data.
6. Okazaki, M., Masujima, M., Murase, H. and Morinaga,
K. Oceanographic conditions in the survey area of
JARPN II coastal; component off Kushiro in September
from 2000 to 2013 using FRA-ROMS data.
7. Murase, H., Hakamada, T., Sasaki, H., Matsuoka, K. andKitakado, T. Seasonal spatial distributions of common
minke, sei and Bryde’s whales in the JARPN II survey
area from 2002 to 2013.
8. Murase, H., Hakamada, T., Matsuoka, K., Nishiwaki,
S., Inagake, D., Okazaki, M., Tojo, N. and Kitakado,
T. 2014. Distribution of sei whales (Balaenoptera
borealis) in the subarctic-subtropical transition area of
the western North Pacific in relation to oceanic fronts.
Deep Sea Research Part II 107: 22-28. [Publishedpaper].
9. Matsuoka, K., Hakamada, T. and Miyashita, T. 2016.
Distribution of blue (Balaenoptera musculus), fin (B.
physalus), humpback (Megaptera novaeangliae) andnorth Pacific right (Eubalaen japonica) whales in thewestern North Pacific based on JARPN and JARPN II
surveys (1994 to 2014).
10. Sasaki, H., Murase, H., Kiwada, H., Matsuoka, K.,
Mitani, Y. and Saitoh, S.I. 2013. Habitat differentiation
between sei (Balaenoptera borealis) and Bryde’s whales(B. brydei) in the western North Pacific. Fisheries
Oceanography 22(6): 496-508. [Published paper].
11. Hakamada, T., Matsuoka, K., Toshiya, K. and Miyashita,
T. The number of the western North Pacific common
minke whales (Balaenoptera acutorostrata) distributedin JARPN II coastal survey areas.
12. Hakamada, T. and Matsuoka, K. The number of thewestern North Pacific common minke, Bryde’s and sei
whales distributed in JARPN II Offshore survey area.
13. Hakamada, T. and Matsuoka, K. The number of blue,
fin, humpback and North Pacific right whales in thewestern North Pacific in the JARPN II Offshore surveyarea.

14. Hakamada, T. and Matsuoka, K. The number of sperm
whales in the western North Pacific in the JARPN II
Offshore survey area.
15. Tamura, T., Konishi, K. and Isoda, T. Updated estimationof prey consumption by common minke, Bryde’s andsei whales in the western North Pacific.
16. Tamura, T., Murase, H., Sasaki, H. and Kitakado,
T. Preliminary attempt of spatial estimation of prey
consumption by sei whales in the JARPN II survey area
using data obtained from 2002 to 2013.
17. Tamura, T., Kishiro, T., Yoshida, H., Konishi, K.,
Yasunaga, G., Bando, T., Saeki, M., Onodera, K.,
Mitsuhashi, M., Yamashita, Y., Funamoto, T. and Kato,
H. Updated prey consumption by common minke
whales and interaction with fisheries in coastal areas of
the Pacific side of Japan.
18. Murase, H., Kawabata, A., Kubota, H., Nakagami,
M., Amakasu, K., Abe, K., Miyashita, K. and Oozeki,
Y. 2012. Basin-scale distribution pattern and biomass
estimation of Japanese anchovy Engraulis japonica
in the western North Pacific. Fish. Sci. 78: 761-73.
[Published paper].
19. Wada, A., Onodera, K., Saeki, M. and Tamura, T.
Estimation of prey species biomass in the Sanriku region
based on 2008 and 2009 JARPN II acoustic surveys.
20. Watanabe, H., Okazaki, M., Tamura, T., Konishi, K.,
Inagake, D., Bando, T., Kiwada, H. and Miyashita, T.
2012. Habitat and prey selection of common minke, sei
and Bryde’s whales in mesoscale during summer in thesubarctic and transition regions of the western North
Pacific. Fish. Sci. 78: 557-67. [Published paper].
21. Murase, H., Tamura, T., Kiwada, H., Fujise, Y.,
Watanabe, H., Ohizumi, H., Yonezaki, S., Okamura,
H. and Kawahara, S. 2007. Prey selection of common
minke (Balaenoptera acutorostrata) and Bryde’s(Balaenoptera edeni) whales in the western North
Pacific in 2000 and 2001. Fish. Oceanog. 16(2): 186201. [Published paper].
22. Murase, H., Tamura, T., Isoda, T., Okamoto, R.,
Yonezaki, S., Watanabe, H., Tojo, N., Matsukura, R.,
Miyashita, K., Kiwada, H., Matsuoka, K., Nishiwaki,
S., Inagake, D., Okazaki, M., Okamura, M., Fujise, Y.
and Kawahara, S. Prey preferences of common minke(Balaenoptera acutorostrata), Bryde’s (B. edeni) andsei (B. borealis) whales in offshore component of
JARPN II from 2002 to 2007.
23. Konishi, K., Isoda, T. and Tamura, T. Decadal changeof feeding ecology in sei, Bryde’s and common minke
whales in the offshore of the Western North Pacific.
24. Tamura, T., Kishiro, T., Yoshida, H., Konishi, K. andKato, H. Relationship between feeding habitat andmaturity status of common minke whale off Kushiro.
25. Ishii, M., Murase, H., Fukuda, Y., Sawada, K., Sasakura,T., Tamura, T., Bando, T., Matsuoka, K., Shinohara,A., Nakatsuka, S., Katsumata, N., Miyashita, K. andMitani, Y. A short note on feeding behaviour of sei
whales observed in JARPN II.
26. Isoda, T., Tamura, T., Konishi, K., Ohizumi, H. andKubodera, T. Feeding habits of sperm whales (Physeter
macrocephalus) in the western North Pacific in springand summer.

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27. Konishi, K. Analyses of body condition in sei, Bryde’s
and common minke whales in the western North Pacificwith JARPN and JARPN II dataset.
28. Murase, H., Tamura, T., Hakamada, T., Watari, S.,
Okazaki, M., Kiyofuji, H., Yonezaki, S. and Kitakado,
T. Ecosystem modelling in the western North Pacificfrom 1994 to 2013 using Ecopath with Ecosim (EwE):
some preliminary results.
29. Kitakado, T., Murase, H. and Tamura, T. Predation
impacts on sandlance population by consumption ofcommon minke whales off Sanriku region.
30. Yasunaga, G. and Fujise, Y. Temporal trend of total Hglevels in three baleen whale species based on JARPN II
data for the period 1994-2014.
31. Yasunaga, G. and Fujise, Y. Temporal trend of PCBlevels in common minke whales from the western North
Pacific for the period 2002-2014.
32. Yasunaga, G. and Fujise, Y. Accumulation features ofPOPs of baleen whales in the western North Pacificbased on samples collected during the 2012 JARPN II
survey.
33. Yasunaga, G. and Fujise, Y. Comparison of total Hglevels in O and J type stock of common minke whales
based on JARPN II coastal samples collected in 2012
and 2013.
34. Yasunaga, G. and Fujise, Y. A note on POPs and Hg
accumulation in sperm whale based on JARPN II
samples collected during 2001-2013.
35. Yasunaga, G. and Fujise, Y. Status of I131, Cs134 andCs137 in baleen and sperm whales from the western
North Pacific during 2011-2015.
36. Niimi, S., Imoto, M., Kunisue, T., Watanabe, M.X., Kim,
E.Y., Nakayama, K., Yasunaga, G., Fujise, Y., Tanabe, S.
and Iwata, H. 2014. Effects of persistent organochlorine
exposure on the liver transcriptome of the common
minke whale (Balaenoptera acutorostrata) from theNorth Pacific. Ecotoxicology and Environmental Safety
108: 95-105. [Published paper].
37. Shimizu, Y., Ohishi, K., Suzuki, R., Tajima, Y.,
Yamada, T., Kakizoe, Y., Bando, T., Fujise, Y., Taru,
H., Murayama, T. and Maruyama, T. 2013. Amino acidsequence variations of signaling lymphocyte activation
molecule and mortality caused by morbilivirus infectionin cetaceans. Microbiology and Immunology 57: 62432. [Published paper].
38. Pastene, L.A., Goto, M., Taguchi, M. and Kitakado,
T. Temporal and spatial distribution of the ‘J’ and ‘O’stocks of common minke whale in waters around Japan
based on microsatellie DNA.
39. Nakamura, G., Kadowaki, I., Nagatsuka, S., Hayashi,R., Kanda, N., Goto, M., Pastene, L.A. and Kato, H.
Morphological differences in the white patch on the
flipper between J and O stocks of the North Pacificcommon minke whale.
40. Pastene, L.A., Goto, M., Taguchi, M. and Kitakado, T.
Updated genetic analyses based on mitochondrial andmicrosetallite DNA indicated no sub-structure of the‘O’ stock common minke whale in the western North
Pacific.
41. Bando, T. and Hakamada, T. Morphometric analysis on
stock structure of the O stock common minke whale in
the western North Pacific.
42. Kishiro, T. and Miyashita, T. 2011. Satellite tracking
of a common minke whale in the coastal waters off

Hokkaido, northern Japan in 2010. Paper SC/63/RMP10
presented to the IWC Scientific Committee, June 2011,
Tromsø, Norway (unpublished). 10pp. [Paper available
from the Office of this Journal].
43. Kitakado, T. and Maeda, H. Fitting to catch-at-age data
for North Pacific common minke whales in the Pacificside of Japan.
44. Pastene, L.A., Goto, M., Taguchi, M. and Kitakado,
T. Updated genetic analyses based on mtDNA
and microsatellite DNA suggest possible stock
differentiation of Bryde’s whales between management
sub-areas 1 and 2 in the North Pacific.
45. Murase, H., Tamura, T., Otani, S. and Nishiwaki, S.
2016. Satellite tracking of Bryde’s whales Balaenoptera
edeni in the offshore western North Pacific in summer
2006 and 2008. Fish. Sci. DOI 10.1007/s12562-0150946-8. [Published paper].
46. Pastene, L.A., Goto, M., Taguchi, M. and Kitakado,
T. Genetic analyses based on mtDNA control region
sequencing and microsatellite DNA confirmed theoccurrence of a single stock of sei whales in oceanic
regions of the North Pacific.
47. Kanda, N., Matsuoka, K., Goto, M. and Pastene, L.A.
Genetic study on JARPN II and IWC-POWER samplesof sei whales collected widely from the North Pacific at
the same time of the year.
48. Kanda, N., Bando, T., Matsuoka, K., Murase, H., Kishiro,T., Pastene, L.A. and Ohsumi, S. 2015. A review of thegenetic and non-genetic information provides support
for a hypothesis of a single stock of sei whales in the
North Pacific. Paper SC/66a/IA09 presented to the IWC
Scientific Committee, May 2015, San Diego, CA, USA(unpublished). 18pp. [Paper available from the Officeof this Journal].
49. Pastene, L.A., Goto, M., Taguchi, M. and Kitakado, T.
A note on the genetic diversity of sperm whales in thewestenr North Pacific as revealed by mitochondrial andmicrosatellite DNA.
50. Pastene, L.A., Goto, M., Taguchi, M., Nishiwaki, S.
and Kitakado, T. A note on the genetic diversity of andphylogeny of western North Pacific and southern rightwhales based on mitochondrial and microsatellite DNA.
51. Pastene, L.A., Goto, M. and Kitakado, T. A note on
mismatch distributions of mtDNA control region
sequences in baleen whales from the western North
Pacific.
52. Maeda, H., Kawamoto, T. and Kato, H. 2013. A study on
the improvement of age estimation in common minke
whales using the method of gelatinized extractionof earplug. NAMMCO Sci. Publ. DOI: http://dx.doi.
org/10.7557/3.2609. [Published paper].
53. Maeda, H., Bando, T., Kishiro, T., Kitakado, T. andKato, H. Basic information of earplugs as age characterof common minke whales in western North Pacific.
54. Fisheries Agency of Japan. Japan’s voluntary
considerations and response to the ICJ judgement inrelation to the adjustment of JARPN II program during
the period from 2014 to 2016.
55. Bando, T., Maeda, H., Ishikawa, Y., Kishiro, T. andKato, H. Preliminary report on progress in earplugbased age determination of sei whales collected during
2002 and 2013 JARPN II surveys.
56. No paper.
57. Addendum of SC/F16/JR29.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

OBSERVER STATEMENTS
SC/F16/O
1. De La Mare, W., Double, M., and Mckinlay, J.
Observers’ Statement to the JARPN II Special Permit
Expert Panel Final Review Workshop.
2. Tamura, T., Kishiro, T., Murase, H., Kitakado, T.
Yasunga, G., Morishita, J. and Pastene, L. A response
to Document SC/F16/O01 ‘Observers statement to the
JARPN II Special Permit Expert Panel Final Review
Workshop’.
FOR INFORMATION DOCUMENTS
Bhuiyan, M.M.U., Suzuki, N., Watanabe, H., Matsuoka, K., Fujise,Y., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2008. Attempts at in vitro
fertilization and culture of in vitro matured oocytes in sei (Balaenoptera
borealis) and Bryde’s (B. edeni) whales. Zygote 17: 19-28.
Bhuiyan, M.M.U., Suzuki, Y., Watanabe, H., Lee, E., Hirayama, H., Matsuoka,
K., Fujise, Y., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2010. Productionof sei whale (Balaenoptera borealis) cloned embryos by inter- and intraspecies somatic cell nuclear transfer. J. Reprod. Develop. 56(1): 131-39.
Birukawa, N., Ando, H., Goto, M., Kanda, N., Pastene, L.A. and Urano, A.
2008. Molecular cloning of urea transporters from the kidneys of baleen
and toothed whales. Comp. Biochem. Physiol. B 149: 227-35.
Glover, K.A., Kanda, N., Haug, T., Pastene, L.A., Oien, N., Goto, M.,
Seliussen, B.B. and Skaug, H.J. 2010. Migration of Antarctic minke
whales to the Arctic. PLoS ONE 5(12).
Glover, K.A., Kanda, N., Haug, T., Pastene, L.A., Øien, N., Seliussen, B.B.,
Sørvik, A.G.E. and Skaug, H.J. 2013. Hybrids between common andAntarctic minke whales are fertile and can back-cross. BMC Genetics 14: 25.
Hiwasa, M., Suzuki, Y., Watanabe, H., Bhuiyan, M.M.U., Matsuoka, K.,
Fujise, Y., Ishikawa, H., Ohsumi, S. and Fukui, Y. 2009. Effects of semen
extenders and storage temperatures on characteristics of frozen-thawed
Bryde’s (Balaenoptera edeni) whale spermatozoa. J. Reprod. Develop.
55(6): 599-606.
Inoue, S., Kishiro, T., Fujise, Y., Nakamura, G. and Kato, H. 2014. Seasonalchanges in the testis of the North Pacific common minke whale. Nippon
Suisan Gakkaishi 80(2): 185-90.
Kanda, N., Goto, M., Kato, H., McPhee, M.V. and Pastene, L. 2007.
Population genetic structure of Bryde’s whales (Balaenoptera brydei) at
the inter-oceanic and trans-equatorial levels. Cons. Genet. 8: 853-64.
Kanda, N., Goto, M., Kishiro, T., Yoshida, H., Kato, H. and Pastene, L.A.
2009. Update of the analyses on individual identification and mixingof the J and O stocks of common mike whales around Japanese waters
examined by microsatellite analysis. Paper SC/61/JR5 presented to the
IWC Scientific Committee, June 2009, Madeira, Portugal (unpublished).
14pp. [Paper available from the Office of this Journal].
Kanda, N., Goto, M., Oikawa, H. and Pastene, L.A. 2014. Update of noteon sampling and laboratory procedure protocols of the genetic work at theInstitute of Cetacean Research (SC/65b/J27rev). Paper SC/65b/DNA01
presented to the IWC Scientific Committee, May 2014, Bled, Slovenia(unpublished). 6pp. [Paper available from the Office of this Journal].
Kanda, N., Goto, M. and Pastene, L. 2006. Genetic characteristics ofwestern north Pacific sei whales, Balaenoptera borealis, as revealed by
microsatellites. Marine Biotechnol. 8: 86-93.
Kanda, N., Park, J.-Y., Goto, M., An, Y.-R., Choi, S.-G., Moon, D.Y., Kishiro, T., Yoshida, H., Kato, H. and Pastene, L.A. 2010. Genetic
analyses of western North Pacific minke whales from Korea and Japan
based on microsatellite DNA. Paper SC/62/NPM11 presented to the IWC
Scientific Committee, June 2010, Agadir, Morocco (unpublished). 13pp.
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Budipitojo, T., Kondoh, D. and Kitamura, N. 2015. Structure and functionsof the placenta in common minke (Balaenoptera acutorostrata) Bryde’s (B.
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Konishi, K., Tamura, T., Isoda, T., Okamoto, R., Hakamada, T., Kiwada,
H. and Matsuoka, K. 2009. Feeding strategies and prey consumption of
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Lee, E., Bhuiyan, M.M.U., Watanabe, H., Matsuoka, K., Fujise, Y.,
Ishikawa, H. and Fukui, Y. 2009. Production of cloned sei whale(Balaenoptera borealis) embryos by interspecies somatic cell nuclear
transfer using enucleated pig oocytes. J. Vet. Sci. 10(4): 285-92.
Maeda, H. and Kato, H. 2012. Technical improvements of the age character(earplug) collections in common minke whales. Bull. Jpn. Soc. Fish.
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Proponents response 2016 (mainly from SC/F16/JR01)

2016 Panel evaluation

Response to this recommendation was given in SC/F16/JR01, p.14.
Addressed in different analyses related to the three research objectives.

Proponents’ general view regarding quantification of objectives and sub-objectives was provided in Pastene
et al. (2015, pp.1-2). Sampling design and sample size estimates were based mainly on main objective 1(feeding ecology and ecosystem studies). As noted by the 2009 review workshop, for the feeding ecology
study determining the appropriate sample sizes is contingent on properly estimating the uncertainty
surrounding the key parameters that are ultimately to be used in the modelling process. The proponents
believe that in the context above, substantial progress was made in addressing the uncertainty in the whale’s
prey consumption estimate, which is one of the key parameters for ecosystem modelling.

Not addressed.
Partially addressed.

Not addressed.

Stock structure
1. The genetic assessments should include a brief description of procedures to ensure data quality (with reference
to IWC guidelines for DNA data quality). (R)
2. Revised papers should include estimates of genetic divergence (along with levels of uncertainty) in addition toprobabilities of homogeneity. (R)
3. P values (and divergence estimates) should be reported for all loci combined rather than for each locus separately.
(R)

Addressed in document Kanda et al. (2014). The IWC SC welcomed this document and agreed that it
Addressed.
responded appropriately to a previous recommendation.
Addressed in papers presented to the IWC SC after 2009. See also SC/F16/JR40 for common minke whales; Partially addressed.
SC/F16/JR44 for Bryde’s whale; SC/F16/JR46 for sei whales.
Addressed in papers presented to the IWC SC after 2009. See also SC/F16/JR40 for common minke whales;
Addressed.
SC/F16/JR44 for Bryde’s whale; SC/F16/JR46 for sei whales.

Being addressed in an ongoing research on the feasibility of non-lethal techniques (2014-16).

Not addressed up to2013.
Efforts were made to maximise the information from sampled animals. A large number of analyses were Partially addressed.
made on the same animals, resulting in a total of 61 peer-reviewed publications (30 of which were not related
to the main objectives of the program), and a large number of IWC SC documents.
5. A full evaluation of relative merits be undertaken as soon as possible after relevant work completed: (1) specified Not applicable.
and quantified objectives; (2) analysis of precision of estimates by technique; and (3) evaluation of practicalities
of field (lab) techniques in context of integrated objectives, sub-objectives and analyses proposed. (R)See response to this recommendation in SC/F16/JR01, p.14. There is an ongoing research on the feasibility Not addressed up to
6. If lethal sampling occurs, design well-specified study to evaluate techniques. (R)of non-lethal techniques (2014-16).
2013.

Comparison of lethal and non-lethal techniques
1. Add it as sub-objective. (R)
2. Analysis of the precision (and any associated biases) of the estimates obtained for the relevant parameters by eachof the lethal and non-lethal techniques. (R)
3. Evaluation of the practicalities and logistics of the field (and, if relevant, laboratory) techniques in the context of
the integrated objectives, sub-objectives and analyses proposed. (R)
4. To maximise the information obtained from animals and re-evaluate the need for such sampling at appropriate
intervals. (R)Sample size and design
For each objective: (a) specify quantities of interest needed to achieve objectives; and (b) for each quantity of interest:- identify/quantify sources of uncertainty; and- determine which are functions of sample size.
The results of such an analysis can be used to determine:- how much progress has been made to achieving objectives;- what further progress can be expected;- effect of changing N on time to achieve objectives; and- whether sampling design is most appropriate and maximises information for a given N. (R)Objectives
1. Development of refined, more quantified sub-objectives for each component of the programme as a priority. (R) Sub-objectives were identified for each main objective of JARPN II (see SC/F16/JR01).
Partially addressed.
2. For any long-term programme such as this, in addition to long-term objectives, proponents should determine Some of the sub-objectives identified above were of a short-term nature. Proponents’ general view regarding
Not addressed.
specific, shorter term objectives that are quantified to the extent possible. (R)quantification of objectives and sub-objectives was provided in Pastene et al. (2015, pp.1-2).

Recommendations (R) [and suggestions (S)] from the 2009 Panel

This summary Table includes the recommendations from the 2009 Panel, the response of the proponents and a summary evaluation by the proponents. Comments explaining thePanel’s summary evaluation are given in the relevant sections of this report.

Summary of Responses to the 2009 Panel Report by the Proponents
and an Evaluation by the Present Panel

Annex D576
REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

Proponents response 2016 (mainly from SC/F16/JR01)
Addressed.

2016 Panel evaluation

6. The authors incorporate into the index of density, the sightability of detected groups (e.g. effective strip half
widths that include appropriate covariates such as weather conditions). As for all modelling exercises, it isimportant to test whether the chosen model is an improvement over a null, uninformative model and to validate
the model results. Approaches to such validation could include: comparison of the modelled results not only withindex of densities from the present study but also with data that were collected from other years (e.g. JARPN orother survey data) and exploration of cross-validation type techniques. (R)

Oceanography and distribution
1. JARPN II data be pooled or compared with other datasets (e.g. JARPN I or other historical surveys) whenpossible. This will increase the sample size and increase the possibility of data covering periods of changing
relationships (e.g. previous regime changes), thus allowing patterns to be detected. (R)
2. To consider conducting future oceanographic surveys over an area larger than at present, not only to further
investigate oceanographic relationships, but also to improve abundance estimates for a variety of species. (R)
3. In the long term, to more fully understand the preferred habitat, prey preferences, niche separation of different
species, functional response, and spatial and temporal trends in local abundance and other biological factors (suchas blubber thickness, pollutants, presence of scars, and stock structure), the oceanographic data collected on thecruises (bottom depth, water column temperature, salinity, and density) and satellite derived data, such as SST,
chlorophyll, and sea surface height be integrated into future analyses. (R)
4. The salinity CTD data must be corrected/calibrated using the water samples that were simultaneously collectedwith the CTD data. (S)5. More of these types of analyses (including using other appropriate modelling techniques such as GAMs or logistic
regressions) be conducted. (R)

5. Provide more details on the analyses involving the program STRUCTURE. (R)Addressed.

Addressed in SC/F16/JR05-08.

The analyses were also recommended as medium to long term. The recommendation was addressed to some Partially addressed.
extent in published documents Watanabe et al. (2012) and Murase et al. (2007) [SC/F16/JR20-21], a
document presented to the 2009 review workshop Murase et al. (2009) and documents presented to thisworkshop (SC/F16/JR07, SC/F16/JR16 and SC/F16/JR27). All these studies but SC/F16/JR07, SC/F16/JR16
and SC/F16/JR27 were conducted at the mesoscale.
In principle, the salinity data recorded under JARPN II were calibrated. Quality checks are carried out for
Not applicable.
input data of the FRA-ROMS.
GLM and GAM has been considered and some of the results were published in scientific literatures. Spatial
Addressed.
distributions of common minke, sei and Bryde’s are attempted and the results are presented to this meeting(SC/F16/JR07). Furthermore, prey consumption of sei whales is preliminary estimated (SC/F16/JR16).
In abundance estimation using standard method (i.e. design based method), Beaufort Sea State was used as a Partially addressed.
covariate of detection function. Beaufort Sea State was selected as one of covariates for some of species (seeSC/F16/JR11-14). SC/F16/JR07 conducted Generalised Additive Model (GAM) for relative abundance
estimation of the common minke, sei and Bryde’s whales. Model selection was conducted based on GCVscores. Sea surface temperature (SST) and seafloor depth was selected as covariates of the GAM for thecommon minke and sei whales. SST, sea surface height anomaly (SSHa), sea surface chlorophyll-aconcentration (Chl-a) and the seafloor depth were selected for the GAM for Bryde’s whales.

Addressed.

Several of the analyses in the documents presented to this workshop used pooled data from JARPN and
JARPN II (see agenda item related to ‘integration’).

More details were provided in documents Kanda et al. (2009) and Kanda et al. (2010). See also SC/F16/JR38
Addressed.
for the application of the STRUCTURE program to individual assignment to J and O stocks common minke
whale.
6. Include a brief discussion of experimental design with respect to sampling (explaining how the design specifically Sampling design in JARPN II was made mainly in the context of main objective 1. For the key species, Partially addressed.
addresses uncertainties related to stock structure, e.g. whether the spatial and temporal coverage of samples of samples obtained by JARPN II were examined for stock structure purposes based on the IWC SC’s designed
minke whales has been sufficient to test adequately the alternative stock structure hypotheses). (R)sub-areas used for management purpose.
7. Redo the Boundary Rank analyses (Taylor and Martien, 2002) with new data. (R)In discussion at the IWC SC after 2009, this task was not assigned to JARPN II workers.
No longer applicable.
Samples from Korean bycatches were incorporated in several genetic studies presented to the IWC SC after
8. Integrate Korean bycatch samples into the new datasets to look at heterogeneity. (R)Addressed.
2009 (Kanda et al., 2010; Park et al., 2010).
9. Undertake the assessments of power to simulate data to evaluate power to detect a specified fraction of a putative Addressed in SC/F16/JR40 for O stock common minke whale; SC/F16/JR44 for Bryde’s whale.
Partially addressed.
stock (e.g. The hypothetical W stock of NP minke whales) in an overall sample using simulated data (withprograms SimCoal, ms, EasyPop9 or other freely available software). (R)
10. Undertake tests for population genetic (drift-mutation-migration) equilibrium. (R)Preliminary analyses presented in SC/F16/JR51.
Partially addressed.
Will be addressed in future following the discussions on SC/F16/JR51. The proponents believe that the IM
11. Undertake using non-equilibrium approaches the estimations of divergence between sample partitions. (R)Not addressed.
approach suggested by the workshop is not useful for population with low effect sizes.
12. Attempt the detection of pairs of individuals that are related (Skaug and Daníelsdóttir, 2006). (R)Work has been started and preliminary results obtained for common minke whale.
Partially addressed.
13. Undertake multivariate analyses of morphological data with respect to stock structure (e.g. PCA, cluster analysis, Addressed in Hakamada and Bando (2009) and SC/F16/JR41.
Addressed.
discriminant analysis, SIMPER and ANOSIM). (R)Previous studies showed that POPs levels in whales could be informative of the stock structure (Fujise et al., Partially addressed.
14. Use of past and present contaminant data should be an integrative study of stock structure. (R)2000; Nakata et al., 2000). Similar studies are planned for future.
Work has been started. Successful experiment were conducted for North Pacific Bryde’s (Murase et al., 2016) Partially addressed.
15. Carry out satellite tagging programmes to narrow the range of plausible stock structure hypotheses. (R)and common minke (Kishiro and Miyashita, 2011) whales.

4. Multiple testing issues: (a) use of the False Discovery Rate could be preferable to the Bonferroni correction; and The recommended False Discovery Rate (FDR) was used in all population genetic structure papers presented(b) to exercise discretion in the number of pairwise comparisons evaluated (e.g. by only comparing samples that to this workshop.
are geographically proximate and hence most likely to be connected demographically). (R)Recommendations (R) [and suggestions (S)] from the 2009 Panel

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017577

Proponents response 2016 (mainly from SC/F16/JR01)
Not addressed.

2016 Panel evaluation

After 2009 survey, design was made so that research areas were covered with sufficient effort by the dedicated Partially addressed.
sighting surveys.

Partially addressed.

Addressed.
Addressed.

7. Need for incorporating information resulting from other studies (e.g. JARPN I, PICES, ESSAS). (R)

8. Present the estimates of consumption by whales in terms of fisheries and prey biomass (this can provide an index Addressed in SC/F16/JR15, SC/F16/JR17.
of relativity and an easy and immediate sense of the magnitudes of the various processes that can affect fish stock
dynamics). (S)Addressed.

Addressed in SC/F16/JR15, SC/F16/JR17.
Addressed in SC/F16/JR15, SC/F16/JR17.

6. Methods used to extrapolate from daily to annual rates and amounts are not clearly explained. (R)

4. Panel recommends that additional analyses be undertaken to identify the greatest sources of uncertainty and to Uncertainties in the number of whales distributed in the research area, body weight of whales, consumption
Not addressed.
models, energy content of prey species, assimilation efficiency and the ratio of low/high feeding intake ofdetermine appropriate sampling and analytical strategies to address them. (R)whales were taken into account in the updated estimation (SC/F16/JR15, SC/F16/JR17).
5. In Ecosystem Modelling, scientific rationale on used modelling formulations and proposed ranges should be Addressed in SC/F16/JR15, SC/F16/JR17.
Partially addressed.
clearly given. (R)

3. For the treatment of uncertainty JARPN II data analyses should:
Addressed in SC/F16/JR15, SC/F16/JR17.
(a) incorporate the use of several reasonable models for estimating daily consumption as a function of body mass
and include the range of possible results in reporting their work;(b) use that range in subsequent analyses (including any ecosystem modelling) that employ these daily/annual
consumption estimates; and(c) undertake sensitivity analyses for the range of parameter values used in the consumption equations. (R)Prey consumption, including biomass estimation of prey species and prey preference, including feeding habits
The rational for the research area was presented in the original research plan of JARPN II (Government of
Not addressed.
1. Fuller rationale for sampling areas is necessary. (R)Japan, pp.22-24, 31). The research area is a high productive area involving the Kuroshio, the Oyashio
Currents, and the Transition Zone between them. The research area covered the distribution of resourcescaught by Japanese fisheries such as Pacific saury. Since 2003, coastal regions such as Sanriku and Kushiro
were added to study the competition between whales and fisheries in a hot-spot area.
2. Characterising uncertainty - steps needing variance estimates:
Addressed in SC/F16/JR15, SC/F16/JR17.
Partially addressed.
- Per capita consumption in area of interest: (1) parameter uncertainty in the relationship between energy
consumption and body mass (multiple and exponent); (2) residual variance of species values around the mean
curve; (3) proportion of annual energy obtained during feeding season; (4) length of feeding season (to getdaily energy requirement); and (5) variance in mean body mass (stratified by sex and life stage, e.g.
mature/immature).
- Diet composition: (1) average undigested biomass of prey group in forestomach; (2) mean residence time ofprey group in the forestomach; (3) average energy content per unit biomass of prey by prey type; (4) average
body weight of undigested prey items by species; and (5) relative frequencies of each species by counts ofindividuals and/or hard parts.
- Abundance (to scale up per capita consumption to population consumption): (1) variance (and possibly
covariance) in estimates of abundance (mean number of whales present) in survey season by sub-area andtime period, including g(0) variance, and process error. (R)Increased effort to obtain better estimates should be a high priority. (R)It also recommends that the photo-identification data be worked up and comparisons made with catalogues elsewhere Photo-identification data for blue, humpback and North Pacific right whales have been collected during Partially addressed.
in the North Pacific. (R)JARPN II surveys. The number of the collected photo-ID data were provided in Annex 3. Collaboration
between Russian and Japanese scientists is being prepared to conduct preliminary analysis of photo-id data
for North Pacific right whales. Also analysis of photo-id data for blue and humpback whales will be
conducted, and data from IWC-POWER surveys will be incorporated (in collaboration with IWC-POWER
specialists).

Abundance
Using the sightings data collected over the 1994-2007 period for the variety of large whales, the Panel recommends Geographic coverage of JARPN II survey was not always the same among the years. For this reason, it would
investigation of whether these data can be used to provide information on trends. (R)be necessary to pool sighting data to obtain full geographical coverage to examine abundance trend. Possible
approach may be that in SC/F16/JR09, in which sighting data were stratified by periods of 7-9 years.

Recommendations (R) [and suggestions (S)] from the 2009 Panel

578
REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

Regarding the collection of data, collaboration with specialists in other disciplines (e.g. oceanography, pelagic Partially addressed.
fish and highly migratory fish [sharks and tunas]) has been strengthened for EwE modelling.

Not addressed.

Addressed.

Addressed.

2016 Panel evaluation

Results of two modelling exercises were presented here (SC/F16/JR28-29) and a third one (MRM) is being Partially addressed.
considered for the near future. Time series data from 1994 to 2013 was fitted in Ecosim to estimate
vulnerability parameters which is one of important parameters in the model.
3. The area covered by JARPN II is not spatially homogeneous, and serious consideration should be given to A quasi sub-model structure is established in EwE considering bottom topography and oceanography of the Partially addressed.
developing separate models for three regions distinguished by the inshore or shelf region, the sub-Arctic oceanic modelled area.
region of the Oyashio current and the sub-tropical region of the Oyashio and Kuroshio transition zone. (R)
4. There is a need to take much wider account of uncertainty at all stages of the modelling process, including that Uncertainty is not addressed fully in EwE although vulnerability parameters are estimated in Ecosim. In
Not addressed.
SC/F16/JR29, both observation and process errors were accounted for using a Bayesian state-space delayassociated with the prey consumption rates of whales. (R)difference model, and estimation uncertainty was of course addressed in posterior distributions for unknown
and latent variables as in usual way.
5. The importance, ultimately, of developing models which incorporate natural variability in dynamic processes was A climate index, the Pacific Decadal Oscillation (PDO) index, is initially considered whether it can be used Partially addressed.
emphasised, although it was recognised that this might not be possible for certain ecosystem modelling as forcing function in Ecosim. However, it is not considered in actual modelling because it appear that
influence of PDO on biomass dynamics in the modelling period (1994-2013) is not strong. But it is still on‘packages’. (R)the to-do list in the further modelling especially if the modelling period is expanded to the past.
6. This is in addition to taking account of uncertainty in model structure and parameter values. (R)To be addressed in future.
Not addressed.
7. If there are other predators making individual contributions to sand lance natural mortality of similar size to that Type II functional response is employed in the state-space delay-difference model in SC/F16/JR29, but any
Not addressed.
estimated for minke whales, their explicit inclusion in this model must be considered. It agrees that Type I predator other than common minke whales has not been taken into account for the assessment of predation
functional relationships are unrealistic and need not be considered further. As noted earlier, any results presented impact because of its potentially biggest consumption in the region.
should distinguish yields of the prey species to predators and the fishery. (R)8. It is important to concentrate first on improving the Ecopath component of this EwE analysis before moving on A series of pre-balance diagnostics, ‘PREBAL’ (Link, 2010) is conducted for both the 2013 and 1994 models
Addressed.
to the next step of extending the modelling effort from a static to a dynamic model such as Ecosim. (R)to evaluate the initial static energy budget of Ecopath before Ecosim modelling is conducted.
9. (a) The species included in the Ecopath analysis should be reviewed giving attention to Ecopath models developed Because EwE mainly focuses on offshore instead of coastal area, it appears that gelatinous zooplankton is not
Addressed.
key species. In addition, such data are scarce so far. Presented EwE modelling mainly focus on interaction
for other regions; in particular the inclusion of gelatinous zooplankton should be considered.
between forage fish and their predators and the number of predators in the model is increased from theprevious one.
(b) Furthermore the values of the parameters of this Ecopath analysis should be compared with values for those Values of basic parameters are compared with other models qualitatively.
Partially addressed.
others, with attention directed towards any instances of major discrepancies. (R)
10. The need to rebalance the Ecopath model. Alternative approaches to doing so should be considered. For example, A series of pre-balance diagnostics, ‘PREBAL’ (Link, 2010) is conducted for both the 2013 and 1994 models Partially addressed.
rather than use values for some parameters drawn from other regions, placing a bound on some relationship (e.g. to evaluate the initial static energy budget of Ecopath before Ecosim modelling is conducted.
P/C<0.6) may lead to an improved result overall. (R)
11. To take full account of the uncertainties associated with model inputs (e.g. using Ecoranger3). (R)Ecoranger is not used in the modelling as its development is little so far.
Not addressed.
12. Further work on Minimal Realistic Modelling approaches is encouraged and should focus in particular on fitting In a MRM off Sanriku region, various kinds of time series data such as CPUE, catch series and age
Not addressed.
composition were used for fitting the model. The model is considered as an integrated model, where weighting
such models to time series of data. (R)to likelihood components is influential to the results. At this moment, this issue has not been dealt well in thepaper, but different effective sample sizes will be used to check the sensitivity/robustness.

Ecosystem modelling
1. Considerably more resources must be allocated to the modelling work – without this, the likelihood that the objectiveof the programme will be reached in a reasonable timeframe will be minimal. The models developed should be used
to identify the areas of uncertainty with the greatest impact on model outputs of relevance to management, and hence
to guide the prioritisation of future data collection and the associated sample size/sampling design. (R)
2. A wider range of models needs to be considered if the objectives of the programme are to be met. Further work
should aim towards fitting dynamic models to time series of data, especially abundance indices. (R)The analyses were recommended as medium to long term. The recommendation was addressed to someextent in published documents (SC/F16/20-21), a document presented to the 2009 review workshop (Murase
et al., 2009) and documents presented to this workshop (SC/F16/JR07, SC/F16/JR16 and SC/F16/JR27).
All these studies but SC/F16/JR07, SC/F16/J16 and SC/F16/JR27 were conducted at the mesoscale.
Addressed in SC/F16/JR16.

9. Combine the oceanographic data, prey distributions and sighting survey data statistically to investigate how prey
and whale distributions are associated with oceanographic conditions, and how whale distributions are related to
distributions of prey - in this regard the sei whale example spatial modelling approach given in Hakamada (2009)
needs to be refined and extended further. (R)
10. Combine data on prey distributions as observed in the area where the whales were caught with the diet of thewhales (referred to as the micro scale) statistically to evaluate how well the whale’s diet reflects prey availability
in the area where it was caught. (R)
11. Compare results from the approaches listed above with the results on selectivity already produced and presented
at the Workshop. (R)To be addressed in future.

Proponents response 2016 (mainly from SC/F16/JR01)

Recommendations (R) [and suggestions (S)] from the 2009 Panel

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017579

Addressed in SC/F16/JR36.

Addressed.

To be addressed in future.
Addressed in SC/F16/JR33, and will be further addressed in future.

19. Undertake power analyses for relationship between N and ability to detect changes at various levels. (R)20. Evaluate covariates (e.g. age and sex) to determine animals chosen for more extensive sampling. (R)

21. In terms of sampling strategy, the value of examining the same individuals for each of the contaminants is To be addressed in future.
emphasised. (R)Addressed in SC/F16/JR30 and SC/F16/JR31.

18. The contaminant results should eventually be linked to the prey consumption studies. (R)

17. Simple mass balance studies (input-output estimates) would contribute to our knowledge of the partitioning and Preliminary attempts were made but results were not satisfactorily at this stage. Further attempts will be madeoffloading of contaminants in these species and the potential impact of changes in exposure. For this, additional in future.
analyses of blood, bile, faeces and urine are required. (R)This was already addressed in Yasunaga and Fujise (2009b). However such kind of analyses have been
stopped because levels of OC isomers in air and water samples were extremely low or under detection limit.

Not addressed.

15. Include fatty acid profiles and stable isotope ratios. This would help discriminate among reasons for temporal Samples from JARPN/JARPN II have not been analysed for stable isotope ratios and fatty acid yet. Futurechanges (i.e. dietary changes or exposure variation with constant diet). (R)analyses are being considered.

16. Air and water samples obtained could have been useful in a ‘fate and behaviour’ study (R)Not addressed.

14. Use of bycaught J-stock animals as comparison group. (R)Not addressed.

Partially addressed.

Not addressed.

Partially addressed.

Not addressed.

Addressed.

Partially addressed.

Age was incorporate as a covariate in the analyses conducted in SC/F16/JR33.
Samples from by-catch are available only for genetic analyses and such samples are not suitable for pollutant
analyses. However J stock animals (24) sampled by JARPN II in Sanriku were used for pollutant analysis(Hg) (see SC/F16/JR33).

13. Priority to have absolute age as an additional covariate for the interpretation of the results. (R)Addressed.

All tissues and organs such as muscle, liver, kidney and blubber of all whales sampled by JARPN II werestored at -20ºC. Unfortunately a substantial number of samples were lost after the 2001 earthquake andtsunami.

12. Tissues should be archived (frozen at –20°C or lower if possible) for future retrospective analyses. (R)

11. Any future contaminant exposure and uptake studies should be based on a balanced, structured study design with In the pollutant studies, the number of samples and strata (location, period, sex, and maturity) examined was Partially addressed.
a specific number of individuals sampled within each strata (e.g. by species, sex, stage, ocean regime and carefully considered in function of the objective of the study.
location). All the necessary data on exposure and confounding variables should be obtained from all of thespecifically targeted individuals and a control or comparison group should be included. In this way a more
powerful and statistically robust study to address clearly stated hypotheses could be designed and carried out. (R)

10. To continue comparative molecular phylogenetic research using mRNA isolated from fresh tissues. (S)Not addressed.

9. Examine T-Hg in brain tissue particularly for comparing the more coastal bycaught animals to the coastal and Sampling other than for genetics is not possible for by-catches.
offshore JARPN II samples, as these would provide a valuable (perhaps more exposed) comparison group. (S)Partially addressed

7. Yasunaga and Fujise (2009a) investigated the accumulation of total and methyl mercury and selenium in baleen The recommendation was mainly for sperm whales, for which samples are limited.
and sperm whales from the western North Pacific. The Panel recommends that in a revised paper submitted to
the 2009 Annual Meeting, greater emphasis is given to this important ecotoxicological finding. (R)Not addressed.

Partially addressed.

8. T-Hg total body burden estimates, additional organs (additional to liver, kidney and muscle) would need to be To be addressed in future.
included (e.g. brain, skin, and blubber). (R)Partially addressed.

Addressed in SC/F16/JR32.

6. Sampling for PCBs and Hg from the same individuals is undertaken to allow combined analyses of these often Addressed in SC/F16/JR34.
co-occurring contaminants. (R)

5. Future studies must be carried out on a lipid weight basis. (R)Not addressed.

Addressed.

Addressed in SC/F16/JR33-34.

4. A GAM fitted to these data would be a better method for determining the change points and examining non-linear To be addressed in a new paper for SC/66b.
trends in the Hg levels. (R)Partially addressed.

Addressed in SC/F16/JR33.

2. Analyses be carried out by age when age data become available. (R)

3. Future studies also should examine levels in the liver to facilitate comparison with other studies. (R)2016 Panel evaluation
Partially addressed.

Proponents response 2016 (mainly from SC/F16/JR01)

Pollution
1. In general, where possible, papers should include a risk assessment statement, summarising the potential risk Addressed in SC/F16/JR35.
from exposure to the various pollutants, based on current toxicology data in model species and other wildlife interms of the health of the animals and dynamics of the stocks. (R/S)

Recommendations (R) [and suggestions (S)] from the 2009 Panel

580
REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

Fujise, Y., Aoki, M., Niimi, S., Hakamada, T., Nakata, H., Honda, K. and Tanabe, S. 2000. An attempt to identify stocks in the western North pacific minke whale (Balaenoptera acutorostrata) using the accumulation levels ofheavy metals and organochlorines as ecological tracers. Paper SC/F2K/J18 presented at the JARPN Review Meeting, Tokyo, Japan, 7-10 February 2000 (unpublished). 18pp. [Paper available from the Office of this Journal].
Hakamada, T. 2009. Examination of the effects on whale stocks of future JARPN II catches. Paper SC/J09/JR36 presented to the Expert Workshop to Review Results of JARPN II, 26-30 January 2009, Tokyo, Japan (unpublished).
51pp. [Paper available from the Office of this Journal].
Hakamada, T. and Bando, T. 2009. Further morphometric analysis on stock structure in the western North Pacific common minke whales (Balaenoptera acutorostrata). Paper SC/61/JR6 presented to the IWC Scientific Committee,
June 2009, Madeira, Portugal (unpublished). 4pp. [Paper available from the Office of this Journal].
Kanda, N., Goto, M., Kishiro, T., Yoshida, H., Kato, H. and Pastene, L.A. 2009. Update of the analyses on individual identification and mixing of the J and O stocks of common mike whales around Japanese waters examined by
microsatellite analysis. Paper SC/61/JR5 presented to the IWC Scientific Committee, June 2009, Madeira, Portugal (unpublished). 14pp. [Paper available from the Office of this Journal].
Kanda, N., Goto, M., Oikawa, H. and Pastene, L.A. 2014. Update of note on sampling and laboratory procedure protocols of the genetic work at the Institute of Cetacean Research (SC/65b/J27rev). Paper SC/65b/DNA01 presented
to the IWC Scientific Committee, May 2014, Bled, Slovenia (unpublished). 6pp. [Paper available from the Office of this Journal].
Kanda, N., Park, J.-Y., Goto, M., An, Y.-R., Choi, S.-G., Moon, D.-Y., Kishiro, T., Yoshida, H., Kato, H. and Pastene, L.A. 2010. Genetic analyses of western North Pacific minke whales from Korea and Japan based on
microsatellite DNA. Paper SC/62/NPM11 presented to the IWC Scientific Committee, June 2010, Agadir, Morocco (unpublished). 13pp. [Paper available from the Office of this Journal].
Kishiro, T. and Miyashita, T. 2011. Satellite tracking of a common minke whale in the coastal waters off Hokkaido, northern Japan in 2010. Paper SC/63/RMP10 presented to the IWC Scientific Committee, June 2011, Tromsø,
Norway (unpublished). 10pp. [Paper available from the Office of this Journal].
Murase, H., Tamura, T., Isoda, T., Okamoto, R., Yonezaki, S., Watanabe, H., Tojo, N., Matsukura, R., Miyashita, K., Kiwada, H., Matsuoka, K., Nishiwaki, S., Inagake, D., Okazaki, M., Okamura, H., Fujise, Y. and Kawahara,
S. 2009. Prey preferences of common minke (Balaenoptera acutorostrata), Bryde’s (B. edeni) and sei (B. borealis) whales in offshore component of JARPN II from 2002 to 2007. Paper SC/J09/JR18 presented to the Expert
Workshop to Review Results of JARPN II, 26-30 January 2009, Tokyo, Japan (unpublished). 31pp. [Paper available from the Office of this Journal].
Murase, H., Tamura, T., Kiwada, H., Fujise, Y., Watanabe, H., Ohizumi, H., Yonezaki, S., Okamura, H. and Kawahara, S. 2007. Prey selection of common minke (Balaenoptera acutorostrata) and Bryde’s (Balaenoptera edeni)
whales in the western North Pacific in 2000 and 2001. Fisheries Oceanography 16(2): 186-201.
Murase, H., Tamura, T., Otani, S. and Nishiwaki, S. 2016. Satellite tracking of Bryde’s whales Balaenoptera edeni in the offshore western North Pacific in summer 2006 and 2008. Fisheries Science DOI 10.1007/s12562-0150946-8. [Published paper].
Nakata, H., Tanabe, S., Niimi, S., Minh, T.B., Sakakibara, A., Fujita, K. and Fujise, Y. 2000. Population structure in minke whale from the North Pacific examined by the persistent organic pollutants as chemical tracers. Paper
SC/F2K/J17 presented at the JARPN Review Meeting, Tokyo, Japan, 7-10 February, 2000 (unpublished). 9pp. [Paper available from the Office of this Journal].
Park, J.Y., Goto, M., Kanda, N., Kishiro, T., Yoshida, H., Kato, H. and Pastene, L.A. 2010. Mitochondrial DNA analyses of J and O stocks common minke whale in the western North Pacific. Paper SC/62/NPM21 presented to
the IWC Scientific Committee, June 2010, Agadir, Morocco (unpublished) 10pp. [Paper available from the Office of this Journal].
Skaug, H. and Daníelsdóttir, A.K. 2006. Relatedness of North Atlantic fin whales. Paper SC/58/PFI9 presented to the IWC Scientific Committee, May 2006, St. Kitts and Nevis, West Indies (unpublished). 8pp. [Paper available
from the Office of this Journal].
Taylor, B.L. and Martien, K. 2002. Interpretation of Boundary Rank results for North Pacific minke whales. Paper SC/J02/NP2 presented to the Scientific Committee Workshop on North Pacific common minke (Balaenoptera
acutorostrata) Implementation Simulation Trials held in Seattle, USA, 19-22 January, 2002 (unpublished). [Paper available from the Office of this Journal].
Watanabe, H., Okazaki, M., Tamura, T., Konishi, K., Inagake, D., Bando, T., Kiwada, H. and Miyashita, T. 2012. Habitat and prey selection of common minke, sei and Bryde’s whales in mesoscale during summer in the subarctic
and transition regions of the western North Pacific. Fisheries Science 78: 557-67. [Published paper].
Yasunaga, G. and Fujise, Y. 2009a. Accumulation features of total and methyl mercury and selenium in tissues of common minke, Bryde’s and sperm whales from the western North Pacific. Paper SC/J09/JR25 presented to theExpert Workshop to Review Results of JARPN II, 26-30 January 2009, Tokyo, Japan (unpublished). 11pp. [Paper available from the Office of this Journal].
Yasunaga, G. and Fujise, Y. 2009b. Temporal trends and factors affecting PCB levels in baleen whales and environmental samples from the western North Pacific. Paper SC/J09/JR24 presented to the Expert Workshop to Review
Results of JARPN II, 26-30 January 2009, Tokyo, Japan (unpublished). 10pp. [Paper available from the Office of this Journal].

REFERENCES

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

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582

REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

Annex
Annex EE

of Panel Recommendations
Summary of Summary
Panel Recommendations(See Main Report for Full
Text forandFull
Explanations)(See Main Report
Text and Explanations)
Sampling design and areas (Item 3.4.2.1)(1) A new paper that in addition to the information on sightings, it should document, for each year and season:(a) the predetermined tracklines for sampling and the rationale for those lines; and(b) the actual coverage of those tracklines and the rationale for any decisions taken to deviate from the predeterminedlines including the rationale for any new lines developed.
It should also address the issue of whether the actual sampling that occurred can be said to be representative of: (a) the
animals in the surveyed area; and (b) those in the biological population(s) and discuss the extent to which this may affectthose objectives/parameters/analyses for which this is or may be important.
(2) Papers using data from the inshore component must fully address the implications of the logistical rather than scientificsampling design.
Sample size (Item 3.4.2.2)(3) A new paper should be developed that:(a) provides a clearer rationale for the changes in sample sizes and any implications for meeting the original objectivesof the programme; and(b) provides the field and analytical protocols for the comparison of using lethal and non-lethal techniques for eachkey parameter taking into account the advice provided in 2009.
Stock structure (Item 4.4.3)(4) All inferences regarding ‘randomness’ of observations (e.g., satellite tracks, mitochondrial DNA haplotypes andunassigned common minke whales) should be substantiated by a statistical assessment of the presumed randomness.
(5) The presence of multiple stocks within sample partitions should be assessed (employing, e.g. STRUCTURE and DAPC).
(6) More explicit information on quality checks be provided in each study as well as study-specific estimates or genotyping
and DNA sequencing error rates.
(7) To facilitate more definitive discrimination between single and multiple stock hypotheses, undertake work to determine
the demographic dispersal rates among areas at which whales in different areas can be managed as a single stock.
Identifying ‘critical’ dispersal rates by specific case and the corresponding levels of genetic divergence, should enable
such discrimination. The approach of Van der Zee and Punt (2014) is commended. This will allow the development ofa working definition of a ‘stock’.
(8) Analytical approaches should be applied that do not assume mutation-drift-migration equilibrium (Hey, 2010).
(9) Serious consideration should be given to using genome-wide SNP genotyping approaches, such as RAD sequencing and
GBS (Elshire et al., 2011; Miller et al., 2007). This will increase the data per sample thereby improving the accuracy andprecision of genetic parameter estimates and facilitate additional analyses (Hey and Machado, 2003; Robinson et al.,2014).
(10) A focussed satellite tagging programme should be developed to greatly increase sample size to assess individualmigration in the context of stock structure hypotheses more thoroughly.
Feeding ecology and ecosystem studies – oceanography (Item 5.4.3.1)(11) Chl-a concentration should be examined as a potential proxy for the food environment for whales.
(12) Oceanographic monitoring is required to compare with prey species distribution and abundance in the new ‘decadalregime’.
Feeding ecology and ecosystem studies – distribution (Item 5.4.3.2)(13) With respect to papers SC/F16/JR7; Murase et al. (2014) [SC/F16/JR08]; SC/F16/JR09; Sasaki et al. (2013)[SC/F16/JR10] and SC/F16/JR16, develop revised versions that: (a) include statistical summaries on model fit (R2 and% deviance explained) and model comparison and spatial covariate selection (e.g. AIC, GCV scores); (b) avoid
extrapolation of the regression models outside to data-poor areas or areas lacking coverage (especially when combining
food consumption with sightings data); and (c) include variance plots of the fitted prediction surfaces in order to address
precision and data sparseness.
(14) Considerable effort be put into the methodological improvement of the spatial modelling in the various analysis relatedwith the objectives on distribution of large whales and oceanography. A particular focus must be on the combination ofsurvey data from the different years to make them more comparable in terms of distribution (and abundance) over time;use of data from other sources (e.g. the IWC POWER programme). This work is not only valuable in itself but is essential
for a better parameterisation of ecosystem models.
(15) Additional effort be placed on fulfilling the 2009 recommendation with respect to the photo-identification data tocontribute to the understanding of large scale movements and whale distribution within and outside the JARPN II survey
area for several species.
Feeding ecology and ecosystem studies – abundance (Item 5.4.3.3)(16) Explore methods to account for sampling differences between areas and years to obtain measures of short- and long-term
variation and trends and estimates the extent of additional variance due to changes over time in spatial distribution(essential for modelling efforts, for example, in food consumption models and ecosystem models).
(17) Compare results from the design-based estimates of abundance with those of model-based estimates to potentiallyaddress problems of unequal sampling coverage between surveys and to potentially account for additional sources orcauses of variability.
Feeding ecology and ecosystem studies - field and laboratory studies (Item 6.4.3)(18) The sampling distribution for the parameters should be used in the assessment of the uncertainty associated with theestimation of consumption.
(19) Clarification should be provided on how density and diet consumption have been extrapolated outside the areas andmonths covered during the surveys and diet studies.
(20) All sources of uncertainty should be quantified and an evaluation of which parameters contribute the most to uncertainty
be conducted and taken into account in the analyses and modelling.

Timeline
2016 Annual Meeting of
the Scientific Committee.

2016 Annual Meeting of
the Scientific Committee.

2016 Annual Meeting of
the Scientific Committee(or 2017 at the latest).
Expected to be completed
2-3 years after the 2016
Annual Meeting.

Within 2 years of the 2016
Annual Meeting.
Several years.
2016 Annual Meeting.

Expected to be completed
2-3 years after the 2016
Annual Meeting.

Expected to be completed
2-3 years after the 2016
Annual Meeting.

2016 Annual Meeting of
the Scientific Committee(or 2017 at the latest).
Expected to be completed
2-3 years after the 2016
Annual Meeting.

583

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017(21) The studies on allometric relationships should be developed further to refine the range of suitable allometric-energy
intake/consumption relationships.
(22) The analyses of diet composition should consider the effect of seasonal changes in energy density of the various preyspecies.
(23) Stable isotope analysis of whale tissues and their prey should be introduced not only into the assessment of diet, but also
to statistically evaluate overlap in distribution and trophic niche between baleen whale species.
Feeding ecology and ecosystem studies – ecosystem modelling (Item 7.4.3)(24) Generic recommendations identified by the 2009 Panel remain.
(25) Establish clear objectives on the ultimate use of the models to make further progress (e.g. better understanding ecosystem
linkages, delivering advice for fishery management) – ecosystem models are not suitable for tactical management.
(26) Use models in concert e.g. use food web modelling to establish key predation linkages for extended single-species ormultispecies models. In such a way the suite of available modelling tools can be used to integrate available knowledge.
(27) Use stable isotopes to provide information on long term feeding patterns and inform models about trophic relationships
between whales and their prey (see also Item 6.4).
(28) With respect to the EwE modelling: (a) evaluate data quality for each input parameter (the ‘pedigree’: e.g. Gaichas et al.
(2015) to characterise uncertainty in model inputs; (b) further evaluate PREBAL and other diagnostics; and (c) present
more clearly and evaluate further the estimated vulnerabilities and other fit diagnostics (including sensitivity analysis
using ranges of consumption estimates).
(29) With respect to extended single-species modelling:(a) ensure that the majority of predation mortality is captured;(b) carry out additional diagnostics: (1) examine the fits to a. fishery-independent survey data; b. proportion
information; and c. trends in fishing mortality; and (2) use posterior predictive checks to evaluate Bayesian model;(c) provide thorough justification for the current spatial boundaries of the model and the use of fishery CPUE as anindex of abundance;(d) focus the model fitting on the fishery-independent survey if CPUE not considered likely to index abundance;(e) examine sensitivity to alternative plausible functional forms of the feeding relationship; and(f) explore the causes of the implausible posteriors (e.g. SC/F16/JR29) by changing the weights assigned to the data
sources and fitting the model.
Monitoring environmental pollutants in cetaceans and marine ecosystem (Item 8.4.3)(30) To improve the statistical analyses based on clear and well-formulated hypotheses.
(31) Recalculate OC concentrations as values on a lipid weight basis, and Hg concentrations on a dry weight basis.
(32) Explore trends in pollutant concentrations using generalised additive models (GAMs) or other non-linear approaches, inaddition to the linear models.
(33) Evaluate the pollutant concentrations found in comparison with data from previous studies conducted in comparable
species and available in the literature.
(34) Since body length is a poor proxy for age, particularly in sexually mature whales, incorporate age data into themultivariate analysis of pollutant concentrations as soon as they become available.
(35) To include stable isotope values in the analyses to investigate the bioaccumulation process of pollutants through the foodchain.
(36) To assess more widely the risk that these chemical pollutants present to the populations’ abundance or distribution.
Ageing (Item 9.1.2)(37) To investigate into whether there is any relationship between age or sex and readability that may affect the
representativeness of the earplugs that can be read.
(38) To age as many of the existing samples as possible and to incorporate age where appropriate in updated analyses (e.g.
see the recommendations on pollutant studies).
Recommendations to the Scientific Committee on process (Item 11)(39) The Panel recommends that the Scientific Committee considers:(a) including a guideline either relating to the minimum time after completion of a programme that a final review cantake place or establishing a small review group to determine whether the materials available are for a review
workshop;(b) adopt guidelines for an integrated final report by the proponents (see Annex F);(c) to consider a mechanism for proponents to provide a short biennial update on progress with recommendations; and(d) develop a mechanism to allow for the completion of expert Panel reviews if a Panel states that its review isincomplete until further information/analyses is provided.

Expected to be completed
2-3 years after the 2016
Annual Meeting.

Within 2 years of the 2016
Annual Meeting.
Within 2 years of the 2016
Annual Meeting2016 Annual Meeting of
the Scientific Committee(or 2017 at the latest).

Expected to be completed
2-3 years after the 2016
Annual Meeting.

Within 2 years of the 2016
Annual Meeting.

At any periodic or final
review.

REFERENCES
Elshire, R.J., Glaubitz, J.C., Sun, Q., Poland , J.A., Kawamoto, K., Buckler, E.S. and Mitchell, S.E. 2011. A robust, simple genotyping-by-sequencing (GBS)
approach for high diversity species. PLoS ONE 6.
Gaichas, S., Aydin, K. and Francis, R.C. 2015. Wasp waist or beer belly? Modeling food web structure and energetic control in Alaskan marine ecosystems,with implications for fishing and environmental forcing. Prog. Oceanog. 138: 1-17.
Hey, J. 2010. Isolation with migration models for more than two populations. Mol. Biol. Evol. 27: 905-20.
Hey, J. and Machado, C.A. 2003. The study of structured populations - new hope for a difficult and divided science. Nature Reviews Genetics 4: 535-43.
Miller, M.R., Dunham, J.P., Amores, A., Cresko, W.A. and Johnson, E.A. 2007. Rapid and cost-effective polymorphism identification and genetyping using
restriction site associated DNA (RAD) markers. Genome Research 17: 240-48.
Murase, H., Hakamada, T., Matsuoka, K., Nishiwaki, S., Inagake, D., Okazaki, M., Tojo, N. and Kitakado, T. 2014. Distribution of sei whales (Balaenoptera
borealis) in the subarctic-subtropical transition area of the western North Pacific in relation to oceanic fronts. Deep Sea Res. II 107: 22-28.
Robinson, J.D., Bunnefield, L., Hearn, J., Stone, G.N. and Hickerson, M.J. 2014. ABC inference of multi-population divergence with admixture from unphased
population genomic data. Mol. Ecol. 23: 4458-71.
Sasaki, H., Murase, H., Kiwada, H., Matsuoka, K., Mitani, Y. and Saitoh, S.I. 2013. Habitat differentiation between sei (Balaenoptera borealis) and Bryde’s
whales (B. brydei) in the western North Pacific. Fish. Oceanog. 22(6): 496-508.
Van der Zee, J.P. and Punt, A.E. 2014. Evaluting ciritical dispersal rates for whale management under the IWCs Revised Management Procedure: an application
for North Atlantic common minke whales. J. Cetacean Res. Manage 14: 125-32.

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REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

Annex F
Proponents’ Response to Panel Request for Information on
Trackline Designs and Realised Tracklines for JARPN II by Yearand Season

Fig. 1. Overview of the survey design under the JARPN II. There are several surveys components under the JARPN II: dedicated sighting survey; whales
sampling surveys (coastal and offshore); concurrent whale/prey surveys in offshore areas; and prey surveys in coastal areas (see details of each survey below*).
In the figure, pre-determined track lines for whale sampling surveys in the offshore component are shown in orange; on effort track lines of dedicated sighting(single line) and whale sampling (parallel lines) surveys in green. Concurrent whale/prey surveys are shown by colour squares. Only the research areas areshown for Sanriku and Kushiro.
[Figures 2-28 are available online only at: http://archive.iwc.int/?c=29].

DESIGN OF TRACK LINES IN JARPN II SURVEYS
Dedicated sighting survey
Sighting surveys were conducted following the IWC
survey guidelines. Zig-zag-shaped track lines were set in
the research area independently from the whale sampling
surveys. The whole research area was covered in early(April to June) and late (July to September).
Whale sampling survey (offshore component)
The survey order of sub-area/strata was decided based on
seasonal distribution of whales and logistics, and zigzagshaped track lines were set in the research area, reflecting
the available information such as surface temperature. The
track line consisted of one main and two parallel courses
established seven n.miles apart from the main course. The
Special Monitoring Survey (SMS) was conducted in areas

where the density of targeted whale species were expected tobe high. Track lines in the SMS were designed independently
from the original track lines. The track lines of SMS
consisted of one main and two parallel courses establishedseven n. miles apart from the main course. Design of tracklines were determined by the cruise leader on the researchbase vessel.
Concurrent whale/prey surveys (offshore component)
For these surveys meso-scale research areas were defined insome years considering physical environmental informationsuch as surface temperature. Zigzag-shaped track lines were
set in the research area and both whale sampling vessels
and prey survey vessels conducted surveys along the same
track lines within a week. Prey survey vessel conducted
quantitative echo-sounder, net sampling and oceanographic
surveys.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

Whale sampling survey (coastal component)
The predetermined course (direction from the port) at anangle of regular intervals (usually 10-15 degree intervals)
were set up, and allocated to each research vessel. The
vessels continued to search along the course until common
minke whales were sighted, or until they reached 30 n.miles
from the port. After 30 n. miles, the vessels changed thecourse freely within a 50 n.miles radius from the port. The
predetermined course is changed every day to cover broad
areas. When whales were caught, the vessels returned to theport to transport the animal to the research land station. Afterlanding the whale, the vessel re-departed to the research
area.

585

Prey survey (coastal component off Sanriku)
Since 2005, the prey survey area was divided into ten blocks(A, B, C, D, E, F, G, H, I, and J) based on bottom depth (20,40, 100, and 200m), and prefectural boundaries (boundary
between Miyagi and Fukushima Prefectures). Because oflogistical constraint, the number of blocks changed in eachyear. In 2008 and 2009, six blocks (B, C, E, D, E and F)
were surveyed. Saw tooth type zigzag lines were used ineach survey. The survey was conducted during the daytime
from an hour after sunrise to an hour before sunset. The
research vessel used was a trawler-type vessel. The preyspecies were investigated using a quantitative echo sounder(EK 500; Simrad, Norway) and net sampling. Prey surveys
were conducted during the survey period of whale sampling
survey and independently from the whale sampling surveys.

Annex G
Some Suggestions for Potential Guidelines for an Integrated
Final Report from a Special Permit Programme
It should be noted that several of the sections should easily
be taken from the original proposal and any periodic reviews(e.g. Chapters 1-3 and early sections of Chapters 4). These
guidelines are intended to assist proponents as well as
reviewers. Electronic copies of the full report, its annexes
and all listed peer-review papers and cited documents should
be submitted to the Secretariat according to the timeline
defined in Annex P.
1. EXECUTIVE SUMMARY
This should be short (usually no more than 3-4 pages)
summary of the results of programme by Objective and Subobjectives with an indication of any limitations and a short
explanation of the contributions the programme in light of
the topics covered by Annex P1.
2. INTRODUCTION
This should include:(a) identification of Objectives and Sub-objectives
and any changes to these over the period of the
programme; and(1) assess the extent of the programme’s scientific output, and whether
this was appropriate in light of the stated research objectives and the time
elapsed; (2) assess the degree to which the programme coordinated its activities with related research projects; this included assessment of whether the
degree of coordination was sufficient to ensure that the field and analyticalmethods were appropriate and best practice to achieve the stated objectives
and whether the degree of coordination was sufficient to avoid unnecessary duplication; (3) evaluate other contributions to important research andinformation needs that were not part of the original set of objectives of theresearch programme; (4) consider any other relevant matters as decided by
the Scientific Committee; and (5) evaluate how well the initial, or revised,objectives of the research were met, and the extent to which results have
led to demonstrated improvements in the conservation and managementof whales, for broad categories of objectives 1 (‘improve the conservationand management of whale stocks’) and 2 (‘improve the conservation andmanagement of other living marine resources or the ecosystem of which thewhale stocks are an integral part’).

1(b) short background as to why they are important andwhy changes were made if they occurred.
3. STUDY AREA(S), SAMPLE SIZE AND SAMPLING
DESIGN
This chapter should contain: (a) a summary of the justification
for sample sizes, design and sampling areas, including any
changes to these over the period of the programme (this may
include logistical as well as scientific considerations); and(b) a summary of how well the achieved sampling matched
the proposed sampling (in terms of design and size).
For programmes with multiple objectives this should
include:(a) specification of the appropriate study areas toaddress each objective;(b) specification of the quantities of interest that need to
be determined to achieve each objective;(c) specification of the sources of uncertainty in theestimation for each quantity of interest and whichof these were functions of sample size;(d) explanation of the calculations used to determine
the optimal sampling design and sample size foreach objective (including consideration of methods
e.g. lethal and non-lethal techniques) and then how
this was integrated into the final sampling designand sample size;(e) an overview of how the achieved sampling
followed the proposed design and numbers (and an
explanation as to why if it did not); and(f) An analysis of the effect of sample size changes (ifthey occurred during the programme) on the ability
to meet objectives and sub-objectives.
Details should be provided as an Annex or Annexes as
described in a later section.

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4. A CHAPTER FOR EACH OBJECTIVECONTAINING:
These should be self-contained2 to the extent possible andcontain sufficient levels of detail (first with sections by subobjective if appropriate and then integrated over the main
objective) to allow a review of:(a) the field methods (and difficulties encountered –any uncertainty arising from this should be covered
under (c) below);(b) the laboratory methods (and difficulties encounteredany uncertainty arising from this should be covered
under (c) below);(c) use of data from other projects or programmes (andany uncertainty arising from this – which should
also be covered under (c) below);(d) the analytical methods (including an explanation ofassumptions, key parameters, how uncertainty was
accounted for);(e) the results;(f) a discussion of the importance of the results(including caveats about conclusions that can be
drawn) and how these add to and/or compare withrelated research from other regions; and(g) an evaluation (for the overall objective) of theresults in light of the topics covered by Annex P3.

ANNEXESThe Final Report should include3 a number of Annexes
including the following.

5. ADDITIONAL RESEARCHThis chapter should contain a summary of any results
and studies that were completed that used data from theprogramme but was not addressing the objectives of the
programme itself.

(6) A list (by objective) of collaborating institutes, expert,
projects or external data sources.
(7) A list (by objective, or for other research, topic) of
published papers that use data from the programme(copies should be archived with the IWC Secretariat).
(8) A list (by objective, or for other research, topic) of
working papers that use data from the programmethat have been presented at international meetings,
including the IWC Scientific Committee (copies should
be archived with the IWC Secretariat).

6. CONCLUSIONSThis should include at least an evaluation for the programme
as a whole in the light of the topics covered by Annex P3 plus
consideration of any other scientific issues that arose fromthe programme.

i.e. Contain a sufficient level of detail that the reader does not have to frequently consult other material to evaluate the work – similar to the level ofdetail provided in a published paper. If a programme has already publishedpapers in peer-reviewed journals comprising all or most of its results thesechapters can be made by the sum of these papers with a short introduction
and an overall conclusion.

(1) Field protocols (and if relevant how these compare with
IWC guidelines).
(2) Laboratory protocols (and if relevant how these comparewith IWC guidelines).
(3) A list of samples and data collected, and samples
analysed by technique.
(4) Analytical details for new approaches or models(including formulae for estimating parameters ofinterest and how uncertainty was dealt with).
(5) For each year (and season if appropriate):(a) the predetermined tracklines for sampling andsightings surveys and the rationale for those lines;(b) the actual coverage of those tracklines and therationale for any decisions taken to deviate from thepredetermined lines including the rationale for any
new lines developed; and(c) an evaluation of how representative the realised
samples may be of the study area and the biologicalpopulations involved.

2

It is assumed that the report will be in electronic format so: (a) links canbe given; and (b) that much of this information will have been developedby the proponents at the start of the programme anyway (e.g. protocols).

3

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587

Annex H
List of Publications Arising out of JARPN II or
Using JARPN II Data
PEER-REVIEWED PRODUCTION OF JARPN II –MAIN OBJECTIVES
1. Kanda, N., Goto, M. and Pastene, L.A. 2006. Genetic
characteristics of western North Pacific sei whales,
Balaenoptera borealis, as revealed by microsatellites.
Mar. Biotechnol. 8:86-93.
2. Kanda, N., Goto, M., Kato, H., McPhee, M.V. andPastene, L.A. 2007. Population genetic structure of
Bryde’s whales (Balaenoptera brydei) at the interoceanic and trans-equatorial levels. Cons. Genet. 8:853864.
3. Kim, E.Y., Iwata, H., Fujise, Y. and Tanabe, S. 2004.
Searching for novel CYP members using cDNA libraryfrom a minke whale liver. Mar. Environ. Res. 58: 495498.
4. Konishi, K. and Tamura, T. 2007. Occurrence of theminimal armhook squids Berryteuthis anonychus(Cephalopoda: Gonatidae) in the stomachs of common
minke whales Balaenoptera acutorostrata in thewestern North Pacific. Fish. Sci. 73: 1208-1210.
5. Konishi, K., Tamura T., Isoda, T. Okamoto, R.,
Hakamada, T., Kiwada, H. and Matsuoka, K. 2009.
Feeding strategies and prey consumption of three baleen
whale species within the Kushiro-Current Extension. J.
Northwest Atl. Fish. Sci. 42: 27-40.
6. Maeda, H., Kawamoto, T. and Kato, H. 2013. A study on
the improvement of age estimation in common minke
whales using the method of gelatinized extraction of
earplug. NAMMCO Sci. Publ. 10:17pp. SC/F16/JR52.
7. Maeda, H.and Kato, H. 2012. Technical improvementsof the age character (earplug) collections in common
minke whales. Bull. Jpn. Soc. Fish. Oceanogr. 76(2):59-65 (in Japanese with English abstract).
8. Matsukura, R., Yasuma, H., Murase, H., Yonezaki,
S., Funamoto, T., Honda, S. and Miyashita, K. 2009.
Measurements of density contrast and sound-speed
contrast for target strength estimation of Neocalanus
copepods (Neocalanus cristatus) in the North Pacific
Ocean. Fish. Sci. 75: 1377-87.
9. Matukura, R., Sawada, K., Abe, K., Minami, K.,
Nagashima, H., Yonezaki, S., Murase, H., Miyashita,
K. 2013. Comparision of measurements and model
calculations of target strength of juvenile sandeel in
Sendai Bay. Nippon Suisan Gakkaishi 79: 638-48 (inJapanese with English abstract).
10. Murase, H., Hakamada, T., Matsuoka, K., Nishiwaki,
S., Inagake, D., Okazaki, M., Toji, N. and Kitakado,
T. 2014. Distribution of sei whales (Balaenoptera
borealis) in the subarctic-sub-tropical transition area of
the western North Pacific in relation to oceanic fronts.
Deep-Sea Res. II 107: 22-28. SC/F16/JR8.
11. Murase, H., Ichihara, M., Yasuma, H., Watanabe,
H., Yonezaki, S., Nagashima, H., Kawahara, S. andMiyashita, T. 2009. Acoustic characterization of
biological backscatterings in the Kuroshio-Oyashio
inter-frontal zone and subarctic waters of the western
North Pacific in spring. Fish. Oceanog. 18(6): 386-401.

12. Murase, H., Kawabata, A., Kubota, H., Nakagami, M.,
Amakasu, K., Abe, K., Miyashita, K. And Oozeki, Y.
2012. Basin-scale distribution pattern and biomass
estimation of Japanese anchovy Engraulis japonicas in
the western North Pacific. Fish. Sci. 78: 761-773. SC/
F16/JR18.
13. Murase, H., Kawabata, A., Kubota, H., Nakagami,
M., Amakasu, K., Abe, K., Miyashita, K. and Oozeki,
Y. 2011. Effect of depth-dependent target strength onbiomass estimation of Japanese anchovy. Mar. Sci.
Technol. 19: 267-72.
14. Murase, H., Nagashima, H., Yonezaki, S., Matsukura,
R. And Kitakado, T. 2009. Application of a generalized
additive model (GAM) to reveal relationships between
environmental factors and distributions of pelagic fishand krill: a case study in Sendai Bay, Japan. ICES J.
Mar. Sci. 66(6): 1417-24.
15. Murase, H., Tamura, T., Kiwada, H., Fujise, Y.,
Watanabe, H., Ohizumi, H., Yonezaki, S., Okamura,
H. and Kawahara, S. 2007. Prey selection of common
minke (Balaenoptera acutorostrata) and Bryde’s(Balaenoptera edeni) whales in the western North
Pacific in 2000 and 2001. Fish. Oceanogr. 16 (2): 186201. SC/F16/JR21.
16. Murase, H., Tamura, T., Otani, S. and Nishiwaki, S.
2016. Satellite tracking of Bryde’s whales Balaenoptera
edeni in the offshore western North Pacific in summer
2006 and 2008. Fish. Sci. 82:35-45. SC/F16/JR45.
17. Nakamura, G., Kadowaki, I., Nagatuka, S., Fujise, Y.,
Kishiro, T. and Kato, H. 2014. Variation in a colour
pattern of white patch on the flippers of North Pacificcommon minke whales: Potential application for theirinteroceanic difference. La Mer 52: 31-47.
18. Niimi, S., Imoto, M., Kunisue, T, Watanabe, M.X., Kim,
E., Nakayama, K., Yasunaga, G., Fujise, Y., Tanabe, S.
and Iwada, H. 2014. Effects of persistent organochlorine
exposure on the liver transcriptome of the common
minke whale (Balaenoptera acutorostrata) from theNorth Pacific. Ecotoxicology and Environmental Safety
108: 95-105. SC/F16/JR36.
19. Niimi S., Kim E.-Y., Iwata, H., Watanabe, M.X.,
Yasunaga, G., Fujise, Y. and Tanabe, S. 2007.
Identification and hepatic expression profiles ofcytochrome P450 1-4 isozymes in common minke
whales (Balaenoptera acutorostrata). Comp. Biochem.
Physiol. B Biochem. Mol. Biol. 147(4):667-81.
20. Niimi, S., Watanabe, M.X., Kim, E.-Y., Iwata, H.,
Yasunaga, G., Fujise, Y. and Tanabe, S. 2005. Molecularcloning and mRNA expression of cytochrome P4501A1
and 1A2 in the liver of common minke whales(Balaenoptera acutorostrata). Mar. Pollut. Bull. 51(812):784-93.
21. Ohishi, K., Fujise, Y. and Maruyama, T. 2008. Brucellaspp. in the western North Pacific and Antarctic
cetaceans: a review. J. Cetacean Res. Manage. 10(1):67-72.

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22. Ohishi. K., Takishita, K., Kawato, M., Zenitani, R.,
Bando, T., Fujise, Y., Goto, Y., Yamamoto, S. andMaruyama, T. 2004. Molecular evidence of new variant
Brucella in North Pacific common minke whales.
Microbes and Infection 6: 1199-1204.
23. Ohishi, K., Takishita, K., Kawato, M., Zenitani, R,
Bando, T., Fujise, Y., Goto, Y., Yamamoto, S. andMaruyama, T. 2005. Chimeric structure of omp2of Brucella from Pacific common minke whales(Balaenoptera acutorostrata). Microbiol. Immunol.
49(8):789-93.
24. Ohishi, K., Maruyama, T., Ninomiya, A., Kida, H.,
Zenitani, R., Bando, T., Fujise, Y., Nakamatsu, K.,
Miyazaki, N. and Boltunov, A. N. 2006. Serologic
investigation of influenza a virus infection in cetaceansfrom the western North Pacific and the Southern
Oceans. Mar. Mamm. Sci. 22(1): 214-221.
25. Ohishi, K., Zenitani, R., Bando, T., Goto, Y., Uchida,K., Maruyama, T., Yamamoto, S., Miyazaki, N. andFujise, Y. 2003. Pathological and serological evidenceof Brucella-infection in baleen whales (Mysticeti) in
the western North Pacific. Comp. Immunol. Microbiol.
Infect. Dis. 26(2):125-36.
26. Sasaki, H., Murase, H., Kiwada, H., Matsuoka, K.,
Mitani, Y. and Saitoh, S. 2013. Habitat differentiation
between sei (Balaenoptera borealis) and Bryde’s
whales (B. brydei) in the western North Pacific. Fish.
Oceanogr. 22 (6): 496-508. SC/F16/JR10.
27. Shimizu, Y., Ohishi, K., Suzuki, R., Tajima, Y.,
Yamada, T., Kakizoe, Y., Bando, T., Fujise, Y., Taru,
H., Murayama, T. and Maruyama, T. 2013. Amino acidsequence variations of signalling lymphocyte activation
molecule and mortality caused by morbillivirus
infection in cetaceans. Microbiol. Immunol. 57:624-32.
SC/F16/JR37.
28. Shishido, R., Ohishi, K., Suzuki, R., Takishita, K.,
Ohtsu, D., Okutsu, K., Tokutake, K., Katsumata, E.,
Bando, T., Fujise, Y., Murayama, T. and Maruyama,
T. 2010. Cetacean Toll-like receptor 4 and myeloid
differentiation factor 2, and possible cetacean-specificresponses against Gram-negative bacteria. Comp.
Immunol. Microbiol. Infect. Dis. 33(6): 89-98.
29. Tojo, N., Matsukura, R., Yasuma, H., Yonezaki, S.,
Watanabe, H., Kawahara, S., Murase, H., and Miyashita,
K. 2010. Spatial analysis of isada krill (Euphausia
pacifica) distribution in frontal environments in the
North Pacific Ocean. GIS/spatial analyses in fisheries
and aquatic science. 4th International Fishery GISSociety 115-38.
30. Watanabe, H., Okazaki, M., Tamura, T., Konishi, K.,
Inagake, D., Bando, T., Kiwada, H. and Miyashita, T.
2012. Habitat and prey selection of common minke, sei,and Bryde’s whales in mesoscale during summer in thesubarctic and transition regions of the western North
Pacific. Fish. Sci. 78:557-67. SC/F16/JR20.
31. Zharikov, K., Fujise, Y., Tamura, T., Kiwada, H.,
Bando, T., Konishi, K. and Isoda, T. 2004. Distribution
and feeding of large cetaceans based on the data of
JARPN II - 2003 May, 17-July, 9. Marine Mammals of
the Holarctic 2004: 207-210.
PEER-REVIEWED PRODUCTION OF JARPN II –OTHERS RESEARCH FIELDS
1. Bhuiyan, M.M.U., Suzuki, Y., Watanabe, H., Hirayama,
H., Matsuoka, K., Fujise, Y., Ishikawa, H., Ohsumi, S.
and Fukui, Y. 2008. Attempts at in vitro fertilization andculture of in vitro matured oocytes in sei (Balaenoptera
borealis) and Bryde’s (B. edeni) whales. Zygote 16:
DOI 10.1017/S0967199408004887.

2.

Bhuiyan, M.M.U., Suzuki, Y., Watanabe, H., Lee, E.,
Hirayama, H., Matsuoka, K., Fujise, Y., Ishikawa, H.,
Ohsumi, S. and Fukui, Y. 2010. Production of Sei Whale(Balaenoptera borealis) Cloned Embryos by Inter- andIntra-Species Somatic Cell Nuclear Transfer. J. Reprod.
Dev. 56: 131-9.
3. Birukawa, N., Ando, H., Goto, M., Kanda, N., Pastene,
L.A., Nakatsuji, H., Hata, H. and Urano, A. 2005.
Plasma and Urine Levels of Electrolytes, Urea andSteroid Hormones Involved in Osmoregulation of
Cetaceans. Zool. Sci. 22: 1245-1257.
4. Birukawa, N., Ando, H., Goto, M., Kanda, N., Pastene,
L.A. and Urano, A. 2008. Molecular cloning of urea
transporters from the kidneys of baleen and toothed
whales. Comp. Biochem. Physiol. Part B 149: 227-35.
5. Fujita, H., Honda, K., Hamada, N., Yasunaga, G.
and Fujise, Y. 2009. Validation of high-throughput
measurement system with microwave-assisted
extraction, fully automated sample preparation device,and gas chromatography-electron capture detector for
determination of polychlorinated biphenyls in whale
blubber. Chemosphere 74:1069-1078.
6. Fukui, Y., Iwayama, H., Matsuoka, T., Nagai, H., Koma,
N., Mogoe, T., Ishikawa, H., Fujise, Y., Hirabayashi,
M., Hochi, S., Kato, H. and Ohsumi, S. 2007. Attempt
at Intracytoplasmic sperm injection of in vitro matured
oocytes in common minke whales (Balaenoptera
acutorostrata) captured during the Kushiro coast
survey. J. Reprod. Dev. 53(4): 945-952.
7. Funasaka, N., Yoshioka, M. and Fujise, Y. 2010. Featuresof the ocular harderian gland in three Balaenopterid
species based on anatomical, histological andhistochemical observations. Mammal Study 35:9-15.
8. Glover, K.A., Kanda, N., Haug, T., Pastene, L.A., Oien,
N., Goto, M., Seliussen, B.B. and Skaug, H.J. 2010.
Migration of Antarctic minke whales to the Arctic.
PLoS ONE 5 (12):9. Glover, K.A., Kanda, N., Haug, T., Pastene, L.A., Oien,
N., Seliusen, B.B., Sorvik, A.G.E. and Skaug, H.J. 2013.
Hybrids between common and Antarctic minke whales
are fertile and can back-cross. BMC Genetics 14:25.
10. Hayashi, K., Nishida, S., Yoshida, H., Goto, M., Pastene,
L.A. and Koike, H. 2003. Sequence variation of the DQB
allele in the cetacean MHC. Mammal Study 28: 89-96.
11. Hiwasa, M., Suzuki, Y., Watanabe, H., Bhuiyan,
M.M.U., Matsuoka, K., Fujise, Y., Ishikawa, H.,
Ohsumi, S. and Fukui, Y. 2009. Effects of semen
extenders and storage temperatures on characteristics
of frozen-thawed Bryde’s (Balaenoptera edeni) whale
spermatozoa. J. Reprod. Dev. 55: 599-606.
12. Inoue, S., Kishiro, T., Fujise, Y., Nakamura, G. andKato, H. 2014. Seasonal changes in the testis of theNorth Pacific common minke whale. Nippon Suisan
Gakkaishi 80 (2): 185-90 (in Japanese with English
abstract).
13. Iwanami, K., Mita, H., Yamamoto, Y., Fujise, Y.,
Yamada, T. and Suzuki, T. 2006. cDNA-derived aminoacid sequences of myoglobins from nine species ofwhales and dolphins. Comp. Biochem. Physiol. Part B
Biochem. Mol. Biol. 45:249-56.
14. Kitayama, C., Sasaki, M., Ishikawa, H., Mogoe,T., Ohsumi, S., Fukui, Y., Budipitojo, T., Kondoh,
D. and Kitamura, N. 2015. Structure and functionsof the placenta in common minke (Balaenoptera
acutorostrata), Bryde’s (B. brydei) and sei (B. borealis)
whales. J. Reprod. Dev. 61(5): 415-421.

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017

15. Lee, E., Bhuiyan, M.M.U., Watanabe, H., Matsuoka, K.,
Fujise, Y., Ishikawa, H. and Fukui. Y. 2009. Productionof cloned sei whale (Balaenoptera borealis) embryos
by interspecies somatic cell nuclear transfer using
enucleated pig oocytes. J. Vet. Sci. 10(4): 285-92.
16. Miyakawa, N. and Kato, H. 2015.Morphology ofinnominate bones (pelvic bones) of Bryde’s whales
Balaenoptera edeni. Honyuurui Kagaku 55(1):1-10 (inJapanese with English abstract).
17. Nakamura, G. and Kato, H. 2014. Developmental
changes in the skull morphology of common minke
whales Balaenoptera acutorostrata. J. Morphol.
doi:10.1002/jmor.20288.
18. Nakamura, G., Kato, H. and Fujise, Y. 2012. Relative
Growth of the Skull of the Common Minke Whale
Balaenoptera acutorostrata from the North Pacific in
Comparison to Other Balaenoptera Species. Mammal
Study 37(2): 105-112.
19. Nikaido, M., Hamilton, H., Makino, H., Sasaki, T.,
Takahashi, K., Goto, M., Kanda, N., Pastene, L.A. andOkada, N. 2006. Baleen whale phylogeny and a pastextensive radiation event revealed by SINE insertion
analysis. Mol. Biol. Evol. 23 (5):866-873.
20. Nishida, S., Pastene, L.A., Goto, M. and Kanda, N.
2003. SRY gene structure and phylogeny in the cetacean
species. Mammal Study 28: 57-66.
21. Nishida, S., Goto, M., Pastene, L.A., Kanda, N. andKoike, H. 2007. Phylogenetic relationships amongcetaceans revealed by Y-chromosome sequences. Zool.
Sci. 24(7): 723-732.
22. Ogawa, G., Ishida, M., Kato, H., Fujise, Y. andUrano, N. 2010. Identification of facultative anaerobic
bacteria isolated from the intestine of the minke whale
Balaenoptera acutorostrata by 16S rRNA sequencing
analysis. Fish. Sci. 76:177-181.
23. Pastene, L.A., Goto, M., Kanda, N., Zerbini, A.N.,
Kerem, D., Watanabe, K., Bessho, Y., Hasegawa, M.,
Nielsen, R., Larsen, F. and Palsboll, P.J. 2007. Radiationand speciation of pelagic organisms during periods ofglobal warming: the case of the common minke whale,
Balaenoptera acutorostrata. Mol. Ecol. 16 (7): 14811495.

589

24. Pastene, L.A., Acevedo, J., Goto, M., Zerbini, A.N.,
Acuna, P. and Aguayo-Lobo, A. 2010. Population
structure and possible migratory links of common
minke whales, Balaenoptera acutorostrata, in theSouthern Hemisphere. Conserv. Genet. 11:1553-1558.
25. Sasaki, T., Nikaido, M., Hamilton, H., Goto, M., Kato,
H., Kanda, N., Pastene, L.A., Cao, Y., Fordyce, R.E.,
Hasegawa, M. and Okada, N. 2005. Mitochondrial
Phylogenetics and Evolution of Mysticete Whales.
System. Biol. 54(1): 77-90.
26. Suzuki, Y., Umatani, M., Bhuiyan, M.M.U., Watanabe,H., Mogoe, T., Matsuoka, K., Fujise, Y., Ishikawa, H.,
Ohsumi, S., Sasaki, M. and Fukui, Y. 2010. Effects ofequilibration steps, type of sugars and addition of whale
follicular fluid on viability and in vitro maturation of
vitrified whale oocytes. Japanese J. Zoo Wildl. Med.
15(2): 65-72.
27. Tsuji, K., Sato, A., Kaneko, H., Yasunaga, G., Fujise, Y.
and Nomata, H. 2009. Comparison of physiologicallysignificant imidazole dipeptides in cetaceans sampled
in Japanese Whale research (short paper). Sci. Rep.
Hokkaido Fish. Exp. Stn. 74: 25-8. 50.
28. Urashima, T., Kobayashi, M., Asakuma, S., Uemura,Y., Arai, I., Fukuda, K., Saito, T., Mogoe, T., Ishikawa,
H., Fukui, Y. 2007. Chemical characterization of theoligosaccharides in Bryde’s whale (Balaenoptera
edeni) and sei whale (Balaenoptera borealis) milk.
Comp. Biochem. Physiol. Part B 146: 153-159.
29. Watanabe, H., Mogoe, T., Asada, M., Hayashi, K.,
Fujise, Y., Ishikawa, H., Ohsumi, S., Miyamoto, A.
and Fukui, Y. 2004. Relationship between serum sex
hormone concentrations and histology of seminiferous
tubules of captured baleen whales in the western North
Pacific during the feeding season. J. Reprod. Dev.
50(4): 419-427.
30. Watanabe, H., Tateno, H., Kusakabe, H., Matsuoka,
T., Kamiguchi, Y., Fujise, Y., Ishikawa, H., Ohsumi, S.
and Fukui, Y. 2007. Fertilizability and chromosomal
integrity of frozen-thawed Bryde’s whale (Balaenoptera
edeni) spermatozoa intracytoplasmically injected into
mouse oocytes. Zygote 15: 9-14.

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Annex I
Annex

A Summary
byJARPN
JARPNIIII(2000-14)(2000-14)
A summaryofofSamples/Data
samples/data Collectedcollected bySome samples and data obtained by JARPN II are not related to the main research objectives of JARPN II or to other main research need, and these items are
not listed here but they are available for research collaboration with ICR under data access protocols of ICR (http://www.icrwhale.org/pdf/appendix2.pdf), outside
the context of the IWC SC review workshop. The tables below show the research items and sample sizes by each item for the period 2000-14.
I-1. Sighting data - Offshore components (SV+SSV)

II-1. Biological data - common minke whale (Offshore component)

Data

Sample size

Weather data (no. observations)
Effort data (searching distance [n.miles])Sighting data (no. of school)
Angle and distance experiments
Photo-ID blue whales (no. of schools photographed)
Photo-ID humpback whales (no. of schools photographed)
Photo-ID North Pacific right whales (no. of schools
photographed)

85,219202,403
15,594
6,031
107
65
50

I-2. Sighting data - Coastal components (Sanriku) (SV)Data
Weather data (no. observations)
Effort data (searching distance [n.miles])Sighting data (no. of school)
Angle and distance experiments

Sample size
1,008
3,048
110
88

I-3. Sighting data - Coastal component (Kushiro SV)
Data
Weather data (no. observations)
Effort data (searching distance [n.miles])Sighting data (no. of school)
Angle and distance experiments

Sample size
2,150
5,155
353
112

I-4. Sighting data - Coastal component (Sanriku) (SSV)
Data
Weather data (no. observations)
Effort data (n.miles)
Sighting data (no. of school)

Sample size
4,769
32,665642

I-5. Sighting data - Coastal components (Kushiro) (SSV)
Data
Weather data (no. observations)
Effort data (n.miles)
Sighting data (no. of school)

Sample size
4,847
33,378985

Note 1=Sighting data (no. of school) are on baleen, sperm, and killer
whales. Note 2=Sighting data of the coastal components surveyed by SSVs
were not obtained by strict line transect surveys. Note 3=Sighting data on
sei and Bryde’s whales obtained during IWC/POWER would be available
depending on progress of data validation by the IWC Secretariat.

Sample size
Male Female Total
861
121
982
861
121
982
861
121
982
861
121
982
847
119
966
861
121
982
861
121
982
861
121
982
861
121
982
180
29
209
861
121
982
861
121
982
861
121
982
15
15
106
106
121
121
861
861
861
121
982
861
121
982
861
121
982
861
121
982
16
51
13
409
55
464
546
0
546
680
0
680
PCBs, DDTs, HCB, HCHs/CHLs concentrations (blubber)
50
5I-131, Cs-134 and Cs-137 concentrations (muscle)82
10
Mitochondrial DNA control region sequences
855
121
976
Nuclear DNA microsatellite (16 loci)
855
121
976
*Analysis of samples is ongoing. **2000-13.
Data and sample
Sampling date
Sampling location
Body length
Body proportion (19 measurements)
Skull (length and width)
Body scar record
Record of external body characters
Sex
Body weight
Organ weight
Blubber thickness (five points)
Girth
Maturity stage
Corpora albicantia/lutea (presence/absence only)
Corpora albicantia/lutea (presence/absence/no.)Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weight
Main prey species in stomach contents
Freshness of stomach contents
Energy contents of prey species
Foetus number, sex, body length, body weight
Aspartic acid isomers ratios (lens of foetus)*
Age (from ear plug)**
Total PCB concentrations (blubber)
Total Hg concentrations (muscle)

II-2. Biological data - sei whale (Offshore component)
Sample size
Male Female Total
Data and sample
Sampling date551
623 1,174
Sampling location
551
623 1,174
Body length551
623 1,174
Body proportion (19 measurements)
551
623 1,174
Skull (length and width)534
603 1,137
Sex
551
623 1,174
Body weight551
623 1,174
Organ weight
77
104
181
Blubber thickness (five points)
551
623 1,174
Girth551
623 1,174
Maturity stage
551
623 1,174
Corpora albicantia/lutea (presence/absence only)
205
205
Corpora albicantia/lutea (presence/absence/no.)418
418
Lactation condition
623
623
Testis weight551
551
Stomach contents (IWS format)
551
623 1,174
Stomach contents weight551
623 1,174
Main prey species in stomach contents
551
623 1,174
Freshness of stomach contents
551
623 1,174
Energy contents of prey species
20
Prey species est. by next generation sequencing (NGS) 8
2
10
Behavioural data using acoustic tags
2
Foetus number, sex, body length, body weight366
Age (from ear plug)*
335
348
683
Total Hg concentrations (muscle)
160
160
PCBs, DDTs, HCB, HCHs/CHLs concentrations (blubber)
50
5I-131, Cs-134 and Cs-137 concentrations (muscle)
12
13
25
Mitochondrial DNA control region sequences
551
622 1,173
Nuclear DNA microsatellite (16 loci)551
623 1,174*2002-13.

591

J. CETACEAN RES. MANAGE. 18 (SUPPL.), 2017
II-3. Biological data - Bryde’s whale (Offshore component)

II-5. Biological data - common minke whale (Coastal component - Sanriku)

Sample size

Sample size

Data and sample

Male

Sampling date
Sampling location
Body length
Body proportion (19 measurements)
Skull (length and width)
Sex
Body weight
Organ weight
Blubber thickness (five points)
Girth
Maturity stage
Corpora albicantia/lutea (presence/absence only)
Corpora albicantia/lutea (presence/absence/no.)Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weight
Main prey species in stomach contents
Freshness of stomach contents
Energy contents of prey species
Prey species estimating by next generation
sequencing (NGS)
Foetus number, sex, body length, body weight
Total Hg concentrations (muscle)
PCBs, DDTs, HCB, HCHs and CHLs
concentrations (blubber)
I-131, Cs-134 and Cs-137 concentrations (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (16 loci)

289
289
289
289
278
289
289
60
289
289
289
289
289
289
289
289
2

391
391
391
391
375
391
391
77
391
391
391
87
304
391
391
391
391
391
4

680
680
680
680
653
680
680
137
680
680
680
87
304
391
289
680
680
680
680
13
6

49
5

0
0

169
49
5

6
284
289

7
387
391

13
671
680

Female Total

II-4. Biological data - sperm whale (Offshore component)

Male Female

Total

Sampling date
Sampling location
Body length
Body proportion (19 measurements)
Skull (length and width)
Body scar record
Sex
Body weight
Organ weight
Blubber thickness (five points)
Girth
Maturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weight*
Main prey species in stomach contents
Freshness of stomach contents
Energy contents of prey species
Prey species estimating by next generation
sequencing (NGS)
Foetus number, sex, body length, body weight
Age (from ear plug)**
Aspartic acid isomers ratios (lens)***Total Hg concentrations (muscle)
Total Hg concentrations (liver)
PCBs, DDTs, HCB, HCHs and CHLs
concentrations (blubber)
I-131, Cs-134 and Cs-137 concentrations(muscle)Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (16 loci)

221
221
221
221
215
221
221
221
7
221
221
221
219
221
205
221
221
1

295
295
295
295
286
295
295
295
10
295
295
295
295
295
295
281
295
295
2

516
516
516
516
501
516
516
516
17
516
516
516
295
295
219
516
486
516
516
3
3

94
46
195
46
5

123
48
48
48
0

44
217
94
280
94
5

7

6

13

221
221

295
295

516
516

*2003-13. Analysis of 2014 samples is ongoing. **2003-13. ***Analysis
of samples is ongoing.

Sample size
Data and sample

Male

Sampling date
Sampling location
Body length
Body proportion (18 measurements)
Skull (length and width)
Sex
Body weight
Organ weight
Blubber thickness (11 points)
Girth
Maturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weight
Main prey species in stomach contents
Freshness of stomach contents
Energy contents of prey species
Foetus number, sex, body length, body weight
Age (from tooth)*Total Hg concentrations (muscle)
PCBs, DDTs, HCB, HCHs and CHLs
concentrations (blubber)
I-131, Cs-134 and Cs-137 concentrations (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (15 loci)

16
16
16
16
16
16
16
10
16
16
13
16
16
16
16
16
4
1
1

40
40
40
40
38
40
40
26
40
40
40
40
40
40
40
40
40
19
4
2

56
56
56
56
54
56
56
36
56
56
53
40
40
16
56
56
56
56
10
10
23
5
3

1
16
16

2
40
40

3
56
56

*2000-13.

Data and sample

Female Total
II-6. Biological data - common minke whale (Coastal component - Kushiro)Sample size
Data and sample

Male

FemaleTotal

Sampling date
Sampling location
Body length
Body proportion (19 measurements)
Skull (length and width)
Body scar record
Sex
Body weight
Organ weight
Blubber thickness (five points)
Girth
Maturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weight*
Main prey species in stomach contents
Freshness of stomach contents
Energy contents of prey species
Prey species estimating by next generation
sequencing (NGS)
Foetus number, sex, body length, body weight
Age (from ear plug)**
Total Hg concentrations (muscle)
PCBs, DDTs, HCB, HCHs and CHLs
concentrations (blubber)
I-131, Cs-134 and Cs-137 concentrations(muscle)Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (16 loci)

438
438
438
438
430
438
438
438
17
438
438
438
438
438
403
438
438
3

219
219
219
219
215
219
219
219
8
219
219
219
219
219
219
203
219
219
3

657
657
657
657
645
657
657
657
25
657
657
657
211
211
429
657
606
657
657
3
6

172
377
5

78
0
0

14
250
377
5

12

13

25

438
438

219
218

657
656

*2002-13. Analysis of 2014 samples is ongoing. **2002-13.

592

REPORT OF THE EXPERT PANEL OF THE FINAL REVIEW OF JARPNII

III. Pollutant data (environmental and prey species samples) - Offshore
components
Data and sample

Sample size

Total Hg concentrations (krill)
Total Hg concentrations (fishes)8
19

IV-1. Oceanographic data - Offshore components

IV-3. Oceanographic data - Coastal components (Kushiro)
Data and sample

Sample size

Temperature and salinity (CTD survey)
Midwater trawl (no. of hawls)
IKMT (no. of hawls)
V-1. Genetic data - North Pacific right whale

Data and sample

Sample size

Data and sample

Temperature and salinity (XCTD survey)
Temperature and salinity (CTD survey)
Midwater trawl (no. of hawls)
MOTH trawl (no. of hawls)
MOCNESS (no. of hawls)
IKMT (no. of hawls)
NORPAC (no. of hawls)
Other nets (VMPS, Ring, BONGO) (no. of hawls)
Echo sounder (2002-07: km)
Echo sounder (2008-13: n.miles)

47
761
262
16
36
34
254
36
12,838
8,098

Mitochondrial DNA sequences

Others nets: VMPS 12, ring net 8, BONGO net 16. Note 4=2008 cruise:
Shunyo-Maru and Kaiko-Maru; 2009, 2012, 2013 cruises: Shunyo-Maru;
2010, 2011 cruises: Hokko-Maru.
IV-2. Oceanographic data - Coastal components (Sanriku)
Data and sample
Temperature and salinity (XCTD survey)
Temperature and salinity (CTD survey)
Midwater trawl (no. of hawls)
Bongo net (no. of hawls)
IKMT (no. of hawls)
Sampling by fishing (no. of stations)
Echo sounder (2005 and 2006 seasons: km)
Echo sounder (2008 and 2009 seasons: n.miles)

109
133
6

Sample size
11
325
109
5
17
2
277.5
354.3

Sample size
20

Note 5=Data of some items for common minke whales are also available
for the JARPN period (1994-99), which were reviewed by the IWC SC in
2000 (IWC, 2001).
Note 6=Genetic data from other sources have been used to complement theprevious genetic analyses on stock structure of baleen whales. These data
would be also available for the review.
REFERENCE
International Whaling Commission. 2001. Report of the ScientificCommittee. J. Cetacean Res. Manage. (Suppl.) 3: 1-76.