J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

405

Report of the Expert Workshop to Review the Ongoing
JARPN II Programme*
The Workshop was held at the National Research Instituteof Fisheries Science (NRIFS) in Yokohama, Japan from 2630 January 2009. The list of Participants is given as
Annex A.
1. INTRODUCTORY ITEMS
1.1 Welcome and opening remarks
Participants were welcomed to NRIFS by its Director, Dr
Kiyoshi Inouye.
Bjørge, the Chair of the IWC Scientific Committee,
welcomed the participants and explained that this was thefirst review meeting of a special research permit programme
to be held under the new rules developed by the IWC
Scientific Committee and approved by the Commission(IWC, 2009b), a summary of which are given as Annex D
to this report. As such, the Workshop was not only
important because of the subject matter, but also because it
was an opportunity to test the new procedure and, if
necessary, to suggest future improvements. He noted that
this was a review of the first six years of an ongoing
research programme. The primary tasks of the Workshop
were to review: (1) the scientific work undertaken thus far
against the stated objectives of the programme and toreview future plans in the context of the likelihood of
meeting those objectives; (2) the techniques used (lethal andnon-lethal); (3) appropriate sample sizes; and (4) the effects
of any catches on the relevant stocks. Where appropriate theWorkshop would highlight those aspects of the programmethat were of most direct relevance to the work of the IWC
Scientific Committee.
1.2 Election of Chair
Bjørge was elected Chair.
1.3 Appointment of rapporteurs
Donovan co-ordinated the production of the report andindividual experts acted as rapporteurs for individual
agenda items, including Butterworth, Cooke, Forcada, Hall,
Reilly and Waples with assistance from others as
appropriate.
1.4 Meeting procedure and time schedule
Pastene outlined the logistical arrangements for the meeting.
Bjørge explained that the format would be as follows:
during the first part of the day, plenary sessions would beheld in which the Japanese scientists would present the
papers on particular agenda items and there would be an
opportunity for the expert panel (hereafter the ‘Panel’) to*Presented to the meeting as SC/61/Rep1.

ask questions of clarification and substance regarding thework that had been undertaken or further work that was
expected to be undertaken. Once this was completed, thePanel would meet in closed session to discuss its
conclusions and begin to draft the report on the individual
items. The objective was to try to leave the Workshop withas complete a report as possible. The draft report would beshown to the Japanese scientists for the purposes ofcommenting on whether there were any technicalmisunderstandings in the text. The report and its
conclusions were the sole responsibility of the Panel. Oncethe report is completed, the process will follow that outlined
in Annex D before submission to the Annual ScientificCommittee meeting in Madeira.
2. ADOPTION OF THE AGENDA
The adopted Agenda is given as Annex B.
The report follows the agreed structure, i.e. a review of
each of the objectives. However, we have also added ageneral item on abundance estimation (Item 7.4) in addition
to more specific comments under the relevant agenda items.
Abundance data have been collected during the course of
the JARPN II research programme. While the estimation ofabundance per se is not a primary objective of the research
programme, the abundance data are required, in conjunction
with data on the diet and body mass of whales, to estimate
the numbers, biomass and energy requirements of whales by
area and season for inclusion in ecosystem models thatelucidate quantitatively the role of whales in the ecosystem.
Abundance estimates are also required to assess theeffects of JARPN II catches on the stocks of whales, inaccordance with the Scientific Committee’s mandate. Thiscan involve abundance data collected outside the JARPN II
research area (see Item 9.3).
Abundance data are also important for the interpretationof data pertaining to stock structure: to infer exchange rates
between subareas from genetic or other data, the relative
size of the putative subpopulations needs to be taken into
account. This aspect is especially important in theconditioning of stock structure hypotheses for
Implementation Simulation Trials.
While sightings from catcher vessels were also recorded,
all abundance estimations conducted in conjunction with
JARPN II were derived from data obtained from dedicatedsurvey vessels not involved in the taking of whales. As
such, this was not research involving Special Permits, andstrictly speaking did not need to be evaluated in the conductof this review.

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3. REVIEW OF AVAILABLE DATA, DOCUMENTS
AND REPORTS
3.1 Workshop documents
The list of Workshop documents (SC/J09/JR1-36) is given
in Annex C1. The Workshop participants thanked theJapanese scientists for fulfilling the difficult time
constraints imposed by the new procedure. Having all of the
documents available at the beginning of December greatly
facilitated the Panel’s work.
3.2 For information papers
Twelve published papers were also highlighted to be ofinterest to the Workshop. These were: Murase et al. (2007);
Niimi et al. (2005); Niimi et al. (2007); Kanda et al. (2007);
Kanda et al. (2006); Watanabe et al. (2004); Watanabe etal. (2007); Urashima et al. (2007); Fukui et al. (2007);
Birukawa et al. (2008); Nishida et al. (2007) and Onbe et al.
(2007).
3.3 Other available documents and data
The meeting had ready access to the relevant ScientificCommittee reports and papers. A list of the available datafrom the programme is given as Annex E. Data are
available to be requested under Procedure B of the IWC
Scientific Committee Data Availability Agreement(http://www.iwcoffice.org/sci_com/data_availability.htm).
4. REVIEW OF JARPN II RESULTS: FEEDINGECOLOGY AND ECOSYSTEM STUDIES
4.1 Formal statement of objectives as given by theGovernment of Japan (based upon SC/J09/JR1)
This component of the JARPN II programme was givenhigh priority by the Government of Japan. The broad
objectives were given as: (a) prey consumption bycetaceans; (b) prey preference of cetaceans; and (c)ecosystem modelling.
A primary motive for the feeding ecology and ecosystem
studies has been the major decrease in catches by Japanesefisheries (Government of Japan, 1999) from 12.8 million
tons in 1988 to 5.8 million tons in 2005. In 1999, theFisheries Agency of Japan announced the principles of its
policy on fisheries and an action program to implement thepolicy. Highest priority was given to science-based
management and sustainable utilisation of fisheries
resources within Japan’s EEZ. In investigating the reason
for decreasing fish resources and fish catches, a number ofpotential factors were recognised including: over-fishing; achanging of the marine environment; and the effect ofconsumption by marine mammals and other animals of fish
resources. To aid the recovery of the fish resources, theGovernment stated that investigations should be carried out
taking into account the management and sustainableutilisation of the whole ecosystem including marine
mammals. Some initial ecosystem model analyses indicatedpossible competition between cetaceans and fisheries andthat the ecosystem of the western North Pacific may beaffected on a large scale by trophic interactions and changesof fishing (Government of Japan, 2002b).
Not only in Japan, but elsewhere in the world, theprinciple of multi-species management has been discussed

by various parties, including many internationalorganisations.
At its 24th Session in 2001, COFI (FAO’s Committee on
Fisheries) agreed that the FAO should conduct studies on
the interaction between fisheries and marine mammals. Thisagreement was endorsed by the 120th Session of the FAO
Council and reaffirmed in the October 2001 Reykjavik
Declaration on Responsible Fisheries in the Marine
Ecosystem (FAO, 2001). The Reykjavik Declaration alsoaffirmed that the incorporation of ecosystem considerations
implies increased attention to predator-prey relationships
and in Point 5 of the Declaration agreed that it is important,among other things:(a) ‘To advance the scientific basis for developing
and implementing management strategies thatincorporate ecosystem considerations and which
will ensure sustainable yields while conserving
stocks and maintaining the integrity of ecosystems
and habitats on which they depend;(b) identify and describe the structure, components
and functioning of relevant marine ecosystems,
diet composition and food webs, species
interactions and predator-prey relationships, therole of habitat and the biological, physical andoceanographic factors affecting ecosystemstability and resilience; and(c) build or enhance systematic monitoring of naturalvariability and its relations to ecosystemproductivity’ (FAO, 2001).
The FAO’s current definition of an ecosystem-based
approach to management is the following:‘An ecosystem approach to fisheries strives to balance diverse societal
objectives, by taking into account the knowledge and uncertainties
about biotic, abiotic, and human components of ecosystem and theirinteractions and applying an integrated approach to fisheries within
ecologically meaningful boundaries’ (FAO, 2003).

In 2001, the IWC agreed to make the study of interactions
between whales and fish stocks a matter of priority (IWC,2002), and in 2006, the IWC Scientific Committee created a
Working Group on Ecosystem Modelling; prototype
ecosystem models from JARPN II were presented anddiscussed in 2007 (IWC, 2008c).
Under the new Japanese fisheries management regime,an ecosystem-based approach is considered important,given the diversity and dynamic inter-relationships of themarine living resources and habitats around Japan.
Understanding of the marine ecosystem around Japan needs
to be substantially strengthened to take the following factors
into account in prescribing fisheries management measures:the decrease in fisheries yields between 1988 and 2005, theexpected increase in whales around Japan since theintroduction of the moratorium, and the historical major
fluctuation of the pelagic fisheries resources in a process of
so-called ‘species replacement’ in the western North
Pacific.
JARPN II was designed to provide data with which toparameterise ecosystem models that are expected to provide
insights into the relationships between different species in
the marine ecosystem and the dynamics of the ecosystem.
This includes information on the extent of the predation ofmarine mammal populations, which will balance the needfor the effective utilisation of fisheries resources and theconservation of marine mammals (Government of Japan,

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

2002a); it is thus consistent with the Reykjavik Declaration(FAO, 2001).
The primary purpose of JARPN II is to study theinteractions between fisheries and cetaceans in the marine
ecosystem of the western North Pacific through ecosystemmodelling. The output from this study can assist in theformulation of effective ecosystem-based management
policies in the future. For this purpose, multidisciplinary,comprehensive surveys are required. The four large whale
species chosen for the study (common minke, Bryde’s, sei
and sperm whales) were chosen because they occupy
important niches in the pelagic zone of the North Pacificand because their populations are relatively abundant.
Available data from other cetacean species are incorporated
into analyses as appropriate.
4.2 JARPN II Coastal component
4.2.1 Proponents’ summary
PREY CONSUMPTION BY CETACEANS

The approach requires estimates of the number of whales ata particular area and time (SC/J09/JR8) for the two coastal
regions chosen, Sanriku (2005 and 2006) and Kushiro(2002-2007). These numbers do not represent estimates ofstock abundance of the common minke whale because thesighting surveys covered only a limited area of the stock
distribution, during a particular time of its migration tonorthern feeding grounds. Furthermore, depending ofchanging environmental factors, the number of whales
sighted in this limited area and time could be different fromyear to year. These values were then used for the estimationof prey consumption by whales using data from the feeding
ecology studies.
The stomach contents of common minke whales sampled
off Sanriku (April-May) and Kushiro (September-October)in 2002-2007 JARPN II were analysed (SC/J09/JR9). In
Sanriku region, the dominant prey species consisted of krill(Euphausia pacifica) and fishes (Japanese sandlance,
Ammodytes personatus and Japanese anchovy, Engraulisjaponicus). In the Kushiro region the dominant prey species
consisted of krill (E. pacifica), fish (Japanese anchovy,
Pacific saury, Cololabis saira and walleye pollock,
Theragra chalcogramma) and squid (Japanese flying squid,
Todarodes pacificus). SC/J09/JR9 used the stomach
contents data to estimate daily consumption for various sex
age classes of animals and then scaled this with theabundance estimates to provide annual consumption bycommon minke whales for the various prey species and thencompared this with the fisheries catch.
PREY PREFERENCES OF CETACEANS

Results of a prey preference study of common minke
whales in the coastal waters of Sanriku are given in
SC/J09/JR10. To estimate prey preference sampling surveys
of common minke whales and their prey, surveys wereconducted in the same area at the same time (April). A prey
preference index, Manly’s α, was used in the analysis.
Common minke whales fed on krill, Japanese anchovy andsandlance (adult). These are important species of local
commercial fisheries. Common minke whales showed
preference for adult sandlance. As previously reported inother regions, krill was not a preferable prey for minke
whales. Ecosystem modelling work (see SC/J09/JR14
below) suggested that changes in the form of the functional
response had a substantial effect on predation impact on

407

sandlance by minke whales. Functional response can be
estimated if long term prey preference data are available.
Results of a prey preference study of common minke
whale in the coastal waters of Kushiro are presented in
SC/J09/JR11. Results suggest that the slope water region ofless than 18°C sea surface temperature (SST) is a rich preyenvironment in both the epi- and mesopelagic zones.
Common minke whales might prefer the rich preyenvironment affected by the Oyashio not only in thecontinental shelf region where walleye pollock, Pacificsaury, and euphausiids are distributed but also in theoffshore region where Pacific saury and euphausiids are
distributed. It is suggested that immature common minke
whales prefer walleye pollock, while mature animals prefer
Pacific saury, although both frequently fed on Japanese
anchovy in some years in the area within 50 n.miles ofKushiro.
OTHER POTENTIAL INFORMATION FOR ECOSYSTEMMODELLING

Relationship between body size, maturity and feeding habitof common minke whales in the Sanriku area in spring
season was reported in SC/J09/JR12. The total number ofwhales examined was 227 (91 males and 136 females).
Three species (krill, Japanese sandlance and Japanese
anchovy were found in stomachs) of which sandlance was
the most dominant prey species, followed by anchovy. All
the whales but two were sighted in waters with a depth of
20-100m. No obvious difference was observed in theirsighting positions between males and females, immatureand mature animals, and the three prey species.
Examination of the frequency of prey species consumed bywhales of different lengths and by whales of different
sexual maturity status showed little difference.
The relationship between body size, maturity and feeding
habit of common minke whales in the Kushiro area in theautumn season was reported in SC/J09/JR13. The total
number of whales examined was 254 (182 males and 72
females). Occurrence of prey species in stomachs differedsignificantly with maturity stage. Smaller and immature
whales tend to feed on walleye pollock and krill whilst
larger and mature whales tend to feed on Pacific saury.
Japanese flying squid was consumed only by mature
whales. Japanese anchovy was equally consumed byimmature and mature whales. For the coastal waters off
Kushiro in the fall, the results suggested that migration andprey preference of common minke whales differed withmaturity stage and that on the continental shelf and slope
regions immature whales showed a greater preference forwalleye Pollock and krill than mature whales.
4.2.2 Expert Panel review of results as presented
The Panel’s views on this section are given under Item
4.3.2, given the similarity in the methods used in the coastaland offshore components.
4.3 JARPN II Offshore component
4.3.1 Proponents’ summary
PREY CONSUMPTION BY BALEEN WHALESThe approach here was similar to that for the coastal
component, i.e. a combination of abundance estimates anddata from stomach contents. It involves sampling of four
species of large whales: common minke whales, Bryde’s
whales, sperm whales and sei whales. ‘Early’ and ‘late’season data were obtained to account for migration. The

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estimates are presented in SC/J09/JR15. The estimates werealso intended as input for ecosystem models in the JARPN
II survey area.
The stomach contents of common minke, sei and Bryde’s
whales sampled in the western North Pacific from May to
September in 2000-2007 JARPN II, were analysed in
SC/J09/JR16. The main prey species of common minke
whale consisted of one copepod, two krill, two squids andeight fish. The main prey species of sei whale consisted of
two copepods, three krill and four fish. The main preyspecies of Bryde’s whale consisted of five krill, one squidand four fish. There were seasonal and geographicalchanges of prey species. The total prey consumption ofJapanese anchovy, mackerels and Pacific saury by the threebaleen whale species was estimated.
PREY CONSUMPTION AND FEEDING HABITAT OF SPERM
WHALES

The stomach contents of sperm whales sampled in thewestern North Pacific from May to September each year
from 2000 to 2007 were analysed in SC/J09/JR17. Thirtyeight prey species consisting of 33 squids, 1 octopus and 4
fishes, were identified. Sperm whales fed mainly on various
deep-sea squids. The most important prey species were foursquids (Taningia danae, Histioteuthis dofleini, Belonella
pacifica borealis and the eight armed squid Gonatopsis
borealis). Sperm whales feed mainly on prey in themesopelagic and/or bottom/benthic zone during daytime.
The seasonal prey consumption (from May to September)by sperm whales in this region was calculated to be nearly
1.2 million tons. The consumption of neon flying squid,Ommastrephes bartrami, was estimated to be 30,000 tons.
Estimated feeding contribution rates of the surface layer topredation by sperm whales in each sub-area were ranged
from 4.7 to 11.4%. The influence on the surface layer of themarine ecosystem resulting from consumption by sperm
whales cannot be disregarded, because the biomass ofsperm whales is large.
PREY PREFERENCE OF CETACEANS

Murase et al. (2007) presented a study on prey selection ofcommon minke and Bryde’s whales in the western North
Pacific based on data collected in the 2000 and 2001
summer seasons. Whale sighting and sampling surveys andprey surveys using quantitative echosounder and mid-water
trawl were carried out concurrently in the study. Biomassesof Japanese anchovy, walleye pollock and krill, which weremajor prey species of common minke and Bryde’s whales,
were estimated using an echosounder. The results suggested
that common minke whales showed prey selection forJapanese anchovy while they seemed to avoid krill in boththe offshore and coastal regions and walleye pollock in thecontinental shelf region. Bryde’s whales showed preyselection for Japanese anchovy in August 2000 and July
2001, while they showed prey selection for krill in May andJune in 2001.
Prey preferences of common minke, Bryde’s and sei
whales at the mesoscale were estimated (SC/J09/JR18)
using data from the concurrent surveys of cetaceansampling and prey of cetaceans. The surveys wereconducted as a part of the offshore component of 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 preyspecies is 1 and prey species with large values of Manly’s α
indicates 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 of threebaleen whale species overlapped, Manly’s α suggested theirtrophic niches were different from each other. Common
minke and sei whales coexisted in the same survey blocks
but their prey utilisation patterns were different.
OTHER POTENTIAL INFORMATION FOR ECOSYSTEMMODELLING

SC/J09/JR19 presented a model for density prediction ofcommon minke, sei and Bryde’s whales in the western
North Pacific during the feeding season. Data used for themodel were densities estimated from dedicated sighting
survey data in JARPN II, and satellite information on
surface temperature, surface height and chlorophyll. The
predicted density distributions by the analysis suggested
spatial distribution patterns of whales and differences in thepattern among whale species.
SC/J09/JR20 examined time trend of blubber thickness
in common minke, sei and Bryde’s whales, and the factors
influencing the energy storage in these whales. Results
suggested that the blubber thickness of the common minke
whale has increased during the JARPN and JARPN II
period; that of sei whales has increased over the five yearsof the JARPN II period, while that in Bryde’s whales havedecreased over seven years. The feeding areas of Bryde’s
and sei whales showed limited overlap, and theirdistribution is separated by SST. Further studies weresuggested to assist in the interpretation of these results.
4.3.2 Panel conclusions and recommendations
The Panel appreciates the notable amount of effort
undertaken and the generally high quality of the sampling
programme, resultant data and information from JARPN II
studies on whale food habits and prey preferences. The
sampling programme was generally well-coordinated acrossa wide range of vessels and platforms, and the degree ofconcurrently collected multi-disciplinary data was laudable.
These efforts have resulted in valuable datasets that havegreat potential for concerted analytical work on a broad
range of topics, not all directly related to the JARPN II
programme objectives.
The collection of diet data is an important, if not themost important potential justification for the lethalcollection of whales in JARPN II. The Panel noted that it istherefore of the utmost importance that if lethal sampling
occurs, the amount of information collected is maximised;
this is discussed further elsewhere in this report. The Panelagrees that resultant diet data have the potential to be ofgreat value in determining whale prey preferences, fordeveloping functional response curves when accompanied
by simultaneous assessments of prey abundance, and fordeveloping estimates of the impacts of whales on their prey.
The collections of diet data from JARPN II are compatible
with these goals and, in addition, may serve as indicators of
the potential prey selection of these whale species in otherregions of the sub-arctic oceans.
However, the Panel had a number of concerns over the
analyses of the data and in addition agrees that the rationale
for the sampling areas chosen required fuller justification.
With respect to the analyses of the data presented, one of
the Panel’s major concerns relates to the lack of a full
treatment of uncertainty. For this reason the Panel does notbelieve that the consumption rates and CVs presented toJ. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

date can be considered reliable without further analysis. For
this to occur, the JARPN II programme would need todevelop and implement a thorough, systematic approach forincluding and representing uncertainty. This applies to eachcomponent involved in estimating the amounts and types ofprey consumed, including the whale and prey abundance
estimates. On the basis of data collected to date, the Panelrecommends that additional analyses be undertaken to
identify the greatest sources of uncertainty and to determine
appropriate sampling and analytical strategies to address
them. The steps in the estimation of consumption rates forwhich estimates of uncertainty are required are given in
Annex F. An essential component of this will be to improve
the precision of the abundance estimates that are used to
extrapolate to population-level rates, for both the coastal(the possibility of regular well-designed aerial surveys
should be considered) and the offshore regions (a fullsynoptic survey of the region should be considered).
Further, it would be productive to focus on the sources orcauses of variability in order to understand the mechanistic
linkages involved.
A particular concern of the Panel is that only a single
model (that of Sigurjónsson and Víkingsson, 1997) wasused in most JARPN II papers for estimating daily
consumption as a function of body mass –. This is just oneof a number of possible parameterisations of the simple
allometric equation originally proposed by Kleiber (1975).
From a recent comprehensive review (Leaper and Lavigne,2007), there appears to be little scientific justification for
selecting any one model formulation from the range of
those possible; in fact, the Sigurjónsson and Vikingsson
approach produces estimates at the upper range ofreasonable values. The Panel therefore recommends that aspart of the treatment of uncertainty, the analyses of the
JARPN II data should: (a) incorporate the use of several
reasonable models 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 usedin the consumption equations. The results could bepresented in revised papers for the 2009 ScientificCommittee meeting. Whatever modelling formulations areused and ranges proposed, the scientific rationale should beclearly given. This is especially important if the results areto be used in ecosystem modelling.
A more specific concern of the Panel was that themethods used to extrapolate from daily to annual rates andamounts were not clearly explained; this should be
addressed in revised papers for the 2009 ScientificCommittee meeting by providing the formulae used to scale
up daily, individual whale consumption rates to annual,
population-level rates.
The Panel noted that in several of the papers presented,
insufficient effort was placed on incorporating information
resulting from other studies. There exists a considerable
body of knowledge from past studies of whale feeding and‘fishery ecology’ from data collected during commercial
whaling and data collected and analysed during JARPN I(IWC, 2001; Kawamura, 1973; 1980; Nemoto, 1957;
Nemoto and Kawamura, 1977), as well as results andinsights arising from other multidisciplinary research409

conducted (e.g. PICES and ESSAS1). Incorporation of thisinformation will strengthen the scientific basis of the workthroughout the range of studies including those of whale
distribution, abundance, food habits, habitats occupied, thedistribution, abundance and population dynamics of theirprey, etc. Whilst the review is of the JARPN II programmealone, it is important to put the JARPN II work in thecontext of what is known from extensive past research. This
will allow a better evaluation of the JARPN II contributions
to the overall state of knowledge of cetacean feeding
ecology. The Panel suggests that, in revised papers for the2009 Scientific Committee Meeting, the presentation of
JARPN II consumption estimates is placed in the context ofthe broader literature on the topic of whale consumption andthe broader literature of mammal consumption. In addition,
it would be valuable to present the estimates ofconsumption by whales in terms of fisheries and preybiomass (this can provide an index of relativity and an easy
and immediate sense of the magnitudes of the various
processes that can affect fish stock dynamics). For example,
revised papers for the 2009 Annual Meeting could show
consumption of whale prey relative to stock biomass andfisheries landings (the latter was done to some extent), as
diagnostics for and in conjunction with the ecosystemmodels (see Item 4.4).
The extensive amount of data collected under JARPN II
thus far makes several approaches available to investigate
interactions between the physical environment, cetaceans
and prey. Using a variety of approaches would increase
understanding of the processes involved in determining the
distribution of prey and whales, the influence ofenvironmental variability on whale diets, and similar suchconsiderations. Some straightforward approaches based on
empirical statistical methods would help to explore thevaluable multidisciplinary data sets collected via JARPN II.
This might lead to the ability to develop predictive modelsof these processes. In that respect, the Panel recommends
pursuing the following medium- to long-term approaches:•combine the oceanographic data, prey distributions andsighting survey data statistically to investigate how prey
and whale distributions are associated withoceanographic conditions, and how whale distributions
are related to distributions of prey – in this regard thespatial modelling approach described in SC/J09/JR36
needs to be refined and extended further;•
combine data on prey distributions as observed in thearea where the whales were caught with the diet of thewhales (referred to as the micro scale) statistically toevaluate how well the whale’s diet reflects preyavailability in the area where it was caught; and•compare results from the approaches listed above withthe results on selectivity already produced andpresented at the Workshop.
Additionally, performing these analyses and comparisons
will also contribute to evaluation of non-lethal alternatives
commonly used to investigate marine mammal – preyinteractions, such as analyses of spatial associations (i.e. theaggregative response) between predators and prey,particularly in coastal regions. This is discussed further
under Item 8.2.
1
PICES, see http://pices.int/; and ESSAS (Ecosystem Studies of Sub-Arctic
Seas), see http://www.globec.org/structure/regional/essas/essas.htm.

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4.4 Ecosystem modelling
4.4.1 Proponents’ summary
Three papers providing examples of ecosystem modelling
approaches for the JARPN II area were presented. The first(SC/J09/JR14) applied to the inshore region and considered
the harvesting-related implications for sandlance of
predation by minke whales. The other two were for theoffshore region and involved the implementation of EwE(Ecopath with Ecosim2) to provide a model of the wholeecosystem (SC/J09/JR21), and the development of an MRM(Minimum Realistic Model) which explicitly included threewhale and two commercially harvested prey species(SC/J09/JR22).
In SC/J09/JR14, the authors developed a preliminary
population dynamics model to investigate the effects ofconsumption by minke whales on sandlances in the Sanriku
region, using a hierarchical Bayes approach. The modelallows for various uncertainties making use of time series
data historically collected by sandlance fisheries andresearches. The impact of predation was examined in termsof MSY. The result was considerably sensitive to the usedfunctional response forms. The authors argued that the
JARPN II data could contribute toward providing necessary
information to estimation of functional response forms andthen explained how the future JARPN II time series data,such as consumption of sandlances by minke whales, should
be used to estimate the parameters of the (global-scale)functional response curves in the Bayesian estimation
framework.
SC/J09/JR21 provided the results of an initial attempt toevaluate the possible impact of whales migrating to the
JARPN II survey area on Japanese fisheries resources, using
the EwE software. The results suggest that in average terms:(1) when minke whales are the only species that are
harvested at 4% of its biomass (catches of other species are
kept constant at current catch rate), depending on thefunctional response form assumed for the species, it is notcertain whether catch of some Japanese fisheries resources
will increase or not; (2) when sei and Bryde’s whales areeach the only species that are harvested at 4% of theirbiomass, regardless of the functional response form
assumed for the species, catch of anchovy, skipjack tuna,and mackerels may increase; (3) when minke, sei and
Bryde’s whales are all harvested at 4% of their biomass,
positive amount of increase in catch is expected for most of
the fish resources (i.e. anchovy, skipjack tuna andmackerels), indicating the effectiveness of harvesting
several whale species simultaneously; and (4) when sperm
whales are the only species that are harvested at 4% of its
biomass, depending on the functional response form
assumed for the species, catch of anchovy, Pacific saury,mackerels and skipjack tuna may decrease, but instead,catch of neon-flying squid may increase. Caveats pertaining
to the results and the use of such ecosystem models in a
management context are also discussed in the paper.
SC/J09/JR22 reported on the initial work to construct aminimum realistic model for the offshore survey area of
JARPN II with difference equations. Pella-Tomlinson
model was applied to three species of cetaceans (minke, sei
and Bryde’s whales) under MSYR1+ = 0.02, 0.04 and 0.06.
The abundance values of cetaceans prior to exploitation and2

http://www.ecopath.org/.

at present were estimated on a yearly basis with abundance
estimates from sighting surveys and catch series data. Twomain prey species of the cetaceans, that were alsocommercially important (Japanese anchovy and Pacificsaury) were age-structured in the model with informationfrom domestic stock assessment. Other supplementary preys(krill, copepods and others) were included in the model as
they had constant biomass with no seasonality. Feeding
season of cetaceans was set in May-September and the daily
predation was assumed to be equal over the all individual
cetaceans to the necessary energy. The proportion of eachprey taken by cetacean was calculated from the product ofprey biomass, overlap of distribution between prey andcetacean, and prey preference of the cetacean. The overlapof distribution between prey and cetacean was estimatedfrom JARPN II sighting surveys. The estimates of prey
preference of the cetaceans were obtained from the JARPN
II cooperated whale/prey surveys. After adjustment for theparameters of the model, natural deaths of Japanese
anchovy and Pacific saury were separated into two parts:
consumption by cetaceans and natural death.
4.4.2 Panel conclusions and recommendations
The Panel emphasises at the outset that the programme
objective (of developing ecosystem models to the level thatthey could contribute in the provision of specificmanagement advice) constitutes a major, complex andambitious undertaking – the difficulties have beenrecognised by many intergovernmental bodies (including
the IWC, FAO and CCAMLR). Any study anywhere in theworld attempting to meet such an objective requires
substantial data collection and analytical efforts; results thatare sufficiently reliable to inform management advice
should not be expected within at least the next few years
and might possibly take considerably longer to obtain.
Ecosystem models can also provide contextual informationof value to managers in some circumstances. The value ofconsidering a wide range of models has been stressed byseveral groups, noting that the various possible objectives
for ecosystem modelling lead to different questions forwhich the best modelling approaches may differ (see, e.g.
table A4 of Plaganyi, 2007).
More specifically related to the approaches presented to
the Panel, the analyses tabled had been appropriately
classified by their authors as preliminary. However, despite
this, the authors generally overstated the certainty
associated with their results (this is in fact a common
practice in this field). The Panel agrees that the models as
developed thus far are not yet at the stage where they can beused to draw even general conclusions and certainly cannot
be used to reliably inform management advice.
Nevertheless, they comprised a substantial and laudable
effort, and an encouraging start to the necessary process ofsynthesising the data collected during the programme.
Although this work might be said to have started relatively
late within the time span of the programme thus far, an early
emphasis more towards data collection has been a common
feature of a number of similar marine programmes
elsewhere. However, the Panel agrees that considerably
more emphasis should be placed on the modelling workfrom now on if the objective of the programme is to have achance of being met within a reasonable timeframe,
especially with regard to the recommendations provided
below.

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

The Panel emphasises that with respect to sperm whales,
the low sample size and the logistical constraints on the sizeof animals taken means that the data obtained provide no
meaningful input to ecosystem models.
GENERIC RECOMMENDATIONS(1) Considerably more resources must be allocated to themodelling work – without this, the likelihood that theobjective of the programme will be reached in areasonable timeframe will be minimal. The models
developed should be used to identify the areas of
uncertainty with the greatest impact on model outputsof relevance to management, and hence to guide theprioritisation of future data collection and the associatedsample size/sampling design.
(2) A wider range of models needs to be considered if theobjectives of the programme are to be met. Further
work should aim towards fitting dynamic models totime series of data, especially abundance indices.
(3) The area covered by JARPN II is not spatiallyhomogeneous, and serious consideration should begiven to developing separate models for three regions
distinguished by the inshore or shelf region, the subArctic oceanic region of the Oyashio current and the
sub-tropical region of the Oyashio and Kuroshiotransition zone.
(4) There is a need to take much wider account of
uncertainty at all stages of the modelling process,including that associated with the prey consumption
rates of whales (e.g. the Bayesian approach of
SC/J09/JR14 should be readily extendable towards thatspecific end, and more generally other approaches suchas sensitivity testing should be employed).
(5) The importance, ultimately, of developing models
which incorporate natural variability in dynamic
processes (e.g. recruitment variability for prey species)
was emphasised, although it was recognised that thismight not be possible for certain ecosystem modelling‘packages’.
This is in addition to taking account of uncertainty inmodel structure and parameter values. The complexity of
ecosystems and the difficulty of modelling species
interactions adequately might mean that management
actions based on such models are more likely to induce
unexpected instabilities than current single-species based
approaches; this suggests a more cautious approach will beneeded on the part of decision makers.
SPECIFIC RECOMMENDATIONSWith respect to the Bayesian analysis of SC/J09/JR14, thePanel agrees that if there are other predators making
individual contributions to sandlance natural mortality ofsimilar size to that estimated for minke whales, their explicit
inclusion in this model must be considered. It agrees thatType I functional relationships are unrealistic and need not
be considered further. As noted earlier, any resultspresented should distinguish yields of the prey species topredators and the fishery.
For the EwE approach of SC/J09/JR21, the Panel agrees
that it is important to concentrate first on improving the
Ecopath component of this EwE analysis before moving onto the next step of extending the modelling effort from astatic to a dynamic model such as Ecosim.

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The species included in the Ecopath analysis should be
reviewed giving attention to Ecopath models developed forother regions; in particular the inclusion of gelatinous
zooplankton should be considered. Furthermore the valuesof the parameters of this Ecopath analysis should becompared with values for those others, with attention
directed towards any instances of major discrepancies.
Inspection of some features of diagnostic plots (e.g. Fig. 1)of the current Ecopath results suggest the need forreconsideration of some of the parameter values, e.g. theplot of log biomass against species does not decline as
rapidly as customary, suggesting perhaps that the abundanceof primary producers is underestimated; for a number of
species, the fraction of production consumed within thesystem (the ecotrophic efficiency parameter EE) is
unrealistically close to the maximum possible of 1; and the
P/C ratio (production by a species relative to its foodconsumption) is unrealistically high for some species.
The features noted suggest the need to rebalance the
Ecopath model. Alternative approaches to doing so should
be considered. For example, rather than use values for someparameters drawn from other regions, placing a bound onsome relationship (e.g. P/C<0.6) may lead to an improved
result overall.
Further analyses must take full account of theuncertainties associated with model inputs, e.g. using
Ecoranger3.
Finally, the Panel noted that the approach in SC/J09/
JR22 was the most preliminary presented. Further work on
MRM approaches is encouraged and should focus inparticular on fitting such models to time series of data.
5. REVIEW OF JARPN II RESULTS: MONITORINGENVIRONMENTAL POLLUTANTS IN CETACEANS
AND THE MARINE ECOSYSTEM
5.1 Statement of objectives as given by the Governmentof Japan
The general objectives for the environmental pollutant
component of the programme were listed as:(a) monitoring environmental pollutants in cetaceans
and the marine ecosystem;(b) pattern of accumulation of pollutants in cetaceans;(c) bioaccumulation process of pollutants through thefood chain; and(d) relationships between chemical pollutants andcetacean health.
Justification and further explanation of this broad
objective was also provided as summarised below (takenfrom SC/J09/JR1).
In 1992, the IWC decided to establish a regular agenda
item for research on the effects of environmental change on
cetaceans (IWC, 1993) and at the 1994 IWC ScientificCommittee meeting, one of the items identified waspollution (IWC, 1995b). In particular, there is concern thatpollutants may have a negative effect on the health ofcetaceans resulting ultimately in a decrease in theabundance of the stocks.
Pollutants such as organochlorines (OCs) and heavy
metals are generally released from land and transported to3

http://www.fishbase.org/ecopath/ecorange.htm.

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Fig. 1. Example of a simple diagnostic plot (see text).

coastal and pelagic waters in run-off as well as byatmospheric transportation and other ways. Higher trophic
animals such as cetaceans generally accumulateorganochlorines and toxic elements through the marine foodweb. The monitoring of pollutants in the marine
environment through the examination of biological tissuesof marine mammals is of importance since marine mammals
can serve as useful biological indicators of environmental
conditions. Large cetaceans may be particularly useful as
they are long-lived animals migrating long-distances.
Therefore pollutants can be monitored in a wide area andpollutant burdens can be integrated over time. This
monitoring can be done through JARPN II surveys.
5.2 Proponents’ summary
Pattern of accumulation of pollutants in cetaceans
PCBs and pesticides were measured in the blubber of spermand sei whales collected from offshore waters of thewestern North Pacific in 2001 and 2002. PCBs and DDTs
were the predominant contaminants in both species.
Organochlorine residue levels in sei whales were one or twoorders of magnitude lower than in sperm whales and werethe lowest reported in whales from the Northern
Hemisphere. Compositions of CHL and DDT compounds
also showed species-specific differences, with sperm whales
retaining a higher percentage of p,p’-DDE and a lower
percentage of oxychlordane compared with sei whales.
These results suggest that the differences in feeding
preferences and metabolic capabilities are the major
contributing factors influencing the accumulation patternsof organochlorines, in different species of marine
mammals4.
4
Results from this study were presented to the First International
Symposium on Environmental Behavior and Ecological Impact of
Persistent Toxic Substances’, Matsuyama, Japan March 18-20, 2004. An
abstract is available.

To investigate temporal changes of mercury (Hg) levelsin the western North Pacific, total Hg concentrations inmuscle samples from common minke, Bryde’s and sei
whales were measured (SC/J09/JR23). Total Hg levels werein the order: mature common minke whales (0.22±0.07 ppm
wet wt.) > mature sei whales (0.052 ±0.009) = mature
Bryde’s whales (0.046±0.008). Yearly changes of total Hglevels in zooplankton and pelagic fishes were not observedin the period 1995-2007. Apart from common minke whales
from sub-area 9, significant yearly changes of levels inwhales were not observed. For minke whales in sub-area 9,levels decreased from 1994 to 1999 but increased from2000 to 2007. Results of a multi linear regression analysis
suggested that changes of Hg levels in sub-area 9 reflectchanges in food habits of minke whale rather than changes
in accumulation levels in the environment. SC/J09/JR24
presented information on PCB levels in blubber samples ofcommon minke, Bryde’s and sei whales from the western
North Pacific. The range of levels in these species was 0.134.0, 0.04-0.21 and 0.03-0.47 ppm wet wt., respectively.
Yearly changes of PCB levels were not observed incommon minke, Bryde’s and sei whales in the period 20022007. Results of previous studies suggested that PCB levels
had been continually decreasing in this oceanic region(1980s-1990s). Results from JARPN II suggest that thelevel has been stable since 2002.
Bioaccumulation process of pollutants through the foodchain
SC/J09/JR23 also examined total Hg levels in prey samplesof whales: two zooplanktons (krill E. Pacifica; copepods
Neocalanus spp. and Calanus sp.), six pelagic fishes(Japanese anchovy; Pacific saury; walleye pollock;mackerels; Pacific pomfret). Total Hg levels in krill andcopepods ranged from <0.001-0.013 and 0.003-0.010 ppm
dry wt., respectively. Total Hg levels in the pelagic fishes
were in the order: Pacific pomfret (0.232±0.027) > walleye

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

pollock (0.045) = Pacific saury (0.039±0.016) = Japanese
anchovy (adult) (0.037±0.025) > Japanese anchovy (larvalfish) (0.005±0.003). No yearly changes of total Hg level
were observed for krill and Japanese anchovy during theperiod 1995-2007. Variation of Hg levels in pelagic fishes
from the western North Pacific was not related to samplingyear.
SC/J09/JR24 also presented information on PCB level in
air and surface seawaters from the western North Pacific.
The range of levels was ND-22 pg/m3 for air samples and1.5-11 ng/L for sea water samples. PCB levels in seawater
decreased from coastal to offshore regions. The trend inlevel of air samples was not clear.
Relationships between chemical pollutants and cetacean
healthTo examine the CYP families, related to immune-toxicity of
PCB and pesticides, full-length cDNA sequences of
CYP1A1, and 1A2, in common minke whales were
determined (Niimi et al., 2005). The deduced full-lengthamino acid sequence of CYP1A1 revealed higher identities
with those of sheep (86%) and pig (87%), and that of
CYP1A2 was most closely related to human (82%) andmonkey CYP1A2 (82%) among species from which
CYP1A2 has been isolated so far. Differences in certain
conserved and functional amino acid residues of CYP1A1
and 1A2 between common minke whale and othermammalian species indicate the possibility of their specificmetabolic function. Concentrations of organochlorinecompounds (OCs) including PCBs and DDTs analysed incommon minke whale liver showed no significant
correlation with hepatic mRNA expression levels of
CYP1A1 and CYP1A2, indicating no induction of these
enzymes by such OCs.
To investigate whether or not CYP expression levels arealtered by organochlorine contaminants (OCs), mRNAlevels of CYPs in the liver of common minke whales were
measured (Niimi et al., 2007). The quantified mRNA levels
were employed for the statistical analysis with the residue
levels of OCs including PCBs, DDTs (p,p’-DDT, p,p’-DDD
and p,p’-DDE), chlordanes (cis-chlordane, trans-chlordane,
cis-nonachlor, trans-nonachlor and oxychlordane), HCHs
(alpha-, beta- and gamma-isomers) and hexachlorobenzene
that have already been reported elsewhere. Spearman’s rankcorrelation analyses showed no significant correlation
between CYP expression levels and each OC level in thecommon minke whale liver, implying that theseenvironmental chemicals have no potential to alter the
expression levels of these CYPs or the residue levels
encountered in the whale livers may not reach theirtranscriptional regulation levels. This suggests that theexpression of individual CYPs in the whale liver may be atbasal level.
SC/J09/JR25 examined the accumulation characteristics
of mercury, a toxic element, and selenium, an antagonist, as
well as the inter-species difference of sensitivity to mercury
toxicity. Total mercury (T-Hg), methyl mercury (MeHg)
and selenium (Se) levels in the liver, kidneys and muscle
were measured. T-Hg and MeHg levels were higher in theorder of sperm whale > common minke whale > Bryde’s
whale. Se levels were higher in the order of sperm whale >Bryde’s whale > common minke whale. The order of the THg and Se levels in the tissues of the common minke and413

Bryde’s whales was kidneys > liver > muscle, and that of
MeHg was liver > muscle > kidneys. The order of the T-Hg
and Se levels in the tissues of sperm whales was liver >kidneys > muscle.
5.3 Panel’s conclusions and recommendations
The Panel concludes that the JARPN II pollutant studies
represent a valuable contribution to our knowledge in thisarea and acknowledged the considerable amount of work
presented. The programme is addressing its objectives andfurther work is recommended (see below). In general, where
possible, papers should include a risk assessment statement,summarising the potential risk from exposure to the various
pollutants, based on current toxicology data in model
species and other wildlife in terms of the health of theanimals and dynamics of the stocks.
Paper-specific comments
SC/J09/JR235 reported on temporal trends and factors
affecting mercury levels in common minke, Bryde’s and sei
whales and their prey species in the western North Pacific,
addressing objectives (a) and (b) above. With respect tocommon minke whales, only ‘putative’ ‘O’ stock animals(based on the genetic analyses discussed under Item 6) were
included in the analyses.
The Panel noted that if the system is now in steady state,as the paper indicated, and redistribution of T-Hg into
muscle reflects ingestion over the previous few months, as
at least studies in laboratory animal models and fish appear
to suggest (Dock et al., 1995; Petersson et al., 1991; Van
Walleghem et al., 2007), then these data might be used toestimate relative bioconcentration factors based on different
prey species for the three predators investigated. However,it was noted that some bias might occur because animals
could also be feeding elsewhere.
In this study, T-Hg levels in muscle not liver were
investigated; this was because the absolute age values forthe whales were not yet available, but only body length
data. In such circumstances, muscle was considered a morestable monitoring tissue, less affected by age differences(and in addition only results from mature male animals wereused in the trend analysis) than liver. However, the majorityof previous studies on mercury trends in cetaceans
investigate changes in liver concentrations rather thanmuscle because the liver is one of the main target organs for
mercury, and levels are often highest in this tissue. The
Panel recommends: (a) that the analyses be carried out by
age when age data become available; and (b) if possible
future studies also examine levels in the liver to facilitatecomparison with other studies. In addition, the authors had
fitted two linear models to the data, one prior to 2000 and
one after 2000 (when there appeared to be no change in theconcentrations with time). However, the Panel suggests thata GAM fitted to these data would be a better method for
determining the change points and examining non-linear
trends in the Hg levels. If possible such an analysis should
be presented at the 2009 Annual Meeting.
5
The Panel’s also received a figure showing the flow of total mercury(THg) in the predators and their prey, not included in the paper. The Panelfound this valuable in linking both parts of the study together and inillustrating how the concentrations found in each species are related to one
another; the Panel recommends that this figure be included in a revised
paper for the 2009 Annual Meeting.

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SC/J09/JR24 reported on temporal trends and factors
affecting polychlorinated biphenyl (PCB) levels in baleen
whales and environmental samples from the western North
Pacific and largely addressed objective (a) above. The Panelnoted that it was not clear whether the study animals
reported in the paper SC/J09/JR24 were all mature males,
although it was clarified that this was the case. This should
be made clear in a revised paper presented to the 2009
Annual Meeting.
The PCB concentrations reported in this study were notgiven on a lipid weight basis, as is the convention with thesecontaminants. Because they are tightly bound to the lipid
fraction in the blubber, which may not be consistent overtime, it is important for comparative purposes to adjust theresults to the amount of lipid in the sample. Lipid
proportion data were not available for all the samples andthe authors had recognised that this was a problem. The
Panel recommends that future studies must be carried out
on a lipid weight basis. It also recommends that in future,sampling for PCBs and Hg from the same individuals isundertaken to allow combined analyses of these often cooccurring contaminants.
SC/J09/JR25 investigated the accumulation of total andmethyl mercury and selenium in baleen and sperm whales
from the western North Pacific. One of the main findings
was that demethylation abilities appeared to be different
among the species, and higher selenium levels, especially in
Bryde’s whales, indicated they were likely to be lessvulnerable to effects of MeHg than the other species. The
Panel recommends that in a revised paper submitted to the2009 Annual Meeting, greater emphasis is given to thisimportant ecotoxicological finding. While the sperm whale
samples (n=5) had high liver T-Hg levels, the molar ratio of
Hg to Se indicated these animals were not at risk of toxic
effects because of the protective effect of selenium. The
Panel emphasises that with respect to sperm whales, thelow sample size and the logistical constraints on the size ofanimals taken means that the obtained data cannot be used
to infer general conclusions.
Although concentrations of T-Hg in liver, kidney andmuscle (i.e. the main target organs) were presented, thePanel agrees that for total body burden estimates, additionalorgans would need to be included. It suggests that theauthors investigate whether the available literature indicates
whether the proportion found in these tissues can be derived
and thus total burden estimates made. With respect to future
work, the Panel suggests examination of T-Hg in brain
tissue particularly for comparing the more coastal bycaught
animals to the coastal and offshore JARPN II samples, asthese would provide a valuable (perhaps more exposed)
comparison group. It was observed that brain tissue is very
important if the newer contaminants are to be investigatedin the future. Additionally, polybrominated compounds
target adrenal glands as well as fat, so contaminant levels
can be as high in these organs as they are in the blubber.
The published studies of Niimi et al. (2007; 2005) report
the molecular characterisation of various cytochrome P450
detoxification liver enzymes in minke whales and how theirmRNA expression levels relate to levels of contaminants inthe liver. These studies addressed the final objective (c)above. The Panel welcomes these results, whilst noting thatthese biomarkers are not necessarily indicative of healthstatus (even though they may be induced by foreigncompounds, this increased activity may not necessarily be
detrimental to the animals’ health). The Panel encourages
the continuation of such comparative molecularphylogenetic research using mRNA isolated from fresh
tissues. Many different hormones, proteins, receptors andenzymes can be characterised using these and othermolecular approaches (such as microarrays) and the utilityof putative biomarkers can be examined at the expression
level. Despite the limitation that the contaminant exposurelevels in these animals might well be too low to induce theactivity of the enzymes above background, the biomarkers
did not correlate with exposure.
Recommendations(1) Any future contaminant exposure and uptake studies
should be based on a balanced, structured study design
with a specific number of individuals sampled withineach strata (e.g. by species, sex, stage, ocean regime
and location). All the necessary data on exposure andconfounding variables should be obtained from all of
the specifically targeted individuals and a control orcomparison group should be included. In this way amore powerful and statistically robust study to address
clearly stated hypotheses could be designed and carriedout.
(2) Tissues should be archived (frozen at –20°C or lower if
possible) for future retrospective analyses.
(3) The importance of having absolute age as an additional
covariate for the interpretation of the results, both thepollutant levels and to provide further information on
population structure, cannot be over-emphasised andevery effort should be made to obtain such data (theProponents informed the Panel that this is being
pursued) .
(4) Consideration should be given to including coastal, ‘J’stock bycaught minke whales in future studies as thesewould provide a valuable (perhaps more exposed)
comparison group.
(5) Future studies should include data on stable isotope
ratios and fatty acid profiles from a variety of tissues(for example muscle, liver, brain, blubber, skin) as
these profiles, also indicative of diet, could help
determine what the whales had been feeding on in thepast (particularly important for assessing predator prey
relationships in blubber PCBs and other persistentorganic pollutants (POPs)) that are integrated in thesetissues over a long timeframe, for an example see Fisk
et al. (2001). This would help discriminate amongreasons for temporal changes (i.e. dietary changes or
exposure variation with constant diet).
(6) The air and water samples obtained could have been
useful in a ‘fate and behaviour’ study but congener
specific data (especially for PCBs) and other elements
would need to have been included to make a substantive
contribution to knowledge in this field. There arevarious modelling approaches that could be
implemented, but more results for air and water,
including the effect of weather using simultaneously
collected data, are required. More resources and effort
need to be allocated to this aspect of the monitoring.
(7) Simple mass balance studies (input-output estimates)
would contribute to our knowledge of the partitioning
and offloading of contaminants in these species and the

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potential impact of changes in exposure. For this,additional analyses of blood, bile, faeces and urine are
required.
(8) The contaminant results should eventually be linked tothe prey consumption studies. For example, theProponents could model the flow of Hg in the marine
ecosystem (using, as one example, the approach taken
by Booth and Zeller (2005), who used Ecotracer, a newroutine in Ecopath, for this) and determine how changes
in the flow of energy within the system might affect theflow of contaminants and their deposition rates.
6. REVIEW OF JARPN II RESULTS: STOCKSTRUCTURE
6.1 Statement of objectives as given by the Governmentof Japan
The general objectives for the stock structure component of
the programme were simply listed as:‘Stock structure of large whales:
common minke whale (Balaenoptera acutorostrata)
Bryde’s whale (B. edeni)
sei whale (B. borealis)
sperm whale (Physeter macrocephalus)’
Justification and further explanation of this broad objective
was also provided as follows (taken from SC/J09/JR1).
Common minke whale
There are the following two remaining issues on stock
structure, which are important in the context of management
under the RMP.
(A) SYSTEMATIC MONITORING OF THE OCCURRENCE OF ‘J’STOCK LIKE ANIMALS IN COASTAL AREAS OF THE PACIFICSIDE OF JAPAN (SUB-AREA 7, FIG. 2) TO DETERMINE THE
DYNAMICS (SPATIAL AND TEMPORAL) OF THEIR
OCCURRENCE

Assignment of individuals to stocks is particularly
important in situations of geographical overlap of stocks(animals of two or more stocks occupying the samegeographical area). Furthermore, individuals assigned toparticular stocks by genetic analysis can be examined forother biological and ecological traits so that differencesamong stocks revealed to non-genetic markers can beinvestigated as well.
(B) PLAUSIBILITY OF THE FOUR STOCK STRUCTURE HYPOTHESES OF THE ‘O’ STOCK USED IN ISTS IN 2003 (FIG. 2)

Hypothesis A: three-stock scenario (‘J’, ‘O’, ‘W’) with the‘W’ stock found only in part of sub-area 9 and only
sporadically (Fig. 2-AB); Hypothesis B: two-stock scenario(‘J’ and ‘O’) with no W stock as a limiting case of Baseline
A (Figure 2-AB); Hypothesis C: four-stock scenario overall,with ‘OW’, ‘OE’ and ‘W’ to the east of Japan. Boundaries
are fixed at 147°E and 157°E and there is no mixing
between the stocks (Fig. 2-C); Hypothesis D: three-stock
scenario (‘J’, ‘O’, ‘W’), with ‘O’ and ‘W’ mixing over
147°E and 162°E, ‘O’ being dominant to the west and ‘W’
to the east (Fig. 2-D).
In 2003, the IWC Scientific Committee gave the samehigh plausibility to these four hypotheses (IWC, 2004b).

415

There was therefore a need to examine new samples(genetic and non-genetic) to evaluate the plausibility ofthese four stock structure hypotheses.
Bryde’s whale(A) PLAUSIBILITY OF THE FOUR HYPOTHESES ON STOCKSTRUCTURE WERE USED IN THE RMP’ ISTS (FIG. 3)

Hypothesis 1: this is a single stock hypothesis under which
only one stock of Bryde’s whale is found in the area from
130°E and 160°W (excluding the area of distribution of theEast China Sea Stock) and there are no sub-stocks.
Hypothesis 2: this is a two-stock hypothesis under which
Stock 1 is found in sub-area 1 and Stock 2 in sub-area 2.
Sub-areas 1 and 2 are divided at 180° longitude. Under thishypothesis there are no sub-stocks.
Hypothesis 3: this is a two-stock hypothesis under which
Stock 1 is found in sub-areas 1 and 2 and Stock 2 only in
sub-area 2. Under this hypothesis there are no sub-stocks.
Hypothesis 4: this is a two-stock hypothesis under which
Stock 1 is found in sub-area 1 and Stock 2 in sub-area 2.
Stock 1 consists of two sub-stocks that mix in sub-area 1.
The IWC Scientific Committee gave different
plausibility to these hypotheses as follows: Hypothesis 1:
high; Hypothesis 2: high; Hypothesis 3: high; Hypothesis 4:
medium (IWC, 2008b). There was therefore a need tocollect and examine new samples to evaluate the plausibilityof these four stock structure hypotheses, in particular to
respond to remaining key questions on (a) whether or not
sub-stocks occur in sub-area 1 and (b) whether or not adifferent stock occurs in sub-area 2. JARPN II attempted torespond to the first question.
Sei and sperm whales
These species are currently not being considered for RMP
Implementation. The IWC Scientific Committee is currently
considering undertaking an in-depth assessment of these
species in the North Pacific to investigate the current
population status in this oceanic region. Estimations ofabundance, biological parameters and examination of thecatch history are conducted on the basis of individual
stocks.
The past information on stock structure of sei whales was
based on marking, catch distribution, sighting, morphologyof baleen plates (Masaki, 1977), and isozymes (Wada andNumachi, 1991). The information is old and limited, and it
was considered non-conclusive by the IWC ScientificCommittee when last examined. Therefore there is the need
to update sampling and analysis on stock structure for thisspecies in the North Pacific.
In the past, the management of sperm whales by the IWC
was based on the assumption of two-stocks, western andeastern stocks divided at 180º. The most recent information
on stock structure was based on analysis of whaling
operation data, movement of marked whales and sighting
distribution (Kasuya and Miyashita, 1988). These authors
suggested two latitudinally segregated sperm whale stocks
in the western North Pacific. Therefore the information on
stock structure of sperm whales in the North Pacific is old
and limited and there is the need to update sampling andanalysis on stock structure for this species in the North
Pacific.

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Fig. 2. Hypotheses on stock structure of North Pacific common minke whale used in the Implementation Simulation Trials of the RMP (IWC, 2004, p. 79).
See text for details.

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417

Fig. 3. Hypotheses on stock structure of North Pacific Bryde’s whale used in the Implementation Simulation Trials of the RMP (IWC, 2008b).
See text for details.

6.2 Proponents’ summary
Common minke whale
SC/J09/JR26 presented the results of a study that attempted
to distinguish minke whales sampled around Japan into
genetically distinct stocks using a combination of
microsatellite analysis and a Bayesian clustering approach.
Samples of 2,542 minke whales were collected during theoffshore component of JARPN and JARPN II from 1994 to
2007, during the coastal component of JARPN II from 2002
to 2007, and from bycatches in the set-net fishery along theJapanese coast from 2001 to 2007. These were analysedusing 16 microsatellite loci. Results of the Bayesianclustering analysis implemented in the computer program
STRUCTURE (Pritchard et al., 2000b) indicated that the
samples came from two genetically differentiated groups ofminke whales. Approximately 91% of the individuals wereassigned into the putative stocks based on their highmembership probability (>90%) obtained from the program.
Spatial distribution of these assigned individuals clearly
indicated that these two-stocks were the ‘J’ and ‘O’ stocks
that have been known to exist around the Japanese coast. In
addition, it was also found that: (1) the ‘O’ stock individuals
appeared to migrate, although rarely, to the Sea of Japan;(2) the ‘J’ stock individuals migrated to the 7W area off theNorth Pacific side of Japan and, very rarely, farther east;and (3) the SA2 (western side of North Pacific coast) was
mainly occupied by the ‘J’ stock. Temporal distribution of
the assigned bycatches collected from SA7 (eastern side ofJapan, North Pacific coast) indicated seasonal movement of
the whales with the number of the ‘O’ stock increased inspring. This study allowed better understanding of thepattern and dynamics of distribution of the minke whales
inhabiting waters around Japan.
A consolidated presentation was made using the resultsfrom a morphometry study (SC/J09/JR27), a mitochondrial
DNA study (SC/J09/JR29) and a microsatellite study(SC/J09/JR30). These three different kinds of analyses wereused to examine the plausibility of the four baseline stock
scenarios proposed during the RMP Implementation for thewestern North Pacific common minke whales (IWC, 2004)
by analyzing samples of minke whales collected during
JARPN II as well as JARPN conducted from 1994 to 2007;

the samples from 2003 to 2007 had not been available in the
Implementation process. All of the studies conducted thestatistical tests in similar fashion to look for evidence ofgenetic heterogeneity in the samples: (1) genetic differences
between the coastal and offshore samples collected in thesame year from the 7W area; (2) among the samples
collected in the different years from the same sub-area; and(3) among the samples divided and compared on the basisof proposed stock divisions from each of the four baseline
scenarios.
These tests were conducted with and without thesuspected ‘J’ stock individuals as well as with only thesuspected ‘O’ stock individuals in the samples assigned
based on SC/J09/JR26. The studies found: (1) whales fromthe ‘J’ stock existed in the 7W with low but large enoughnumber to cause genetic heterogeneity observed in the 7W
samples as well as between the 7W and other samples; (2)except the ‘J’ stock whales, the survey area was mainly
occupied by ‘O’ stock; and (3) the baselines C and D were
not supported because no evidence of distinct coastal stock
was observed. These studies supported the baseline scenario
B as the most plausible.
SC/J09/JR28 examined the status of scars on the skin ofminke whales, and whether or not it is possible to identify
the stock of the individual animal based on external
morphological scars using samples of western North Pacificcommon minke whales collected by JARPN II. This study
was assisted by the genetic assignment to stocks shown in
SC/J09/JR26. Assignments of the number of scars were not
a complete diagnostic for the minke whales samples for ‘J’
and ‘O’ stocks. However, at least there appeared a strong
likelihood that animals which have no scars on the body
were ‘J’ stock animals.
The main conclusions of papers dealing with North
Pacific minke whales (SC/J09/JR26-30) can be summarised
as follows.
(1) A great deal of new information has been developed,
drawing on both genetic and non-genetic methods.
(2) New data provide strong support for, and additional
insights regarding, what have previously been referred
to as ‘J’ and ‘O’ stocks.

418

REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II(3) New data do not provide support for the existence ofmore than these 2 stocks in the study area. In particular,
no evidence was found to support hypotheses C or D
considered by the IWC (IWC, 2004a).
Bryde’s whale
Kanda et al. (2007) investigated the pattern of genetic stock
structure in the Bryde’s whale at the inter-oceanic and transequatorial levels, using microsatellites (17 loci) and mtDNA
control region sequences (299bp). Samples were available
from the western North Pacific (JARPN II), South Pacific(historical) and Indian Ocean (historical). While nosignificant differentiation was found within the western
North Pacific, marked genetic differentiation was foundamong oceans and between South and North Pacific.
SC/J09/JR31 examined genetic variations at 17
microsatellite loci and 299bp of mitochondrial DNA(mtDNA) control region were analysed to investigate theexistence of genetically differentiated sub-stocks of Bryde’s
whales in the sub-area 1 (stock hypothesis 4 in figure 5 of
SC/J09/JR1). No evidence was found of genetic
differentiation between the samples from the 1W and 1E(separated at 153ºE), indicating these JARPN II samples
came from a genetically same group of Bryde’s whales. The
same result was found when historical samples from thecentral western North Pacific and around the Ogasawara
Islands were incorporated into the analysis.
Sei whale
SC/J09/JR32 presented an analysis of genetic variation at 17
microsatellite loci and 487bp of mitochondrial DNA(mtDNA) control region sequences in samples of sei whales
in order to describe their stock structure in the North
Pacific. The samples consisted of 489 whales collectedduring JARPN II from 2002 to 2007 in the area between
143°E and 170°E and 301 whales collected from the 1972
and 1973 commercial whaling conducted at the North
Pacific from 165°E to 139°W. Due to the condition of the
DNA extracted from the archived blood samples, 14 of the
17 loci were analysed in the commercial whaling samples.
Conventional hypothesis testing was conducted to look forany evidence of genetic differences among the samples. Allof the tests found no evidence of genetic differences within
as well as between the JARPN II and commercial whaling
samples. Both females and males showed the same patternof stock structure. This study showed that the open water of
the North Pacific appeared to be mainly occupied byindividuals from a single stock of sei whales.
Sperm whale
SC/J09/JR33 analysed genetic variations at 15 microsatellite
DNA loci and mitochondrial DNA (mtDNA) control region
sequences in samples of sperm whales collected during
JARPN II from 2000 to 2007 in order to examine theeffectiveness of these genetic markers for stock structure
study of the species. Analyses of mtDNA and microsatellite
markers in the total of 45 sperm whales demonstrated thatthese genetic markers were variable enough to explore stock
structure of sperm whales. Statistical tests found noevidence of deviation from the expected Hardy-Weinberg
genotypic proportion at all of the 15 microsatellite loci. At
this point no signal of multiple stocks in the research area
was found.

6.3 Panel’s conclusions and recommendations
The Panel acknowledged the substantial scope of thegenetic analyses undertaken under JARPN II, which
provides a uniquely large data set for testing hypotheses
regarding stock structure in the target species. Analyses
conducted with the genetic data under JARPN II were in
general sound and of a nature common to other genetic
analyses within and outside the IWC Scientific Committee
framework. The inclusion of morphological andmorphometric studies as well as genetic information helps
to provide a more well-rounded picture of stock structure.
Although genetic data can provide valuable insights
regarding stock structure, they have some limitations forweakly differentiated populations, as is the case for many of
the species targeted by JARPN II.
Particularly in the context of weakly-differentiatedpopulations, the Panel acknowledges the generaldifficulties in examining questions of stock structure, notthe least of which is that there is no formal IWC definitionof ‘stock’ (see the extensive deliberations in recent years
within the Working Group on Stock Definition). Similarly,
none of the stock structure papers provides a definition of a‘stock’ or describes an objective method for determining
whether a group of individuals constitutes a ‘stock’. This
complicates the work of the Panel in evaluating theeffectiveness of the JARPN II programme in identifying
stocks. In practice, however, the objective of the JARPN II
programme is to assist the IWC Scientific Committee inproviding information that can be used within the context of
the RMP Implementation process and more specifically, tonarrow the number of hypotheses used for western North
Pacific common minke whale and Bryde’s whale
Implementation Reviews, and to assist in the proposed indepth assessment of sei whales that may ultimately lead tothe inauguration of the Implementation Process for these
species in the North Pacific. The papers presented had
focussed on this objective.
The Panel emphasises that with respect to sperm whales,
the low sample size means that the obtained data provide no
meaningful input to stock structure discussions.
Recognising the general difficulties, the Panel identified
a number of limitations to the analyses presented as listed
below. It later provides detailed suggestions for addressingthese limitations.
(1) The genetic analyses rely heavily on hypothesis testing
and P values, with limited consideration and discussionof effect sizes (e.g. FST values or other measures of thedegree of genetic differentiation). Most of the statisticaltests used evaluate the null hypothesis that allindividuals come from a single panmictic population.
This is a reasonable (and common) starting point forevaluations of genetic data but rarely is sufficient tofully inform questions related to stock structure.
Informed
management
decisionsabout
theconsequences of harvest generally depend on at least aqualitative assessment of the degree of demographic
linkage between putative populations or stocks. Thisrequires additional information and analysis beyond
that provided by tests of heterogeneity. See Waples andGaggiotti (2006) and Palsbøll et al. (2007) for
discussion of these issues.
(2) The main conclusions in the papers presented are
based on a failure to find evidence of population

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

differentiation or stock structure. To evaluate how
strongly negative results support a single stock
hypothesis, statistical power to detect heterogeneity, as
well as alternate explanatory hypotheses, need to beassessed. Although large amounts of genetic data havebeen collected under JARPN, the absence of anassessment of statistical power makes it impossible toreduce the plausibility of any of the current stock
hypotheses proposed for minke and other baleen whalesin the North Pacific (although an initial examination ofthis problem for western North Pacific Bryde’s whales
was presented by Kitakado et al., 2005a; 2005b).
Inclusion of an assessment of statistical power would
add substantially to the analyses conducted and aid in
the selection of stock hypotheses proposed by the IWC
Scientific Committee for this area. The use of
STRUCTURE to estimate relative probabilities for thenumber of stocks present was noted, although it is
known that STRUCTURE can produce unreliable
results under some circumstances (IWC, 2007).
(3) The low levels of genetic structure identified by the
JARPN II analyses might be due to substantial
deviations
fromthemutation-drift-migration
equilibrium assumed in most of the analyses
undertaken. The mtDNA data yielded low estimates ofnucleotide diversity but high haplotypic diversity,
which is consistent with a recently [in evolutionary
terms] exponentially expanding population (slatkin andHudson, 1991), and thus non-equilibrium. The data
should be subjected to assessments of possible
deviations from population genetic equilibrium, andmethods to estimate gene flow that allow for nonequilibrium conditions should be explored.
(4) Most of the assessments involve a large number of tests
conducted separately for each locus and/or various
combinations of samples. Although the authors of thevarious papers have recognised this issue and madeformal adjustments for multiple testing, the Bonferroni
correction (Rice, 1989) used is known to beconservative, with the result that true departures can goundetected. This issue also affects the strength of theconclusions one can draw from the failure to findsignificant differences.
In the light of this, the Panel agrees that it is not possible
to conclude, as the Proponents did, that the number ofhypotheses proposed during the Implementations forwestern North Pacific common minke whales and Bryde’s
whales has been unequivocally reduced. However, this does
not mean that such conclusions may not be the ultimate
outcome of the additional analyses recommended below;that remains to be seen.
While recognising the considerable value of the stock
structure work already undertaken, the Panel has thefollowing specific recommendations/suggestions for ways
in which the scientific benefits from JARPN II stockstructure research can be enhanced. Some of theserecommendations should be easy to implement; others willtake more time and effort.
6.3.1 Simple issues(1) The genetic assessments should include a brief
description of procedures to ensure data quality. This

419

section should refer to the recently discussed IWC
guidelines for DNA data quality (IWC, 2009a).
(2) The revised papers should include estimates of genetic
divergence (along with levels of uncertainty) in addition
to probabilities of homogeneity6.
(3) P values (and divergence estimates) should be reported
for all loci combined rather than for each locus
separately. In addition to providing more useful
information and increasing statistical power, this will
help reduce issues related to multiple testing.
(4) Multiple testing issues will still arise in some cases. In
general, use of the False Discovery Rate (Benjamini
and Yekutieli, 2001) could be preferable to theBonferroni correction, as it is less conservative anddoes not sacrifice as much power. Another strategy thatcan be useful is to exercise discretion in the number of
pairwise comparisons that are evaluated – for example,
by only comparing samples that are geographicallyproximate and hence most likely to be connected
demographically. See Økland et al. (2008) for an
example of this approach.
(5) Provide more details7 on the analyses involving the
program STRUCTURE (Pritchard et al., 2000a).
(6) Include a brief discussion of experimental design withrespect to sampling. Although the rationale for the
sampling design is discussed in other papers (especially
SC/J09/JR3 and JR4), it would benefit these evaluations
to have a short discussion 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 under consideration by the IWC.
6.3.2 More extensive matters – some of which might ideallybe addressed in time for the 2009 Annual Meeting8(1) The original justification for considering hypothesis C(and to some extent D) for common minke whales wasprimarily based on results from the Boundary Rank
analyses (Taylor and Martien, 2002) – it would be
informative to redo those analyses using the new data(including taking into account information on
assignment of animals to ‘J’ stock) to see if evidence
for a narrow coastal stock remains.
(2) Previous analyses of 1999 and 2000 Korean bycatchsamples suggested that they differed from other ‘J’
stock samples. It is important to integrate these samples
into the new datasets to see if this heterogeneity still
exists.
6
Some information on this issue was kindly developed by the proponents
during the Workshop.
7
Specific examples include: color figures showing assignment probabilities
for each individual (for k=2 and k=3); indicators of convergence of MCMC
estimations including information about repeatability of assignment
probabilities of individuals to the putative ‘J’ and ‘O’ stocks (andunassigned categories) in different runs (e.g. CV of assignment
probabilities for the same individuals across multiple MCMC estimations
and comment on whether the same individuals consistently end up being
unassigned or if there is a different mix of individuals in each run); moredetailed geographical plots of the distribution of ‘J’, ‘O’, and unassigned
individuals; include unassigned individuals in plots.
8
The issues concerning deviations from population genetic equilibrium andstatistical power also apply to the Bryde’s and sei whale. Note: Points (1)and (2) are important but it is not necessarily the responsibility of JARPN
II to undertake them.

420

REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II(3) Assessments of power using simulated data should beundertaken (see Annex G). It should be relatively
straightforward to simulate data to evaluate power todetect a specified fraction of a putative stock (e.g. thehypothetical W stock of NP minke whales) in an overall
sample. This analysis would require specifying a range
of genetic divergence values (e.g. FST values) for theputative stock. More challenging but still feasible
would be simulations to evaluate the power of
STRUCTURE (Pritchard et al., 2000a) to detect various
mixture fractions of closely related stocks. Althoughstrictly speaking power is a frequentist concept and
STRUCTURE uses Bayesian methodology, it should bepossible to construct a power analogue that reflects
robustness to delivering what, with simulated data, is
known to be the correct answer. In this case, thespecific question could be: how large a proportion of
the samples could be from another population and still
result in a situation in which K=1 is favoured with highprobability? These simulations could be carried out
using the programs SimCoal (Laval and Excoffier,2004), ms (Hudson, 2002), EasyPop9 (Balloux, 2001) orother freely available software. The statistical power todetect a second stock that is under-represented in asample will depend upon the fraction of these
individuals in the total sample and how geneticallydivergent they are. This implies that simulations either
are conducted as a sensitivity test (i.e. assessing a rangeof combinations of the sample proportion and degree ofgenetic divergence) or by deciding upon a minimum
case (e.g. the minimum detectable fraction should be
X% at a degree of divergence equivalent to a migration
rate at Y migrants per generation).
(4) Tests for population genetic (drift-mutation-migration)equilibrium should be undertaken. High haplotype
diversities coupled with low nucleotide diversities
indicate
deviations
frompopulation
genetic
equilibrium.
(5) Estimations of divergence between sample partitions
should be undertaken using non-equilibrium
approaches. An example of such an approach is IM by
Hey and Nielsen (2004); the estimates of dispersal
emerging from IM may be used in the powerassessment simulations suggested above. It is probably
advantageous to conduct initial estimations between thepotentially most divergent sample partitions (e.g. mostextreme parts of the range). These methods improve theapproach previously used to estimate dispersal rates forthe common minke whale Implementation Simulation
Trials (Taylor and Martien, 2004).
(6) With genotypes from 17 microsatellite loci in 2,500
individuals, it may be possible to detect pairs ofindividuals that are related, as was the case among asmaller set of samples genotyped at the same number ofloci in the North Atlantic fin whale (Skaug and
Daníelsdóttir, 2006). The spatial distribution of related
individuals can provide information directly relevant tostock structure considerations; Økland et al. (2008)
have demonstrated the use of such an approach.
Notably, such analyses provide information about

contemporary stock structure and do not rely uponassumptions of population genetic equilibrium.
(7) Multivariate analyses of morphological data could be
informative with respect to stock structure. Many
multivariate classification methods (such as cluster
analysis, discriminant analysis, SIMPER and ANOSIM)
now include permutation tests. Another option would
be to use a principle components (or similar) analysis ofindividuals that does not require a priori decisions
about group membership. PCA does not attempt todefine groups but can reveal patterns in the data related
to time or place of sampling.
(8) Data on contaminants in western North Pacific minke
whales have been reported (Fujise, 1996) and were used
as further support for the baseline C stock structure
hypothesis (Taylor and Martien, 2004). The use of pastand present contaminant data should continue to be
pursued as part of an integrative study of stock
structure.
6.3.3 Longer term
Notwithstanding the practical difficulties associated with
attaching satellite tags to minke whales, the increasing
success of satellite tagging programmes for several whale
species (e.g. see Weller, 200810) suggests that efforts should
be made to establish such a programme for western North
Pacific common minke whales. Information such a
programme might produce could be very valuable inallowing the IWC to narrow the range of plausible stockstructure hypotheses.

7. REVIEW OF JARPN II RESULTS: REVIEW OFOTHER CONTRIBUTIONS TO IMPORTANT
RESEARCH NEEDS
7.1 Oceanography
7.1.1 Statement of objectives and Proponents’ summary
The Government of Japan stated (Government of Japan,
2002b, p.34) that during JARPN II, oceanography surveys
will be obtained using XCTD (Expendable Conductivity,
Temperature and Depth Sensors), CTD and EPCS (Electric
Particle Counting and Sizing System).
More specific objectives included:(1) clarify the geographical distribution patterns of preyspecies of common minke whale in relation to
oceanographic features; and(2) clarify the geographical distribution patterns ofcommon minke whale in relation to prey environmentand oceanographic features (SC/J09/JR11).
Oceanographic data collected during the cruises included:
on the sighting and prey sampling vessels in the coastal andoffshore study areas, water column profiles of temperature,salinity, and density using XBTs (expendable
bathythermographs), XCTDs, and CTDs; and only on the
Shonan Maru No. 2, the continuous measurements ofsurface water temperature, conductivity, chlorophyll,
dissolved oxygen and particles using EPCS. Oceanographic
data were not collected on the whale sampling vessels.
SC/J09/JR34 summarised the oceanographic conditionsof the western North Pacific from 2002 to 2007 in the9

http://www.unil.ch/webdav/site/dee/shared/softs/EASYPOP_201_userguid
e.pdf (Balloux, 2001).

10

http://www.mmc.gov/pdf/final_tagging_82608.pdf.

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

JARPN II coastal and offshore study areas using the CTD
and XCTD data collected during the cruises as well as datafrom the Global Temperature and Salinity Profile Program(GTSPP) that were incorporated in FRA JCOPE, an
operational ocean forecast system. Oceanographic fronts
and water bodies were classified by surface currents andsubsurface temperature, salinity and density gradients. Yearto year variations in the spatial distributions and strengths ofthe various fronts, warm core rings, and water bodies were
described.
In addition to this overall description of theoceanographic characteristics in the study areas, a few otherstudies investigated relationships with respect tooceanographic characteristics. SC/J09/JR11 described
preferred temperature ranges and distributions of thepotential prey species and common minke whales inrelationship to the major water bodies, fronts and bottom
depths in the Kushiro coastal study area.
SC/J09/JR13 compared the distributions of potential preyspecies and maturity status of common minke whales
relative to their distribution on the continental shelf,
continental slope, and offshore waters in the Kushiro coastal
study area.
SC/J09/JR19 developed density prediction models on amonthly 1 × 1° degree grid for common minke, sei and
Bryde’s whales in the Western North Pacific in theirfeeding season using NPMR (non-parametric multiplicative
regression models) which included satellite derived SST,
chlorophyll, and sea surface height as potential predictor
variables.
SC/J09/JR35 identified the SST ranges where most blue,
fin, and North Pacific right whales were seen during the
JARPN and JARPN II sightings surveys during 1994 to2007.
SC/J09/JR36 estimated the abundance of North Pacificsei whales using SV and JSV sightings data in a hierarchicalGAM of the interactions between school density and thefollowing environmental factors: satellite derived SST, sea
surface height anomalies, and sea surface chlorophyll
concentration, and CTD/XCTD derived water temperaturesand salinities at 50, 100 and 200m depths.
Murase et al. (2007) briefly discussed the distribution
patterns of Japanese anchovy, walleye Pollock and krill in
relationship to the Kuroshio-Oyashio transition zone and thecontinental slope and continental shelf zones.
The Proponents recognised that their studies which relate
oceanographic features to distributions of prey and whales
are in progress. Before these relationships can be identified,the underlying data need to be collected and processed. At
the time of this review, most of the initial oceanography
data processing has been completed, which will now allow
future analyses that relate oceanographic features toimportant aspects of the whale ecosystem.
7.1.2 Panel’s conclusions and recommendations
The Panel congratulates the Proponents for simultaneously
collecting in situ sea surface and water column
characteristics while conducting the whale and preysurveys, recognising the practical challenges ofcoordinating these sampling methods on the same ship at
the same time. Such simultaneous collections represent thebest way to obtain a direct comparison between
oceanographic features and the distributions of the various
trophic level components. The Panel welcomes these421

analyses as a good initial attempt at investigating relations
with oceanographic features and they encourage theanalyses to be continued and expanded. The programme isaddressing its objectives and continued work is
recommended.
At the practical level, the Panel concurred with thesuggestion of the authors of SC/J09/JR34 that the salinity
CTD data must be corrected/calibrated using the water
samples that were simultaneously collected with the CTD
data.
The Panel welcomed SC/J09/JR19, which predicted thedensity spatial-temporal patterns of common minke, sei and
Bryde’s whale distributions, as a good initial example ofattempting to integrate several data types to investigate therelationships between whale densities and oceanographic
features. The use of NPMR modelling techniques isappropriate, particularly if there are complex interactions
among the ecological factors.
To improve this approach, the Panel recommends thatthe authors incorporate into the index of density, thesightability of detected groups (e.g. effective strip half
widths that include appropriate covariates such as weather
conditions). As for all modelling exercises, it is important totest whether the chosen model is an improvement over anull, uninformative model and to validate the model results.
Approaches to such validation could include: comparison of
the modelled results not only with index of densities fromthe present study but also with data that were collected fromother years (e.g. JARPN or other survey data) and
exploration of cross-validation type techniques. The Panelrecommends that more of these types of analyses(including using other appropriate modelling techniquessuch as GAMs or logistic regressions) be conducted. The
shipboard oceanographic data that were collected should beconsidered in future models, as was suggested by theauthors of SC/J09/JR19. Potential additional oceanographic/
biological features that could be investigated include
modelling the satellite or in situ measurements ofchlorophyll to estimate primary productivity. Other
examples of potential future analyses that investigate
relationships with oceanographic features are described in
Item 8.2.
GENERAL RECOMMENDATIONS

To further investigate oceanographic relationships, thePanel recommends that the JARPN II data be pooled orcompared with other datasets (e.g. JARPN I or otherhistorical surveys) when possible. This will increase thesample size and increase the possibility of data covering
periods of changing relationships (e.g. previous regimechanges), thus allowing patterns to be detected.
The Panel also suggests 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.
In summary, the Panel recommends that in the longterm, 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 (such as blubber
thickness, pollutants, presence of scars and stock structure),
the oceanographic data collected on the cruises (bottom
depth, water column temperature, salinity and density) and422

REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

satellite derived data, such as SST, chlorophyll, and sea
surface height be integrated into future analyses.
7.2 Distribution of large whales
7.2.1 Statement of objectives and Proponents’ summary
The Government of Japan stated that during JARPN II,
sighting records will be collected for all cetaceans and in
addition to sighting records, photo-ID, observations ofwhale behaviour, especially feeding behaviour, will berecorded (Government of Japan, 2002b, p.33).
Results related to the distribution and abundance of thefour target species are discussed elsewhere in the Panelreport. SC/J09/JR35 reported the Density Index (DI:individuals/100 n.miles) and monthly distribution pattern ofblue, fin, humpback and right whales from May to
September in the western North Pacific based on JARPN(1994-1999) and JARPN II (2000-2007) sighting data.
Among four species, fin whales were most frequently
sighted, and next were blue, humpback and right whales inorder. Northward migration patterns of whales wereobserved for these species. Additionally, sighting areas ofthese species were spread out compared to the previous
information except for right whales. SC/J09/JR1summarised the photo-identification data collected during
the programme: 24 schools of humpback whales, 22 schoolsof North Pacific right whales and 65 schools of blue whales
had been photographed.
7.2.2 Panel conclusions and recommendations
The Panel welcomes SC/J09/JR35 and the analyses of the
distribution. To investigate relationships with oceanographic variables and improve abundance estimates of avariety of species, the Proponents could consider
conducting future surveys that cover an area larger than thatof the present JARPN II sampling area.
Using the sightings data collected over the 1994-2007
period for the variety of large whales, the Panelrecommends investigation of whether these data can beused to provide information on trends. It also recommends
that the photo-identification data be worked up andcomparisons made with catalogues elsewhere in the North
Pacific.
7.3 Other research
7.3.1 Proponents’ summary
REPRODUCTIVE BIOLOGY AND PHYSIOLOGY

Watanabe et al. (2004) presents a study to obtain new
information on relationships among serum testosterone (T),estradiol-17β (E2), follicle-stimulating hormone (FSH), andluteinizing hormone (LH) concentrations and histology ofseminiferous tubules in captured common minke andBryde’s whales during the feeding season. Results indicatedthat the low serum T concentrations reflect the inactivity ofspermatogenesis in both baleen whales, and that it is notpossible to assess gonadal activity in either species using
serum sex hormone concentrations during the feeding
season.
Watanabe et al. (2007) investigated whether spermatozoa
of Bryde’s whale can retain the capacity for oocyte
activation and pro nucleus formation as well as
chromosomal integrity under cryopreservation by using
intra cytoplasmic sperm injection (ICSI) into mouse
oocytes. Results showed that motile Bryde’s whale

spermatozoa are competent to support embryonicdevelopment.
Urashima et al. (2007) examined samples of milk from a
Bryde’s whale and a sei whale. Milk samples of these
species contained 2.7 g/100 mL and 1.7 g/100 mL ofhexose, respectively. Both contained lactose as thedominant saccharide. The dominance of lactose in the
carbohydrate of these milks is similar to that of minke
whale milk and bottlenose dolphin colostrum, but theoligosaccharide patterns are different from those of these
two species, illustrating the heterogeneity of milkoligosaccharides among the Cetacea.
Fukui et al. (2007) presented a study aimed at producing
common minke whale embryos. The study was based onminke whale samples collected in the coastal researchcomponent off Kushiro. Results indicated that a 40 h IVM(in vitro maturation) culture produces significantly higher
rates of in vitro maturation than a 30 h IVM culture forcommon minke whale oocytes. Following ISI (intracytoplasmatic sperm injection), some oocytes cleaved to the
16-cell stage, but no further development was observed.
Birukawa et al. (2008) examined kidney samples ofcommon minke, sei, Bryde’s and sperm whales to determine
the nucleotide sequences of mRNAs encoding UT (ureatransporters). Urea transport in the kidney is important forthe production of concentrated urine. It was speculated thatdifferent phosphorylation sites found in whale UT-A2s may
result in the high concentrations of urinary urea in whales,by reflecting their urea permeability.
GENETICS

Nishida et al. (2007) reconstructed cetacean phylogeny
using a 1.7-kbp fragment of the non-recombining Y
chromosome (NRY), including the SRY gene and aflanking non-coding region. The topology of the Ychromosome tree is robust to various methods of analysis
and exhibits high branch-support values, possibly due to theabsence of recombination, small effective population size,and low homoplasy. The Y-chromosome tree indicatesmonophyly of each suborder, Mysticeti and Odontoceti,with high branch support values.
Onbe et al. (2007) examined the amino-acid sequencesof the T-domain region of the Tbx4 gene, which is requiredfor hindlimb development. Cetaceans have lost most of theirhindlimb structure, although hindlimb buds are present invery early cetacean embryos. They investigated whether the
Tbx4 gene has the same function in cetaceans as in othermammals. The study concluded that the Tbx4 gene
maintains its function in cetaceans, although full expression
leading to hind limb development is suppressed.
7.3.2 Panel conclusions and recommendations
The Panel welcomes the results of these additional research
projects, noting that they have resulted in publications inpeer-reviewed journals.
7.4 Abundance
7.4 1 Summary of results to date
Survey procedures and summaries of the surveys conductedto date are described in SC/J09/JR2. The Sanriku coastal
area was surveyed in April-May each year, the Kushiro
coastal area in September-October, and the offshore area in
May-August. The entire JARPN survey was surveyed inJ. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

two years in 2002-03 and 2004-05, and in each year 2006
and 2007.
Abundance estimates for the JARPN II area have beenobtained for minke, sei and Bryde’s whales using bothconventional stratification (SC/J09/JR8, JR15) and spatialmodelling using NPMR (SC/J09/JR19) and GAMs
(SC/J09/JR36, Appendix 2). Distribution maps have been
produced for fin, humpback and right whales(SC/J09/JR35).
In addition to the surveys in the JARPN II area, theOkhotsk Sea was surveyed in 2003 and the Russian EEZeast of Kuril and Kamchatka was surveyed in 2005. The
estimates of abundance that were used for the assessment of
the effects of JARPN II catches on the minke whale ‘O’stock include estimates for the Okhotsk Sea andKamchatka-Kuril, on the assumption that these animals arefrom the same stock as the JARPN II area (SC/J09/JR36).
The Sea of Japan has also been surveyed during the
JARPN II period, including a survey in the Russian EEZ in2006. The results of the surveys have been used to obtain
estimates of abundance of the ‘J’ stock, which are used in
the assessment of the effect of JARPN II catches on the ‘J’stock (SC/J09/JR36).
7.4.2 Analysis and comment
While the Panel did not examine this in great detail, thededicated survey procedures appear to be in accordancewith IWC Scientific Committee guidelines for abundance
data for use in the RMP (IWC, 2005); g(0) has been
estimated from independent observer (IO) data for minke
whales (Okamura et al., 2008b), and is assumed to be 1.0
for larger baleen whales.
The timing and distribution of survey effort has changed
from year to year (see maps in SC/J09/JR2) due to weather
and logistic constraints, and also to fill in holes left inprevious surveys. This means that monthly and annual
effects may be confounded to some extent in the data series.
This might limit the value of the data to analyse suchvariations.
7.4.3 Recommendations
Recommendations for the conduct of sightings surveys andmethods of analysis are regularly made by the IWC
Scientific Committee, and the Panel did not attempt toduplicate this work.
The abundance data collected in the JARPN II area areprimarily for the purpose of determining the density anddistribution of whales with respect to their consumption of
resources. The Panel emphasised the great importance of
the abundance data for the consumption estimates. As noted
under Item 4, the confidence intervals for the abundance
estimates are generally wide, especially in the coastal area;
the Panel recommends that increased effort to obtain better
estimates should be a high priority.
In addition, total abundance estimates are an essential
component of estimating total population sizes for thedetermination of the expected effect of catches on thestocks. As discussed under Item 9.3, the Review Panelcautions against extrapolations from the JARPN II area forthis purpose.

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8. REVIEW OF JARPN II RESULTS: THE
RELATIONSHIP OF THE RESEARCH TO
RELEVANT IWC RESOLUTIONS ANDDISCUSSIONS
A summary of the relevant recent resolutions is given in
Annex H, based on Zeh et al. (2005).
8.1 Research on the ecosystem and environmental
change8.1.1 Summary of Proponents’ view (from SC/J09/JR1)
Resolution 1994-13 (IWC, 1995a) encouraged Contracting
Governments/Scientific Committee to study environmental
changes and impact on cetaceans. Resolution 1995-10(IWC, 1996b) encouraged Contracting Governments tostudy the effects of pollutants on cetaceans as recommended
by the Bergen workshop. Resolution 1997-7 (IWC, 1998a)
encouraged Contracting Governments to continue toprovide information on environmental changes and potential
effects on cetaceans. Resolution 1999-4 (IWC, 2000)
requested Contracting Governments to provide the
Scientific Committee with data on contaminants incetaceans.
JARPN II included the monitoring and assessment ofchemical pollutants in whales and their environment.
Information on feeding ecology is important for examining
the effects of environmental changes. Therefore JARPN II
is investigating several of the topics mentioned in these
Resolutions.
Resolution 1998-7 (IWC, 1999) invited Japan to take fulladvantage of the existing mechanisms for cooperation
between national research programs and the ScientificCommittee’s Standing Working Group on environmental
concerns.
Japan has accepted this invitation positively and will
cooperate by providing the information on environmental
research obtained in JARPN II.
8.1.2 Panel conclusions and recommendations
The Panel agrees that many of the objectives of JARPN II
are relevant to Resolutions of the Commission and thatscientific results have been submitted to the ScientificCommittee, as requested in several of the Resolutions.
8.2 Utility of the lethal techniques used by JARPN II
compared to non-lethal techniques
8.2.1 Summary of Proponents’ view (from SC/J09/JR1)
Resolution 1995-9 (IWC, 1996a) had recommended
Contracting Governments to use non-lethal methods andinstructed the Scientific Committee to review scientificpermit research in the light of the use of non-lethal methods.
One of the main characteristics of JARPN II is the
combination of both lethal and non-lethal surveys andanalyses, which is important for achieving the main
objective of the research programme, i.e. feeding ecology
and ecosystem studies as discussed under Item 4; somebiological information from whales can be obtained only
through the lethal approach.
General discussions on lethal versus non-lethal
approaches in whale research under special permits haveoccurred several times in the IWC Scientific Committee inthe past (IWC, 1998b; 2008a).
To estimate prey preferences of cetaceans, examinationof stomach content data is the only way to identify prey

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REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

species consumed and quantify prey consumption (e.g.
SC/J09/JR9; 16 and 17). However, the feeding ecology ofbaleen whales can be studied using several methods otherthan examination of stomach contents. Barros and Clarke(2002) categorised those methods as follows: direct
observation of feeding; traditional methods (analysis ofvomit, scat, stomach and intestine contents); fatty acids;
stable isotopes; genetic identification of scat; andvideotaping of feeding behaviour.
In recent years, telemetric studies (e.g. satellite tags,
time-depth recorders) have also been used for feeding
ecology studies (e.g. Croll et al., 1998). Direct observationof feeding is limited to above the sea surface or shortduration underwater observation by scuba survey. Given thediversities and vertical distribution patterns of prey speciesof minke, Bryde’s and sei whales, the direct observation
method is not applicable to those species. Haug and
Lindstrøm (2002) compared the traditional methods withthe rest of the new methods. They concluded that the new
methods have not proven to provide detailed quantitative
information on the diet of individual animals and must be
supplemented with traditional methods. Identifying andmeasuring items in gastrointestinal contents with a
combination of methods could reinforce the reliability of
any conclusions.
Regarding environmental studies (see Item 5) somepollutants are organ-specific and therefore studying
different organs for different pollutants will provide
valuable information. Furthermore, studies on the effect ofpollutants on the health of whales require examination oftarget organs. The level of some lipophilic pollutants can bemeasured from blubber samples obtained by biopsy
sampling. However, biopsy sampling is a difficult method
for collecting samples over the whole research period andarea of JARPN II. This is because sampling efficiency isaffected by wind force, the particular research area and thetargeted school size. The difficulty of obtaining biopsy
samples could be also different among whale species,
depending on body size, swimming speed and pattern of
movement.
To understand the pattern of accumulation of pollutantsin whales, it is important to have access to some biologicalinformation about the whales under investigation such assex, reproductive status, body length, weight of stomach
contents, some of which can be obtained only by using thelethal approach. A previous study attempted to assess
reproductive status by examining hormone metabolites infaecal samples of the right whale (Rolland et al., 2005).
However there are practical difficulties for obtaining faecal
samples.
Studies on stock structure under JARPN II (see Item 6)
are based on both genetic and non-genetic approaches. The
non-genetic approach used in JARPN II (for common minke
whale at this stage) is morphometrics, which requires that
accurate body measurements be obtained from whales
sampled. Genetic analysis on stock structure based on DNA
can be carried out by using biopsy samples. As explained
above, there are some practical problems with the use ofbiopsy sampling.
8.2.2 Panel conclusions and recommendations
Both lethal and non-lethal methods are often used to address
the same question (for example stock structure, diet andcontaminant exposure), even if they do not always provide

precisely the same data. The number of new non-lethal
techniques being developed is increasing apace and thePanel recommends that a proper evaluation of the use of
non-lethal and lethal techniques in any long-term
programme such as this should occur periodically atappropriate intervals – this applies both to the developmentof new analytical techniques and to technical developmentsthat allow appropriate samples to be collected from freeranging animals. Clearly, it also involves regular carefulanalysis of appropriate sample sizes (see Item 9.2) over along-term programme, since additional data from lethal
sampling may become less important/unnecessary, in termsof meeting objectives, as the programme progresses.
A full comparison of the various lethal and non-lethal
techniques requires inter alia an analysis of the bias andprecision of the estimates obtained using the different
approaches in the light of stated quantitative objectives or
sub-objectives; often the data to allow such a comparison
are not available (see recommendation below) and/or
objectives are insufficiently stated/quantified (see Item
9.2.1). In addition, for a complex multi-disciplinary
research programme such as JARPN II, an evaluation of
appropriate techniques (lethal or non-lethal) must include an
integrative analysis to ensure maximised efficiency fromboth a scientific and logistical perspective. The Panel was
not in a position to evaluate this in detail in the absence ofknowledge of available resources and the outcome ofanalyses detailed under Item 9.2 to assist with an evaluationof sampling strategies and sizes. Any comparative analysis
must also take into account any biases arising out of therelevant sampling strategies.
Given these important difficulties and information gaps
(not all of which are specific only to this particular
programme), the Panel does not consider its evaluation
below to comprise a full quantitative evaluation as
described in the preceding paragraph; that is not possible
given the available data. Rather, it presents a very brief
initial evaluation/commentary on the available techniques in
the context of the primary stated objectives of JARPN II
that are presently addressed with lethal techniques.
FEEDING ECOLOGY

A suite of non-lethal methods are available for studying dietand foraging ecology of marine mammals (e.g. Barros and
Clarke, 2002). However, proper evaluation of their value
relative to lethal sampling is in many cases difficult, as few
studies have properly and quantitatively compared theresults of the different approaches; in fact for many
approaches appropriate measures of precision are as yetlacking. The primary rationale for stomach sampling, which
requires killing the animal, is the qualitative andquantitative information on prey composition (e.g. species,age) and stomach fullness that it provides, all of which arevaluable when studying consumption and functional
relationships between marine mammals and their prey.
However, as has been noted elsewhere (See Item 4 andreviews such as those of Leaper and Lavigne, 2007), suchdata also have their limitations and these must berecognised in any analyses. Combinations of several
methods applied simultaneously to the same animals are aparticularly powerful approach (Karnovsky et al., 2008).
The following list very briefly summarises the mostcommon non-lethal methods used in feeding ecology
studies and the information they provide:J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010
•

•

•

•

•

•

Direct observations of feeding. Primarily giveinformation only on surface feeding whales and eventhen may rely on inference. Cameras mounted on
marine mammals can provide information on preyencounters (e.g. Bowen et al., 2002a; Bowen et al.,
2002b), but the difficulties in deploying cameras on
fast-moving whales such as the rorquals, even if itproved possible, would probably result in substantiallyreduced sample sizes in terms of individuals, althoughpotentially increasing the number of feeding events
observed per individual.
Analyses of faeces is a common approach to studying
diet in seals (e.g. Casper et al., 2007) and theexamination of DNA in faeces to determine thepresence or absence of important prey groups has been
applied to several cetacean species (e.g. Gales andJarman, 2002; Jarman et al., 2006). The feasibility ofobtaining sufficient samples, particularly on the highseas for the JARPN II target species, would need to beevaluated.
Analyses of stomach samples from bycaught orespecially stranded individuals is possible but care mustbe given to using the results due to inter alia nonrandom sampling, reduced spatial coverage and thecompromised state of stranded animals that may haveaffected their ability to feed before stranding.
Analysis of fatty acid composition from biopsy samples
is a promising method both in terms of qualitative andquantitative assessment of diet, which reflects the diet
integrated over a period of around 1.5-3 months(Nordstrom et al., 2008). Providing quantitative (i.e.
proportional) contribution of different prey species, thismethod could be used for quantifying functional
relationships. However, the method may still need
improvements in separating contributions from related
species, and in calibrating tuning levels (Nordstrom etal., 2008). Practical aspects of taking biopsy samples in
the JARPN II context are discussed under the section on
genetic studies.
Analyses of stable isotopes on tissues from biopsy
samples quantify trophic level and indicate relative
contributions to the diet of different potential primarysources in a trophic network. This method integrates
diet information over weeks/month (e.g. Das et al.,2003a; 2003b). While stable isotope analysis does notquantify the diet, it provides a valuable tool for groundtruthing the trophic level estimated obtained from othermethods. Again, practical aspects of taking biopsy
samples in the JARPN II context are discussed below.
Analyses of spatial associations between whales andtheir prey from synoptic surveys simultaneously
recording whale and prey distribution may provide
information on which prey species are targeted,particularly in areas/systems with few, spatiallysegregated prey species (Mauritzen et al., In press).
Such an approach could yield quantitative informationon the proportion of whales targeting the different
species, but should be combined with more quantitative
methods for estimating prey composition or trophic
level.

POLLUTANT STUDIES

A number of pollutant analyses can be undertaken on
samples obtained non-lethally (i.e. biopsy samples) that can

425

be used to monitor temporal and spatial trends in the levelsof certain contaminants that are deposited in skin andblubber, although the range of contaminants is limited. In
addition, many persistent environmental contaminants cooccur, so monitoring a subset of elements and POPs
(persistent organic pollutants) should be sufficient to detectchanges in several pollutants. However, this would not be
possible for all contaminants. In particular, studies on the
newer groups of emerging compounds and the more
complex toxicological, mass balance and health studies
could not be undertaken using data from biopsy samplingalone at least at present. Biomarkers (such as metabolising
enzymes) are expressed in skin as well as liver so biopsy
samples could be used once the relationship between skinexpression and liver expression has been established.
Validation of the use of biopsy samples was an importantcomponent of the IWC’s POLLUTION 2000+ programme
and was undertaken successfully for some small cetacean
species (Reijnders et al., 2007). The limitations of bycaught
and especially stranded animals for pollutant studies havebeen well documented elsewhere (Reijnders et al., 1999).
GENETIC STUDIESFor genetic studies, skin and outer layer blubber samples
may be collected by non-lethal remote biopsy sampling
methods from free-ranging baleen whales. Biopsy samplingfrom free-ranging baleen whales is now commonplace andthousands of biopsies have been collected in this manner;
this includes (although relatively few) minke whales.
Current methods consist of relatively light equipment (e.g.
150lbs draw-weight crossbows) and the collection ofsamples is typically conducted from light open boats at
close range. Heavier, rifle-based, long-range biopsy
equipment for high seas conditions have been developed aswell (e.g. the ‘Larsen gun’ used inter alia on the IWC
SOWER cruises).
Analyses of genomic DNA are readily conducted on skinbiopsy samples, which yield more than adequate amounts ofgenomic DNA. In contrast, gene expression analyses (which
are based upon analyses of mRNA levels) may be tissuespecific and thus not possible using skin biopsies.
PRACTICALITIES OF BIOPSY SAMPLING IN THE CONTEXT OFJARPN II11

The effort required for collecting skin biopsies in principle
should be comparable to (or less than) the effort required forlethal sampling, provided the biopsy equipment has beendeveloped to allow efficient collection on the high seas.
However, a significant proportion of the sampling carriedout during JARPN II is conducted on the high seas, and thus
under conditions that are appreciably more challenging thanthe majority of current skin biopsy sampling projects, which
typically are undertaken in coastal areas. The IWC
Antarctic SOWER cruises (large vessels on high seas) havesuccessfully obtained samples from a number of large
whales including southern right whales, blue whales, fin
whales and humpback whales, and have also undertaken an
experiment with Antarctic minke whales. It would bevaluable to examine the available biopsy sampling andeffort data from the SOWER cruises to examine theefficiency of this further.
11
As detailed later in this section, biopsy samples can be used to address
aspects of a number of the objectives of JARPN II. This discussion ofpracticality is equally relevant there. Of course, the mass of the sample
may become relevant if multiple analyses are required.

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The main limitation of biopsy collection on the high seasfrom large vessels is the limited range and low mass ofcurrent projectile units (which carry the biopsy tip). A
heavy projectile is less sensitive to high winds and thus
development of biopsy equipment for use on the high seas
would need to aim at developing heavier projectile units
than those currently employed in most biopsy systems. The
use of heavier projectile units would necessitate the use of
more powerful delivery units in order to obtain thenecessary range and trajectory. However, adding mass andpower to the projectile unit increases the risk ofunwarranted penetration and damage to the target animal;
the collar (preventing penetration beyond the depth of thebiopsy tip) needs be of an appropriate size. If further lethal
sampling is undertaken during JARPN II, this would
provide an opportunity to test the potential impact of heavy,
high-powered biopsy system upon whole animals.
In summary, provided that the practical difficulties in
obtaining biopsy samples on the high seas can be overcome,then analyses of genomic DNA are readily conducted onskin biopsy samples which yield more than adequate
amounts of genomic DNA.
GENERAL CO-VARIATES

Many analyses related to all of the objectives of JARPN II
benefit from more detailed knowledge of the individuals
from which the samples come (e.g. age, reproductive status,
sex and health). Some of these can only be obtained fromdead animals, but techniques to obtain such informationfrom biopsy samples (and perhaps faecal samples) are
increasing (Jarman et al., 2006; Jarman et al., 2003; Rolland
et al., 2005). At present, sex is the easiest to obtain frombiopsy samples but others, such as age are in development
and appear very promising, at least for some species(Herman et al., 2007; Herman et al., 2008).
GENERAL CONCLUSIONS AND RECOMMENDATIONSThe Panel recommends that a full evaluation of theapplicability of lethal and non-lethal techniques be
undertaken as soon as possible after the relevant work
recommended elsewhere in this report has been undertaken.
Such a full evaluation inter alia requires information on thefollowing:(1) Specified and quantified objectives and sub-objectives(see recommendation under Item 9.2.1);(2) Analysis of the precision (and any associated biases) of
the estimates obtained for the relevant parameters byeach of the lethal and non-lethal techniques (seerecommendations under Item 4 and at the end of thissection);(3) Evaluation of the practicalities and logisitics of the field(and, if relevant, laboratory) techniques in the context
of the integrated objectives, sub-objectives andanalyses proposed.
The Panel recognises that at present, certain data,primarily stomach content data, are only available via lethal
sampling. However, if lethal sampling is deemed necessary
by Proponents for a research programme, the Panelconsiders that there is an obligation to maximise theinformation obtained from those animals and to re-evaluate
the need for such sampling at appropriate intervals.
As noted above, the ability to fully evaluate and compare
non-lethal methods in a quantitative manner is severely
limited by a lack of appropriate data. In addition,

examination of the effects of post-mortem time andconditions on various analyses of tissue samples would
allow an evaluation of the suitability or otherwise of tissue
from stranded or bycaught animals (e.g. see the
POLLUTION 2000+ programme). The Panel recognisesthat such an objective (i.e. to quantitatively compare lethal
and non-lethal research techniques) is not part of the present
JARPN II programme. However, it considers that a welldesigned and implemented programme to evaluate andcompare lethal and non-lethal methods, especially those
directly related to the existing JARPN II objectives
summarised above, would provide extremely valuable andimportant scientific information that will greatly assist
scientific research worldwide as well as improve the abilityof future Panels and Proponents to objectively address thisTerm of Reference. It will provide a unique opportunity tocompare results of stomach analyses with non-lethal
estimates that can be obtained from the same time andplace. This provides a way of evaluating the reliability andefficiency of the non-lethal methods.
The Panel therefore strongly recommends that Japanconsiders the addition of an objective to quantitatively
compare lethal and non-lethal research techniques if it
decides to continue a lethal sampling programme.
Appropriate samples can be archived for future analysis if
necessary. Whilst recognising the sensitivities surrounding
this issue, the Panel respectfully requests that if lethal
sampling programmes occur, the IWC or an appropriatescientific body or bodies may wish to consider collaboratingin the design of a well specified study to fully evaluate
lethal and non-lethal techniques.
9. ADVICE ON ONGOING SPECIAL PERMITRESEARCH
9.1 Practical and analytical methods, including nonlethal methods, that can improve research relative tostated objectives
The report details a number of practical and analytical
recommendations and suggestions that will improve theresearch, under each of the relevant agenda items.
9.2 Appropriate sample sizes to meet the statedobjectives, especially if new methods are suggested
under Item 9.1
9.2.1 Proponents’ view (from SC/J09/JR1)
Sample sizes of minke, sei and Bryde’s whales were set for
estimating prey consumption with good precision(coefficient of variation, CV=0.2), in the same way as in thecase of the Norwegian research programme (NMMRP,
1992). The sample size of sperm whale was set as aminimum level necessary for obtaining qualitative
information for a feasibility study.
The CV of stomach content weight of three baleen
whales from the first six years of JARPN II can besummarised as follows (see SC/J09/JR1, Annex 6 fordetails):(a) Target CV was satisfactory in most cases forminke whales. This means that the sample sizes ofminke whales for both offshore and coastal
component seem to be appropriate.
(b) CVs were larger than the target in most cases forsei whales. This means that the sample size was

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

smaller than the appropriate number, which could
reflect diversity of prey species.
(c) CVs were not satisfactory in more than half of thecases for Bryde’s whales. This means that the
sample size might be slightly smaller than theappropriate number.
9.2.2 Panel conclusions and recommendations
GENERAL COMMENTThe Panel notes that many of the parameter estimates theresearch is designed to obtain, such as the prey consumptionof whales, are subject to various sources of uncertainty not
all of which are related to sample size. A pre-specified levelof precision will not necessarily be achievable with any
sample size. Furthermore, sample size is necessarily
constrained by considerations of the effect of catches on thestocks. Sampling design can be as or more important thansample size.
For each objective of the research, it is necessary to
specify the quantities of interest that need to be determinedto achieve those objectives. For each quantity of interest, all
the sources of uncertainty in its estimation should be
identified and quantified, and in particular it should bedetermined which of these are functions of sample size.
This analysis should be conducted for each objective.
The results of such an analysis can be used to determine:
(i)
(ii)
(iii)
(iv)

how much the research has contributed, inquantitative terms, to achieving the objectives;
what further quantitative progress towards theobjectives can be expected from completing the
programme;
the extent to which increasing/decreasing
sample size would enhance/reduce the rate ofprogress towards achieving the objectives; andthe extent to which the sample design is themost appropriate for achieving the objective
and in particular for maximising theinformation gained from the chosen sample
size.

The analysis is also required for a quantitative comparisonof the performance of lethal or non-lethal methods.
Although this issue was briefly addressed by the
Proponents in SC/J09/JR1, this was not undertaken
sufficiently; a much more thorough approach is warranted
and should be carried out as soon as possible. Until this iscompleted, the Panel is not able to provide appropriatescientific advice on the appropriateness of the sample sizes.
The Panel recognises that a full evaluation of sample
sizes for an integrated study is a major undertaking andprovides the following guidance to the Proponents to assist
in this process. In each case, the Panel recommends that thedevelopment of refined, more quantified sub-objectives foreach component of the programme should be undertaken asa priority; this lack of such sub-objectives is a general
weakness of the present JARPA II programme and limits
the Panel’s ability to review it more thoroughly.
FEEDING ECOLOGY

Determining the appropriate sample size is contingent on
properly estimating the uncertainty surrounding the key
parameters that are ultimately to be used in the modelling
process (e.g. see Item 4). Annex F outlines an appropriate

427

approach to do this. Some but not all of the sources of
uncertainty are related to sample size.
In addition, the spatial and temporal variation in dietsand more generally whale food habits must not be lost,when estimating the impacts of whales on prey stocks. In
this regard, the presentations of JARPN II results that take
into account coastal and offshore regions is valuable.
However, in regions where many whales are foraging on acommon prey, there is probably less need for extensive
sample sizes than where diets are diverse. To meet its
objectives, the programme should try to maximise theinformation obtained over the entire JARPN area. Thus,
considering ways to enhance, improve and ultimately arriveat the optimal sampling design will require considering suchcovariates as stock, season, sex, size, oceanographic
conditions, prey field, and relative location. Additionally,issues of pseudoreplication of whale sampling should beconsidered in this context (Okamura et al., 2008a).
STOCK STRUCTURE

Determining the appropriate sample size and strategy will
depend on the results of the power analyses discussed under
Item 6 and summarised in Annex G. This will also informon the geographical and temporal distribution of samples
required. It is already clear (see discussions under Items 6
and 9) that it is important that samples be obtained from the
Okhotsk Sea with respect to determining the proportions of‘O’ and ‘J’ stock animals, given the implications for theabundance estimates of the assumptions regarding ‘O’stock.
POLLUTANT STUDIESGiven the results obtained thus far, it would be valuable toundertake power analyses to determine the relationshipbetween sample size and the ability to detect changes atvarious levels should they occur. Further evaluation ofcovariates such as age and sex is important to determine
which animals should be chosen for more extensive
sampling. In terms of sampling strategy, the value of
examining the same individuals for each of thecontaminants is emphasised (see Item 5).
9.3 Effects on stocks in light of new knowledge on statusof stocks
In discussing this item, the Panel noted that there is no
specific guidance from the IWC Scientific Committee as tothe appropriate way in which to provide advice of theeffects of scientific permit catches on stocks, particularly
for ongoing programmes that do not have an official
endpoint. Although it notes the past difficulties in trying todevelop an agreed method, the Panel notes that advice on
this matter would be valuable for both future expert panel
reviews under the Annex P protocol and for the Proponents
themselves. As a minimum, the Panel recommends that forcomparison, results should be provided for model runs inwhich research catches are equal to zero. This is particularly
relevant to cases where there is other anthropogenic
mortality (e.g. bycatches), as is the case for western North
Pacific minke whales. An expression of objectives undervarious circumstances would go some way toward
addressing this (e.g. with respect to rates of increase of
populations believed to be below some given percentage of
unexploited population size).

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9.3.1 Proponents’ analyses
SC/09/JR36 examined the effect on whale stocks of futureplanned JARPN II catches. The general approach used wasapplication of the HITTER procedure: given a time series ofpast catches, trajectories were computed to pass through a
recent estimate of abundance for values of MSYR1+ of 1%,
2%, 3%, 4% and 5%, and then projected for 20 years into
the future under the continuation of JARPN II catches at thecurrent sample size. These computations were repeated forthe lower 90% confidence limit for the abundance estimate.
With one exception (the ‘J’ stock of minke whales for
MSYR1+ = 1% for the lower 90% limit on the recent
abundance estimate), the trajectories showed an increaseover the next 20 years for the catches proposed.
COMMON MINKE WHALESThe calculations presented assumed two-stock scenarios, A
and B, involving ‘J’, ‘O’ and ‘W’ stocks, without further
structure, motivated primarily on the absence of indicationsof such further structure in hypothesis tests making use ofgenetic data. However there remain complications as to how
abundance estimates and past catches are best to be split
between these two-stocks. Some smaller parts of the
Okhotsk Sea were treated as containing only ‘J’ stock
animals, but historic commercial catches were considered toconsist of ‘O’ stock whales only, unless taken in the Sea ofJapan or Yellow Sea and so assigned to the ‘J’ stock. A g(0)estimate of 0.732 (Okamura et al., 2008b) was applied
throughout.
BRYDE’S WHALESThe HITTER calculations assumed either a single stock in
the western North Pacific, or one stock in each of subareas 1
and 2, in accordance with the hypotheses developed by the
Scientific Committee for Implementation Simulation Trials.
In addition to the HITTER runs, runs were performed at the
IWC Secretariat of each of the eight base-case IST operating
models for western North Pacific Bryde’s whales. The
HITTER runs and the ISTs used the catch series andabundance estimates that have been accepted by the IWC
Scientific Committee.
SEI WHALES

Based on the results of genetic analyses reported in
SC/J09/J32, it was assumed that the western North Pacific(west of 180°) contained a single stock of sei whales. In fact
no differences were found throughout the whole North
Pacific. The Proponents therefore considered restricting theanalyses to the western North Pacific alone to be suitably
precautionary. Abundance estimates from dedicated surveys
in the JARPN II area were extrapolated to the western North
Pacific using JSV (Japanese scouting vessel) data collectedduring 1972-1988. The extrapolation was conducted
separately for the early (May-June) and late (July-August)periods. The extrapolation factor (the ratio of theextrapolated estimate to the JARPN II survey estimate) was
2.83 and 3.04 for the early and late periods respectively,
resulting in extrapolated estimates of 21,612 and 16,341.
An extrapolation of the JARPN II estimates to the entire
North Pacific using a GAM analyses with various
geographic covariates was also conducted. This yielded asimilar abundance estimate for the entire North Pacific asthat obtained when extrapolating to the entire North Pacificusing JSV data (~60,000 animals). These estimates were notused for assessing the effects of catches on the stock. No
GAM estimate was computed for the area west of 180°.
SPERM WHALESThe Proponents noted that the take level at 10 animals wasless than 0.1% of the abundance estimate and hence it canbe assumed that the planned catches will have no negative
effect on the stock.
9.3.2 Panel comments and recommendations
GENERAL

Although the appropriate lower bound to use for MSYR in
RMP trials is currently under review, the current situation
remains that a value of MSYRmat=1% is accorded medium
plausibility by the Scientific Committee. The Panel thus
recommends that calculations of the effect of catches
should also include results for this value of MSYR. The
Panel noted that the choice of MSYR1+ or mat is an ongoing
matter being discussed within the IWC ScientificCommittee.
The Panel further recommends that in circumstanceswhere Implementation Simulation Trials (ISTs) haverecently been developed for a species in a region, theseprovide the best basis for evaluating the effect of catches on
stocks, as: (1) they constituted the Scientific Committee’s
best appraisal of the range of plausible dynamics for thestocks; and (2) they were based on all appropriate
population abundance and related data. Calculations of theeffect of scientific catches need not be carried out for every
IST for a particular stock (or group of stocks), but should atleast include the baseline trials covering the major stock
structure hypotheses, together with any robustness trials
reflecting stock status appreciably worse than for any ofthese baseline trials. This approach was followed for
Bryde’s whales in SC/J09/JR36. Note that this is not thesame as using the RMP to provide catch advice.
COMMON MINKE WHALES

Ideally the IST approach should also be followed forcommon minke whales, for which ISTs have beendeveloped (IWC, 2004b), but there is the difficulty thatthose trials are now somewhat dated. Considerable further
data that could be used in conditioning and in reconsidering
the plausibility of the existing alternative stock structure
hypotheses has become available since those ISTs weredeveloped by the Scientific Committee.
SC/J09/JR36 considers only two of the four primarystock structure hypotheses of the ISTs for minke whales,arguing that the new genetic evidence excludes the othertwo. As discussed under Item 6.2, the Panel considered thatfurther analyses needed to be tabled before such a definitive
conclusion might be drawn. Until that work has beenpresented, the Panel recommends that the effect of catches
is examined for all four hypotheses.
The Panel noted that of the abundance estimate of 37,170
animals for the ‘O’ stock used in the computations of
SC/J09/JR36, most (88%) come from Russian waters of the
Okhotsk Sea based on a survey conducted in 2003, forwhich there are no genetic samples (by comparison theabundance estimate for the JARPN II area is less than
3,000). The assessments of the effect of JARPN II catcheson the ‘O’ stock are thus highly dependent on theassumption that the whales in the Okhotsk Sea (apart fromthe SW part, which was assigned in SC/J09/JR36 to the ‘J’stock) belong to the ‘O’ stock. In quantitative terms, theJ. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

single factor that could make the greatest contribution toenhancing confidence in the assessment of the effect of
JARPN II catches on the ‘O’ stock would be new data to
support this assumption. The Panel thus recommends that a
new full survey of the Okhotsk Sea is undertaken with aconcerted effort being made to obtain biopsy samples ofcommon minke whales for genetic comparison with the
JARPN II samples (recognising the difficulties in obtaining
biopsy samples). Should unexpected genetic differences befound12, a re-examination of the IST stock structure
hypotheses would become necessary.
Given the above, the Panel considered that the approach
adopted for minke whales in SC/J09/JR36 required further
sensitivity tests, including alternative plausible assignments
of incidental catches and of the abundance estimate for the
Okhotsk Sea between the ‘O’ and ‘J’ stocks. Further
computations were kindly carried out for the Panel by the
Proponents to address these two specific issues (the results
are appended to SC/J09/JR36). The two alternative
approaches to dividing the Okhotsk Sea abundance
estimates were (1) to assume 50% of the whales in sub-area
11 and 25% of those in sub-area 12 belonged to the ‘J’stock, and (2) to assume 5% and 0% respectively for these
allocations. The Panel agrees that the results of the
HITTER runs under these assumptions showed no instancesof conservation concern for the ‘O’ stock. However, thePanel notes with concern that for the ‘J’ stock for the latterof these options and use of the lower 90% confidence limit
for abundance, there was a decline in abundance for
MSYR1+ = 2% which became severe for MSYR1+ = 1%. It
notes that the primary source of the anthropogenic removalsfor ‘J’ stock is bycatches, not scientific permit catches. This
reiterates the general comment made earlier of the value inproducing the results of runs for scientific permit catches =0
for comparative purposes.
The Panel concludes that the information available did
not constitute a sufficient basis to provide advice of theeffect of planned JARPN II catches on common minke
whale stocks. It also noted that the approach used in
SC/J09/JR36 required ad hoc assumptions concerning
disaggregation of ‘J’ and ‘O’ stock animals, and the results
were in one sense overly conservative because restriction toa single set of abundance estimates led to confidence
intervals that were wider than if all available abundance
estimates had been taken into account. Use of the ISTs for
projections is the preferred approach, but as elaborated
above the existing ISTs are now dated.
Given this, the Panel thus recommends that the
hypotheses underlying these ISTs and their conditioning
should be reviewed and updated by the ScientificCommittee as a matter of urgency, given the extensive
new information made available from the JARPN II
programme. Such updated ISTs should form the basis for
projections of stock abundance under JARPN II catches
which could reliably inform appraisals of the effect of the
JARPN II catches on stocks. Such projections should be
carried out both including and excluding JARPN II catches
12

Historical biological data had been used to decide that there were no
additional stocks to the ‘O’ and ‘J’ stocks during the IST development e.g.
sex ratio data (Wada, 1989) and migration patterns (Hatanaka andMiyashita, 1997). In addition, the Panel was informed that genetic analysis
showed no genetic difference in the microsatellite allele frequencies
between the ‘O’ stock individuals from the SA11 and SA7-9 (N. Kanda,
ICR, personal communication). Hence the use of the term ‘unexpected’.

429

so that the contributions of the JARPN II and incidental
catches to any negative trends in abundance can be
distinguished.
BRYDE’S WHALES

Apart from the generic issues of the use of MSYR1+ vsMSYRmat, the Panel found no problems with the assessmentof the effects of JARPN II catches on the Bryde’s whale
stock(s) provided in SC/J09/JR36.
The Panel noted that a recent attempt (Kitakado, 2009) toextract trend information from the Bryde’s whale data(which in principle might help to narrow down the range for
MSYR) yielded a trend estimate with very wide confidence
intervals (-3% to +11% p.a.).
SEI WHALESThe Panel noted that an important part of the evaluation
involves the historic catch series. Given the well known
problems with respect to the identification of sei and
Bryde’s whales in the earlier catch period, it believes it
would have been appropriate to take advantage of the work
undertaken during the Bryde’s whale Implementation on
catch series. The Panel recommends that the Secretariat berequested to produce the corresponding catch series for sei
whales based on the work conducted for the Bryde’s whale
series, and that this be used in the assessments of the sei
whale stock.
While welcoming the presented analyses of the genetic
data that suggested little stock structure in North Pacific sei
whales, as discussed under Item 6.2, the Panel believes
further analyses are required before firm conclusions can bereached with respect to stock structure.
However, the Panel’s greatest concern over the analysis
in SC/J09/JR36 is related to the extrapolation of theabundance estimate outside the survey area (this represented
an increase in abundance by a factor of about three with a
low CV to the 180° boundary). The Panel noted that the CVof the extrapolation factor was based on jacknife samplingof the JSV data using year as the sampling unit. Thisapproach can underestimate the uncertainty in theextrapolation in at least three respects:(1) it does not take into account the inter-annual process
variance in the abundance ratio between the two areas,
which may be important because the JARPN II timeseries is still quite short;(2) it assumes no serial correlation in sei whale distribution
from year to year (a phenomenon that the Panelconsidered a priori to be likely); and(3) it ignores the possibility that there may have been asubstantive regime shift between the 1970s-1980s(when the JSV data were collected) and the current
decade when the JARPN II surveys were conducted,
which could render the relative abundances as found by
the JSV inapplicable to the present.
In addition, apart from the possibility of ecologicalchanges, the distribution of sei whales in the 1970s and1980s may have been distorted by the then recent heavy
catching.
Inspection of the GAM plots in SC/J09/JR36 suggested
that a three-fold increase in abundance when moving fromthe JARPN II area (north of 35°N and W of 170°E) to thefull western north Pacific (north of 30°N and W of 180°) is
greater than one would expect (indeed it would have been

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REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

instructive to compute an abundance figure for the western
North Pacific using the GAM approach for comparison).
In the absence of recent survey data for the whole area,
the Panel recommends that the assessment of the effect on
stocks be repeated without the extrapolation, based on the
JARPN II boundary at 170°E, using an assumed range for
MSYR(mature) of 1-4%, recognising that this might beconsidered conservative. The catch series could berecomputed for this boundary, although this is notconsidered essential. The Panel is thus unable to provide acomplete scientific review of the effects of catches upon
western North Pacific sei whales until this additional work
is undertaken.
The Panel suggests that the sei whale sightings data be
examined for evidence of trend, as has been done for
Bryde’s whales (see above), while recognising that resulting
confidence intervals might be too wide to draw muchinference.
SPERM WHALESThe Panel concurs that the effect on the stock of the small
JARPN II takes is negligible. However, given the comments
made elsewhere in the report, the scientific value of thesesmall and unrepresentative takes of sperm whales isseverely questioned.
9.4 Time of further review
Given the comments and additional analyses that the Panelhas recommended with respect to calculations of sample
size (Item 9.2) and the effect of catches on some of thestocks (Item 9.3), the Panel notes that the present review
cannot be fully completed at this time. The IWC should
consider the most appropriate way to ensure that this review
is completed.
With respect to a further review, the Panel believes thatthe timing should be related to the establishment of interim
objectives for the programme; following ScientificCommittee discussions of the review process, this further
review would probably occur within about six years.
10. SUMMARY AND CONCLUSIONSThe Panel stresses that its primary task at this meeting wasto provide an objective scientific review of results of the
JARPN II programme thus far and plans for future work, inthe light of the stated objectives, including consideration oflethal and non-lethal methods; its task was not to provide
either general condemnation or approval of lethal sampling
research under scientific permit in general - consideration ofthis would require discussion of some issues well beyond
the purview of a scientific panel.
The Panel emphasises that this summary andconclusions section should not be seen as a replacement forthe more comprehensive discussions in the body of thereport.
The Panel thanks the Proponents of JARPN II for theextensive documentation of both published and unpublishedwork that was presented to it well in advance of themeeting, as well as the thorough presentations given during
the Workshop summarising the work undertaken. It also
appreciated the courtesy and openness with which questions
and queries were answered.
The Panel recognises that an enormous amount ofscientific work has been undertaken in the field, laboratory
and in analysis during the first six years of the programme.

It also notes that this is the first part of an ongoing research
programme and that assessing progress against what are, insome cases at least, rather general long-term objectives ischallenging. The Panel therefore recommends that for any
long-term programme such as this, in addition to long-term
objectives, proponents should determine specific, shorterterm objectives that are quantified to the extent possible.
Lack of such objectives hinders any thorough review and isa weakness of the programme. This is also relevant tosample size considerations as indicated below.
10.1 Review of work undertaken to date
The first stage of the review process was to examine resultsof the first six years of the JARPN II programme.
The first broad objective concerns feeding ecology andecosystem modelling, with the ultimate goal of being able toprovide multispecies management advice. The Panel
recognises that this is an extremely ambitious task from adata collection and an analysis/modelling perspective. As
has been stated within and outside the IWC, obtaining
results sufficiently reliable to inform management advice
should not be expected within at least the next few years
and could require considerably more time. The review isthus one of ongoing work and, while progress has been
made by the Proponents, considerably more work is
required, particularly with respect to estimates ofparameters for non-cetacean components of the ecosystem
and analytical and modelling techniques.
With respect to prey consumption and prey preferencesof baleen whales, the Panel recognises the high quality of
the field and laboratory work undertaken. The data collectedhave the potential to be of great value in informing
ecosystem modelling in both a generic and quantitative
manner. However, when reviewing the analyses presented,
the Panel was concerned that insufficient work has been
undertaken to address the full level of uncertainty (this istrue for a number of the objectives, not merely those
relevant to feeding ecology); recommendations to remedy
that are included in the report (e.g. see Item 4.3.2). In
conclusion, therefore, although progress has been made, thePanel does not believe that the estimates of cetacean
consumption rates presented to the Workshop can beconsidered reliable until further analyses have been
undertaken.
With respect to the ecosystem modelling work, varying
degrees of progress have been made using three modelling
approaches. The Panel welcomes this work and notes that,as the authors stated, in all cases the modelling was in the
exploratory stage. The Panel emphasises that this is
generally true of all ecosystem modelling work currently
underway throughout the world, and agrees that the models
as developed thus far are not yet at the stage where they
could be used to draw even general conclusions; theycertainly cannot be used to reliably inform management
advice. Nevertheless, they comprised a substantial andlaudable effort, and an encouraging start to the necessary
process of synthesising the data collected during the
programme. However, the Panel agrees that if there is to bea reasonable chance of meeting the programmes objectives
in a reasonable time-frame, it is essential that considerably
more emphasis be placed on the modelling work from now
on, and it made a number of recommendations in that regard(see Item 4.4.2).

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

The second broad objective relates to monitoring ofenvironmental pollutants in cetaceans and the marine
ecosystem. The Panel concludes that the JARPN II
pollutant studies represent a valuable contribution toknowledge in this area and acknowledges the considerable
amount of work presented. The ongoing programme has
been addressing its objectives, and further work has been
recommended (see Item 5.3). In general, where possible,
papers should include a risk assessment statementsummarising the potential risk to cetaceans based on current
toxicology data in ‘model’ species and other wildlife interms of the health of the animals and dynamics of thestocks.
The third broad objective relates to stock structure issues.
The Panel acknowledges the large amount of new genetic
data collected and the substantial number of analyses
undertaken under JARPN II, which provide a uniquely large
data set for testing hypotheses regarding stock structure in
the target species. Analyses conducted with the genetic data
under JARPN II were in general sound and of a naturecommon to other genetic analyses within and outside the
IWC Scientific Committee framework. The inclusion ofmorphological and morphometric studies as well as genetic
information helps to provide a more well-rounded picture ofstock structure. Although genetic data can provide valuable
insights regarding stock structure, they have some
limitations for weakly differentiated populations, as is thecase for many of the species targeted by JARPN II.
Particularly in the context of weakly-differentiatedpopulations, the Panel acknowledges the generaldifficulties in examining questions of stock structure.
Recognising these general difficulties, the Panel
identified a number of limitations to the analyses presented
and made some detailed suggestions for addressing these(see Item 6.2). In light of this, the Panel agrees that it is notpossible at this time to conclude, as did the Proponents, thatthe number of hypotheses proposed during the
Implementations for western North Pacific common minke
whales and Bryde’s whales has been reduced. However, thePanel noted that such conclusions might be possible after
considering the outcome of the additional analyses
recommended; but that remains to be seen. The Panelagrees that the genetic and other analyses do (and will in
the future) assist in the formulation/narrowing of
hypotheses for use in RMP Implementation Simulation
Trials.
The Panel welcomes the information on the simultaneouscollection of in situ sea surface and water column
characteristics obtained while conducting the whale andprey surveys. It recognises the practical challenges ofcoordinating these sampling methods on the same ship at
the same time. The Panel made a number ofrecommendations for future work including, in the longer
term, that the oceanographic data collected on the cruises(bottom depth, water column temperature, salinity, anddensity) and satellite derived data (such as SST,
chlorophyll, and sea surface height) be integrated into future
analyses.
The Panel also welcomes the collection of sightings data
for non-target species and the analyses of their distribution,
along with photo-identification studies. To investigate
relationships with oceanographic factors and improveabundance estimates for a variety of species, the Proponents

431

should consider conducting future surveys that cover an
area larger than the present JARPN II research area. The
Panel also made recommendations with respect to trend
analyses and the photo-identification data.
A number of other published research papers were
presented that were in addition to the primary work of
JARPN II on reproductive biology, physiology, andcetacean phylogeny.
10.2 Relationship of the programme to the IWC andCommission resolutions
After concluding its review of the ongoing work, the Panelthen considered the relationship between the JARPN II
research and the IWC. With respect to ecosystem andenvironmental change research, the Panel agrees that manyof the objectives of JARPN II are relevant to Resolutions of
the Commission and that, as requested in several
resolutions, scientific results have been submitted to the
Scientific Committee on a number of relevant issues
including those of feeding ecology, pollutant studies,abundance and stock structure.
As is well known, the issue of lethal versus non-lethal
research is one that remains controversial within and outside
the IWC. A major contributory factor to this is that the issue
is not only a scientific question. However, the Panel’s
expertise is of a scientific nature and its comments are
confined to scientific matters. A full comparison of various
lethal and non-lethal techniques requires an analysis of theinformation content of the estimates obtained using
different approaches, in the context of stated quantitative
objectives or sub-objectives; often the data to allow such a
comparison are not available (see recommendations under
Item 8.2.2) and/or objectives are insufficiently
stated/quantified (see Item 9.2.1). In addition, for a complex
multi-disciplinary research programme such as JARPN II,
an evaluation of appropriate techniques (lethal or nonlethal) must include an integrative analysis to ensure
maximised efficiency from both a scientific and logisticalperspective. The Panel was not in a position to evaluate thisin detail in the absence of knowledge of the logisticalresources required and the outcome of analyses detailed
under Item 9.2 to assist with an evaluation of samplingstrategies and sizes.
Given these important difficulties and information gaps
(not all of which are specific to this particular programmeonly), the Panel was not in a position to complete this itemon its Agenda. It nevertheless made a number ofrecommendations in this regard. The Panel recommends
that a full evaluation of the relative merits of lethal and nonlethal techniques be undertaken as soon as possible after the
relevant work recommended elsewhere in this report has
been completed. Such a full evaluation inter alia requires
information on the following.
(1) Specified and quantified objectives and sub-objectives(see recommendation under Item 9.2.1).
(2) Analysis of the precision of the estimates obtained forthe relevant parameters by each of the lethal and nonlethal techniques (see recommendations under Item 4
and at the end of this section).
(3) Evaluation of practicalities of field (and, if relevant,laboratory) techniques in the context of the integrated
objectives, sub-objectives and analyses proposed.

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REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

The ability to fully evaluate and compare non-lethal
methods in a quantitative manner is severely limited by alack of appropriate data. The Panel therefore strongly
recommends that Japan considers the addition of an
objective to quantitatively compare lethal and non-lethal
research techniques if it decides to continue a lethal
sampling programme. Whilst recognising the sensitivities
surrounding this issue, the Panel respectfully requests that if
lethal sampling programmes occur, the IWC ScientificCommittee, together with other appropriate scientificbodies, might wish to consider collaborating in the design ofa well specified study to fully evaluate lethal and non-lethal
techniques.
10.3 Sample size
An evaluation of sample sizes depends on each of theobjectives being better specified, with an identification of
those quantities that need to be estimated to achieve theobjectives. For each such quantity, the sources of
uncertainty to which the estimate is subject should be
identified, including both those which are sample-related
and those which are not sample-related. The precision of theestimate and its relation to sample size and sampling design
should be determined. Such an analysis is a pre-requisite foran evaluation of the appropriateness of the sample size andsampling design.
Although this issue was briefly addressed by the
Proponents in SC/J09/JR1, this was not undertaken
sufficiently; a much more thorough approach is warranted
and should be carried out as soon as possible. Until this iscompleted, the Panel is not able to provide scientific advice
on the appropriateness of the sample sizes.
The Panel recognises that a thorough review is a major
undertaking and it provided guidance to the Proponents toassist in this process. The Panel recommends that thedevelopment of refined, more quantified sub-objectives
should be undertaken as a priority; the lack of suchobjectives is a weakness of the present JARPA II
programme and limits our ability to review its future plans
adequately.
10.4 Effects on the status of the stocks
In discussing this item, the Panel noted that there is no
specific guidance from the IWC Scientific Committee as tothe appropriate way to provide advice on effects ofscientific permit catches on stocks. Advice from the
Scientific Committee on this matter would be valuable forboth future expert panel reviews under the new Protocol forreviewing special permits (IWC, 2009b) and for the
Proponents themselves. As a minimum, the Panelrecommends that for comparison, results should also be
provided for projections for which research catches areequal to zero as well as for catches equal to the proposed
catches. This is particularly relevant to cases where otheranthropogenic mortality occurs (e.g. bycatches), as is thecase for western North Pacific minke whales. An expressionof performance against possible conservation objectives by
Proponents and/or the IWC under various situations would
go some way to addressing this (e.g. with respect to rates ofincrease of populations believed to be below some given
percentage of unexploited population size).

Although the appropriate lower bound to use for MSYR
in RMP trials is currently under review, the current situationis that a value of MSYRmat=1% is accorded medium
plausibility by the Scientific Committee. The Panel thus
recommends that calculations of the effect of catches
should also include results for this value of MSYR. The
Panel noted that the choice of MSYR1+ or mat is an ongoing
matter being discussed within the IWC ScientificCommittee.
The Panel further recommends that in circumstanceswhere Implementation Simulation Trials (ISTs) haverecently been developed for a species in a region, theseprovide the best basis for evaluating the effect of catches on
stocks; they constitute the Scientific Committee’s best
appraisal of the range of plausible dynamics for the stocks,
having been based on all appropriate population abundance
and related data. Note that this is not the same as using the
RMP to provide catch advice.
10.4.1 Common minke whales
The Panel concludes that the information available did notconstitute a sufficient basis to provide advice on the effectof planned JARPN II catches on common minke whale
stocks. Use of the ISTs for projections is the preferredapproach, but as elaborated above, the existing ISTs are now
dated.
Given this, the Panel thus recommends that the
hypotheses underlying these ISTs and their conditioning
should be reviewed and updated by the ScientificCommittee as a matter of urgency, given the extensive
new information made available from the JARPN II
programme. Such updated ISTs should form the basis for
projections of stock abundance under JARPN II catches,
which might reliably inform appraisals of the effect of the
JARPN II catches on stocks. Such projections should be
carried out both including and excluding JARPN II catches
so that the contributions of the JARPN II and incidental
catches to any negative trends in abundance can be
distinguished.
In addition, although not strictly part of a review of
JARPN II, the Panel emphasises its concern that the resultsof some of the HITTER runs involving the depleted ‘J’stock presented by the Proponents revealed a decline inabundance for MSYR1+=2% which became severe for
MSYR1+=1%. It notes that the primary source of theanthropogenic removals for ‘J’ stock is bycatches, notscientific permit catches. This provides further support for
the need to complete the in-depth assessment of ‘J’ stock assoon as possible, along with a full Implementation Review
for western North Pacific minke whales.
10.4.2 Bryde’s whales
The Panel accepts the assessment of the effects of JARPN
II catches on Bryde’s whales provided by the Proponents
and agreed that this level of take does not pose a problem tothe stocks.
10.4.3 Sei whales
The Panel had a number of concerns over the analysis on
the effect on sei whale stocks provided by the Proponents particularly extrapolation of the abundance estimate outside
the survey area (this represented an increase in abundance
by a factor of about three with a low CV) to the 180°boundary. In the absence of recent survey data for the wholearea, the Panel recommends that the assessment of the

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

effect on stocks be repeated without the extrapolation, basedon the JARPN II boundary at 170°E, and using an assumed
range for MSYR(mature) of 1-4%, while recognising that
this might be considered conservative. The catch series(adjusted in light of the Bryde’s whale Implementation
approach to the historical problems regarding the species
identification of sei and Bryde’s whales) should berecomputed for this boundary, although this is notconsidered essential. The Panel is unable to provide acomplete scientific review of the effects of catches upon
western North Pacific sei whales until this additional work
is undertaken.
10.4.4 Sperm whales
The Panel concurs that the effect on the stock of the small
JARPN II takes is negligible. However, given the comments
made elsewhere in the report, the scientific value of thesesmall and unrepresentative takes of sperm whales isseverely questioned.
10.5 Further review
Given the comments and additional analyses that the Panelhas recommended with respect to calculations of sample
size and the effect of catches on some of the stocks, thePanel agrees that the present review cannot be fullycompleted at this time. The Scientific Committee should
consider the most appropriate way to ensure that this review
is completed.
Until this review has been completed, it would bepremature to advise when a further review should be
conducted.
11. ADOPTION OF REPORTThe report was adopted by e-mail.
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Annex A
List of Panel Members
Arne Bjørge
Institute of Marine Research
Gaustadalleen 210349 Oslo
Norwayarne.bjoerge@imr.no
Doug Butterworth
Dept of Mathematics and Applied
Mathematics
University of Cape Town
Rondebosch 7701
South Africa
Doug.Butterworth@uct.ac.za
Justin Cooke
Center for Ecosystem Management
Studies
Alexanderstr. 10 79261Gutach
Germany
jgc@cems.de
Greg Donovan
IWC Secretariat
The Red House
135 Station Road
Impington, Cambridge
CB24 9NP, UK
greg.donovan@iwcoffice.org
Jaume Forcada
British Antarctic Survey
High Cross
Madingley Road
Cambridge CB3 0ET, UK
jfor@bas.ac.uk

Ailsa Hall
Scottish Oceans InstituteEast Sands
University of St. Andrews
St Andrews
Fife KY16 8LB
UKajh7@st-andrews.ac.uk
George Hunt
School of Aquatic and Fishery
Sciences
Box 355020
University of Washington
SeattleWA 98195
USA
geohunt2@u.washington.edu
Jason Link
Northeast Fisheries Science Center
166 Water St
Woods Hole
MA 02543
USA
jlink@mercury.wh.whoi.edu
Debra Palka
Northeast Fisheries Science Center
166 Water St
Woods Hole
MA 02543
USA
debra.palka@noaa.gov

Per Palsbøll
Department of Genetics,
Microbiology and Toxicology
Stockholm University
106 91 Stockholm
SwedenPer.Palsboll@gmt.su.se
Steve Reilly
Southwest Fisheries Science Center
PO Box 271
La Jolla, CA 92038
USA
steve.reilly@noaa.gov
Mette Skern-Mauritzen
Institute of Marine Research
PB 1870 Nordnes
5817 Bergen
Norwaymette.mauritzen@imr.no
Ichii Taro
National Research Institute of Far
Seas Fisheries
2-12-4 Fukuura, Kanazawa-ward
Yokohama-city, 236-8648
Japanichii@affrc.go.jp
Robin Waples
Northwest Fisheries Science Center
2725 Montlake Boulevard East
Seattle, WA 98112
USA
robin.waples@noaa.gov

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437

Annex B
Agenda1. Introductory items
1.1
Welcome and opening remarks
1.2Election of Chair
1.3
Appointment of rapporteurs
1.4
Meeting procedure and time schedule
2. Adoption of the agenda
3. Review of available data, documents and reports
3.1
Workshop documents
3.2
For information papers
3.3
Other available documents and data
4. Review of JARPN II results: Feeding ecology andecosystem studies
4.1
Formal statement of objectives as given by theGovernment of Japan (based upon SC/J09/JR1)
4.2
JARPN II Coastal component
4.2.1 Proponents’ summary
4.2.2 Expert Panel review of results presented
4.3
JARPN II Offshore component
4.3.1 Proponents’ summary
4.3.2 Panel conclusions and recommendations
4.4
Ecosystem modelling
4.4.1 Proponents’ summary
4.4.2 Panel conclusions and recommendations
5. Review of JARPN II results: Monitoring Environmental
Pollutants in Cetaceans and the Marine Ecosystem
5.1
Statement of objectives as given by theGovernment of Japan
5.2
Proponents’ summary
5.3
Panel’s conclusions and recommendations
6. Review of JARPN II results: Stock structure
6.1
Statement of objectives as given by theGovernment of Japan
6.2
Proponents’ summary
6.3
Panel’s conclusions and recommendations
6.3.1 Simple issues
6.3.2 More extensive matters – some of whichmight ideally be addressed in time for the2009 Annual Meeting
6.3.3 Longer term
7. Review of JARPN II results: Review of other contributions to important research needs
7.1
Oceanography
7.1.1 Statement of objectives and Proponents’summary
7.1.2 Panel’s conclusions and recommendations
7.2
Distribution of large whales

7.2.1 Statement of objectives and Proponents’summary
7.2.2 Panel conclusions and recommendations
7.3
Other research
7.3.1 Proponents’ summary
7.3.2 Panel conclusions and recommendations
7.4
Abundance
7.4 1 Summary of results to date
7.4.2 Analysis and comment
7.4.3 Recommendations
8. Review of JARPN II results: the relationship of theresearch to relevant IWC resolutions and discussions
8.1
Research on the ecosystem and environmental
change8.1.1 Summary of Proponents’ view (from
SC/J09/JR1)
8.1.2 Panel conclusions and recommendations
8.2
Utility of the lethal techniques used by JARPN II
compared to non-lethal techniques
8.2.1 Summary of Proponents’ view (from
SC/J09/JR1)
8.2.2 Panel conclusions and recommendations
9. Advice on ongoing special permit research
9.1
Practical and analytical methods, including nonlethal methods, that can improve research relative
to stated objectives
9.2
Appropriate sample sizes to meet the statedobjectives, especially if new methods aresuggested under Item 9.1
9.2.1 Proponents’ view (from SC/J09/JR1)
9.2.2 Panel conclusions and recommendations
9.3
Effects on stocks in light of new knowledge on
status of stocks
9.3.1 Proponents’ analyses
9.3.2 Panel comments and recommendations
9.4
Time of further review
10. Summary and conclusions
10.1 Review of work undertaken to date
10.2 Relationship of the programme to the IWC andCommission resolutions
10.3 Sample size
10.4 Effects on the status of the stocks
10.4.1 Common minke whales
10.4.2 Bryde’s whales
10.4.3 Sei whales
10.4.4 Sperm whales
10.5 Further review
11. Adoption of report

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Annex C
List of Documents
SC/J09/JR
1. Pastene, L.A., Hatanaka, H., Fujise, Y., Kanda, N.,
Murase, H., Tamura, T., Miyashita, T. and Kato, H. The
Japanese Whale Research Program under Special
Permit in the western North Pacific Phase-II (JARPN
II): origin, objectives and research progress made in the
period 2002-2007, including scientific considerations
for the next research period. 73pp.
2. Kiwada, H., Kumagai, S. and Matsuoka, K.
Methodology and procedure of the dedicated sighting
surveys in JARPN II – Offshore and coastal componentof Sanriku and Kushiro. 16pp.
3. Kishiro, T., Yoshida, H., Goto, M., Bando, T. and Kato,
H. Methodology and survey procedure under the
JARPN II – coastal component of Sanriku and Kushiro,with special emphasis on whale sampling procedures.
27pp.
4. Tamura, T., Matsuoka, K. and Fujise, Y. Methodology
and survey procedure under the JARPN II - offshore
component- with special emphasis on whale sampling
procedures. 16pp.
5. Yonezaki, S., Nagashima, H., Murase, H., Yoshida, H.,
Bando, T., Goto, M., Kawahara, S. and Kato, H.
Methodology and procedures of surveys of prey ofcommon minke whales JARPN II - Coastal componentof Sanriku. 6pp.
6. Watanabe, H., Yonezaki, S., Kiwada, H., Kumagai, S.,
Kishiro, T., Yoshida, H. and Kawahara, S.
Methodology and procedures of common minke
whale’s prey surveys in JARPN II – Coastal componentof Kushiro. 12pp.
7. Murase, H., Watanabe, H., Yonezaki, S., Tamura, T.,
Matsuoka, K., Fujise, Y. and Kawahara, S.
Methodology and procedures of cetacean prey surveys
in JARPN II – Offshore Component. 11pp.
8. Hakamada, T., Matsuoka, K. and Miyashita, T. The
number of western North Pacific common minke
whales (Balaenoptera acutorostrata) distributed in
JARPN II coastal survey areas. 12pp.
9. Tamura, T., Konishi, K., Goto, M., Bando, T., Kishiro,T., Yoshida, H., Okamoto, R. and Kato, H. Prey
consumption and feeding habits of common minke
whales in coastal areas off Sanriku and Kushiro. 18pp.
10. Murase, H., Kawahara, S., Nagashima, H., Onodera, K.,
Tamura, T., Okamoto, R., Yonezaki, S., Matsukura, R.,
Minami, K., Miyashita, K., Yoshida, H., Goto, M.,
Bando, T., Inagake, D., Okazaki, M., Okamura, H. andKato, H. Estimation of prey preference of common
minke whales (Balaenoptera acutorostrata) in a coastal
component (off Sanriku) of JARPN II in 2005 and2006. 15pp.
11. Watanabe, H., Yonezaki, S., Kiwada, H., Kumagai, S.,
Kishiro, T., Yoshida, H. and Kawahara, S. Distribution
and abundance of prey species and prey preference ofcommon minke whale Balaenoptera acutorostrata inthe coastal component of JARPN II off Kushiro from2002 to 2007. 37pp.

12. Yoshida, H., Kishiro, T., Goto, M., Bando, T., Tamura,T., Konishi, K., Okamoto, R. and Kato, H. Relationshipbetween body size, maturity, and feeding habit ofcommon minke whales off Sanriku in spring season,
from 2003-2007 whale sampling surveys under the
JARPN II coastal component off Sanriku. 20pp.
13. Kishiro, T., Yoshida, H., Tamura, T., Konishi, K.,
Kanda, N., Okamoto, R. and Kato, H. Relationshipbetween body size, maturity, and feeding habit ofcommon minke whales off Kushiro in autumn season,
from 2002-2007 whale sampling surveys under the
JARPN II coastal components off Kushiro. 25pp.
14. Okamura, H., Nagashima, H. and Yonezaki, S.
Preliminary assessment of impacts on the sandlance
population by consumption of minke whales off
Sanriku region. 20pp.
15. Hakamada, T., Matsuoka, K. and Miyashita, T.
Distribution and the number of western North Pacificcommon minke, Bryde’s, sei and sperm whales
distributed in JARPN II Offshore component survey
area. 18pp.
16. Tamura, T., Konishi, K., Isoda, T., Okamoto, R. andBando, T. Prey consumption and feeding habits ofcommon minke, sei and Bryde’s whales in the western
North Pacific. 36pp.
17. Tamura, T., Kubotera, T., Ohizumi, H., Konishi, K. andIsoda, T. Feeding habits of sperm whales and theirimpact on neon flying squid resources in the western
North Pacific. 22pp.
18. 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. 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. 31pp.
19. Konishi, K., Kiwada, H., Matsuoka, K., Hakamada, T.
and Tamura, T. Density prediction modeling andmapping of common minke, sei and Bryde’s whales
distribution in the western North Pacific using JARPN
II (2000-2007) data set. 20pp.
20. Konishi, K., Tamura, T., Goto, M., Bando, T., Kishiro,T., Yoshida, H. and Kato, H. Trend of blubber thickness
in common minke, sei and Bryde’s whales in thewestern North Pacific during JARPN and JARPN II
periods. 4pp.
21. Mori, M., Watanabe, H., Hakamada, T., Tamura, T.,
Konishi, K., Murase, H. and Matsuoka, K.
Development of an ecosystem model of the western
North Pacific. 49pp.
22. Kawahara, S. A minimum realistic model in the JARPN
II offshore survey area. 22pp.
23. Yasunaga, G. and Fujise, Y. Temporal trends andfactors affecting mercury levels in common minke,
Bryde’s and sei whales and their prey species in thewestern North Pacific. 13pp.

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

24. Yasunaga, G. and Fujise, Y. Temporal trends andfactors affecting PCB levels in baleen whales andenvironmental samples from the western North Pacific.
10pp.
25. Yasunaga, G. and Fujise, Y. Accumulation features of
total and methyl mercury and selenium in tissues ofcommon minke, Bryde’s and sperm whales from thewestern North Pacific. 11pp.
26. Kanda, N., Goto, M., Kishiro, T., Yoshida, H., Kato, H.
and Pastene, L.A. Individual identification and mixingof the J and O stocks around Japanese waters examinedby microsatellite analysis. 9pp.
27. Hakamada, T. and Bando, T. Morphometric analysis on
stock structure in the western North Pacific common
minke whale (Balaenoptera acutorostrata). 13pp.
28. Goto, M., Kanda, N., Pastene, L.A., Bando, T. andHatanaka, H. Differences in cookie cutter sharkinduced body scar marks between J and O-stocks ofcommon minke whales in the western North Pacific.
7pp.
29. Goto, M., Kanda, N., Kishiro, T., Yoshida, H., Kato, H.
and Pastene, L.A. Mitochondrial DNA analysis on
stock structure in the western North Pacific common
minke whales. 10pp.
30. Kanda, N., Goto, M., Kishiro, T., Yoshida, H., Kato, H.
and Pastene, L.A. Microsatellite analysis of minke
whales in the western North Pacific. 14pp.

439

31. Kanda, N., Goto, M. and Pastene, L.A. Stock structureof Bryde’s whales in the western North Pacific as
revealed by microsatellite and mitochondrial DNA
analyses. 8pp.
32. Kanda, N., Goto, M., Yoshida, H. And Pastene, L.A.
Stock structure of sei whales in the North Pacific as
revealed by microsatellite and mitochondrial DNA
analysis. 14pp.
33. Kanda, N., Goto, M. and Pastene, L.A. Genetic
characteristics of sperm whales sampled during JARPN
II from 2000 to 2007 as revealed by mitochondrial
DNA and microsatellite analyses. 5pp.
34. Okazaki, M., Inagake, D., Masujima, M., Murase, H.,
Watanabe, H., Yonezaki, S., Nagashima H., Matsuoka,
K., Kiwada, H. and Kawahara, S. Oceanographic
conditions of the western North Pacific based on
oceanographic data collected during the JARPN II.
13pp.
35. Matsuoka, K., Kiwada, H., Fujise, Y. and Miyashita, T.
Distribution of blue (Balaenoptera musculus), fin (B.
physalus), humpback (Megaptera novaeangliae) andNorth Pacific right (Eubalaena japonica) whales in thewestern North Pacific based on JARPN and JARPN II
sighting surveys (1994 to 2007). 12pp.
36. Hakamada, T. Examination of the effects on whale
stocks of future JARPN II catches. 51pp.

Annex D
Extract from Guidelines for Review of Scientific Permits*
2. THE REVIEW PROCESSIntersessional specialist workshop
The initial review of a new proposal, or interim and final
reviews, shall take place at a small specialist workshop witha limited but adequate number of invited experts (who may
or may not be present members of the ScientificCommittee). A limited number of scientists associated withthe proposal should attend the workshop in an advisory role,
primarily to present the proposal and answer points ofclarification. It is important that the composition of thespecialist group is considered balanced and fair. The choiceof experts shall be made by the Chair, Vice-Chair and Headof Science in conjunction with a Standing Steering Group(SSG) established by the Chair at an Annual Meeting, withspecial emphasis on the field and analytical methods
provided in the proposal and estimation of the effect ofcatches on the stocks(s). The SSG shall be selected by the
Chair, Vice-Chair and Head of Science, such that it
represents an appropriate range of experience and expertise*Excerpts from IWC (2009).

within the Scientific Committee. The selection process forthe specialist group shall occur in the manner described
below.
Procedure for periodic and final reviews
For ongoing research without a defined final year, aperiodic review shall take place in accordance with either
the advice provided under Item (5) of the workshop (seebelow) to review new proposals or on the advice of aperiodical review workshop and taking into account theavailability of the proponents. The final review shall takeplace no later than three years after the final take underSpecial Permits. The periodic and final reviews shall bebased on documents provided by the proposers and othermembers of the Scientific Committee six months before theAnnual Meeting at which the Workshop report is to bepresented. Information on the analytical methods likely to
be used in documents presented to the Workshop that might
assist with the selection of appropriate experts shall becirculated nine months before the Annual Meeting.

440

REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN IIThe Chair shall circulate the information on theanalytical methods to the Vice-Chair, Head of Science and
SSG, normally within 1 week of receipt.
(1) The SSG shall examine the information available on thefield and analytical methods and, normally within 2
weeks, suggest names for consideration for theSpecialist Workshop (if these experts are not membersof the Committee they shall include a rationale for theirchoice) and the suggestions will be available to all SSG
members.
(2) The Chair, Vice-Chair and Head of Science willdevelop a proposed final list (with reserves) for
consideration by the SSG within 2 weeks and begin theprocess of establishing the time and venue of theWorkshop taking into account the availability of the
proposed experts and experts associated with theproposal.
(3) The SSG will send final comments within 1 week.
(4) The Chair, Vice-Chair and Head of Science will agree a
final list (with reserves); the proposal (with a note
concerning any restrictions) will be sent to the selected
experts and reserves - the process thus far will havetaken about 6 weeks since the information on analyticalmethods has been received.
(5) The full documents shall be circulated no later than 6months before the Annual Meeting.
(6) Responses to those documents shall be submitted nolater than 1 month before the Workshop.
The Workshop will take place at least 100 days before
the Annual Meeting. In addition to the selected experts it
will include at least one of the Chair, Vice-Chair and Headof Science, one of whom shall chair the workshop.
Availability of data relevant to the periodic or final
review
Applications for the access to data for the purpose of
periodic or final review, should follow the recommended
approach of Procedure B of the IWC Scientific Committee
Data Availability Agreement (IWC, 2004). For data
provided under the DAA, the conditions for data recipientsare outlined in the agreement. Applications made bymembers of the Scientific Committee and other participantsat the Specialist Workshop should be considered promptly
and normally accepted within two weeks of the application.
Terms of reference of the Specialist Workshop for
periodic and final reviews
The primary objective of the specialist workshop will be toreview the scientific aspects of the research under Special
Permits in the light of the stated objectives following theguidelines in the pro forma provided by the Secretariat. In
particular, the Specialist Workshop shall evaluate:(1) how well the initial, or revised, objectives of theresearch have been met;(2) other contributions to important research needs;(3) the relationship of the research to relevant IWC
resolutions and discussions, including those dealing
with the respective marine ecosystem, environmental
changes and their impact on cetaceans and Committee
reviews of special permit research;(4) the utility of the lethal techniques used by the Special
Permit Programme compared to non-lethal techniques;and(5) in case of periodic review, provide advice on:
(i)
(ii)
(iii)
(iv)

practical and analytical methods, including non
lethal methods, that can improve research
relative to stated objectives;appropriate sample sizes to meet the statedobjectives, especially if new methods aresuggested under item (i);effects on stocks in light of new knowledge on
status of stocks; andwhen, in the case of ongoing programmes, afurther review should occur.

Reports of Workshops (applies to new proposals,
periodic reviews and final reviews)
The Chair is responsible for the level and nature ofparticipation of the scientists involved in the proposal,
which should be limited to: (1) providing information to theinvited experts in addition to that contained in the proposal
or research results; and (2) answering questions posed by
the invited experts. The specialist group should attempt toreach consensus on the individual issues referred to above,
but where this is not possible, the rationale behind thedisagreement should be clearly stated in the Workshop
report. The final report of the Workshop shall be completed
at least 80 days prior to the Annual Meeting and will bemade available to the proponents.
Circulation to the Scientific Committee
The original special permit proposal, or the original result
documents from ongoing or completed special permit
research, the report of the specialist workshop, and any
revised permit proposal (following the agreed protocol), or
any revised results, from the Contracting Government shallbe submitted to Scientific Committee members no later than
40 days before the Annual Meeting. The revised proposal,
or revised results, will also be submitted to the members of
the specialist group and they will be invited to submit joint
or individual comments on that revision to the Annual
Meeting.
REFERENCES
International Whaling Commission. 2004. Report of the ScientificCommittee. Annex T. Report of the data availability working group. J.
Cetacean Res. Manage. (Suppl.) 6:406-08.
International Whaling Commission. 2009. Report of the ScientificCommittee. Annex P. Process for the review of special permit proposals
and research results from existing and completed permits. J. Cetacean
Res. Manage. (Suppl.) 11:398-401.

441

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

Annex E
Summary of Available Data for JARPN II
Taken from SC/J09/JR1 (*= database complete)

I. Sighting data – coastal and offshore components
Total sample
size

DB
*
*
*
*
*

Angle and distance experiment data (no. of experiments)
Photo ID humpback whales (no. of schools photographed)
Photo ID right whales (no. of schools photographed)
Photo ID blue whales (no. of schools photographed)
Sighting data (no. of schools)
Effort data (n.miles)
Weather data (no. observations)

3,807
24
22
65
9,426
439,047
66,260II-1. Biological data – common minke whale – offshore component
Number of whales
DB Data and sample

Male Female Total

*
*
*
*

661
661
661
661
656
661
661
661
661
661
158
661
661
661
661
661
661
661
661
661
347
77
35
35
35
655
654

*
*
*
*
*
*
*
*
*
*
*
*

*
*
*
*
*
*
*

Sampling date
Sampling location
Body length
Body proportion
Skull (length and breadth)
Body scar record
Parasites, external occurrence record
Parasites, internal occurrence record
Sex
Body weight
Organ weights
Blubber thickness
GirthMaturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weights
Main prey species in stomach contents
Freshness of stomach contents
Foetus, number
Foetus, sex
Foetus, body length
Foetus, body weight
Ear plug
PCB concentrations (blubber)
Total Hg levels (liver)
Total Hg, methyl Hg and Se levels (liver)
Total Hg, methyl Hg and Se levels (kidney)
Total Hg, methyl Hg and Se levels (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (16 loci)

79
79
79
79
78
79
79
79
79
79
23
79
79
79
79
79
79
79
79
79
79
79
5
5
5
79
79

740
740
740
740
734
740
740
740
740
740
181
740
740
740
79
79
661
740
740
740
740
79
35
35
35
740
347
77
40
40
40
734
733

442

REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

II-2. Biological data – sei whale – offshore component
Number of whales
DB Data and sample

Male Female Total

*
*
*
*

238
238
238
238
232
238
238
238
238
238
63
238
238
238
238
238
238
238
238
238
15
30
238
238

*
*
*
*
*
*
*
*
*
*
*
*

*
*
*
*
*
*
*

Sampling date
Sampling location
Body length
Body proportion
Skull (length and breadth)
Body scar record
Parasites, external occurrence record
Parasites, internal occurrence record
Sex
Body weight
Organ weights
Blubber thickness
GirthMaturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weights
Main prey species in stomach contents
Freshness of stomach contents
Foetus, number
Foetus, sex
Foetus, body length
Foetus, body weight
Ear plug
PCB concentrations (blubber)
Total Hg levels (liver)
Total Hg, methyl Hg and Se levels (liver)
Total Hg, methyl Hg and Se levels (kidney)
Total Hg, methyl Hg and Se levels (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (17 loci)

251
251
251
251
243
251
251
251
251
251
64
251
251
251
251
251
251
251
251
251
251
—
—
—
251
251
251

489
489
489
489
475
489
489
489
489
489
127
489
489
489
251
251
238
489
489
489
489
251
115
115
115
489
15
30
489
489

II-3. Biological data – Bryde’s whale – offshore component
Number of whales
DB Data and sample

Male Female Total

*
*
*
*

166
166
166
166
161
166
166
166
166
166
56
166
166
166
166
166
166
166
166
166
15
20
21
21
21
166
166

*
*
*
*
*
*
*
*
*
*
*
*

*
*
*
*
*
*
*

Sampling date
Sampling location
Body length
Body proportion
Skull (length and breadth)
Body scar record
Parasites, external occurrence record
Parasites, internal occurrence record
Sex
Body weight
Organ weights
Blubber thickness
GirthMaturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weights
Main prey species in stomach contents
Freshness of stomach contents
Foetus, number
Foetus, sex
Foetus, body length
Foetus, body weight
Ear plug
PCB concentrations (blubber)
Total Hg levels (liver)
Total Hg, methyl Hg and Se levels (liver)
Total Hg, methyl Hg and Se levels (kidney)
Total Hg, methyl Hg and Se levels (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (17 loci)

227
227
227
227
222
227
227
227
227
227
71
227
227
227
227
227
227
227
227
227
227
227
22
22
22
227
227

393
393
393
393
383
393
393
393
393
393
127
393
393
393
227
227
166
393
393
393
393
227
127
127
127
393
15
20
43
43
43
393
393

443

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

II-4. Biological data – sperm whale – offshore component
Number of whales
DB Data and sample

Male Female Total

*
*
*
*

13
13
13
13
13
13
13
13
13
13
8
13
13
13
13
13
13
13
13
13
3
3
3
13
13

*
*
*
*
*
*
*
*
*
*
*
*

*
*
*
*
*
*
*

Sampling date
Sampling location
Body length
Body proportion
Skull (length and breadth)
Body scar record
Parasites, external occurrence record
Parasites, internal occurrence record
Sex
Body weight
Organ weights
Blubber thickness
GirthMaturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weights
Main prey species in stomach contents
Freshness of stomach contents
Foetus, number
Foetus, sex
Foetus, body length
Foetus, body weight
Ear plug
PCB concentrations (blubber)
Total Hg levels (liver)
Total Hg, methyl Hg and Se levels (liver)
Total Hg, methyl Hg and Se levels (kidney)
Total Hg, methyl Hg and Se levels (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (15 loci)

32
32
32
32
30
32
32
32
32
32
22
32
32
32
32
32
32
32
32
32
32
32
2
2
2
32
32

45
45
45
45
43
45
45
45
45
45
30
45
45
45
32
32
13
45
45
45
45
32
7
7
7
45
5
5
5
45
45

II-5. Biological data – common minke whale – coastal component off Kushiro
Number of whales
DB Data and sample

Male Female Total

*
*
*
*

182
182
182
182
181
182
182
182
182
11
182
182
182
182
182

72
72
72
72
72
72
72
72
72
3
72
72
72
72
72
72

254
254
254
254
253
254
254
254
254
14
254
254
254
72
72
182
254

182
182
181
46
181
182

72
72
72
—
—
—
71
72
71

254
254
72
4
4
4
252

*
*
*
*
*
*
*
*
*
*
*
*

*
*
*
*
*
*
*

Sampling date
Sampling location
Body length
Body proportion
Skull (length and breadth)
Body scar record
Parasites, external occurrence record
Parasites, internal occurrence record
Sex
Body weight
Organ weights
Blubber thickness
GirthMaturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weights18272254
Main prey species in stomach contents
Freshness of stomach contents
Foetus, number
Foetus, sex
Foetus, body length
Foetus, body weight
Ear plug
PCB concentrations (blubber)
Total Hg levels (liver)
Total Hg, methyl Hg and Se levels (liver)
Total Hg, methyl Hg and Se levels (kidney)
Total Hg, methyl Hg and Se levels (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (16 loci)

46
253
253

444

REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

II-6. Biological data – common minke whale – coastal component off Sanriku
Number of whales
DB Data and sample

Male Female Total

*
*
*
*

91
91
91
91
89
91
91
91
91
4
91
91
91
89-89
91
89
91
91
91
29
91
91

*
*
*
*
*
*
*
*
*
*
*
*

*
*
*
*
*
*
*

Sampling date
Sampling location
Body length
Body proportion
Skull (length and breadth)
Body scar record
Parasites, external occurrence record
Parasites, internal occurrence record
Sex
Body weight
Organ weights
Blubber thickness
GirthMaturity stage
Corpora albicantia and lutea (number)
Lactation condition
Testis weight
Stomach contents (IWS format)
Stomach contents weights
Main prey species in stomach contents
Freshness of stomach contents
Foetus, number
Foetus, sex
Foetus, body length
Foetus, body weight
Ear plug
PCB concentrations (blubber)
Total Hg levels (liver)
Total Hg, methyl Hg and Se levels (liver)
Total Hg, methyl Hg and Se levels (kidney)
Total Hg, methyl Hg and Se levels (muscle)
Mitochondrial DNA control region sequences
Nuclear DNA microsatellite (16 loci)

136
136
136
136
133
136
136
136
136
5
136
136
136
135
136

227
227
227
227
222
227
227
227
227
9
227
227
227
135
136

136
133
136
136
135
—
—
—
136
136
136

227
222
227
227
135
30
30
30
227
29
227
227

III. Pollutant data (environmental and prey species samples) – offshore component
DB Item
*
*
*
*
*
*
*
*
*
*

Number of samples

Organochlorine compounds (air)Organochlorine compounds (sea water)
Total Hg compounds (copepods)
Total Hg compounds (Krill)
Total Hg compounds (Larval of anchovy)
Total Hg compounds (Adult of anchovy)
Total Hg compounds (Pacific saury)
Total Hg compounds (mackerels)
Total Hg compounds (Pacific pomfret)
Organochlorine compounds (walleye pollock)

6
7
5
17
6
20
41
5
3
2

IV-1. Oceanographic data – offshore component
DB Item
*
*
*
*
*
*
*

Marine debris (sighting survey, days)
Temperature and salinity (XCTD survey: 2000-07)
Temperature and salinity (CTD survey: 2000-07)
Midwater trawl (no. of hawls)
MOCNESS (no. of hawls)
IKMT (no. of hawls)
NORPAC (no. of hawls)
Echo sounder (km: 2002-07)

Number of samples
56
38
593
141
24
30
75
12,838

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010445

IV-2. Oceanographic data – coastal component off Sanriku
DB Item
*
*
*
*
*
*
*

Number of samples

Temperature and salinity (XCTD survey: 2003-07)
Temperature and salinity (CTD survey: 2003-07)
Midwater trawl (no. of hawls)
Bongo net (no. of hawls)
IKMT (no. of hawls)
Sampling by fishing (no. of stations)
Echo sounder (km; 2005 and 2006 seasons)

11
149
57
5
17
2
2,775

IV-3. Oceanographic data – coastal component off Kushiro
DB Item*****Number of samples

Temperature and salinity (CTD survey: 2002-07)
Midwater trawl (no. tow)
MOCNESS survey (no. tow)
IKMT survey (no. tow)
NORPAC net survey (no. tow)

109
133
6
-

Annex F
Characterising the Uncertainty in the Estimation of Food
Consumption of Baleen Whales by Prey Type
SC/J09/JR9 describes the approach used for estimating
consumption from diet samples of minke whales collectedin the coastal part of JARPN II. Ranges are given for theestimates of consumption, but these account for only part of
the uncertainty in the estimates. It is important to estimate,at least approximately, the overall level of uncertainty inconsumption estimates.
This Annex aims to list the main sources of uncertainty
in the estimates that should be taken into account whenestimating the overall CV of consumption estimates.
The estimation of total consumption by prey species inthe study area involves the steps listed below. The
uncertainty associated with each step should be quantified.
Due to lack of a sufficiently strong diurnal signal instomach fullness that might enable stomach throughput
times to be determined, consumption estimates rely on
considerations of energy requirements. The stomach content
data are used to quantify the proportional contribution ofeach prey species or species group to the diet.
‘Prey type’ is by species of prey for the main preyspecies, otherwise by groups of species.
Where an item is based on data, CVs should be estimatedby sub-sampling the appropriate units. Where data arelacking and an item has to be guessed, a plausible range
should be given with a rationale. Nominal CVs can be
estimated by assuming that the plausible range represents± 2 SE on an appropriate scale.
Estimates of consumption can only be evaluated wheneach of the listed sources of variance has been taken into
account. When this has been done, estimates ofconsumption from different methods (e.g. lethal vs nonlethal) can be compared.

When comparing two estimation approaches, thecalculation steps should be divided into those which are (a)common to both methods (i.e. same data and method); (b)different for each method.
STEPS FOR WHICH VARIANCE ESTIMATES AREREQUIRED
1. Per capita consumption in area of interest
1.1 Parameter uncertainty in the relationship between
energy consumption and body mass (multiple and exponent)
The quoted SEs of ‘best’ published curve can be used, or ameta-analysis of several published curves.
1.2 Residual variance of species values around the meancurve
The nominal variance around the curve (calculated from thedata used if not given) can be used as a conservative
approach, although it is in principle an overestimate ofprocess error, due to the contribution of observation error.
1.3 Proportion of annual energy requirement obtained
during summer feeding season
1.4 Length of summer feeding season (to get daily energy
requirement)
Due to lack of data, plausible ranges should be given foreach of the above, with a rationale. In considering plausible
ranges, the energy storage capacity of the species should beconsidered (e.g. Blix and Folkow, 1995, for common minke
whales). The less the weight gain in the main feeding
season, the greater the required intake during the rest of the
year.

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REPORT OF THE EXPERT WORKSHOP TO REVIEW JARPN II

Residence time in the study area is not required if
abundance data are expressed in terms of the average
number of whales present in the study area during the study
season.
1.5 Variance in mean body mass (stratified by sex and life
stage, e.g. mature/immature)
Alternatively, the allometric formula for energy requirement
can be applied to each individual and the variance of
nominal energy requirement calculated.
2. Diet composition
The first step is to identify appropriate sampling units andsubunits, for the purpose of variance estimation. The
sampling units could be time-space cells, such as 1° square
by week, or cells more appropriate for the data set. The
subunits should be the individual whales.
The relative proportions of each of the main prey groups(fish, squid, zooplankton) can be estimated only from freshmaterial, subject to assumptions on relative stomach
residency times.
The steps in the calculation that are subject to varianceinclude:2.1 Average undigested biomass of each main prey group in
the forestomach
2.2 Mean residence time of each main prey group in theforestomach
2.3 Average energy content per unit biomass of prey by preytype
For refining the estimates of the relative proportion of eachspecies within the main groupings (a) fish and (b) squid, thefollowing data can be used:

2.4 Average body weight of undigested prey items by
species
2.5 Relative frequencies of each species by counts ofindividuals and/or hard parts
Because the majority of stomachs contain only one preytype, the estimation of relative proportions in mixed-species
stomachs probably accounts for a relatively small part of theuncertainty in total consumption estimates.
3. Abundance (to scale up per capita consumption to
population consumption)
3.1 Variance (and possibly covariances) in estimates ofabundance (mean number of whales present) in survey
season by sub-area and time period, including g(0)variance, and process error
Standard stratum-specific variance estimates can be used,
but if a common g(0) factor is applied to all strata, then firstthe variance of the uncorrected total estimate should be
calculated, and then the g(0) factor and its variance applied
to the total estimate.
Alternatively, a multi-year/area estimation procedure canbe applied, for example for Bryde’s whales (Kitakado et al.,2008).
REFERENCES
Blix, A.S. and Folkow, L.P. 1995. Daily energy expenditure in free living
minke whales. Acta Physiol. Scand. 153(1): 61-6.
Kitakado, T., Shimada, H., Okamura, H. and Miyashita, T. 2008. CLA
abundance estimates for western North Pacific Bryde’s whales and theirassociated CVs with taking the additional variance into account. Paper
SC/60/PFI3 presented to the IWC Scientific Committee, June 2008,
Santiago, Chile (unpublished). 27pp. [Paper available from the Office of
this Journal].

Annex G
Conducting Power Tests using Coalescent Simulations
In this example we illustrate the use of coalescentsimulations to assess the ability of a genetic analysis (in thiscase STRUCTURE - Pritchard et al., 2000) to detect thepresence of multiple populations.
Coalescent simulations work back in time from thepresent. Accordingly, if one population diverged into two x
generations ago, this is coded in the simulations as two
populations merging x generation before the present. A
population expansion in forward time becomes a population
contraction in backward time.
Coalescent approaches are in general based upon theassumption of ‘ideal’ Wright-Fisher populations, where themost notable deviations from cetaceans are non-overlapping
generations, and random mating. For the specific purpose ofthis assessment, these deviation are unlikely to have a major
effect. Both aspects essentially reduce the effective
population size (Ne), and thus reducing population size inthe simulation accordingly would capture the effects ofthese deviations from a Wright-Fisher population.
One user friendly coalescent simulation program is
SimCoal 213 (Excoffier et al., 2000), which has the
additional advantage that the data output is in the Arlequinformat. One aspect that sets SimCoal apart is that multiple
coalescent events per generation are possible (which would
be the case if a large part of the population has beensampled). Most coalescent simulation programs only
accommodate one coalescent event per generation.
For the purpose of assessing the ability of STRUCTURE
to detect multiple populations, this example will aim at thecase where a sample contains individuals from two
populations in different proportions.
In order to conduct simulations for this purpose under
SimCoal 2 the following input parameters are required:13

http://cmpg.unibe.ch/software/simcoal2/.

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J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

Sample size from each population. Sample sizes are in
number of gene copies, which in a haploid system (e.g.
mtDNA) equals the number of sampled individuals, but istwice the number of sampled individuals for diploid loci.
Population sizes. This is the effective and not census,
population size (i.e. Ne). As above in gene copies.
Migration rates. These are written as full matrices. The
values entered are the probability that a gene copy is an
immigrant (m). The product of population size andmigration rates mNe is the number of immigrants per
generation into the target population (which may be derived
from the observed FST estimates assuming population
genetic equilibrium).
Historical events. Changes in migration rates andpopulation sizes. In a two population model, the two
populations have to be merged into one at some point in thepast or the simulation will run infinitely long. The further
back in the past the populations merge, the less thedeviation from drift-mutation equilibrium will be.
In addition, the kind of loci (e.g. DNA sequence data or
microsatellite loci), the degree of linkage among loci andthe mutation rate (u) need be specified. In addition, formicrosatellite loci one can also specify some additional
mutational parameters.
In most cases neither the effective population size nor the
mutation rate is known. However, what is often termed the‘scaled mutation rate’ may be estimated from the observed
data. The scaled mutation rate (θ) is the expected number ofmutations between two gene copies in a population and is
the product of the effective population size (Ne) and themutation rate (u). For mtDNA θ=2Nef μ, where Nef denotes
the effective population size of females. For autosomal
diploid loci, θ=4Nef μ , where Ne is the effective population
size (male and females). θ may be estimated from different
aspects of the data (most based upon an assumption of
population genetic equilibrium). For instance, if one
assumes an infinite site [mutation] model then θ equals thenucleotide diversity. For microsatellite loci θ is equal totwice the variance in the number of repeats.
Accordingly the product of Ne and u used in thesimulations should equal that inferred from the data (e.g. thenucleotide diversity for mtDNA or twice the variance inrepeat size for microsatellite loci). One should checksimulated data if they generate the expected outcomes, e.g.
observed hetetrozygosity, number of alleles and genetic
divergence.
Data sets are then generated by SimCoal 2 for eachcombination of parameter values, and the data are thenanalysed using the relevant program (i.e. STRUCTURE in
this case).
In the file above data are simulated from two
populations, each of an effective population size of 6,500
gene copies. The number of 6500 gene copies translates into
an effective population size of 3,250 diploid individuals,
which in turn translates into a census population size (Nc) of~20,000 if the ratio of Ne/Nc is ~1/6. In this case themigration is symmetrical, with equal rates in both
directions. The rate was set at 0.01, which then translates
into ~32 immigrants per generation. The samples sizes are
125 diploid individuals from one population and 2,375 fromthe other population, yielding a ratio of 1:20 of the two
populations in the combined sample of 2,500 individuals.
Here we have encoded a total of 16 microsatellite loci each

Table 1
Example of a parameter input file for SimCoal 2.
//Input parameters for the coalescence and recombination simulation
program : simcoal2.exe
2 samples to simulate//Deme sizes (haploid number of genes)
6500
6500
//Sample sizes
250
4750//Growth rates
00//Number of migration matrices: If 0 : No migration between demes
1//Migration rates matrix 0:0.00000.01000.01000.0000//Historical event: time, source, sink, proportion of migrants, new demesize, new growth rate, new migration matrix
1
1000
0
1
1
1
0
0
//Number of independent (unlinked) chromosomes, and “chromosome
structure” flag: 0 for identical structure across chromosomes, and 1 fordifferent structures on different chromosomes.
160//Number of contiguous linkage blocks in chromosome 1:
1//Per Block: Data type, No. of loci, Recombination rate to the right side
locus, plus optional parameters ***see detailed explanation here***MICROSAT
1
0.00000.0005 0[EOF]

in a separate linkage block (i.e. all loci are unlinked) andeach with a mutation rate of 0.0005. The mutation rate neednot be the same for each locus but different blocks withdifferent numbers of loci, recombination rates and mutation
rates may be specified.
The genotype data produced by SimCoal 2 are in the
Arlequin (Excoffier et al., 2005) format and need to bereformatted into the STRUCTURE format. Software iscontinuously being published to convert between formats,or if none is available it is a relatively simple matter to writea small routine for this purpose.
Simulations differ from analyses of observed data, by thefact that a large number of data sets need be analysed.
Hence software that may run in batch mode is preferable.
SimCoal is able to run in batch mode. If the publishedversion of a piece of software cannot run in batch mode, it
is often useful to contact the author who may have a batch
version (these are often command-line versions and lessuser friendly, which is why they are often not distributed
widely).
In the above example the ability of STRUCTURE todetect the presence of two populations will depend upon therelative proportions of the two populations in the sample,
their effective population sizes and the degree of migration
between the populations (assuming no change in divergencetime and that the number of loci and samples is given).
Simulations should cover reasonable ranges of theseparameters and for each combination of parameter values asufficient number of simulations need be conducted toobtain a reasonable of the probability of detection. In thepresent case it will mainly be a matter of detecting underwhich parameter values STRUCTURE fails to detect thepresence of two populations, i.e., the fraction of the first
population is low and the migration rates high when thenumber of samples and loci is given.

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The procedure for assessing the necessary amount ofdata (loci and samples) for tests of homogeneity andcharacterising proportions of different stocks in areas of
mixing is similar to that outlined above, adding sample size
and number of loci to the parameters that are assessed andusing the appropriate analytical approach to generate the
distribution of estimates from the simulated data sets.
There are a number of simulation programs in addition to
SimCoal 2, some of which are individual-based such as
EASYPOP14.

REFERENCES
Excoffier, L., Laval, G. and Schneider, S. 2005. Arlequin ver. 3.0: an
integrated software for popultion genetics data analysis. Evol. Bioinf.
Online 1: 47-50.
Excoffier, L., Novembre, J. and Schneider, S. 2000. SIMCOAL: a general
coalescent program for the simulation of molecular data in interconnected
populations with arbitrary demography. J. Hered. 91: 506-09.
Pritchard, J.K., Stephens, M. and Donnelly, P. 2000. Inference of
population structure using multilocus genotype data. Genetics 155: 94559.

Annex H
Recent Resolutions of the Commission Relevant to this Review*
This Annex does not repeat the full text of the Resolutions(full references are given) but rather focuses on those action
paragraphs that may have some relevance.
Resolutions dealing with the Antarctic marine ecosystem
and/or environmental change including pollution
Resolution on the Environment and Whale Stocks (IWC,
1996b)
ENCOURAGES Contracting Governments to continue tocooperate in providing information on the potential effects
both direct and indirect of pollutants on cetaceans as these
become known by forwarding them to the Secretariat.
Resolution on Environmental Change and Cetaceans (IWC,1998)
ENCOURAGES Contracting Governments to continue toprovide available information on environmental changes as
identified above and their known or potential ecologicaleffects on cetaceans through annual Progress Reports andattendance of experts at meetings of the ScientificCommittee;
DIRECTS the Scientific Committee, through the
SWGEC, to provide regular up-dates to the Commission on
environmental matters that affect cetaceans and, inparticular, those that relate to non-natural mortalities
relevant to implementation simulation trials or future RMP
catch limit calculations or that require the action of theCommission within future five year periods of validity ofcatch limit calculations;
ENCOURAGES Contracting Governments to carry out
relevant non-lethal research within domestic andcollaborative
multinationalandmulti-disciplinary
programmes and also to provide new and additional funds
to support the work of the Scientific Committee and
SWGEC in this regard.

14

Resolution on Health Effects from the Consumption of
Cetaceans (IWC, 2000b)
REQUESTS the Scientific Committee to receive, review
and collate data on contaminant burdens in cetaceans … andto report on this matter to the Commission.
Resolutions dealing with Scientific Committee reviewsof Special Permit research
Resolution on Whaling under Special Permit (IWC, 1996a)
RECOMMENDS•
that scientific research intended to assist thecomprehensive assessment of whale stocks and theimplementation of the Revised Management Procedure
shall be undertaken by non-lethal means;•that scientific research involving the killing of
cetaceans should only be permitted in exceptional
circumstances where the questions address criticallyimportant issues which cannot be answered by the
analysis of existing data and/or use of non-lethal
research techniques;

REQUESTS the Scientific Committee, with respect to allSpecial Permit research programmes:
•

•
•

•

to undertake a comprehensive review of all existing
programmes notified to it and report its views on
whether such programmes remain justifiable in the lightof the recommendations above and, in particular, on
whether any lethal scientific research substantiallycontributes to answering critically important questions
which cannot be answered by other means;
to consider all new programmes submitted to it in thelight of the above recommendations;to undertake annual reviews of all programmes and toundertake more intensive reviews of all long-term
programmes at five year intervals;
to structure its reviews of programmes to:

Available at: http://www.unil.ch/webdav/site/dee/shared/softs/EASYPOP_201_userguide.pdf.
*Based on Zeh et al. (2005).

J. CETACEAN RES. MANAGE. 11 (SUPPL. 2), 2010

Identify the relationship between programme objectives
and research needs previously identified by ScientificCommittee;•
evaluate the likelihood of the programme meeting its
objectives by providing reliable answers to thequestions posed;•
identify, where a proposal specifies lethal methods,
non-lethal methods and alternative sources of data
which might be used in meeting the research objectives;
AGREES, should a continuing or proposed special
permit research programme not, in the view of theCommission, satisfy the criteria specified in this Resolution
to so notify the Contracting Government concerned;
RECOMMENDS that Contracting Governments, inproviding the Secretary with proposed special permits andin submitting reports on research programmes to the
Scientific Committee for review, specify how each
proposed special permit or programme satisfies the above
recommendations;
REQUESTS each Contracting Government to ensure thatall scientific information and data available to it withrespect to whales and whaling, including results of research
conducted pursuant to Articles IV and VIII of theConvention, are submitted promptly to the ScientificCommittee for review, analysis and consideration;
Resolution on Special Permits for Scientific Research (IWC,
2000a)
REQUESTS the Scientific Committee, with respect to allSpecial Permit Research Programmes, to provide advice to
the Commission, on the research to be undertaken pursuant449

to any proposed Special Permit or that has been undertaken
in respect of any Special Permit, as to whether the
information sought in the research programme under eachSpecial Permit is:••required for the purposes of management of the species
or stock being researched; andwhether the information sought could be obtained by
non-lethal means.

REFERENCES
International Whaling Commission. 1996a. Chairman’s Report of the
Forty-Seventh Annual Meeting. Appendix 10. IWC Resolution 1995-9.
Resolution on whaling under special permit. Rep. int. Whal. Commn
46:46-47.
International Whaling Commission. 1996b. Chairman’s Report of the
Forty-Seventh Annual Meeting. Appendix 11. IWC Resolution 1995-10.
Resolution on the environment and whale stocks. Rep. int. Whal. Commn
46:47-48.
International Whaling Commission. 1998. Chairman’s Report of the FortyNinth Annual Meeting. Appendix 7. IWC Resolution 1997-7. Resolution
on environmental change and cetaceans. Rep. int. Whal. Commn 48:4849.
International Whaling Commission. 2000a. Chairman’s Report of the FiftyFirst Annual Meeting. Appendix 3. IWC Resolution 1999-2. Resolution
on Special Permits for Scientific Research. Ann. Rep. Int. Whaling
Comm. 1999:52.
International Whaling Commission. 2000b. Chairman’s Report of the
Fifty-First Annual Meeting. Appendix 5. IWC Resolution 1999-4.
Resolution on health effects from the consumption of cetaceans. Ann.
Rep. Int. Whaling Comm. 1999:53.
Zeh, J., Brownell, R.L., Childerhouse, S., Fujise, Gales, Hatanaka, Pastene,
L. and Schweder, T. 2005. Progress Report of the JARPA Review
Planning Steering Group. Paper SC/57/O2 presented to the IWC
Scientific Committee, June 2005, Ulsan, Korea (unpublished). 11pp.
[Paper available from the Office of this Journal].