JA/J05/JR12

First attempt of estimation of feeding impact on krill standing
stock by three baleen whale species (Antarctic minke,
humpback and fin whales) in Areas IV and V using JARPA
data
HIROTO MURASE, TSUTOMU TAMURA, KOJI MATSUOKA, TAKASHI HAKAMADA AND KENJI KONISHI
The Institute of Cetacean Research, 4-5 Toyomi-cho, Chuo-ku, Tokyo 104-0055, Japan
Contact e-mail: murase@cetacean.jp
ABSTRACTTo assess the magnitude of the interspecific competition among three baleen whale species,
Antarctic minke, humpback and fin whales, for their major prey krill, impact of feeding of
three baleen whales on krill was estimated in Areas IV and V form 1999/2000 to 2002/2003
using JARPA data. Three data sets, whale abundance, daily krill consumption rate of whales
and krill biomass, were used in the analysis. All data used in this analysis were collected insitu except daily krill consumption rate of humpback and fin whales because of lack of
biological information. Three baleen whales consumed 10-21% and 30-35% of krill standing
stock in Area IV and V, respectively. In Area IV, humpback whales consumed krill about
twice of Antarctic minke whales. The results indicated that krill surplus for Antarctic minke
whales could be now ended because of increase of humpback whales in Area IV though thispoint should be investigated quantitatively in future using multi-species model including
both baleen whales and krill.
INTRODUCTION
In the Antarctic, intensive commercial whaling on large baleen whales, blue (Balanoptera
musculus), fin (B. physalus) and humpback whales (Megaptera novaeangliae) between early andmid 20th century resulted in increasing in amount of available food, Antarctic krill (Euphausia
superlba), to other krill feeder such as Antarctic minke whales (B. bonaerensis). This phenomenon
is called as “krill surplus” (Laws, 1977) and it has been a central theorem of the Antarctic
ecosystem study. In response to the krill surplus, the decline of mean age at sexual maturity ofminke whales was observed (Kato 1987). But after the ban of commercial whaling of large whales
in 1987, abundance of those species appears to increase in recent years. For example, abundance of
blue whales increase 8% per year at the circumpolar level (Branch et al. 2004) though theabundance was still low comparing with pre-exploitation population size. Humpback whale
Breeding stock D (Area IV) have been showed remarkable increasing rate of abundance (Matsuoka
et al. 2005) and recover to near pre-exploitation level (Jhonston and Butterworth, 2005). Given suchinformation of recovery of abundance of large baleen whales, we have to revisit the krill surplus
theorem and reconsider the current magnitude of interspecific competition of baleen whales for krill.
To assess the magnitude of interactions between krill and baleen whales in multi-species model,
estimation of feeding impact of baleen whales on krill is critically important. Recently, feeding
impact of baleen whales to standing stock of krill was estimated in the South Atlantic region of theSouthern Ocean (Reilly et al., 2004) but it lacked the in situ biological information to estimate thekrill consumption rate of baleen whales.
Data from the Japanese Whale Research Program under Special Permit in the Antarctic(JARPA) provided us opportunity to investigate the feeding impact of baleen whales on krill using
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in situ cetacean sighting and biological data as well as krill biomass data which were collectedconcurrently. JARPA has been conducted every year since the 1987/88 in compliance with Article
VIII of the International Convention for the Regulation of Whaling (ICRW). One of the primaryobjectives of the JARPA is elucidation of the role of whales in the Antarctic marine ecosystemthrough the study of whale feeding ecology. To achieve the study objectives, cetacean sighting andbiological survey has been conducted from the beginning of JARPA. The JARPA interim review
meeting took place in May, 1997. In the meeting, it was pointed out that concurrent studies on the
distribution and abundance of prey species was required to achieve the objective. In response to thatpoint, echo sounder survey to examine distribution and abundance of krill has been conductedconcurrently with cetacean survey in later years of JARPA.
This paper presented the results of estimation of feeding impact of three baleen whale
species, Antarctic minke, humpback and fin whales, on krill standing stock in Area IV and Area V
from 1999/2000 to 2002/2003.
MATERIALS AND METHODS
GeneralTwo baleen whale management area defined by the International Whaling Commission (IWC),
Area IV (70°E-130°E) and Area V (130°E-170°W), were surveyed alternative years in JARPA. In
addition to two Areas, eastern part of Area III and western part of Area VI were also surveyedalternative years. The echo sounder data from 1999/2000 to 2002/2003 were considered as suitable
for this analysis after the biomass estimation of krill using the echo sounder data was conducted.
Intra-annual variability in krill abundance was well known around the Antarctic. For example, krill
abundance early (December) and late (March) austral summer was significantly lower than that in
January around South Georgia (Brierley, et al., 2002). Though echo sounder data were corrected in
1998/1999 in Area V, the survey timing and coverage was different from rest of two years(2000/2001 and 2002/2003). Higher Krill biomass in 1998/1999 than rest of two years could
indicate such intra-annual variability. Eastern part of Area III and Western part of Area VI weresurveyed in either/both December or/and March. Because intra-annual variability of krill biomass
could be one of the confounding factors to interpret inter-annual differences of feeding impact ofbaleen whales on krill, data from Area V 1998/99, Eastern part of Area III and Western part of Area
VI were not considered in this paper.
Abundance estimation
Methodology of abundance estimation of whales used in this study was described by Burt and Stahl(2000) which is the standard methodology adopted by IWC. The program DISTANCE (Buckland etal., 1993) was used for abundance estimation. Details of abundance estimation methods of Antarctic
minke whales and, humpback and fin whales were described in Hakamada et al. (2005) andMatsuoka et al. (2005), respectively.
Prey consumption estimation
We estimated the total prey consumption of krill by three baleen whale species. For Antarctic minke
whale, we estimated the total prey consumption of krill consumed by different maturity stages ofAntarctic minke whales in Areas IV and V based on the abundance data of Antarctic minke whaleand composition of maturity stages of Antarctic minke whales sampled during JARPA surveys(Tamura et al., 2005). For humpback and fin whale, we estimated the total prey consumption of krill
using Method 2 of Tamura et al. (2005). We assumed mean body masses of 30,400 kg and 55,600
kg for humpback and fin whale, respectively (Trites and Pauly, 1998). The average field metabolic
rate used in these calculations was obtained from Blix and Folkow (1995). The value of 80 kJ/kg
per day is based on indirect determination of oxygen consumption from studies of the respiratory
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rates. The total muscle, internal organs fat and blubber masses of humpback and fin whale did not
calculated. We assumed that they need 71.3 % of field metabolic rate as muscle, internal organs fat
and blubber deposition. We assumed that humpback and fin whales spend about 120 days during
the austral summer in the Antarctic (Lockyer, 1981).
Krill biomass estimation
Echo sounder surveys to estimate the krill biomass were conducted concurrently with cetaceansighting survey. An EK500 scientific echo sounder (Simrad, Norway) with software version 5.30
operating frequency at 38 and 120 kHz on board the cetacean survey vessel was used to collect data.
We applied the acoustic data analysis described by Hewitt and Demer (1993) and Demer and
Hewitt (1995). Details of the analysis were described in Murase (2005).
RESULTS
Results of the abundance estimation and the krill consumption of Antarctic minke, humpback, andfin whales and the biomass of krill were summarized in Table 1. Humpback whales fed krill morethan Antarctic minke whales in Area IV. In 2001/02, humpback whales fed krill twice of Antarctic
minke whales. Fin whales fed about same amount of krill as Antarctic minke whales in 2001/2002.
In contrast, Antarctic minke whales fed about 25% of krill standing stock in Area V whereas
humpback and fin whales fed less than 10%. Three baleen whales fed 10-21% and 30-35% of krill
standing stock in Areas IV and V, respectively.
DISCUSSION
Matsuoka et al. (2005) and Jhonston and Butterworth (2005) suggested that humpback whales in
Area IV have increased rapidly in recent years. In conjunction with it, long term trend of age atsexual maturity of Antarctic minke whales were remained constant at 7-8 years in recent years in
Area IV (Zenitani et al. 2005). As the number of predators increase, intraspceicifc and interspecificcompetition for krill would increase. As the results, the decreasing trend of age of sexual maturityof Antarctic minke whales could be suspended. In addition to the constant trend of age at sexual
maturity, it was indicated that blubber thickness of Antarctic minke whales getting thinner andthinner in recent years in Area IV (Konishi et al. 2005). The present analysis suggested that amountof krill consumed by humpback whales was about twice of Antarctic minke whales. Together withthose analysis results, interspecifc competition between humpback and Antarctic minke whales forkrill in Area IV could already occur but the magnitude would not be too high because the trend ofage at sexual maturity of Antarctic minke whales were remained constant at this stage.
Aside from yearly trend, it was indicated that year to year food availability fluctuation could
be occurred. The blubber thickness of Antarctic minke whales in 1999/2000 was higher than
2001/2002 (Konishi and Tamura, 2005). In 1999/2000, abundance of humpback whales was lower
than 2001/2002 (Matsuoka et al. 2005). As the results, feeding impact of humpback whales on krill
was lower in 1999/2000. Such a year to year abundance fluctuation of humpback whales could
affect the biological parameters of Antarctic minke whale though actual mechanism is still open to
the question.
In contrast, because abundances of large baleen whales were still in low levels in Area V, Antarctic
minke whales appeared to be dominated over krill but blubber thickness of mature males ofAntarctic minke whales getting thinner and thinner in recent years (Konishi and Tamura, 2005). The
result could indicate the intraspecific competition of Antarctic minke whales in Area V.
Though feeding impact of fin whales on krill appeared to be low in survey area, one shouldbear in mind that main distribution area of fin whales is far north of the survey area. Though there
are many krill feeders other than baleen whales such as seals and sea birds it is difficult to quantity
the impact of their feeding impact on krill because of luck of precise abundance estimates of those3

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species in the survey area.
Production to biomass (P/B) ratio of Antarctic krill was ranged from 0.8 to 2.27 (Siegel,2000 for review). Given the P/B ratio of Antarctic krill, feeding impact of baleen whales on
standing stock of krill reported in this paper could not be too high. But krill density showed both
short and long term changes. In short term, krill biomass showed large year to year fluctuation at
decadal scale in response to environmental variability such as sea ice extent and oceanographic
conditions (e.g. Pakhomov, 2000; Hewitt and Demer, 2003). It was reported that krill biomass haveshowed statistically significant decreasing trend in the southwest Atlantic since 1976 (Atkinson et al.
2004) though the magnitude of decrease should be studied further to draw the conclusion because
wide varieties of net types were used in the analysis. Because the P/B ratio could change in
response to the environmental variables in both short and long term, feeding impact of baleen
whales on standing stock of krill could be different year to year.
Recent study suggested that baleen whale consumed 4-6% of krill biomass in the South
Atlantic sector of the Southern Ocean (50°S-65°S, 20°W-70°W) (Reilly et al., 2004). Large
discrepancy between their result and this result could be explained by following reasons. As pointed
by Reilly et al. (2004), large proportions of Antarctic minke whales in the South Atlantic sector of
the Southern Ocean may migrate to the ice edge zone during the survey period. It was well knowthat the ice edge zone is important feeding habitat of Antarctic minke whales (Ichii, 1990; Murase
et al., 2002). There was no pack ice in the South Atlantic sector of the Southern Ocean in australsummer. Thus, few presences of Antarctic minke whales could contribute lower krill consumption
rate. Inclusions of sightings in poor weather conditions and ambiguous species identificationsightings in abundance estimation in Reilly et al. (2004) added large uncertainty to estimation ofamount of krill consumed by cetaceans. It was reported that Antarctic minke whales distributed in
the pack ice region where normal cetacean sighting survey vessels could not access (e.g. Naito,
1982; Ensor, 1989). Krill biomass under sea ice was significantly higher than that in open sea(Brierley et al., 2002). Because sea ice is not retreated to the Antarctica in most of portion of Area
IV and Area V, within pack ice distribution of Antarctic minke whales and krill could affect theestimation of krill consumption rate by baleen whales in both negative and positive directions. Krill
consumption by humpback and fin whales could have either positive or negative biases because theestimates had many assumptions. To assess the adequacy of estimated krill consumption rate ofbaleen whales, simulation study of multispecies model such as Mori and Butterworth (2004) could
be useful to predict the dynamics of both krill and baleen whales.
A combination of several analyses including presented in this paper indicated that krill
surplus situation could be changed. But it is not clear whether the trajectory of the Antarctic marine
system could go to the pristine state or to the new equilibrium at this stage. Long term datacollection of baleen whales and krill is necessary to monitor the change in the Antarctic marine
ecosystem. Following points should be considered to improve the estimate of the rate of krill
consumption by baleen whales in future: 1) long term concurrent cetaceans and their prey survey
should be conducted in peak abundance season of krill and whales (January and February) tominimize seasonal effect, 2) the survey should be conducted in same area in same survey timing tointerpret yearly changes and 3) biological information of humpback and fin whales will improve thekrill consumption estimation by those two species.
ACKNOWLEGEMENTS
Authors express thank you to the crews and the researchers who dedicated to collect data in harshenvironmental condition in the Antarctic. We also thank Drs. Hiroshi Hatanaka, Seiji Ohsumi andother colleagues at the Institute of Cetacean Research who made critical comments to improve thismanuscript.

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Table 1. Summary of abundances and krill consumption of Antarctic minke, humpback and fin
whales and, krill biomass in Area IV and V in each survey year.

Area

Year

Abundance Krill consumption
Krill biomass(inds.) （ten thousand ton） （ten thousand ton） (Whale ％）・Antarctic minke whaleIV
1999/00
44,572
2001/0261,463

134.7
185.0

3,642
3,812

3.7
4.9

480.7
521.2

1,867
2,104

25.8
24.8

215.7
439.2

3,642
3,812

5.9
11.5

4,720
2,735

62.8
36.4

1,867
2,104

3.4
1.7

・Fin whale (2,027.9 kg/day)IV
1999/00
1,162
2001/02
7,642

28.3
186.0

3,642
3,812

0.8
4.9

116.9
73.8

1,867
2,104

6.3
3.5

1999/00
2001/02

378.7
810.2

3,642
3,812

10.4
21.3

2000/01
2002/03

660.3
631.4

1,867
2,104

35.4
30.0

V

2000/01
2002/03

148,509
163,593

・Humpback whale (1,108.8 kg/day)IV
1999/00
16,211
2001/0233,010V

V・TotalIV

V

2000/01
2002/03

2000/01
2002/034802
3031

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