Satellite-Monitored Movements of the Northern Right Whale

Author(s): Bruce R. Mate, Sharon L. Nieukirk and Scott D. Kraus

Source: The Journal of Wildlife Management, Vol. 61, No. 4 (Oct., 1997), pp. 1393-1405
Published by: Wiley on behalf of the Wildlife Society
Stable URL: http://www.jstor.org/stable/3802143
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J. Wildl. Manage. 61(4):1997 FOREST ELEPHANT NUMBERS * Barnes et al. 1393

1993. The WCMC biodiversity map library: WILKINSON, L. 1990. SYSTAT: the system for statis-
availability and distribution of GIS datasets. tics. SYSTAT Inc, Evanston, Ill. 677pp.
World Conserv. Monit. Cent., Cambridge, U.K. WILKS, C. 1990. La Conservation des Ecosystemes
61pp. Forestiers du Gabon. IUCN, Gland, Switzerland.
WESTERN, D. 1986. The pygmy elephant: a myth and 215pp.
a mystery. Pachyderm 7:4-5. WING, L. D., AND I. O. BUSS. 1970. Elephants and
WHITE, F. 1983. The vegetation of Africa. UNESCO, forests. Wildl. Monogr. 19. 92pp.
Paris, France. 356pp.
WILKIE, D. S., J. G. SIDLE, AND G. C. BOUNDZANGA. Received 22 August 1995.
1992. Mechanized logging, market hunting, and Accepted 1 May 1997.
a bank loan in Congo. Conserv. Biol. 6:570-580. Associate Editor: Lebreton.

SATELLITE-MONITORED MOVEMENTS OF THE NORTHERN RIGHT

WHALE

BRUCE R. MATE, Coastal Oregon Marine Experiment Station, Fisheries and Wildlife, Hatfield Marine Science Center, Ore
State University, Newport, OR 97365, USA
SHARON L. NIEUKIRK,1 Coastal Oregon Marine Experiment Station, Fisheries and Wildlife, Hatfield Marine Science Cen
Oregon State University, Newport, OR 97365, USA
SCOTT D. KRAUS, New England Aquarium, Boston, MA 02110, USA

Abstract: The northern right whale, Eubalaena glacialis, remains the most critically endangered of the lar
cetaceans despite international protection since 1936. We used satellite-monitored radiotags to identify
late-summer and fall habitat use patterns of right whales in the western North Atlantic. We tagged 9 wh
in the Bay of Fundy (BOF) and successfully tracked them for a total of 13,910 km (f = 1,546 km) in
whale-tracking days (range 7-42 days each, i = 21.7 days). Individuals tracked for more than 12 consecu
days (N = 6 whales) left the BOF at least once and had higher average speeds (Y = 3.5 km/hr) than th
that stayed within the bay (Y = 1.1 km/hr). Three of the tagged whales not only left the BOF, but travel
more than 2,000 km each before returning to the general tagging area. One adult female with a calf went
New Jersey and back to the BOF (3,761 km) in 42 days. Most locations were along bank edges, in basin
along the continental shelf. Eighty percent of locations were in water <182 m (100 fathoms [F]) deep. All
the tagged whales were located in or near shipping lanes. Right whale distribution coincided with areas in
sively used by humans for fishing, shipping, and recreation. Individuals moved rapidly among areas previo
identified as right whale habitat. Whale locations plotted on sea surface temperature (satellite infrared) im
suggest that one whale spent time at the edge of a warm core ring and others spent extended period
upwellings. Observations of whales surfacing with mud on their heads suggest that these whales fed near
BOF seafloor. Satellite telemetry is a useful means of tracking cetacean species that are difficult to view, m
long distances, and might be too expensive to monitor by other means.
J. WILDL. MANAGE. 61(4):1393-1405

Key words: Bay of Fundy, Eubalaena glacialis, Gulf of Maine, habitat, marine mammals, radiotracking, rig
whale, satellite imagery, satellite telemetry, whale.

tection since 1936, fewer than 300 northern

The northern right whale is the most critically
right whales remain in the North Atlantic
endangered of the large cetaceans (Aguilar
1986). Right whales were heavily exploited(Knowlton
by et al. 1994). There is strong evidence
that collisions with ships and entanglement in
early whale fisheries because of their slow-mov-
fishing gear are major sources of injury and
ing, coastal nature, and high oil yield (Aguilar
mortality for this species, and may hinder the
1986, Brown 1986). Despite international pro-
recovery of this population of right whales
(Kraus 1990). The National Marine Fisheries
1 Present address: Battelle Marine Science Labo-
ratory, 1529 W. Sequim Bay Road, Sequim,Service,
WA under section 4(f) of the Endangered
98362, USA. Species Act (ESA) of 1973, has developed a re-

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1394 RIGHT WHALE MOVEMENTS * Mate et al. J. Wildl. Manage. 61(4):1997

covery plan to protect and enhance populationstag attachment. Until now, large tag size and
of northern right whales. Two of the primary lack of adequate attachment methods for un-
objectives of the recovery plan are to reduce restrained animals were the principle obstacles
to using satellite technology to track large ce-
interactions of whales and ships and to identify
taceans. The purpose of this investigation was
and protect critical habitat of the northern right
whale (Natl. Mar. Fish. Serv. 1991). Thesetoob- use satellite-monitored radiotags to clarify
jectives require the delineation of high-usefurther
ar- habitat-use patterns of individual North
eas and the identification of migratory move- Atlantic right whales. Herein we describe the
ments among these areas. movements of remotely-monitored animals
Aside from whaling records, what is known within the western North Atlantic, and identify
the areas in which interactions with human ac-
of right whale distribution in the western North
Atlantic has been deduced from visual sightings
tivities are likely.
during the last 20 years. Right whales are seen We thank Minerals Management Service for
in March through May in Cape Cod Bay funding
and during development of these tech-
the Great South Channel. In May through July, niques and unique datasets; R. Mesecar for tag
individuals are scattered offshore throughout design and construction; M. L. Mate, T. Martin,
the Gulf of Maine (GOM), and from July the New England Aquarium right whale group,
through October, right whales are seen in J. theOwen and D. Stahlke for field assistance; T.
lower BOF and on the Nova Scotian shelf Martin for tag program and data analyses ad-
(CeTAP 1982, Kraus et al. 1986, Mitchell et V.
vice; al.B. Barry for manuscript assistance; Te-
1986, Schevill et al. 1986, Winn et al. 1986, lonics, Inc. (Mesa, Ariz.) for technical assistance
Mayo and Marx 1990). Between November and with tags; C. P. Fairfield for Television Infra-red
March, only the distribution of females andObservation Satellite (TIROS) infrared sea sur-
calves on the calving ground off Georgia and face temperature (SST) images from the Uni-
northern Florida is known. Little is known versity of Rhode Island National Oceanic and
about the detailed movements of individual
Atmospheric Administration (NOAA) facility; J.
Clark
right whales throughout the year (Kraus et al. for additional sea surface temperature
1986, Mead 1986). summaries and interpretation, and S. C.
Beavers for editorial review. Hewlett Packard
Obtaining habitat-use data for right whales
Company employees (Corvallis, Oreg.), Hayes
traditionally has been difficult because they
spend so much time underwater, and individu-
Corporation (Atlanta, Ga.), Fred Biller (Bloom-
ingdale, Ill.), Memorex Inc.(Portland, Oreg.),
als are often difficult to re-identify. In addition,
data collected from visual observations alone Toshiba Inc.(Sunnyvale, Calif.) and Zenith Data
Systems, Inc.(Buffalo Grove, Ill.) provided
are restricted to daylight and periods of fair
weather. High frequency (HF) and very high products for our research. The Oregon State
frequency (VHF) radiotelemetry can overcome University Endowed Marine Mammal Research
some of these constraints (Evans 1974, Watkins Program provided additional financial assistance
et al. 1981, Mate and Harvey 1984, Goodyear for this project. This research was conducted
1989), but require that a vessel stay within the under U.S. National Marine Fisheries Service
limited transmission range of each tagged indi- permit #678 and Canadian Department of Fish-
vidual. Thus, long-term monitoring with con- eries and Oceans permit #116.
ventional telemetry is logistically difficult and
METHODS
expensive.
Recently, satellite-monitored ultra high fre- We tagged right whales from the R/V Nereid,
quency (UHF) radiotags have been used suc- a diesel-powered 10-m fiberglass boat, during
cessfully to track small marine mammals such cruises in the BOF and off the southern tip of
as harbor seals (Phoca vitulina; DeLong and Nova Scotia August through October in 1989
Stewart 1991), a pilot whale (Globicephala me- and 1990, and late September through early
laena; Mate 1989), a white-sided dolphin (La- October in 1991 (Fig. 1). To identify, sex, and
genorhynchus acutus; Mate et al. 1994), nar- age individuals, we photographed callosity pat-
whals (Monodon monoceros; Martin et al. 1994), terns and scars of whales and compared these
and bottlenose dolphins (Tursiops truncatus; photographs with those in the New England
Tanaka 1987, Mate et al. 1995). Capture of Aquarium catalog of North Atlantic right whales
these species was required to achieve a suitable (Crone and Kraus 1990). We drove the boat on

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J. Wildl. Manage. 61(4):1997 RIGHT WHALE MOVEMENTS * Mate et al. 1395

NEW
WESTERN NORTH ATLANTICNS C
STUDY AREA ND OA 0
......................... ...U N DY
0 50 100 150 200
Kilometers

)EALD/

NHL7EDGE1
UL G UF OF BRO S
IBAPK

GEORGES BANK N]RTH

Ri A Tl AN TIG
CT 07EAN

39 00-.:?i?:l.::?':~_:?::i'.?:~';?~ 1 ~ i ~

74 00 73 0 72 0 71 0 70 0 6900 6 00 U uu 6 00 5 00 64 0

741 0 7300 7200 7100 70 OO o6 O 6o0 67 00 6600 6500 6400

Fig. 1. Map of the 1989-91 right whale study area showing the 50 fathom

a course parallel to the whale transmitter varies with latitude.

to obtain photo- The amount of
graphs and attempt tagging. timeWe did is
the satellite not tag
within rangean-
during a "pass"
imals if they were calves, depends had pronounced
on the elevation of the skin
satellite above
sloughing, were visibly underweight, or were
the horizon. In the study area (45'N), each of
entangled in fishing gear. We the 2 attempted
satellites is overhead
toabout
min- 5-6 times each
imize the disturbance of females with calves day for an average of 10 minutes on each pass
during approaches for tagging. (Argos 1990). As a satellite passes over the tag,
transmissions are received (an "uplink") when
The Argos Data Collection and Location the whale is on the surface and the transmitter
System
clears the water. Service Argos uses the Dopp-
We used the Argos Data Collection and Lo- ler shift data (resulting from the speed of the
cation System, a satellite-based tracking system satellite) to calculate the location of a transmit-
administered by Service Argos (Argos 1990, ter when there is a sufficient number of trans-
Fancy et al. 1988, Harris et al. 1990). Argos- missions (Fancy et al. 1988). Service Argos lim-
certified transmitters used an UHF signal of ited our transmissions to no more than 1 every
401.650 MHZ. Argos transmissions are moni- 40 seconds.
tored by any of 4 receivers on each of 2 polar- The accuracy of Argos-acquired locations de-
orbiting, sun-synchronous, TIROS-N weather pends primarily on the number and distribution
satellites. The number of times per day a sat- of transmissions received during a pass (Argos
ellite passes within the reception range of a 1990). Locations are categorized by location

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1396 RIGHT WHALE MOVEMENTS * Mate et al. J. Wildl. Manage. 61(4):1997

class (LC). Locations can be calculated with as

few as 2 transmissions (LC = 0), but the ac-
curacy of such a location is unknown. Argos has
estimated that at least 68% of locations (? 1 SD)
classified as LC 1, 2, and 3 are within 1,000,
350, and 150 m, of the true location (Argos,
1990). We conducted experiments through the
Argos system to determine how these estimates
compared to the actual performance of our
transmitters.

Tag Design
In response to technological advances, both
the hardware and software of the satellite-mon-
itored radiotags evolved during the course of
the study. A reduction in transmitter size from
1989 to 1990 resulted in a radical change in
attachment methodology.
O 5 10cm
In all years of the study (1989-91), we used
a cylindrical tag housing as the best shape to
accommodate the pressures from deep dives. In
1989 the cylinder (7 cm diam, 15 cm length)
contained the transmitter, controller board, 6
Altus? C-cell lithium batteries (no longer avail-
able) and a programmable 1-Watt Telonics ST-
3 transmitter (Mesa, Ariz.). The marine mam-
mal group at Oregon State University devel-Fig. 2. The 1990-91 satellite-monito

oped the housing, a microprocessor controller,

and appropriate software. One endcap con-
tained pressure and temperature sensors. Theconsisted of a stainless steel
other endcap had a 17-cm, vertically-mounted, ameter and 14 cm long, m
flexible UHF antenna. We attached a small of the tag perpendicular to
VHF transmitter (3.0 x 5.0 x 2.5 cm) to cylinder
the (Fig. 2). Double-e
distal end of each rod pen
outside of the tag housing to facilitate shipboard
relocation of tagged individuals. skin and 2 pairs of folding t
from
We lowered the 1989 tag from the end of a backing out easily. W
Barnett International, Inc. (
4.5-m pole onto the back of a free-swimming
compound
whale. A spring coil wrapped around the tag crossbow to app
terminated in 2 arched, stainless steel, 0.6-cm
tances up to 10 m. An alum
diameter sutures that secured the tag to the
"C"-shaped cup held the tag
whale. A mechanical trigger at the base ofandthe fell away after tag attac
tag released the spring tension and applied kg
the test line was attached to
and recovered the tag if it m
sutures 8 cm deep in the epidermis and blubber
of the whale. A conductivity sensor (sa
In 1990, Telonics, Inc. developed a smaller, the endcap allowed transm
0.4-Watt transmitter (ST-6) that enabled us to above the surface to conse
reduce the tag volume and weight considerably. We placed tags with a vert
The 1990/1991 cylindrical tag (5.6 cm diam X tation high on each whale
12.5 cm length, wt = 0.57 kg) contained an ST- behind the blowhole, to ens
10 transmitter powered by 4 Saft? lithium A- A software program limited
cells and a VHF transmitter powered by a single sion schedule ("duty cycle")
Saft? A-cell (San Diego, Calif.). We mounted that most closely coordinat
antennas for the VHF and UHF transmitters on of the 2 active NOAA satell
opposite ends of the cylinder. Tag attachments tive portion of this duty cy

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J. Wildl. Manage. 61(4):1997 RIGHT WHALE MOVEMENTS * Mate et al. 1397

Table 1. Distances between Argos-calculated locations and

2, and LC 3 locations (Argos relies on the user
actual test location for different location classes. Oregon State
University estimates were computed by calculating the great
to estimate the error associated with LC 0 lo-
circle distances between Argos-acquired locations and the lo-
cations). The 78 LC 0 locations ranged from
cation of the tag in our laboratory. One Class 0 location was
264 km from the test location and was not included in the <.01 to 59.6 km from the true location (? = 8.3
analysis. km). Sixty-eight percent of the LC 0 locations
were within 7.5 km and 95% were within 22.5
Argos esti- Oregon State University estimates km of the reference point. Thus, the accuracy
matesa

Loca- 68% of lo- 68% of 95.4% of of any specific LC 0 location cannot be well-
tion cations locations locations Mean Median No. of lo- defined and reinforces the need for additional
class (km) (km) (km) (km) (km) cations
screening criteria. Less than 33% of locations
0 NAb 7.5 22.5 8.28 4.82 78
1 1 2.3 7.3 2.38 1.61 102
met the Argos estimated distance criteria for
2 0.35 0.9 1.7 0.79 0.64 71
LC 1, LC 2, and LC 3.
3 0.15 0.3 0.9 0.32 0.26 72 The animals showed little or no obvious re-
action to projectile tag attachment. Often
"Argos sample sizes and tag test conditions were not available.
tagged
b Not Available; Argos relies on the animals
user towould swim away
estimate from
the our ves- of
accuracy
Class 0 locations.
sel, as did many whales that we closely ap-
proached but did not tag. In one instance we
approached and tagged a whale that had been
sent a 64-bit message (no more than once every
resting quietly at the surface. The whale re-
40 sec) that was 0.34 seconds long whenever the
sumed resting within 10 minutes after tagging.
whales were at the surface.
Whales tagged by pole (1989 only) required a
We plotted Argos-acquired locations with closer
a approach than the crossbow deployment
mapping software (CAMRIS; Ecol. Consulting and had stronger reactions to tagging attempts.
Inc. Portland, Oreg.) and digitized data from This result may have been due primarily to our
NOAA chart #13003 and Canadian Hydro-close approach as some whales reacted even
graphic Service Chart #L/C 4003 and calculatedwithout tagging. On 2 occasions, whales tagged
speeds (km/hr) between adjacent locations. Weby pole made a forceful dive, in one case with
have observed right whales swimming up to 15 a lateral tail slash.
km/hour. We added an additional 20% (3 We tried visually to relocate tagged individ-
km/hr) to this speed and reevaluated all location
uals (during subsequent tagging attempts with
pairs that resulted in speeds exceeding 18 other whales) to document the condition of
km/hour. We examined how speeds wouldboth be the tag and the animal before and after
affected by the elimination of one or the other
tag loss. In 1990, we observed 2 instances where
of the locations and deleted the one apparently
attachments fixed to the endcaps had broken
responsible for the excessively high speed. Lo-
and resulted in some localized swelling around
cations on land were also eliminated. In 1989-
the attachments. In 1991, we redesigned the at-
90, we used infrared sea surface temperature
tachments to go through the wall of the metal
images (SST) from NOAA TIROS-N satellites cylinder which eliminated this problem. We ob-
(Univ. Rhode Island Remote Sensing Lab., Nar-
served all but 1 of the tagged whales (#1421)
ragansett) to examine the movements of tagged
after tagging, including all tagged females with
whales with respect to identifiable oceano-
calves. These resightings demonstrated that
graphic surface thermal features. there was no apparent effect on the close as-
sociation between mother and calf. Whale
RESULTS
#1140 was seen 58 days after tagging (16 days
To estimate location accuracy, we examinedafter the last received transmission). She was
323 locations acquired from experiments still with her calf and had lost the tag. There
in our
laboratory (Table 1). The great circle distances
were no signs of swelling or infection and only
between Argos-acquired locations and athe tag 1-cm-diameter white scar where the tag
single
test location were computed. Frequencyhad tabu-
been. A photograph of whale #1140 5 years
lations were used to determine distances for after tagging showed the scar surrounded by a
68% and 95.4% of the data because the calcu- slightly raised mound in the skin. The area of
tag attachment has been photographed for 6
lated distances were not normally distributed.
Frequency tabulations also facilitated compari-
other individuals as well. Typically, scarring was
son with Argos accuracy estimates for LC 1, minimal
LC and included small, white, depressed

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1398 RIGHT WHALE MOVEMENTS * Mate et al. J. Wildl. Manage. 61(4):1997

Table 2. Satellite-acquired location data from 9 radiotagged right whales.

Total # Total # Distance traveledd Speed

NEA#a Statusb Tag date Tag days XMITSe locations (km) (km/hr) Left BOFe?
#1146 Adult 6 15 Oct '89 24 304 72 1,523 2.7 Yes
#1421 Adult 612 Sep '90 42 665 137 3,056 3.0 Yes
#1629 Adult Y+ Calf 26 Aug '90 10 30 8 196 0.8 No
#1981 JUV ? 25 Aug '90 12 112 26 433 1.5 No
#1140 Adult 9 +
Calf 24 Aug '90 42 528 110 3,761 3.7 Yes
#1135 Adult 9,
PREG 24 Aug '90 7 86 16 779 4.6 Yes
#1243 Adult 9+ Calf 27 Sep '91 10 29 6 218 0.9 No
#1608 JUV 9 28 Sep '91 24 162 42 2,012 3.5 Yes
#1406 JUV 9 5 Oct '91 24 228 48 1,932 3.4 Yes
Total: 195 2,144 463 13,910
Mean: 21.7 238.2 51.7 1,545.6 2.7
(SD): 13.34 224.33 46.27 1,233.09 1.33

a New England Aquarium (NEA) right whale catalog no.

b JUV = juvenile animal; ? = unknown sex; PREG = pregnant female.
c XMITS = satellite-acquired transmissions.
d Straight-line distance between tagging location and subsequent Argos-acquired lo
e BOF = Bay of Fundy.

scars at the site of tyneMean tracking time and

attachment for the 9 animals was 21
pink
cyamids in one case. Overall, ? 13.3 days
there
(rangewas
= 7-42no
days),
vis-and a total of
ible evidence of adverse 2,144 health effects
transmissions was(heavy
received. Eighty-seve
external parasite loads,percent skinofsloughing, etc.) (456 ) m
the 525 original locations
from tagging. our editing criteria with a range of 5-136 loca
During the 3 years of the study,
tions per we tagged
whale, resulting in an overall daily a
19 whales and successfully eragetracked
of 2.1 ? 0.9 9 (1 ofper
locations 7 whale
in (range =
1989, 5 of 9 in 1990, and 3 of 3 in 1991) via 0.6-3.3) during the maximum 8 hours of daily
satellite for more than 7 days each (Table 2). transmission. Ninety-five percent of the spee
calculated between locations were less than 10

Table 3. Precision of satellite-acquired locations of right km/hour. Seventy-one percent of the 456 edited
whales. Number and percentage of locations recorded for locations were LC 0; 18%, LC 1; 10%, LC 2;
each whale are categorized by location class. and 1%, LC 3 (Table 3).
Location classa Movements
NEA # 0 1 2 3 Totalb
Although all of the whales we tagged were
1421 86 31 17 2 136
in the BOF, subsequent use of the bay, dis-
(63%) (23%) (13%) (1%)
1629 6 1 0 0 7 tance traveled, and average speed differed
(86%) (14%) widely among tagged whales. Six whales left
1981 22 2 1 0 25 the BOF within 1-7 days after tagging. We
(88%) (1%) (1%) tracked these whales 27.2 ? 13.2 days ( =
1140 71 24 14 0 109
(65%) (22%) (13%)
2,177 + 1,072 km). They either took a route
1135 13 2 0 0 15 southwest along the New England coast, trav-
(87%) (13%) eled south and east around the southern tip of
1146 52 11 8 0 71 Nova Scotia, visited both coastal areas, or in
(73%) (16%) (11%)
one instance traveled south into very deep wa-
1243 5 0 0 0 5
(100%) ter before returning north to the Scotian shelf.
1608 37 3 1 0 41 All 5 whales tracked for more than 12 consec-
(90%) (7%) (3%) utive days left the BOF at least once. Three
1406 34 9 4 0 47 whales did not leave the BOF; these whales
(72%) (20%) (8%)
All animals 326 83 45 2 456
were tracked an average of 10.7 ? 1.2 days and
(71%) (18%) (10%) (1%) 282 ? 131 km and traveled 1.1 + 0.4 km/hour.
Nineteen percent (n = 86) of all locations were
a Location classes were assigned by Argos, and reflect more than
estimated quality100 km from shore; most of these
and quantity of transmissions (see Table 1).
b Total does not include tagging location. locations were acquired from whale #1421 (n

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J. Wildl. Manage. 61(4):1997 RIGHT WHALE MOVEMENTS * Mate et al. 1399

Table 4. Water depth of Argos-acquired right whale locations.

Water depth
Fathoms 0-10 10-50 50-100 100-500 >500
Meters 0-18 18-91 91-182 182-194 >914
NEA# Total locations % % % % %

1421 137 0.73 36.50 25.54 5.84 31.39

1629 8 12.50 50.00 37.50 0.00 0.00
1981 26 3.85 3.85 65.38 26.92 0.00
1140 110 9.09 44.54 42.73 3.64 0.00
1135 16 6.25 37.50 43.75 12.50 0.00
1146 72 0.00 8.33 66.67 25.00 0.00
1243 6 0.00 33.33 66.67 0.00 0.00
1608 42 13.50 29.70 43.3 13.50 0.00
1406 48 2.08 18.75 64.58 12.50 2.08
All animals 465 4.52 29.89 45.16 10.97 9.46

= 61) and whale locations

#1406 (n =
at an average speed19). Thir
of 3.5 km/hour.
percent of all locations
Tagged in the BOF, were
he went southin to wat
Brown's
deep (Table 4). An
Bank additional 45%
within 2 days of tagging and remained of
there
were in water between 91-182 m. for 7 days. He then crossed the Gulf of Maine in
Movement of Males.-The 2 tagged adult 2 males
days, and spent the last 10 days of tag operation
(#1146 in 1989 and #1421 in 1990) left the onBOF
Jeffrey's Ledge, where SST images showed
almost immediately after tagging (Fig. 3). strong
During upwelling. The other adult male (#1421)
24 days whale #1146 traveled 1,523 km among 72 further offshore than any other whale.
traveled

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Kilometer
il:illiiiiili~ id. i ~i~ilii~iii~'l:iii ill~il.:~::ii0 # 14 2 1j ;l:i~~::.i1:~::i:I?': iiiiiiil~llli'~iiiiii
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74 iiiilIiii:.ii~iiiil O 5 00 72 0 70 00 0 0 /?..?:? IXVI111 16 1 - ?-?l-d

Fig. 3. Satellite-monitored movements of 2 male right whales radiotagged in the Bay of Fundy. The locations of the Gulf Stream
and Warm Core Ring (WCR) were approximated from NOAA/TIROS-N satellite sea surface temperature images. The WCR
pictured was designated WCR #41 and originated at 40ON x 60OW on 25 July 1990.

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1400 RIGHT WHALE MOVEMENTS * Mate et al. J. Wildl. Manage. 61(4):1997

............ .....
-
..............
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.... .
. . . .. . ...... . . . .. . . . .. . . . . .. . . -. .. . . ..
.. .. -- -- - - -- -
.. . . . . . -.............. .. . . . . . .. . . . .
. . . .. .. .. . . .

. ...... -------------------- .... .. ................ ...............

..... ... .. .. . ... .. .. . ... .. ...... .. .. .. ... ..
.......... . .......

... .. ... .. ... ... ... . ... .. . . .. .. . . . .. . . . . .. . . . . . . . .

. . . .. .. . .. .. . .. . . .. . .. . .. .

. ...... .. . .... .... .. .

.... .... .... ...

.... ................
.. . .. ... .. . .. . .. . . .. . .. . .. ........
. .. . .. . .. . . ...

- - - - - - - - - - . . .

.... . . .... . .......i~ ii~i?;

. . .. . ...........

... .. .. .. ... .. .. ..

.. . .. . . .-- - - .. . .. . . .. . .

. . . .. .. ....

74 00 72 0 1. . . ......... . . 0

Fig. 4. Satellite-monitored
(#1135), and 3 females wi

Tagged in Movemen
the BOF i
south calves
through the (#16 G
of km in 1990 Bank
George's and 218 km in 1991. Both whales
to th
Stream, 500
remained within km the BOF during soutthe 10 days
4,200 m each
deep.
was tracked (Fig. 4). Calculated
The average
George's Bank and
speeds for these 2 whales th
(f <1 km/hr) were low-
crescent er than the other whales. Fouras
shape, of the 6 tagged
di
female whales (#1140,
northbound moveme #1135, #1406 and #1608)
the moved out of the BOF (Table 2). In 1990, whale
crescent-shaped
the #1140, a female with
western and a calf, was tracked
east for 42
mass (18-20
days and traveled 3,764 C) bein
kmn (f= 89.6 km/day or
along the 3.7 km/hr)
eastern among 110 locations (r =sid 2.6/day)
#41 (J. Clark,
(Fig. 4). She spent the firstNOAA
2 weeks after tagging
parting the
in the BOF shippingWCR channel east of Grand Ma- a
nearly nan Island and in the shallower waters
straight north to the
ifax, south of the island. north
further During the next 3 weeks, shein
tracked took a coastal route (all locations
whale. He within 120
th km
and basins
of shore) south
off to Cape Cod Bay,
the where she ands
SST images
her calf wereshowed
sighted (D. Matilla, Cent. Coastal t
ized Stud., pers. commun.). Whale
upwelling #1140 then contin-
during
Overall, we
ued to New Yorktracked
and New Jersey waters, and
returned to the locations
Forty-three BOF in late September during
than 900 the
m (Table
last week of tracking. 4).
was locatedThe female whale
(once) #1135 was sighted with
ina

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J. Wildl. Manage. 61(4):1997 RIGHT WHALE MOVEMENTS * Mate et al. 1401

45 0

------ - -----------------

44 00~

430

oo
?8
42 00 _____

0 20 40 60 80 100120140160180

Kilometers

41 00 * 41406
V1 41608
0 4981

70 00 69 00 ~ 68 00 67 00 66 00 65 00 64 00

Fig. 5. Satellite-monitored movements of 3 juvenil

and #1608), and one of unknown sex (#1981).

calf in September 8 1991 days (Fig. 5). and

The whale'swas
route was therefo
similar to
nant when tagged thein1990 track
Augustof a female with a calf (whaleWhe
1990.
sighted during the 1990
#1140) research
and also returned very near its BOFseason
tag-
fore tagging, she ging was unusually
location. This round trip movement active,
was
ming rapidly in a accomplished corkscrew in 13 days (I = 94fashion
km/day). After at
face. The tag transmitted only 2 days in the BOF, forwhale #1608only
returned 7 d
during this time to whale
Cape Cod Bay in#1135 swam
just 5 days (z? = 130 km/day, fr
BOF to Brown's Bank, Baccaro Bank and Em- or 5.4 km/hr). Overall, this whale moved 2,012
erald Basin, a minimum of 770 km (n = 16 km in 24 days among 42 locations at an average
locations; Fig. 4). Her overall mean swimming speed of 3.5 km/hour. Another juvenile female
speed, 4.6 km/hour (0.6-12.8 km/hr), was the (#1406) tagged in 1991 moved out of the BOF
highest recorded during the study. 3 days after tagging. After 6 days in the upper
Movement of Related Whales.-In 1990, we GOM, she moved 175 km E.S.E. of the tip of
tracked a pregnant adult female (#1135), and in Nova Scotia, further east than any other tagged
1991 both her daughter (#1243) and one of her whale, into an area with no previous reported
granddaughters (#1608). These 3 whales used sightings. A juvenile of unknown sex (whale
all of the areas that have been identified to date
#1981) traveled 433 km in 12 days among 26
as right whale feeding areas (BOF, Brown's locations, but never left the BOF.
Bank, GOM, and Cape Cod Bay). However,
DISCUSSION
there was no precise overlap of movements to
suggest that related whales have stereotypic Previously, the right whale was considered a
movement patterns (Figs. 4 and 5). slow-moving, nearshore species that tends to
Movement of Juvenile Whales.-In 1991, a occur in restricted areas for well-defined peri-
juvenile female (#1608) traveled a coastal route ods of time (CeTAP 1982, Natl. Mar. Fish. Serv.
southwest from the BOF through Stellwagen 1991). This pattern was not observed in the
Bank, to the northern edge of Cape Cod Bay in whales we monitored. Instead, most individuals

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1402 RIGHT WHALE MOVEMENTS * Mate et al. J. Wildl. Manage. 61(4):1997

were highly mobile. On average, the 9 whales Florida and Georgia, (2) the Great South Chan-
moved 68 km/day, and 2 individuals each trav- nel, (3) Cape Cod Bay and Massachusetts Bay,
eled more than 3,000 km in 6 weeks during (4) thethe BOF, and (5) Brown's and Baccaro
study. All animals tracked more than 12 days Banks. Right whales are observed routinely in
the latter 2 areas in the late summer and fall
left the BOF and several left within a few days
of tagging. Two individuals returned to the (Natl.
BOF Mar. Fish. Serv. 1991). Tagged whales
one or more times. In the past, "residency traveled through areas 3, 4 and 5. Six of the 9
times" in the BOF were estimated by firstwhalesand used 2 or more of these areas while
tracked
last dates of visual sightings. From this study it and often moved quickly between
appears that those visual techniques overesti-them. Spatial and temporal use of these areas
mated residency times in the BOF. Movements varied among individuals. Two animals (#1421,
in and out of the BOF may be routine for many,an ad M, and #1406, a juv F) used the banks,
or even most, right whales. basins and shelf edge east of Nova Scotia, a re-
gion previously not considered a high use area
Currently, Argos is the only satellite-linked
for right whales. Whale #1421 unexpectedly
tracking system available to civilians. The Argos
system has been used in numerous studies to
traveled more than 500 km offshore into deep
monitor the movements of wildlife (Craighead water, where right whales have not been re-
and Craighead 1987, Fancy et al. 1988), but ported.
it A female (#1140) and her calf spent 2
has limited ability to provide accurate and weeks
de- in coastal waters off New Jersey and New
tailed movement data for free-ranging marine York, an area previously not identified as a feed-
animals that spend short periods of time at ingthe area or used by calves. Continuous moni-
surface. For example, 71% of our 456 edited toring of the movements of tagged right whales
locations were Class 0 (i.e., unknown accuracy).
suggests that the summer/fall high use area for
Quantifying the error associated with an indi-these whales is the BOF and the entire GOM
vidual Class 0 location obtained from a free-
(Fig. 6).
ranging cetacean was not possible. Increased
Tagged whales spent time along banks, ba-
confidence in Class 0 locations was achieved sins, upwellings, thermal fronts, and the edges
when they occurred frequently enough thatofsev- WCRs. Zooplankton are typically found in
eral were clustered in a small area during a high
rel-concentrations in association with these
atively short period of time. Based on ourphysical
lab- features (Owens 1981, Wroblewski and
Cheney 1984, Kenney and Winn 1987, Brown
oratory tests, we believe Class 0 locations were
adequate for describing the far-ranging move-and Winn 1989, Mayo and Marx 1990, Herman
ments of right whales. Locations of higher et ac-
al. 1991). As a WCR turns clockwise, organ-
curacy were not obtained more often because isms are concentrated at the frontal boundary
between the WCR and the cold shelf and slope
of the long dives and short surfacings of right
water entrained by the ring (Olson and Backus
whales. In a study of a bottlenose dolphin (Tur-
1985). Therefore, the copepods upon which
siops truncatus), a cetacean that surfaces more
frequently than a right whale, only 19% ofrightlo- whales feed may have been quite abun-
dant in the 16-18 C water to the east of WCR
cations were Class 0 (Mate et al. 1995). In ad-
dition, in our control tests and those of others,
#41. This concentration mechanism may explain
the error associated with locations of class 1,
why 2, the southward and northward movements
and 3 often exceeded Argos error estimates
of whale #1421 closely paralleled the edges of
this cold water mass.
(Fancy et al. 1988, Stewert et al. 1989, Keating
et al. 1991). Although the combined impreci- The extensive whale movements documented
sion of both Argos locations and SST imageshere likely represent searching for food. The
(the resolution of SST images was limited to 1.1 degree of overlap among the movements
high
km) makes determination of whale tracks along of tagged whales in different years suggests that
small-scale thermal fronts difficult, Argos loca- most whales are quite mobile and may use
tions and SST images were sufficiently accurate many of the same areas routinely. The large
to place whales in the vicinity of large-scalenumber
fea- of whales in the BOF shipping channel
tures such as WCRs and areas of upwelling.also is likely related to food availability. The
The National Marine Fisheries Service has channel is the deepest portion of the bay, which
defined 5 areas in the North Atlantic as seasonal is well known for its unusually large tides. We
"high-use areas" for right whales: (1) coastal saw whales in this region surfacing in water 200

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J. Wildl. Manage. 61(4):1997 RIGHT WHALE MOVEMENTS * Mate et al. 1403

ii-:j~:il::rs:i j?::- iiiiii i litil i i +i 1 + Ii n;::j:j + I +1 i l i +i! E ii i iii +++i+++++ii+

!iiii i i~S~ TU Y A E iii iiii .:~~~l i ii }l:-#:: ri:: }i~iii:iiiii::::: :;riii~ l ! Y" OF" ii
... . . .... ... ...... . :., ;: ::,: j .I
... . .... .. ....... . ... . .. .. .. .... .
. 1. .......... N EW:I8i~r ?8
;fbf.......... . . D10 MR: ...........S~
...... . . ............ .... . ....?:iiilli ~ ??ij:il'jc

sliiii...
WESTERN iO T ATLANTI :Iiiiiil;?i:i ili~~r~ ii i ,,,,,,,,,,,,,,,,,,,,,++ -0 , "
S:vv

,ii ....
45
.... . . - _: ....- o . .
.......... ...... 0 1'i~~i! l~i?:::::::
..............

00
. ... .... ..............

:'..+.'-.i-.:l:;illllli'lll::iB+ii:i,8
~

_..++ii ! .I. ........ . . .. .....$

.................. O0 7 O0 7O6O60
0 :::::::::::::::::::::::::::
50 100 ili +,:i!,+++ :150
+::+,,,+ ,,, +,+,,,,+,,,+200~i'i:ii:ijiiii ~~W~~j JL F f~ - ~
::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::::++:;:::+
+,,,+++ +,:+,,,,+ + ++,+ ........................ .... .....++,+ : - ? J E ,: , , _ l ., , w
ii::iiiiiii::i~ii~i~iiiiii~ i:i::i::ii iiii::i:iiii::iiiiiiiiii::iiiiiiiiiiiii.!ii::i:iiiiiiiiiiiiiiiiii::iiiiiii iiiiii:iii:.+ii::::Y:ii:iiii iiai::iiiiiiii:i:iiii::iiiiii i:::::: !i~ii!i!.i.::iiili+:i~l " Ij U / t ! ' " ' Ii<
i llii!+i+ii~i!ii
,X.- + ii?! _ :?i:!iii+~iiiiii~!!?MEN
I ;i i;iii !ii++!iiiii~iii!!+i+++ii `` ?
. .......

:.'l?:-i::ll?:iI?i~l_:::~i~;:: :8::`:~ia::::-::::::?:::i~:I':-
:~i:::~:: :~i?jai::i:_:::::::::::::-:::-iiil::i''~i'~ j~Iir;;::::iiii ? ~ BANK
0 AERALD:'":?'~ii~iriiiiaii~
. ..... ....
~ i:p-i~ii-'~i ll-iil' Il~ll~p:il iijiil(iii

.......... ..... E D G E
........... Bl~iii~iii;'
AN''''--I:-'i~i B A
.............. .. .. .:ji::::A I;A T
........... .........''' ': :~ iii ii ii ii 'i 'l B
..........i:i.~~~iiiiii I~ii~;iii: j

+ + + +++ +++::++ ++++++ ++++::+ + ++ ++++ I

-ilil'_iiii':~:. :: ::: ::::::::~i::-::?M A INEi:- B )~i
USTELLWAC E

#,~i~ic:- ".0 %o
:-.1 .. . ........... . .... . -BAN
.. .....

.. . . . . .. . . . . .

m ;:::m m j::.::.. :
I/I I I I I K A ~
42NVA 1?pl_ o

......... .. ..?BA_ GERGES BAK N TJ

..............0 7 0 6 0 8 00 67 0 6 0 5 0 64 0

Fig. 6. A composite of all satellite-acquired locations for 9 right whales tagged in

m deep with mud on the topput ofthem

their heads.
in danger As collisions. Despite
of ship
copepods are often found in our denseextensive searching, right whales were
aggregations
near the sea floor (Kann and found Wishner 1995), we
most frequently and in greatest concen-
speculate that right whales may be diving
trations to the
in the deepest part of the BOF, which
bottom of the channel to feed. Our observations is also the main shipping channel for the area.
of whales in areas of known or likely high con- Tagged whales also were exposed to ship traffic
centrations of food (WCR, multiple upwelling when 3 whales (#1140, #1146, and #1608) swam
regions, the basins between offshore banks, and through the ship traffic area off Boston, whale
deep channel areas), suggest that right whales #1146 was in the shipping lanes east of Port-
are extremely good at either identifying such land, Maine, whale #1140 swam through the
opportunities or remembering them from pre- New York shipping lanes twice, and whale
vious experience. #1421 was in the shipping lanes east of Halifax,
Nova Scotia. In contrast to the movements re-
MANAGEMENT IMPLICATIONS
corded in the BOF shipping lanes, tagged
Nearly one-third of known right whale mor- whales appeared to travel through shipping
talities are caused by collisions with ships,lanes
and in U.S. waters, rather than remain within
at least 12% of the photographed population them for extended periods. In addition, because
show scars from ship collisions (Kraus 1990, right whales move long distances, have varied
Kenney and Kraus 1993). Our data suggest movement
that patterns, and appear to use much of
right whales use areas and behave in ways the thatwestern North Atlantic coastline (Fig. 6),

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All use subject to http://about.jstor.org/terms
1404 RIGHT WHALE MOVEMENTS * Mate et al. J. Wildl. Manage. 61(4):1997

adjusting shipping lanes to reduce ship/whaleAtlantic Ocean. Pages 121-128 in R. L. Brownell,

P. B. Best and J. H. Prescott, eds. Right whales:
collisions may not be effective. Similarly, the
past and present status. Int. Whaling Comm.
seasonal and regional fishing restrictions pro-
Spec. Issue 10.
posed to minimize interactions with rightCETAP. 1982. A characterization of marine mammals
and turtles in the mid-and north Atlantic areas of
whales (Natl. Mar. Fish. Serv. 1991) also may
be ineffective because right whales tagged inthe U.S. outer continental shelf. Final Rep. Ce-
tacean and Turtle Assessment Progr. to U.S. Dep.
this study did not remain in discrete areas forInt. Contract AA551-CT8-48. 570pp.
well-defined periods of time. Changes in fishing
CRAIGHEAD, D. J., AND J. J. CRAIGHEAD. 1987.
methods or fishing gear modifications may re- Tracking caribou using satellite telemetry. Natl.
duce whale entrapments, but if such changesGeogr. Res. 3:462-479.
are to be successful, more data are neededCRONE,
to M. J., S. D. KRAUS. 1990. Right whales (Eu-
balaena glacialis) in the western North Atlantic:
define when, where, and why these entrap-
a catalog of identified individuals. New England
ments occur. Seasonal speed limits in the high-
Aquarium, Boston, Mass. 208pp.
DELONG, R. L., AND B. S. STEWART. 1991. Diving
use right whale areas and educational programs
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EVANS, W. E. 1974. Radio-telemetric studies of two
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FANCY, S. G., L. F. PANK, D. C. DOUGLAS, C. H.
time (up to 42 days). Even with this small sam-
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more effective attachments, it should be possi-
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