MARINE MAMMAL SCIENCE, l6(4):811-8 14 (October 2000)
0 2000 by the Society for Marine Mammalogy

DIVING BEHAVIOR AND SWIMMING SPEED OF A
FREE-RANGING HARBOR PORPOISE,PHOCOENA PHOCOENA
Our understanding of cetacean diving behavior has recently increased (Westgate et al. 1995, Heide-Jgrgensen and Dietz 1995, Davis et al. 1996). However, there are few reports which measured swim speed of free-ranging cetaceans (Hanson and Baird 1998). Measurement of swim speed during diving
helps to understand the dive function (Boyd et al. 1995, Crocker et al. 1997)
and the swimming effort or energetics (Feldkamp 1987). We report here thefirst record of the diving behavior and swimming speed of a free-ranging
harbor porpoise.
The set of instruments to measure the diving behavior consisted of a micro
data logger, a time-scheduled releaser, a float, and an Argos transmitter (PTT).
The micro data logger (UWE-200PDT, developed by National Institute ofPolar Research and Little Leonardo Co., Tokyo; 20 mm in diameter, 90 mm
long, 42 g in weight, 2 MB memory size) was set to record depth, numberof rotations of the propeller, and water temperature at intervals of one second.
The propeller was placed in front of the logger and rotated in response towater flow. Swim speed was calculated from the number of rotations of thepropeller according to a linear relationship. The logger could detect speeds
faster than 0.5 m s e c 1 . Depth and temperature sensors were placed on opposite sides of the logger. Absolute accuracy of the logger was 0.5 m in depth,O.lÂ° in temperature, and 0.5 m s e c 1 in speed; resolution was 0.05 m,
O.OlÂ°Cand 0.04 m s e c 1 , respectively. The releaser was set to activate one
day after the animal was released. The float had a mass of 195 g in air, andthe dimensions were 170 X 90 X 40 mm. The PTT (T-2038, TOYOCOM
Co., Ltd., Tokyo, 120 X 35 X 15 mm, 116 g) was implanted into the float
except for its antenna. This was left exposed in order to facilitate transmission
after the instruments were released from the porpoise. The total weight of theinstruments was 486 g in air and - 50 g in sea water. Details of instruments
and attachment techniques were described in Otani et al. (1998).
The study was carried out at Usujiri (41 Â °57'N 140Â°58'E)Hokkaido, Japan, in July 1997. A female harbor porpoise was accidentally caught in a setnet off the coast of Usujiri (134 cm in body length and 37 kg in body mass).

MARINE MAMMAL SCIENCE, VOL. 16, N O . 4,2000

812

0:OOPM

6:OO PM

5 July 1997

0:00 AM

6:00 AMTime of day

Figure 1. Total records of dive depth (upper) and swim speed (>0.5 m s e c 1 )(bottom) for female harbor porpoise over period of 22.8 h.

After the instruments were attached to the porpoise, it was released about 2km off the coast of Sawara (42'09'N, 140Â°41'E)which is located about 30
km northwest of Usujiri. The release point was within Funka Bay; the depth
was less than 100 m. After release the package was recovered using ARGOSsupplied PTT locations coupled with a board-based automatic direction-finding receiver (Gonio 400, Cubic-I Co., Ltd., Tokyo). The recorded data weredownloaded from the logger to a personal computer. Horizontal speed wascalculated from swim speed and depth change rate (vertical speed). Only divesdeeper than 0.5 m were analyzed, because the absolute accuracy of the logger
was 0.5 m.
We obtained 2,878 dives with speed data over the 22.8 h of continuousrecording from the free-ranging harbor porpoise (Table 1, Fig. 1). The harbor
porpoise dived continuously and repeatedly, as Otani et al. (1998) reported.
More than 90% of the dives were shallower than 10 m and more than 80%
were shorter than one minute in duration. The harbor porpoise swam at relatively slow speed. Its mean swim speed was 0.9 m s e c 1 during the whole
recorded period and more than 90% of records of swim speed were slower
than 1.5 m s e c 1 . The fastest speed recorded in this study was 4.3 m s e c 1 .

Table 1.

Summary statistics of diving records for female harbor porpoise.
Mean 2 SD

Dive depth (m)Dive duration (sec)
Surface time (sec)Descent speed (m s e c 1 )Ascent speed (m s e c 1 )Descent angle (deg.)
Ascent angle (deg.)

3.8 % 6.8
26.2 2 31.0
3.9 5 7.2
0.8 % 0.3
0.9 % 0.3
31 % 20
17 L 10Max

Min.
0.6
210.0
0.0
11Median813

NOTES0E

1020g 30

2 40a5060
04080 120 160
Time (S)040

80 120 160
Time (S)04080 120 160
Time (S)Figure 2. Example of dive depth (upper) and swim speed (>0.5 m s e c 1 ) (bottom)
of dives with bottom time for harbor porpoise.

Leatherwood et al. (1988) reported that harbor porpoises can swim at about
12 kn. (about 6.2 m sec-l).
Horizontal speed was significantly faster than vertical speed (Wilcoxon
signed rank test, P < 0.001). Both horizontal and vertical distance increased
during the 22.8 h of the recording period. Total horizontal distance (50.5 km)
was about 2.9 times longer than the vertical distance (17.7 km). Distancetraveled by the porpoise was estimated to be about 53 km per day. This value
was within previous records (32-57 km per day) obtained using radio tracking(Westgate et al. 1995). In that case the horizontal distance was based on thedistance calculated between two points where the radio signal was detected
and was reasoned to underestimate total distance. Our value would also be anunderestimate, because the logger does not detect speeds slower than 0.5 ms e c 1 . Even so, this is the first measurement of the gross movement of cetaceansusing such a method.
Descent speed (Spearman rank correlation test, r 2 = 0.57, P < 0.001, n =2,451), ascent speed (r2 = 0.41, P < 0.001, n = 2,311), descent angle (r2 =0.20, P < 0.001, n = 2,470), and ascent angle (r2 = 0.66, P < 0.001, n =2,294) were positively correlated with dive depth. Because the angle of shallow
dives was smaller than that of deep dives, shallow dives had a relatively larger
horizontal component.
Dives both with and without bottom time were observed in this study.
Otani et al. (1998) assumed that porpoises foraged during the bottom phaseof dives. For the dives with bottom time, swim speed at the bottom either
decreased to less than the recording limit or it alternated between increase
and decrease (0-2.0 m s e c 1 ) (Fig. 2).
Initial descent speed (r 2 = 0.27, P < 0.001, n = 2,451) and initial descentangle (r2 = 0.56, P < 0.001, n = 2,470) were also positively correlated withdive depth. Otani et al. (1998) hypothesized that harbor porpoises anticipate
the depth of their dives before initiating the dive process, because the initialdescent rate increased as the dive depth increased. However, it was unclear
whether the change of initial descent rate was caused by the change of descent
speed or descent angle. In the present study we analyzed the swim speed data
to determine whether porpoises changed the swim speed or dive angle when
diving deeply. When the tagged porpoise dived deeply, it increased both its

814

MARINE MAMMAL SCIENCE, VOL. 16, NO. 4,2000

dive speed and dive angle. It could therefore reach the bottom faster than ona slow dive or a dive with a less abrupt angle.

The authors would like to thank Dr. H. Munehara, Mr. Y. Arashida, Mr. K. Nomura,
Dr. K. Nakamura, and all the fishermen of Nomura Fishing Company and Usujiri
Fisherman's Association in Usujiri, Hokkaido for their cooperation in collecting andkeeping specimens, and Mr. R. Kawabe, Mr. S. Kaga, Mr. Y. Kaga, Mr. T. Kawamura,Mr. T. Matsui, and Mr. T. Fukuda for helping with the field work. We also thankthree anonymous referees for valuable comments on the manuscript. This work waspartly supported by a grant from the Sasakawa Scientific Research Grant from theJapan Science Society and a Grant-in-Aid for Scientific Research (No. 08660227) fromthe Japan Ministry of Education, Science and Culture.

BOYD,I. L., K. REIDAND R. M. BEVAN. 1995. Swimming speed and allocation of time
during the dive cycle in Antarctic fur seals. Animal Behavior 50:769-784.
CROCKER,D. E., B. J. LE BOEUFAND D. P. COSTA. 1997. Drift diving in female northernelephant seals: Implications for food processing. Canadian Journal of Zoology 75:27-39.
DAVIS, R. W., G. A. J. W ORTHY, G. WURSIG,S. K. LYNNAND F. I. TOWNSEND. 1996.
Diving behavior and at-sea movements of an Atlantic spotted dolphin in the Gulfof Mexico. Marine Mammal Science 12:569-58 1.
FELDKAMP, S. D. 1987. Swimming in the California sea lion: Morphometrics, drag andenergetics. Journal of Experimental Biology 131:117-1 35.
HANSON, M. B., AND R. W. BAIRD. 1998. Dall's porpoise reactions to tagging attempts
using a remotely-deployed suction-cup tag. Marine Technology Society Journal
32:18-23.
HEIDE-J~RGENSEN,
M. P., AND R. DIETZ.1995. Some characteristics of narwhal, Monodon
monoceros, diving behavior in Baffin Bay. Canadian Journal of Zoology 73:2 1202132.
LEATHERWOOD, S., R. R. REEVES, W. F. PERRINAND W. E. EVANS. 1988. Whales,dolphins, and porpoises of the Eastern North Pacific and adjacent arctic waters.
A guide to their identification. Dover Publications, New York, NY.
OTANI, S., Y. NAITO, A. KAWAMURA,
M. KAWASAKI, S. NISHIWAKI AND A. KATO. 1998.
Diving behavior and performance of harbor porpoise, Pbocoena phocoena, in Funka
Bay, Hokkaido, Japan. Marine Mammal Science 14209-220.
WESTGATE, A. J., A. J. R EAD, P. GERGGREN, H. N. KOOPMAN AND D. E. GASKIN.1995.
Diving behaviour of harbour porpoises, Pbocoena pbocoena. Canadian Journal ofFisheries and Aquatic Sciences 5 2:1064-107 3.

SEIJIOTANI,~Department of Polar Science, The Graduate University for Advanced Studies, National Institute of Polar Research, Itabashi, Tokyo 1738515, Japan; YASUHIKONAITO and AKIKO KATO, National Institute of Polar
Research, Itabashi, Tokyo 173-8515, Japan; AKITO K AWAMURA , Departmentof Bioresources, Faculty of Bioresources, Mie University, Tsu, Mie 5 14-8507,
Japan.
Current address: The Institute of Cetacean Research, Tokyo Suisan Building 4-18, Toyomicho, Chuo, Tokyo 104-0055, Japan; e-mail: ohtani@i-cetacean-r.or.jp. Received 11 May 1999.
Accepted 8 February 2000.