Marine Biology 114, 175-183 (1992)
Marine
Depth and temperature of the blue marlin, Makaira nigricans,
observed by acoustic telemetry
B. A. Block 1, D. T. Booth 1 and E G. Carey 2
1 The University of Chicago, Department of Organismal Biology and Anatomy, 1025 East 57th Street, Chicaog, Illinois 60637, USA
2 Woods Hole Oceanographic Institution, Department of Biology, Woods Hole, Massachusetts 02543, USA
Date of final manuscript acceptance: July 1, 1992. Communicated by J. Grassle, New Brunswick
Abstract. Multiplex acoustic transmitters were used to
monitor the depth, swimming speeds, body temperature
and water temperature preference of six blue marlin,
Makaira nigricans (Lacrprde), near the Hawaiian islands
in July and August 1989. The blue marlin ranged in size
from 60 to 220 kg and were tracked for 1 to 5 d. All of the
fish moved away from the point of capture and were
followed up to 253 km from the island, of Hawaii. The
blue marlin tracked remained in the top 200 m of the
water column, spending half the time in the upper 10 m,
and rarely ventured below the thermocline. In the near-
surface waters the temperature was uniformly warm (25
to 27~ The coldest water temperature, 17~ was en-
countered on the deepest descent recorded (209 m).
D e p t h changes occurred rapidly and excursions below
10 m were usually less than 60 rain in duration. Muscle
temperature was similar to water temperature except for
a 2~ elevation in muscle temperature observed at the
beginning of tracking one individual. This initial rise in
body temperature was associated with the anaerobic
muscle activity during capture and is an indication of the
physiological stress involved in capture.
Introduction
Blue marlin, Makaira nigricans, inhabit the open ocean
and are among the largest teleosts. They range over trop-
ical and warm temperate waters of the Pacific, Atlantic,
and Indian Oceans and make seasonal migrations which
may carry them thousands of miles across ocean basins
(Squire 1974). As apex predators they feed on a wide
assortment of organisms including other large epipelagic
fishes (Scombridae, Istiophoridae, Carangidae), small
fishes (Acanthuridae, Mullidae, Tetraodontidae), Om-
mastrephid squids, octopods and decapods (Baker 1966,
Brock 1984). Blue martin are sought by anglers for their
spectacular acrobatics when caught on rod and reel but
they are also a valuable species in the commercial pelagic
longline fishery. Recently, with the rapid world-wide de-
BiOlOgy
9 Sprlngor-Ver[ag 1992
pletion of blue marlin stocks, researchers have focused
attention on the biology and conservation of this species.
Laboratory-based research on blue martin and other is-
tiophorids (marlins, spearfish, sailfish), has rapidly in-
creased in the past decade (for reviews see Davie 1990,
Stroud 1990) and has revealed numerous physiological
and morphological specializations for a nomadic lifestyle
in the open sea. Acoustic telemetry studies of free ranging
marlin in the open ocean provide insights as to how mar-
[in may actually use such specializations.
Acoustic telemetry has successfully been used to study
free-swimming billfishes (Yuen et al. 1974, Jolley and
Irby 1979, Carey and Robison 1981, Carey 1990, Holland
et al. 1990, Holts and Bedford 1990). Holland et al.
(1990) reported on tracking six blue martin during the
summer of 1989 in the same area of the Hawaiian Islands
as in the present report. Results from 1 to 2 d tracks of
martin fitted with depth transmitters showed that the fish
preferred the warm mixed layer. They concluded that
blue martin had a daily depth cycle, swimming deeper
during the day while remaining closer to the surface at
hight, and that the tag-release procedure had little mor-
tality associated with it. In the present study we present
further blue martin tracks of longer duration which in-
clude data from temperature, depth and swimming speed
sensors. A thorough analysis of the speedometer data has
appeared in a previous publication (Block et al. 1992).
Methods
Transmitters were attached to six Makaira nigricans (Lacrp&te)
caught off the Kona coast of Hawaii during July and August 1989.
Five fish were released within a 4 km radius of Kealakekua Bay. A
sixth fish was released outside the harbor in Kailua-Kona. Depth,
body temperature, and water temperature were monitored from
marlins 1 and 2, depth only from marlin 3, and depth, speed and
water temperature from marlins 4, 5 and 6. The fish were caught on
rod and reel using a surface trolled artificial lure. Professional char-
ter boat captains were instrumental in the tag and release of marlin
1, 2 and 6; marlin 3, 4 and 5 were captured and released with the aid
of anglers participating in the Kona Hawaiian and Hawaiian Inter-
national Billfish Tournaments.
176
Multiplex transmitters were 18.0 cm long, 4.0cm wide, and
2.2 crn thick. They weighed 190 g in air and 78 g in seawater and
were capable of withstanding hydrostatic pressures to depths of
2000 m. A thermistor mounted in an aluminium stub on the trans-
mitter measured water temperature. Muscle temperature was mea-
sured by a thermistor on a harpoon dart in the epaxial muscle.
Pressure sensors (Keller PA-2-20) were used to measure depth with
an accuracy of + 8 m over a 400 m range and were insensitive to
changes in temperature. Sensors were multiplexed with a 4521 crys-
tal oscillator clock and 4020 octal divider controlling 4066 analog
switches. A dual slope integrator encoded information from the
sensors as a pulse rate that varied between 0.3 and 2.5 Hz and was
proportional to pressure or temperataa-e. The pulses were 20 ms
bursts of 32 768 kHz ultrasound from a class D output stage that
drove a TCD5 transducer through a transformer and inductor. The
circuitry was insensitive to variations in temperature and battery
voltage. The signal could be detected 1 to 2 km away depending
upon sea state, and battery life was approximately 6 d. Marlin 3 was
fitted with a commercially available depth transmitter (VEMCO
V4P4).
Data was time-multiplexed by the transmitter clock so that 32 s
of information on depth, 16 s of water temperature and 16 s of body
temperature were broadcast in sequence over a continuously repeat-
ing 64-s cycle. A similar crystal clock on shipboard stayed syn-
chronous with the transmitter clock and allowed the computer to
identify the parameter currently being broadcast. During data anal-
ysis, a mean was calculated for the parameters of interest for each
data period within a 64-s cycIe.
The acoustic signal was detected with two independent receiving
systems aboard the tracking vessel. A narrowly directional hydro-
phone was mounted on a shaft in a sonar well near the bow of the
ship. We followed the fish by rotating the hydrophone to the direc-
tion of the strongest signal and steering the boat on that course. A
second set of hydrophones consisted of a square array of four
broadly directional elements mounted on a tow body suspended
from the bow at a depth of ca. 7 m on a multiconductor electrome-
chauical cane. The element receiving the best signal was switched
to a PC computer-based data collection system which displayed the
various parameters on screen, produced a paper printout for ship-
board review, and stored data on disk for later analysis. Vessel
position was determined at approximately hourly intervals by satel-
lite navigation or a combination of Loran-C and radar and was
used to approximate the fish's course. Temperature prof'des through
the water column were obtained at 4 to 6 h intervals using expend-
able bathythermograph probes (Sippican %6 and T-10).
Length of time between hooking a fish and attaching the
transmitter varied and was an important factor in determining the
condition of marlin at release. A stzinless steel dart either
1.00 cmx 3.50 em x 0.65 mm (marlin 1, 4, 5) or t.20 x 5.50 x 0.65 cm
(marlin 2, 3, 6) in size was attached to the transmitter by a loop of
either monofilament or stranded steel leader. Transmitters were
strapped onto a tagging pole with rubber bands and the dart insert-
ed so that it toggled 4 to 5 em deep into the expaxial muscle directly
below the first dorsal fm. An adjustable stop on the tagging pole
limited dart penetration. Weights of fish were estimated by the
captains of sport fishing boats. Fish estimated to be larger than
140 kg were assumed to be females (Hopper 1990).
Results
Vertical m o v e m e n t s
Six blue m a r l i n r a n g i n g in size f r o m 60 to 220 k g were
t r a c k e d for p e r i o d s o f 25 to 120 h d u r i n g the m o n t h s o f
July a n d A u g u s t in 1989. F i s h size, d u r a t i o n o f track, a n d
t o t a l d i s t a n c e travelled are s u m m a r i z e d in Table 1. M a r -
lins 1, 3 a n d 6 were in excellent c o n d i t i o n (Table 1) a n d
B.A. Block et al.: Blue marlin depth and temperature
Table 1. Makaira nigricans. Summary of tracks
Marlin Esti- Date Condition Dura- Distance
mated tagged tion travelled
weight
0~g) 0a) (~)
1 220 10 July 1 9 8 9 Excellent 67 253
2 80 16 July 1 9 8 9 Good 42 43
3 70 23 July 1 9 8 9 Excellent 71 84
4 70 31 July 1989 Fair 72" 71
5 60 4 August 1989 Fair 25 18
6 125 6 August 1989 Excellent 120 100
9 Blue marlin was attacked by a pelagic whitetip shark 15 h into
track. The shark was tracked for 57 h
s w a m r a p i d l y a w a y f r o m the p o i n t o f release. T h e o t h e r
three m a r l i n h a d a m o r e stressful time d u r i n g the c a p t u r e
a n d t a g g i n g process. D e p t h r e c o r d s t y p i c a l o f m a r l i n
m o v e m e n t s are p r e s e n t e d in Fig. 1 to 3. A l l m a r l i n
t r a c k e d , except m a r l i n 4 e n t e r e d into a s t e r e o t y p i c "re-
c o v e r y b e h a v i o r " i m m e d i a t e l y after t a g g i n g (see Figs. 2
a n d 3). T h e s e fish d e s c e n d e d to b e t w e e n 50 a n d 120 m
a n d b e g a n a 4 to 6 h p e r i o d in w h i c h t h e y s w a m a t a t
g r e a t e r d e p t h a n d g r e a t e r speed t h a n for the r e m a i n d e r o f
the track.
M a r l i n s p e n t m o s t o f their time w i t h i n 10 m o f the
surface (Fig. 4). H o w e v e r , r a p i d descents a n d ascents to
v a r i a b l e d e p t h s w i t h i n the t o p 130 m o f the w a t e r c o l u m n
were c o m m o n . D e s c e n t s f r o m the n e a r surface waters
were u s u a l l y o f s h o r t d u r a t i o n ( < 60 min), a n d fish o f t e n
r e t u r n e d to the s a m e d e p t h in the w a t e r c o l u m n f r o m
w h i c h the descent was m a d e . D e s c e n t s were i n v a r i a b l y
a c c o m p a n i e d b y a n increase in s w i m m i n g speed (Block
et al. 1992), b u t s l o w e d u p o n r e a c h i n g the n e a r surface
w a t e r s again.
Blue m a r l i n h a v e a m a r k e d preference for the w a r m
m i x e d layer a b o v e the t h e r m o c l i n e , s p e n d i n g h a l f o f their
time in the u p p e r 10 m o f the w a t e r c o l u m n (Fig. 4) where
w a t e r t e m p e r a t u r e v a r i e d b e t w e e n 26 a n d 27 ~ M a r l i n 1
m a d e the d e e p e s t dive d u r i n g o u r 340 h o f r e c o r d i n g s
r e a c h i n g a d e p t h o f 209 m (Fig. 1). M a r l i n 3 m a d e a sim-
ilar short, r a p i d dive b e l o w the t h e r m o c l i n e to a d e p t h o f
180 m (Fig. 3). T h e 17 ~ w a t e r t e m p e r a t u r e e x p e r i e n c e d
d u r i n g these dives was the lowest r e c o r d e d . T h e w a r m e s t
w a t e r t e m p e r a t u r e r e c o r d e d , 27.2 ~ was e n c o u n t e r e d on
the surface d u r i n g t r a c k s o f m a r l i n s 4, 5, a n d 6.
Muscle temperature
T h e r m i s t o r s were i m p l a n t e d in the e p a x i a l m u s c u l a t u r e (a
m o s a i c o f fast oxidative, fast glycolytic a n d slow o x i d a -
tive fibers) o f two m a r l i n s , a n d 6 d o f m u s c l e t e m p e r a t u r e
d a t a were o b t a i n e d . T h e t h e r m i s t o r in m a r l i n 1 was im-
p l a n t e d a p p r o x i m a t e l y 4 c m i n t o the m u s c u l a t u r e w i t h a
small d a r t (1.00 • 3.5 • 0.65 cm). M u s c l e t e m p e r a t u r e o f
m a r l i n 1 was 0.5 ~ h i g h e r t h a n w a t e r t e m p e r a t u r e w h e n
we first b e g a n r e c o r d i n g it 4 h after release ( n o t shown),
a n d t h e r e was a lag b e t w e e n muscle a n d w a t e r t e m p e r a -
ture d u r i n g descents a n d ascents. M u s c l e t e m p e r a t u r e
B.A. Block et al.: Blue marlin depth and temperature 177

Makaira nlgrlcans 1. July 10-13, 1989

I
o

1 oo ~
'-'-- j
/
15o

200" , , | i = ! i i | | | = 'C ! | i 'l i '"| , i ,

6 12 18 24 6

O-
o

5O-

lOO-
g --
,- 150-
t'~ ~

r~ 200- , . . . . I , ~ , ' , I ',

6 12 18 24 6

I I I

gig. 1. Makaira nigricans. Depth

record for marlin I superimposed on a
1~ isotherm plot drawn from expend-
able bathythermographcasts. Dark
bands indicate night. Equipment mal-
function prevented data collectiondur-
200[ . . . . . ,' ~" , , , ~ , , '. . . . I . . . . ' ' I
ing initial 4 h of this track
6 12 18 24 6

Time (hours)

approached water temperature 12 h into the track and (Fig. 6). The fish then followed consistent headings for
the time lag disappeared by Day 2, presumably because the remainder of their tracks. Marlin 1 headed NNE
the probe was slipping out of the muscle and moving through the rough waters of the Alenuihaha Channel
closer to the body surface. between the Islands of Hawaii and Maui, traveling
The probe in marlin 2 was implanted with a larger dart 253 km in 67 h. Marlin 2 swam 43 km to the NW in 42 h.
(1.20 x 5.50 x 0.65 cm) at least 5 cm into the musculature. Marlin 3 moved south past the southern tip of the island
Muscle temperature of marlin 2 was 2.1 ~ greater than of Hawaii and then headed east, swimming 84 km in 71 h.
water temperature immediately after tagging, but gradu- Marlin 5 stopped moving offshore eight hours into the
ally decreased and approached ambient water tempera- track and only moved a short distance in the following
ture 5 h after release (Fig. 5). No other activity during the 16 h. Marlin 6 followed a similar path to martin 3 but
course of the track caused a similar rise in muscle temper- continued south after passing the southern tip of Hawaii.
ature. Drift of the thermistor probe caused muscle tem- This last fish spent many hours stationary or moving
perature to appear 0.1 ~ colder than water temperature slowly on the surface and only moved 100 km in 120 h.
during the remainder of the track.

Course Individual variation

After release all marlin initially moved offshore (west- The following is a brief description of each of the blue
ward) then changed course to either north or south marlin tracked. Martin 1, (220 kg, female), was captured
178 B, A. Block et al,: Blue marlin depth and temperature

Mak~ira nlgrlcans 3. July 23-26, 1989

O-

50-

100-

150-

200-
12 18 24 6 12

0-

1oo
~150
D
2
12 18 24 6 12

5O-

100-
. Fig. 2. Makaira nigricans. Depth
record of marlin 3 with 1 ~ isotherms.
150- Recovery period lasted 5 h. A cliel
change in diving pattern was most ap-
200 . . . . . t * t t ] i t i t I t t i
parent during the f'mal day
12 ll8 ' 24- 6
Time (hours)

and released in less than 15 min. No obvious diel behav-
ior was apparent, although there was a trend toward
more frequent and deeper dives during daytime than at
night. An increase in speed of marlin 1 is apparent from
the separation of the hourly tics in Fig. 6. Numerous
other fishermen reported a brisk northerly running cur-
rent in the region during the period of the track. The
current probably accounts for this fish having the highest
average track speed, 3.8 k m h -1. This track ended at
Hour 67 when the transmitter went to the bottom sinking
at a rate of 50 cm s- 1.
Marlin 2 (80 kg, sex unknown) spent 15 rain on line
and was harpooned with a larger, 5.5 cm long metal dart.
The capture and tagging may have injured this fish which
at first made only shallow descents, but began making
deeper descents late on the second day of the track. There
was no day-night difference in the frequency of dives. The
track was terminated due to bad weather in the Alenuiha-
ha Channel.
Marlin 3 (70 kg, sex unknown) struggled for 45 rain on
light tackle during capture, leaping five times before be-
ing brought along side the boat. The fish appeared un-
harmed, however, and swam away vigorously upon re-
lease. After the recovery period, this fish assumed a pat-
tern of repeated descents of short duration. There was a
distinct diel trend in depth preference, with 48% of night-
time hours but only 18% of daytime hours spent near the
surface (0 to 10 m). This track ended when the transmit-
ter sank to the bottom at Hour 71, sinking at a rate of
6 9 c m s -1
Marlin 4 (70 kg, sex unknown) was captured after an
18 rain fight and appeared fatigued when tagged. It was
towed slowly behind the boat to irrigate its gills to assist
its recovery before release. This marlin did not exhibit the
stereotypic recovery behavior. Its poor condition was
probably a factor in causing its death by shark attack
15 h after tagging (Fig. 7).
Marlin 5 (60 kg, sex unknown) slapped the side of the
boat several times during tagging. It appeared to be in
fair condition on release although its speed fluctuated
erratically. This fish assumed a westerly course and spent
most of its time in the upper 10 m, both day and night.
B.A. Block et al.: Blue marlin depth and temperature 179

Mokoiro nigricons 6. August 6-11, 1989

9 II
o
9

50-

100-

150- =~
lJ

200 i" i" ! , , , ", , , , I , . . . .

6 72 . . . . 1'8 24

O.
--~ . ~ ~ ' 7, n ~ r, ~ , ~ , ~ nr ~

100-

6 12 18 24 6

! m II I
0

5O-
r

E 100-
- - S
150-

2OO | l l l l ; 2 1 | | J I l | l l l l l l l l l l
6 18 24

0 - . II

50-

100-

150-

200 i , , , i , , ~ , ,
6 12 18 24

Fig. 3. Makaira nigricans. Depth

record of marlin 6 with 1 ~
isotherms, Recovery period lasted
10 6 h. This is the only fish that
showed a clear diurnal cycle of
behavior with a strong preference
150 t for the surface during the clay,
Activity at night was punctuated
200| ", . . . . = , , , , , ! i | i" i I = = i by frequent dives
6 12 8 24 6
Time (hours)
180 B.A. Block et al.: Blue marlin depth and temperature

The track ended abruptly at Hour 26 after several min- Discussion

utes of erratic activity with the fish reaching speeds in
excess of 270 cm s- 1, followed by the transmitter sinking
to the bottom at a speed of 46 cm s- 1. Diving and behavior of Makaira nigricans
Marlin 6 (115 kg, sex unknown) was released in excel-
lent condition after a 15 min struggle. In contrast to the In Hawaiian waters blue marlin prefer to remain in warm
other fish, a clear diel pattern of diving behavior was 22 to 27 ~ surface waters and spend most of their time
observed in this fish. Day time hours were spent almost above the thermocline in the 26 to 27~ mixed layer
entirely at the surface at which time forward movement which is 70 to 100 m deep during the summer months.
often ceased. Around sunset on four of the five days that While they are capable of ranging through the thermo-
this marlin was tracked it made a 80 to 100 m descent of cline, they rarely to so. The results obtained in our study
2 to 4 h duration before ascending again. Frequent verti- are consistent with those of Holland et al. (1990), and
cal movements occurred for remainder of the night together the two studies provide strong evidence of a
(Fig. 3). An increase in swim speed was correlated with preference of blue martin for the warm surface waters
the increase in vertical movements (Block et al. 1992). while off the west coast of Hawaii. Other Istiophorids
This track was intentionally broken off after 5 d. (striped and white marlins, sailfish) also prefer to stay
close to the surface. Seven striped marlin, Tetrapturus
audax, tracked offthe Californian coast spent 85% of the
time in the upper 50 m where the water temperature
ranged between 11 and 21 ~ (Holts and Bedford 1990).
Blue and striped marlin both appear to occupy the near-
surface habitat, but the surface waters off the Southern
60 California coast are colder than in Hawaii. Thus, striped
marlin experience colder temperatures, range through the
50 thermocline, and may, in some locations, forage deeper
0~
than the blue marlin. Blue martin in other locations may
_E 40 exhibit a broader temperature range, however, a 12-h
I--
track of a blue marlin caught off the Bahamas showed a
r
similar preference for the warm mixed layer (Block 1990).
30 Blue martin off North Carolina are commonly caught in
e- waters with surface temperatures ranging from 24.4 to
,,
r 20 27.2~ (Block unpublished data). Two sailfish, Istiopho-
rus platypterus, tracked in the Straits of Florida also re-
10 mained in 22 to 26 ~ water within 30 m of the surface,
although in this case a bottom depth of 50 m limited
vertical movement (Jolley and Irby 1979).
0-10 11-20 ~-~11 31-40 r ~ 11-70 ~ 81.40 91-100101-110111,1~ M20
Four of the marlin in the present study showed no
Depth(meters) significant difference in diel depth preferences. While
Holland et al. (1990) state that marlin prefer shallower
depths at night, we find their data to be similar to our
Hg. 4. Makaira nigricans. Summary of depth data for six martin
indicating proportion of time spent at a given depth. Cumulative own, with considerable individual variation in diel be-
data from all marlin were divided into 10-m bins and the % time havior. For example, the descents made by marlin 3 on
spent at each depth calculated Days 2 and 3 were to greater depths during the day than

Maka|ra nlgricans 2. duly 16-17, 1989

3O
Fig. 5. Makaira nigricans. Water
and body temperature record for
marlin 2. A 2.1 ~ elevation in
o
o 28- muscle temperature evident imme-
diately after capture. Muscle tem-
== 2e- perature (M) stabilized with water
temperature (W) 5 h after release.
a) 24- A drift in calibration of the
thermistor was responsible for
I.-
E
22-" muscle temperature appearing
0.l ~ lower than water tempera-
i i ture
20
6 ';2 ' ' ' ' ' ~ 8 ' ' ' ' ' 1 ' ' ' 2 4 6

Time (hours)
B.A. Block et al.: Blue marlin depth and temperature 181

at night (Fig. 2). In contrast, marlin 6 clearly spent most

1 ' ' O' of its time on the surface during the day, descending more
frequently and to greater depths at night (Fig. 3). Al-
though some martin may show a change in depth with
day and night, there is much individual variation.
21 The relatively shallow depth and short duration of
descents seen in telemetry studies of istiophorids are in
sharp contrast to the large diel vertical migrations ob-
served in the sister group to all istiophorid bilIfishes, the
swordfish Xiphias gladius (Carey and Robison 1981,
Carey 1990). Although swordfish and other billfishes
share a common ancestry and have many convergent fea-
tures in external morphology, the behavior of swordfish
is very different from other bill_fishes (Block 1990). In the
open sea, swordfish make large vertical excursions, com-
ing close to the surface at night and going as deep as
" "i 600 m during the day. During these vertical dives, sword-
20 ~
fish encounter water temperature changes as great as
d

!,". .~, I 19 ~ in 30 to 60 min, but their brain temperature is reg-

ulated by a thermogenic organ, so that changes in brain
temperature are relatively small compared to changes in
e;,,[+, Hawaii
water temperature (Carey 1982, 1990). The blue marlin
also has a heater organ with high oxidative capacity lo-
>.i
cated close to the brain (Tullis et al. 1991). Although the
oxidative capacity of the blue marlin and swordfish
heater tissues are similar, there is much less of the heater
phenotype expressed in blue martin extraocular muscle as
19 ~ compared to the swordfish. This correlates well with the
thermal ecology shown here with very small excursions in
J I
temperature for the blue marlin ( < 5 ~ It is possible
that in other regions of their large range, or at other
seasons of the year, blue martin may enter colder water
I , | I where thermoregulation of the brain and eyes becomes
157 ~ 156 ~ ' ~ 155 ~
more important.
Fig. 6. Makaira nigncans. Courses taken by blue marlins 1 to 6. Our muscle telemetry data for free-swimming blue
Hourly intervals indicated by small black circles and triangles. Scale marlin gives no indication that these fish use behavioral
bar equals 50 km or physiological mechanisms to regulate body tempera-

loo

1
i
200 -
~

WltW TernlpO~tul~
I " ' 28

27

26
o~

~.

Fig. 7. Makaira nigricans. Shark at-

E" 1 0 0 - tack on marlin 4. Arrow indicates
presumed time of shark attack at
50-
05:00 hrs. See "Discussion - Shark
0 , attack" for details
12 .... . . . . . .... '.d . . . . . . . . . . 1'8 . . . . .
182
ture. The absence of a significant band of internalized
red muscle (Davie 1990), the dispersal of the slow oxida-
tive fibers throughout the myomeres, and the lack of a
vascular heat exchanger within the circulatory system
suggest that marlin, unlike swordfish, have little ability to
maintain an elevated muscle temperature. Muscle tem-
peratures in excess of 3 ~ above water temperature were
only recorded immediately after the intense period of
anaerobic activity during capture on hook and line. At all
other times muscle temperature was at near equilibrium
with water temperature. Because blue marlin experience
temperatures of 17 to 27~ it may not be necessary for
them to employ the thermoregulatory strategies used by
swordfish to elevate their muscle temperatures while for-
aging in cold (6~ waters beneath the thermocline
(Carey 1990). Our telemetry data corroborate the/n vitro
findings of Johnston and Salamonski (1984) who found
blue marlin contractile proteins function independently
of temperature over the range of 15 to 25 ~
Stress of capture, tag and release
One of the disadvantages of working with fish as large as
martin is the unavoidable stress associated with capture
and tagging. Studies on blue marlin (Holland et al. 1990,
present study) and striped marlin (Holts and Bedford
1990) indicate that immediately after tagging there is of-
ten a behavioral response in which the fish descend and
remain at depth for many hours. Holland et al. (1990)
suggested that this "recovery period" was a response to
the stress of capture. A period of recovery was evident in
most of the blue marlin tracked in both studies (nine out
of 11 records examined). It emphasizes the need to track
fish long enough for activity to stabilize and to avoid
conclusions based on behavior during the first few hours
of a track. For instance, during recovery behavior blue
marlin swim at an elevated speed (approximately
100 em s- 1) for up to 6 h (Block et al. 1992).
The recovery period probably represents a physiologi-
eal response to the severe anaerobic debt acquired during
the initial fight on hook and line (Wood et al. 1983, Hol-
land et al. 1990). Blood lactate concentrations are tightly
correlated to fight time. Blood obtained from Pacific blue
marlin captured under similar conditions to those used in
this study displayed a pronounced acidosis (Dobson et al.
1986). The rise in body temperature of marlin 2 during
capture and tagging could be caused by intense activity
during such a struggle. Because such recovery lasted for
several hours and thus can be characterized as sustained
swimming, it must have been powered primarily by aero-
bic metabolism. Increased swimming speed during recov-
cry would increase ram ventilation of the gills facilitating
oxygen transfer to the blood. Increased oxygen transfer is
needed to meet higher than normal oxygen demand
caused by metabolism of accumulated lactate. Variation
in length of the recovery period between individual mar-
lin presumably relates to the size of the anaerobic debt
(and hence amount of lactate accumulated) incurred dur-
ing capture.
B.A. Block et al.: Blue marlin depth and temperature
Shark attack
A shark attacked marlin 4 15 h after tracking began.
Fig. 7 presents records of depth, water temperature and
swimming speed immediately before and after the attack.
At 05:10 hrs a rapid acceleration occurred immediately
before the speedometer signal stopped permanently. A
sudden 0.5 ~ increase at the water temperature probe
occurred at this time although there was no change in
depth. From that time on water temperature lagged no-
ticeably behind depth changes, and changes in water tem-
perature were much smoother indicating a thermal
damping effect. A pelagic whitetip shark, Carcharinus
limbatus, was observed multiple times on the surface
when the transmitter was indicating zero depth and the
signal came from the direction of the shark. Two days
after the event it became obvious to us that we had been
tracking a shark. At this time, two divers determined that
the marlin was not swimming beneath the shark. The
abrupt stop of the speedometer, the slight rise in trans-
mitter temperature and the time lag in the water temper-
ature probe would be expected for a transmitter that had
been swallowed and entered the shark's stomach.
Marlin 4 needed to be revived by slowly towing behind
the boat before release and it did not display the stereo-
typic recovery behavior observed in the other marlin. Its
poor condition probably contributed to the shark attack.
A shark attack on an ultrasonically tagged sailfish has
been reported previously (Jolley and Irby 1979). Similar-
ly, Yuen (1974) suggested that shark attack may have
been reponsible for the high mortality in the first at-
tempts to study the movements of blue marlin by teleme-
try. In our observations, three transmitters sank to the
bottom resulting in termination of the tracks. The events
surrounding the transmitters falling to the bottom re-
main uncertain, but the falling speeds of the transmitters
were similar (marlin 3 had a heavier transmitter hence the
faster rate of falling was not anomalous). The body tem-
perature data (not shown) indicate that the transmitter in
marlin 1 probably fell out, but shark attacks cannot en-
tirely be ruled out as the termination events for marlins 3
and 5. In some fishing areas a freshly released marlin may
be in danger of shark attack, particularly if it has strug-
gled violently during capture and incurred a severe meta-
boric acidosis. Techniques that shorten struggle and min-
imize trauma will aid in survival of released fish.
Marlin course
The movements of marlin in this study were very similar
to those reported by Yuen et al. (1974) and Holland et al.
(1990). None of the blue martin that have been studied
made any of the cyclical inshore-offshore movements
that have been reported for swordfish (Carey and Ro-
bison 1981, Carey 1990) or yellowfin tuna off Hawaii
(Holland et al. 1990). Because of the sustained move-
merits away from the island after tagging, it appears that
these fish are itinerant visitors following migratory routes
to more distant places and are not part of a resident
population. Recent genetic studies indicate that blue
B.A. Block et al.: Blue marlin depth and temperature
marlins f r o m the Pacific basin have a single p r e d o m i n a n t
c y t o c h r o m e b D N A h a p l o t y p e (Finnerty and Block
1992). This is consistent with o u r finding that martin
show no residential b e h a v i o r a r o u n d the island o f
Hawaii, and with tag and release studies that indicate
long distance m o v e m e n t s (Witzell and Scott 1990). Re-
cent recaptures o f blue martin off Hawaii indicate move-
ments f r o m Hawaii to the S o u t h Pacific and f r o m Mexico
to Hawaii, j o u r n e y s o f over 4000 kilometers. A l t h o u g h
martin are travelling at a p p a r e n t l y low speeds (2 to 3 k m
h -1, Block et al. 1992)'these fish are continuous swim-
mers and cover relatively large distances, 40 to 70 km in
a day.
Substantial declines in the populations o f billfishes
have occurred worldwide due in large p a r t to b y c a t c h
( h o o k i n g o f non-target species) by tuna and swordfish
longhnes and drift net operations. International m a n a g e -
m e n t strategies for blue marlin are p o o r l y developed and
are complicated by the m i g r a t o r y nature o f the species
and the resulting multinational fishing pressure. The spe-
cies is considered threatened in the Atlantic, and p o p u l a -
tions estimates f r o m the Pacific Ocean indicate substan-
tial declines in n u m b e r s over the past two decades
(Stroud 1990). O u r study, and the similar study o f Hol-
land et al. 1990, provide striking evidence that blue mar-
tin spend over 50% o f the time within 10 m o f the surface.
Thus a n y drift nets set in the top o f the water c o l u m n will
inadvertently target the blue marlin. M a n a g e m e n t strate-
gies for blue martin should consider their preference for
the surface and examine closely fishing practices (e.g. gill
nets) that indirectly target this species.
Acknowledgements. This work was supported by a grant from the
Billfish Foundation and by grants DCB-8958225 to B.A.B. and
OCE-8811421 to F.G.C. from the National Science Foundation.
The Pacfific Ocean Research Foundation and the Hawaiian Big
Game Fishing Club also provided support. This work would not
have been possible without the skill and patience of Captains M.
Hind and S. Miyamoto of the F. V. "Heola" and Captains F. and
M. Rice who provided the opportunity to tag blue marlins 1, 2 and
6 aboard the E V. "Ihu Nui". Tagging of marlin was also made
possibleby B. Brown, B. Casey, J. Fay, T. Hanlon, B. Hoey, J. Judd,
P. Nelson, B. Skimp, I. Takahashi, G. Testa and the participants of
the 1989 Kona Hawaiian and Hawaiian International Billfish Tour-
n~ments. Data processing was expedited by the careful work of K.
Mazurkiewiez and S. Conova. Many others helped to bring this
project to fruition and we are indebted to the following for their
effort and encouragement: O. Brazier, O. and J. Butler, P. Fithian,
J. Long, and W. Rockefeller.
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