JPRS ID: 8948 USSR REPORT POLITICAL AND SOCIOLOGICAL AFFAIRS

• Document Type: CREST
• Collection: JPRS
• Document Number (FOIA) /ESDN (CREST): CIA-RDP82-00850R000200020050-2
• Release Decision: RIF
• Original Classification: U
• Document Page Count: 92
• Document Creation Date: November 1, 2016
• Sequence Number: 50
• Content Type: REPORTS

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Figure 13. Schematic Depiction of the "Ve~tical" and "Horizontal"
Carousels of the Bottlenose Dolphins
35
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Figure 14. Schematic Depiction of the Catching of Fish by Bottlenose~ ~
Dolphins, Called a "Boiler" ~
The "carousel" could be closed up (the bottlenose dolphins would swim in ~
a circle, usually counter-clockwise) and not closed up. The "carousel"
type hunt could be performed by the whole herd or part of it, The school
of fish could be surrounded by several methods: ~
1, The dolphins would curve around the school of fish from one side
and would close the ring with a"carousel" (6 October, 8 October (season ~
II) Fig. 15)~. 2, The dolphins would encircle the fish from both sides
s imultaneously, and would take them into a"fork," after which the
closed-up "carousel" was formed (for example, on 7 October (season II)
Fig. 16), The dimensions of the dolphins' "carousel" would reach 150 m
in diameter (12 June (season II)), but could be 50-70 m(6 June (Season
III)) or even 25-50 m(17 August (season III)), which was most often ~
determined by the initial area of the school of fish. As has already been '
noted, it might be that not all the dolphins in the herd would participate ~
in the "carousel" (see Table 9), ;
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rU}t OFrLCZAL US[. oNi,Y
~r~~i, ~ ~ s
Number oE
- Date Time dolphins Nature of Activity
,~1.........J....~~~......._.,.~..~........~.~...._:._.._..
...._.._..._........~1.__..r._..........,.4...,....,._........
Season II "Carousel"
- ~i.8. 12.: 6 22(1 I) "Vertical carousel"
18,8. 06.35 8-10 3"carousels" of 2, 3,
4 dolphins `
- 24 �A� 14.52 12-1~ Same, of 2, 3, 5
dolphins
1:1,:a, 10.3~ 10 "Carousel" twice
4 E;
Season III -
05.6. 10.26 2 "Carousel" twice
05.6. 15.10 3
~ 07.6. ~J7.19 8 "Carousel"
09.6. 16.45 8_9 ~
- 10.7. 6.30 11 "Vertical carousel"
of 5-6 dolphins -
10.7. , 16.15 8-10 Same, of 6-8 dolphins
, 16.7. 09.30 3(1 j) "Vertical carousel"
28.7. 11.12 2
[Continued on Following Page]
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- Table 8 (cont'd)
~+~r-r~tir .rr,. w.~~�~. ~.rr....rw~r~.rr..~.~wr~wr~
- 1 r 2--~..- 3..,.~____ 4 -
_ _ .,.r.~___._.._...
Season II "Boiler"
19.6. 5.06 5 "Boiler"
- 02.8. 5.38 7
19.8. 13.05 15 "Boiler" of 8 dolghins
23.9. 11.56 10~ "Boiler" of 4 dolphins
' 06.10. 16.25 10 "Boiler" of 3 dolphins
01.10. 13.03 12 Two "boilers," then three ~
13.10. 10.38 11 "Boiler"
17.10. 10.30 8 ~ "
17.10. 14.56 8 "
= Season III
" 06.6~ 15.05~ 10-12 "Boiler" of 5-6 dolphins
09.6. 16.45 8-8 Several "boilers"
12,6. 14.03 8 Two "boilers" of 4-S -
dolphins each
10.7. 13.26 12 "Boiler" of 4-5 dolphins
04,8. 13.43 10(15 lj ) "Boiler"
20.8. 16.05 10-12 Two groups, two "boilers"
26.8. 11.41 10-12 "Boiler of then
" "boiler" of 3 dolphins
27.8. 07.45 8-10 "Boiler of S-6 dolphins
27.9. 14.20 12(11)
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Figure 15, Schematic Depiction of Consecutative Phases (A-D) of -
Bottlenose Dolphins Encircling a School of Fish
[Continued on Following Page]
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~igure 15 (cont'd)
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Table 9
Number of dolphins
Date Total Participating in
"carousel"
01,8,75 (season II) 13 5 _
17.8.75 (season II) 8(li) 4
18.8.76 (season III) 8-10 4-6
19.8.75 (season II) 8 3-4; 7-8'
07,10.75 (season II) 12 $
b) "The wall." In addition to the encirclement of the fish, the dolphins
would use the device of driving the fish, for example, to the shore,
or the nets placed in the water, or to a"wall" made up of one or several
dolphins. One of these methods--driving them to the shore--was quite
often practiced by bottlenose dolphins (Fig. 17C) (see Table 10),
Table 10
Date Time Number of Nature of activity
dolphins
Season II Drive.to the shore
24,7 16,20 9-10 (2j) "Front"
30.7 12.44 9 "Front"
O1 ,8 10 ,34 13 "Double Front"
19 .8 12 .57 15 "Front"
17.9 04,29 12 3 groups of 3-4-2
19.9 . 10,20 8 2"atCacks" in a"front"
28.10 13.25 13 "Corridor" -
Season III
11,7 10,34 11 "Attack," 3 dolphins
28.7 1L.12 2 Series of "attacks"
4.8 13.43 10 Entire herd; "in a
front" ~
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The drive in to the enclosure might be participated in by the whole herd,
which would most often line up in a"front," and sometimes--by individual
groups, It is possible that this was determined by the species and
number of f ish. A t thia final phase of the hunt the dolphins m~~ved
rapidly, and caught the fish in the foam and spray at the very shore.
For example, a hunt on 24 July (se~son II) was described as follows:
"The dolphins fly completely out of the water, and more fly through the
" air than swim. The fish from a school of large gray mullets, which they -
are chasing, also fly completely out of the water and fly along for 4-7 m.
Often the dolphins ar.d the fish fly together, and sometimes the dolphins
appear to be under a cluster of falling fish." In some cases (30 July
(season II)), the school would prove to be large, and many fish would
go off near the shore because they could not be caught. Therefore, cases
were observed of a new drive to the shore (19 September (season II)). .
It has already been indicated that a group of dolphins serve as a barricade
for the fish. Therefore, it is expedient to present data on this hunting
device as well, when the dolphins would use each other as a"wall" (see
Table 11).
Cases were noted of a hunt of the "wall." type, when the bottlenose dolphins
chased the fish to the shore in two almost parallel "columns"; here,
_ apparently, both ~he shore and the parallel groups of dolphins were used as -
the "walls" (Fig, 17a), The dolphins were functionally equ.al, regardless
of their number in the group, as when a pair of dolphins who, not having
managed to press the fish near the shore (24 July (season II)), drove them
to the herd and then the whole herd "as a front" pinned down the fish, _
or when groups approximately equal in number drove a school of fish against
each other (for example, on 30 July, and 27 June (season II)), (13 June
(season III)). Even one dolphin could fulfill the function of a"wall,"
since it inhibited the movement of the fish schoal or did not let a
single large fish get away to the side. This was noted repeatedly in
seasons II and III (see Table 12).
The wing of the stationary net was also used as a barrier to drive the
fish into it. The group of dolphins in this case often drove the fish
along the wing of the net to the shore. Some animals sometimes jumped
over the net (26 May (season II)). (See Table 13).
c) Pursuit, As a form of capture by a group of dolphins and single
dolphins we singled out the pursuit of the fish, when, without restricting _
the mobility of the school, the dolphins simply overtook the fish. Noted
for the group of dolphins was pursuit "in a front" and "diffused,!'
apparently with spawned fish in small schools, and this was als~o noted
_ simultaneously, i.e., one part of the herd would.pursue the fish collectively
("in a front," "on a course") and the other would swim separately at this
time.
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Table 11.
Date Time Number of Nature of drive
dolphins
Season II
20.6 11.05 9(2j) 2 groups
27.6 17.44 9(2j) 2 groups
24.7 16.20 9-10 (2j) 2 dolphins drive fish from shore to
herd, and herd then drives to shore
25.7 09.00 14 (2j) 2 groups (4 and 4) and
2 groups (3 and 3)
_ 30.7 12.44 9(j) "front to front"
17.8 10.53 8(j) triangle .
19.8 10.31 8 ~ -
_ 19.8 11.50 13 2 "fronts" .
19.8 12.52 15 2 groups
17.10 10.30 8 groups of 2 and 6 dolphins
Season III
7.6 14.20 7
9.6 16.45 8-9 2 dolphins drive fish to herd of
7 dolphins
I3.6 5.38 9-10 "wall to wall"
Table 12
-
Number of Number of
Date Time dolphins Date Time dolphins
~ ,
Season II Season III
5.6. 15t10 3 26.6. 12.09 4. .
6.6. 11.0? 7..8 9.7. 13.42 6 ;
. .
11.6. 09.10 3 10.7. 05.30 11
12.6. 05.18 5 18.7. 6.0? 6
. 12.6. 14.03 8 31.7. 5.51 12-13
1.3.6. 15.08 8 4.8. 13.43 10 . ~ :
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Figure 17. [Caption, Continuation on Following Page] _
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Figure 17. Schematic Depiction of Hunting Behavior of Bottlenose Dolphins,
Driving Fish Against Barrier--"A Wall." A~B--between two
groups of dolphins, C--toward the shore _
Table 13 ~
Date Time Number of Nature of activity
- dolphins
Season II
27.6 12.49 9(2 j) 2 groups ~
Season III
8.6 11.07 . 4-5 In ari "arc" and "front"
11.6 9.10 3 Alone and in group -
11.6 12.33 3 "
' 12.6 14.03 8 "In a front" twice
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Pursuit of the fish "in a front" was noted in the following cases (see
Table 14),
Tab le 14
Date Time Number of Nature of pursuit
dolphins
Season II
27.6 5.30 10 "Front." with graup of
3 dolphins
05.7 14.30 7(lj) "On a course"
. 31.7 OS.Sl 12-13 "Front" in a group of
5-6 toward shore, then
in sea
02.8 05.38 7 "Front"
18.8 06.35 8-10 Rushes of 2-3 dolphins
in synch ronization
18.8 07.45 10-12 "Front" of entire herd
in sea
19.9 1~.20 8 "Front"
Pursuit "in a diffused manner" was also noted several times (see Table
15) .
Pursuit "in a diffused manner" was also noted several times (see Table 15).
Tab le 15
- Date Time Number of dolphins
Season II
27.6 05.30 10
05.7 14.30 7
02.8 05.38 7 _
Season III
26.8 11.41 10-12
~ 17.8 14.27 2-3.
08.9 13.55 20
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rux urrl~itw u5~; U1VLY
Characteristic of a herd's hunting "in a front," "in a spiral," and "in _
a circle" is the diving of the animals for a considerably longer time
than usual--2-3 minutes. It appeared to us that this was connected with
the natural localization of the food. This type of benthonic search
(spawned fish) was also noted for other groups of bottlenose dolphins
(see Table 16) .
Table 16
Date Time Number of dolphins Length of dive, min.
Season III
7,6 14.20 7(2 gr.) 1 -
12,6 07.31 6(1-2 gr) 1-2
(3-4 gr,) 20 sec,
29 .7 09 .32 12-15 1-3
18.7 06.00 5-7 1.5-2
Hunting Alone
We discussed above the hunting procedures for a herd and groups of
bottlenose dolphins. In these cases too, however, it would happen that
the dolphin, although it found itself not far from a group of kinscnen,
acted alone. It attempted either to drive the fish to the shore or to
the wing of a s tationary net, or to drive it up to the surface of the
- water. These are also variants of the "wall" type hunt, but only for a
single animal. The data on hunting by a single dolphin using the "wall" -
method are given in the following table:
_ a) ttack" to the shore (see Table 17). -
b) We repeatedly noted the use of the wing of a stationary net as a
"wall." In addition to this method ot hunting, sin~gle dolphins pursued
fish in a number of cases (see Tab1e 18),
The most widespread procedur~ for hunting by a single dolphin is the
"attack to the shore." The dolphin suddenly made a sharp change in the
direction of its movement and rushed headlong toward the shore on the
very surface of the water, with its dorsal fin thrust out of the water.
At a distance of a few meters from shore the dolphin turned belly upwards
and snatched the fish (most often a gray mullet), then iaunediately turned
around and swam back (sometimes on its back), tossing up the catch once,
or else two or three times, in order to grasp it again at the moment when
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Table 17
Date Time Number of Number of "attacks"
dolphins
- 1 2 3 4
- Season II
17.6 10.00 4 1
21.6 10.98 7 1
25.6 08.03 8 2
26.7 19.00 1
30.7 12.44 9(2 gr.) Several
31.7 05.51 12-13 "Attacks" of
2 dolphins
- 31.7 12.I/+ 18-22 . Several ~
_ 3.10 10.40 10 1 _
Season III
4.6 18.29 2
18. 33
10. 6 16. 00 4 1
11.7 10.34 7-8 1
4. 8 13. 43 10 2
20. 8 5. 45 2 2
Table 18
Date Time Total dolphins
19.6 (season II) 16,55 8-9
12,6 (season III) 07.31 1 .
4.8 (season III) 13.43 ~4
24.8 (season III) 16.08 5-7
26,8 (season III) 11,41 10-12
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it came into contact with the water, Functionally, tossing the fish up
ensured a convenient orientation of the fish for swallowing--head first.
What is involved in the procedure, constantly noted~ of turning belly
upward during the attack to the shore? Several ideas may be expressed
with respect to rhis . One of them is tha.t, under the conditions of
shallow water, good light and clear water, the dolphin does not use the
location apparatus, The fish are detected by means of the acoustic
channel, and then during the pursuit and catch the dolphin uses its
sight, Since its field of vi.sion is directed downward, and the fish is
overhead, near the surface of the water--it turns over, apparently to
see it better. Another assumption is that the dolphin is protecting its
pectoral fins from injury, and a third--it is more convenient this way
to snatch the fish at the frons level. Most probably all of these and
perhaps other reasons as well, operate simultaneously, because sometimes
dolphins seize the fish in deep water too in the pQSition "on the side,"
or "on the back." '
Leaps When Hunting
Dolphins' leaps were often observed during hunting. Of the five types
of leaps recorded by us among bottlenose dolphins during the hunt, only
two types were noted:
1, A low horizontal leap (simple)--the dolphin jumped 30-50 cm out of
the water in a steep arc (to a height no moxe than the thickness of the ~
_ body from the surface), entered the water without splashes (Fig. 20b).
Such leaps were particularly characteristic of the peaceful movement of
common dolphins, and among bottlenose dolphins they were observed during
rapid movement. Individual animals or a whole herd at once could leap,
obviously when driving in the tish. It was these leaps that were observed
during movement "in a line," with the entire herd jumping out in syn-
chronization.
2, "Log" leap--th~ dolphins would fly out of the water at a slant and
- drop into the water flat~ Often in this case the dolphin would thrust
itself out of the water only with two-thirds or three-fourths of the
body--the tail would remain in the water (Fig. 20d). Individual animals
could leap like a"log," but more often--the entire herd~ Such leaps
were sometimes observed during the hunt; obviously, the dolphins drove the
fish with blows of the body agains t the water.
"Vertical climbs" could be observed among the dolphins--vertical surfacing -
from the water with the head upwards and iaunediately--submersion on the -
vertical or with the fall on the sids or on the back. This was often
observed after a successful hunt, when ma.ny fish had been devoured. In
the opinion of V. M, Bel'kovich, the possibility is not ruled out that the _
"vertical climbs" help to "shake down" the devoured fish in the belly.
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Some d~lphins would make 5-8 "vertical climbs" each, in a row, in this
case surfacing wirh the back along the course of the movement, and once
13 "vertical climbs" were recorded~ with one vertical climb taking no
more than one s~cond.
Sometimes af ter the hunt, unusual submersions were observed, w:lth up to
half the body protruding from the wat:er tail up. For example, on 14
July (season III), after the hunt, every third to fourth submersion of
the dolphins was "tail up." V, M. Bel'kovich feels that perhaps, just
as in the case of the "vertical climbs," in this case the devoured fish
was being "shaken down." _
_ Porpoises .
It is more difficult to mal;e a strict differentiation of the behavior of
azovki [Phocoena phocoena Linnaeus] porpoises during hunting and migration
than it is among bottlenose dolphins. The reason for this lies in the
periodic long surfacings (up to 6 minutes) of thsse animals, during which
they could swim far away from the former location, and the frequent hunting
alone, Under conditions of good visibility and with only slight swell of
the sea, however, such a classification proved possible. At the same
time, the hunting behavior was much more easily identified than the
migrational. The movement of the Azov porpoise consisted of a series of
several dives (each one for 5-10 seconds), after which it wou~d disappear
under the water for 1-6 minutes, etc. During the clearly marked
' migrational movement, the Azov porpoises would move along a straighC
line at a speed of approximately 10 km/hour. During the hunt the animals
moved more slowly (2-3 km/hour),
The slow movement ot the animals through a water area several hundred ~
- meters in diameter served as a sign of hunting and searching behaviox.
_ Often during Chis movement, schools of icthyophagous birds would follow
, the porp~ises: small orns, cormorants and herring gulls. In some
cases, however, the birds did not react in any way to the dolphins. In
~ this cases apparently, search behavior occurred, The Azov porpoises
could move through the water area during the hun.t or search ~by several
means. Most often the animals moved "like a shuttle" with varying
range (Fig. 18). The animals often swam back and forth along the current
belt, after diving a few dozen meters in one direction, af ter which Chey
turned back. In a group of 2-3 specimens, the Azov porpoises moved in
synchronization several meters from each other~ Solitary gorpoises
during the hunt might make "loops" or move in a circle (Fig. 18, c, d).
5
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Figure 18. Movement and Hunting of Azov Porpo~.ses: a) sinusoid; '
b) "shuttle"; c) loop; d) circle, e-g) carousel formation, '
h) boiler; i) "flower" ~
52 -
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Sometimes m~~re complex types of porpoise hunts took place--the interaction
- oF individual specimens in groups was observed, As a rule the complex
~ types of hunting were carried out by groups of 3-5 and more animals, Such -
a group might separate into two small groups, which would swim in "shuttle"
fashion toward each other, For example, on 27 July (season III), three
porpoises separated and one animal swam to meet two others, moving to-
gether, On 25 September (season II), we observed a group of six porpoises,
which twice separated into groups of two and four specimens (possibly, to
search for fish), and after a little while came together again. After the
merging of these groups a new separation of a single animal occurred, who ~
moved "like a s huttle" to meet the five, It is possible that the dolphins
used this method to drive fish toward each other (in accordance with the -
"wall" type), "Carousels" and "boilers" of porpoises were noted repeatedly
_ (Fig. 18 e, f). Sometimes the surfacing of several animals from the same
place in diverging directions was observed (Fig, 18, h), On 5 October
- (season III) a"wall" of porpoises was observed--after lining up in a
row, three of the animals quickly moved toward shore.
- The Azov porpoises remained from a few dozen mi~zutes to several hours in
the hunting region, with their behavior and disappearance from this region
. being very difficult to record, As the result of several successful
observations it was established that the hunting behavior among Azov
porpoises aLternated with the migrational. A group of dolphins would move
For a long ~ime in some direction and then suddenly cease this "purposeful"
movement and begin to hunt. In the hunting region individual specimens -
could move "on their own." Then the animals would gather into a group
again, which would swim away from the region of the observations.
- Fewer individual behavior pattern~ were noted among the Azov porpoises
than among the bottlenose dolphins, ~ao types of leaps were observed:
a lo~a horizontal leap and a"log" leap. Exposure of the tail and slapping
the tail against the water were repeatedly observed. Noted twice was the
turning over of the porpoise belly upwards, and once it seized a fish in
this position. Leaps toward each other were noted among the group leaps
(the animals usually leaped alternately),
On 5 August it was possible to observe in quite detailed ma.nner the sexual
behavior of the Azov porpoises in a group of three animals, one of which
was a young porpoise with a"marking" (see above). The two adult dolphins
often leaped out of the water, leaped over each other and swam for several
seconds, with their abdomens pressed to each other (Fig. 19). The sexual
behavior was observed for over 5 minutes. During this entire time the
young porpoise swam first behind and then in front of the adults.
Common Dolphins
The migrational movement of a herd of belobochki [Delphinus delphis -
Linnaeus] white-sided dolphins was dis tinguished by its great swif tness.
54 ~
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The animals moved at high speed, often leaping from the water, During
the migrational passage the groups could be seen quite clearly--they as
a rule consisted of 1-2-4 animals, In each group the dolphins would
leap out in turn, one after the other, The hunting of white-sided
dolphins was described in relatively great detail in the section on
accompaniment in the catamaran. From time to time individual, slowly
swimming white-sided dolphins were also seen in the biopolygon. This
behavior was also, apparently, nutritive, In season II the similarity
- to the "carousel" and "boiler" was observed among the white-sided
- dolphins. The series of observations make it possible to assume the
existence among white-sided dolphins of the "scout" phenomenon. On 21
August, near one of the bays, small groups of 2-4 animals were observed
several hundred meters from the main herd, On 14 September 4 animals
detached themselves from a herd of 25, behind the herd~ they made a pass
toward the shore and 3 minu.tes later again joined the herd. During this
maneuver~ the entire herd rarely slowed its movement and the diving of
the animals became longer, The leaps of the white-sided dolphins, ap-
parently, are similar in type to those of the bottlenose dolphins. We
observed low horizontal leaps "like a log" and of "vertical climb." During
the leap, however, the white-sided dolphins curved their bodies less than
the bottlenose dolphins, The leaps of white-sided dolphins are described
in detail in the article by Pilleri and Knuckey, 1967,
Therefore, a detailed analysis of several hundred situations in dolphin
hunting makes it possible to reveal the great variety of the searching
and hunting behavior of these animals~ We singled out and described
certain formerly unknown basic types of hunting behavior of dolphins in
the natural habitat, These behavioral reactions of the search, detecting
and catching of fish combine with each other and alternate during the ~
hunt, depending on the type and number of the fish, the meteorological
conditions, the presence of interference, and the number of dolphins
taking part in the hunt, The abundance of procedures for hunting is -
impressive and, mainly, the great plasticity of the behavior of the
dolphins, permitting them to react adequately to a change in circumstances.
BIBLIOGRAPHY
1. Bel' kovich, V. M, ; Andreyev, F. V.; Vronskaya, S. D.; and Cherdantsev,
A, I,, 1975. "A Study oF the Behavior of the Bottlenose Dolphin in
Nature," in the book, "Morskiye mlekopitayushchiye" [Marine Mammals],
Matarials of the Sixth All-Union Conference on the Study of Marine
Mammals," Kiev, Naukova Dumka, Pt 1, 24-25.
. 2, Bel'kot~ich, V, M, and Dubrovskiy, N. A., 1976, "Sensornyye osnovy
oriyentatsii kitoobraznykh" [Sensory Bases of Cetacean Orientation],
_ Leningrad, Nauka,
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3, Morozov, D. A,, 1970. "The Dolphins Are Hunting," RYBN, KHOZ,~ Vol
46, 16-17.
4. Tomilin, A, G,, 1957. "Zveri SSSR i pri?ezhashchikh stran" [Wild
Animals of the USSR and Adjacent Countries] Vol 9, Cetaceans, Moscow,
Izd, AN SSSR.
' S. Caldwell, M, C, and Caldwell , D, V. , 1965 ,"Individualized Whis tle
Contours in the Bottlenose Dolphin (T, Truncatus)," NATURE, Vol 207,
- No 4995, 434-435.
6, Evans, W, E, and Bastian, J., 1969. "Marine Ma.mmal Communication:
Social and Ecological Factors," in "The Biology of Marine Mammals,"
N, Y,-London, Acad, Press, 425-476,
7. Pilleri, G, and Knuckey, J., 1968. "The Distribution, Navigation and
- Orientation by the Delphinus delphis L, in the Western Mediterranean,"
EXPERIMENTIA, Vol 24, No 4, 394-396.
- 56
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!
UDC 534.7+612.85
DESCRIPTION OF PULSED SIGNALS OF BOTTLENOSE DOLPHINS IN THE OPEN SEA
Moscow POVIDENIYE I BIOAKUSTIKA DEL'FINOV in Russian 1978 pp 104-116 ~
[Article by V. M. Bel'kovich, V. V. Kaznadzey, S. A. Kreychi, E. A.
Khakhalkina] _
[Text] Among the numerous acoustic signals of dolphins, the majority are
of a pulsed nature and are emitted in series of varying length and repe- _
tition frequency. The primary and principal biological function of pulsed '
signals is echolocation.
In an evaluation of the spectral characteristics of signals made by
various researchers in t-i:a 1960's, broad use was made of a verbal
description of them, while listening on the actual time scale, or with
a delay. For example: a"bark" and "yelp" correspond to relatively
short series of pulses with a repetition frequency of 400-500 pulses/sec,
_ (Schevill, 1964); "squeak of door hinges"--to series with an increasing
pulse repetition frequency 2-10 sec, long, etc. A more detailed classi-
fication of pulsed signals is also well known (Titov, Tomilin, 1970;
Titov, Yurkevich, 1971; Markov et al., 1974), including tha.t according to
- acoustic analogy: clicks--1-3 msec; resounding booms (4-9 msec); pro-
longed booms--up to 60 msec; crackling--from 100 to 1200 pulses/sec;
quacks--series of pulses with the noise duty cycle in a range of,0.4-
80 kHz, with a variable position of the energy pe~ak in the range of 1.6-
32 kHz and the length of the series 0,1-0.8 sec; bursts--series 30-180
msec long, with the noise spectrum of the pulse duty cycle in the range
of 10-30 kHz; howl--with the length of the series 0.5-0,8 sec; a spectrum
of the pulse duty cycle of 10-80 kHz with the energy peak within a range
of 32-40 kHz; roar--a sharply increasing pulse frequency., and a wail--
variable pulse frequency, changing according to a random law. Also
displayed were complex signals, formed by imposing pulses on whistles,
which is possibly related to the participation of the pulses in the
. structure of the messages during communication along with the whistle-like
signals (Kaznadzey et al., 1975; Khakhalkina et al., 1977).
A great deal of attention was paid to the study of these questions by
Vincent, R, G. Busnel, A. Dziedzic, D. Caldwell, M. Caldwell, J. Dreher,
W. Schevill, W. Watkins, K. Norris and other researchers, references to
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which are contained in recent surveys (Dierks et al., 1973; Ayrapet'yants,
Konstantinov, 1974; Bel'kovich, Dubrovskiy, 1976).
In the 1970's the researchers' attention was shifted s trictly to the
_ individual pulse. As a result, two opposing points of view were formed.
. According to one, the pulsed signals as a whole are quite stereotyped
and standard, and according to the other--the pulsed signals are variable:
the energy peak in the spectrum of pulsed series "floats," drawing on
spectrograms structural (forma.nt) bands of varying configuration (Schevill,
1964), the frequency in the series changes from pulse to pulse (Romanenko,
1964) and the peak of the spectrum, the pulse repetition frequency and -
duration vary to equal the task (Akopian et al., 1973; Ayrapet'yants,
Konstantinov, 1972). All these data, however, were obtained under the
conditions of a tank or pen. Therefore, the data on adaptation and _
- variability of the pulses among dolphins in nature is of great practical
interest, In performing bioacoustic research under the conditions of the
open sea, we naturally attempted to gather and analyze material on the
pulsed signals of dolphins under the conditions of their natural habitat,
Ma.terial and Method +
We recorded the acoustic activity of the Black Sea bottlenose dolphin
from May through September in the course of two (II and III) seasons,
during observations of their beha.vior. The basic beha.vioral situaCion
consisted of different stages of hunting fish by single delphins and
groups of 1, 2, 3... 12 dolphins in a biopolygon. Equipment with a
- passband of 0,1 - 100,kHz (Fig. 23) was used to record the signals.
The recordings obtained were heard with an 8-fold lag, and analyzed on
a 48-channel dynamic spectroanalyzer, describ~d above.
' Results and D iscussion
_ In the auditory analysis of the pulsed signals, as well as the analysis
of their spectral-temporal characteristics, obtained from the spectro-
analyzer, it was noted that the classification of the pulsed signals
by acoustic analogy, which was mentioned above, in general is retained,
although within each type ~there are a number of variations in the pulsed
signals, which can be clearly differentiated by ear (auditory) when
transposed, and give rise to various auditory image repreaentations.
For example, Figure 30.1 shows the spectral-temporal characteristics of
four series of pulsed signals, classified as clicks, which upon auditory
analysis evoke the following associations: -
a--the crack of a whip--short (3 msec), relatively infrequently spaced
(115-145 msec) broadband pul:,es (0-80 kHz). More frequent pulses in the
low-frequency area are noise interference;
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b--tapping a ntiil against glass--duration--3 msec, but the period of
_ repetition is 20-25 msec and the energy peak is in the range of 18-80 kHz.
On the spectrogram interference of a pulaed nature could also be seen,
the source of which consists of musaels and shrimps (pulsea of 10 msec
and longer with a broad spectrum and energy peak in the low-frequency
and high-frequency ranges);
c--rapping a rod against dry wood. Duration--3 msec, repetition period
15-20 msec, and they have a discontinuity in the duty cycle spectrum
(the low-frequency part of the spectrum is 0-3 kHz; the hig?1-frequency--
12-80 kHz), Most of the energy is concentrated in the high-frequency
area of the spectrum;
d--rolling sounds resembling pile-driving--a series of pulses with the
repetition period up to 100 msec and the duration up to 3 msec in the
_ low-frequency range and three times as long (up to 10 msec) in the high-
frequency range (20-80 kHz).
Figure 30,2 shows the spectral-temporal characteristics of other pulsed
- signals, classified as booms. Characteristic of the entire group is a
greater change in the time of the various sections of the spectrum than -
~ in the pulses shown in Figure 46,1, due to which the pulse on the spectro-
gram ha.s the shape of a wedge with the point underneath. In an auditory ~
analysis with transposition there are differentiated: -
a--blows with a hammer along an iron plate--pulses 3 msec long in the low-
frequency area of the spectrum and 15-20 msec in the high-frequency
(spectrum up to 80 kHz), with the repetition period 20-50 msec;
b--gunshots--duration up to 30 msec in the high-frequency range of the
spectrum (15-~0 kHz) and 3 msec in the middle frequencies, with the
repetition frequency 95-115 msec. Present at the same time is interference
from shrimps in the form o� chaotic crackling in the high-frequency
range;
c--bursts--duration up to 60 msec, spectrum--18-80 kHz, The great inten-
siveness and high percentage of encounter rate in hunting situations
permit the assumption to be made tha.t they are used to deafen the fish;
d--rumbling of a tractor motor, alternating with discharges from an
exhaust pipe. A series of location pulses increasing in frequency, which
indicates the dolphin's comtng close to the target (Norris, Turner, 1967;
Bel'kovich, Reznikov, 1971) and usually ends with a"quack" or "roar"
charactPristic when catching the fish. _
As was already noted above, the pulsed signals are emitted by the dolphins
usually in series, which are distinguished by the duration, as well as by
the mutual spacing of the pulses within the series. For example, for, -
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series classified as crackling a duration of up to several seconds is
- characteristic, with the length of each of the pulses being 1-3 msec,
the pulse repetition frequency 5-15 msec and the change according to a
random law of the spectral density during the series, which from the
auditory standpoint ma.kes them similar to the operating sounds of various
mechanisms and machines. Figure 30,3 shows certain characteristic
cracklings which, when transposed 8-fold, can be easily distinguished
by acous tic analogies: _
a--a circular saw--pulses with the r~petition period about 8 msec, and
have a broad "slump" in the spectrum in the middle frequencies, which by
the end of the series narrows down and disappears;
_ b--a tractor moving alor.g where there is no road. The energy of the
pulses is concentrated in the high-frequency range (18-80 kHz) and the
~ repetition period of the pulses cha.nges sharply from 15 to 2 msec;
c--motorcycles picking up speed, The repetition frequency varies from
- ZO msec to 5 msec. The basic energy of the pulses is concentrated in the ~
low-frequency and high-frequancy ranges of the spectrum, while in the .
middle frequency ranges the spectrum changes according to a random law;
d--a short machine-gun .burst, The pulse repetition period decreases from
5 msec to 1 msec, at the beginning of the series the energy peak is
concer~trated in the high-frequency range, and by the end expands sharply,
All the time intervals and frequency values of tha spectral components
analyzed above the pulsed signals are given on an actual scale, while the
description using the auditory analogy of individual pulses and their
series was made with an 8-fold lag. Therefore, the relative scale of
all types of signals is retained, It must be remembered~ however, that
o n the real time scale all the pulsed signals are dis tinguished for us
only by the repetition frequency. For example, Figure 30.4 shows the
spectrograms of the sounds of the dolphins perceived by the ear as �a .
" roar" (a), "wail" (b) and "quack" or "mew" (c). These sounds are a
series of short pulses (1-3 msec) with a relatively uniform density of
the spectrum of 0-80 kHz and very short interpulse interval (pulse
repetition frequency from 500 to 1200 pulses/sec). As a rule these
sounds conclude the locating when catching the fish. ;
_ Here it should be said that the frequency boundaries of the energy -
distribution in the spectrum of these series may be modulated, describing
a certain contour, similarly to the envelope of the time-frequency '
characteristics of the basic tone of the whistle-like signals.
W hen analyzing Che sounds of a pair of dolphins in a tank, we also recorded '
the series of pulsed signals, the spectrum of which was concentrated in a
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- relatively narrow band of frequencies and was modulated so as to describe
a certain contour, analogous to the contour of the envelope of the whistle-
like signsls (Figs, 31, 32). These changes in the spectrum in the series
oL pulsed si~nals, upon comparison with the behavior at the moment of
emission, then permitted the assumption of the possibility of their uae
in communication (Kaznadzey et al., 1976; Khakhalkina et al., 1975), and
the spectral-temporal analysis revealed certain special features that
indicate the possibility of arbitrary control of their spectral-temporal _
structure (Fig. 31) by means of several simultaneously operating sources,
as well as resonators rearranging the structure of the signals.
The above comparisons of objective spectral-temporal characteristics and
subjective auditory analysis of the pulsed signals of dolphins, when
transposed, show that man can easily distinguish subtle changes in the
spectrum, duration and repetition frequency of these signals. This
permits the assumption tha.t the dolphin also differentiates the above-
noted changes in the parameters of the "stereotype" pulses, since the -
dolphins' acoustic anal,~zer possesses a higher resolving capacity for
the procedure (Vel'min, Dubrovskiy, 1975; Bel'kovich, Dubrovskiy, 1976)
and it is poss ible that this is used as an additional source of infor-
matio~n. -
One must, however, take into consideration the fact that there is still
a great deal that is unclear with respect to the ques tion of the variability _
in the structure of the pulsed signal. As was noted above, some people
- feel that the dolphins' sounding pulses are always stereotyped~ and the
variations in the spectral-temporal characteristics recorded are brought
about by a change in the dolphin's directivity diagram, due to turning
the head, Others rhink it is possible tha.t there is arbitrary control -
of the spec tral-temporal characteris tics of the pulses, on the basis of
the series recorded in the s ituation of the animal's fine adjustment to
the conditions of tha experiment (see the surveys, Ayrapet'yants,
Konstantinov, 1974; :~ubrovskiy, 1975). Probably, the truth lies somewhere
between, namely, depending on the situation and the conditions of the
study of the sounding signals, the dolphin uses the discontinuity of its
pattern for sufficientl.y fine tuning (Bel'kovich, Dubrovskiy, 1976).
If this does prove to be eff icient for recognition, the o ther animals
may receive quite a large amount of additional information on both the _
~ rarget and the source. On the other hand, the reason for the change,
. in the signals with respect to duration and position of the energy peaks
in the spectrum in the pulse series may be related merely to the bioLogical
nature of the transmitter, which for some reason is forgotten about and
which ensures both a certain stability in the parameters of the pulsed
' signals and a certain dispersion of them. The above-noted characteristics
of the pulsation are apparently inherent in the transmitter of each ~
animal, and this individuality is.undoubtedl,y important for recognition
of individuals, and for,orientation, determa,:ning the mutual positioning,
e tc. This biological instability of each transmitter may prove to be
61 ~
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Dolphins in Nature '
62 ~
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Figure 31, Spectrograms of Pulsed Signals of Dolphins Under Conditions
of Captivity -
~ 63
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_ _ . , . . .
. - ~ ~ , , ~.f _ y~� ,...~~,a,..~ ~ ;
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~ + T 1 S '~i ~ ~ ~
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r - e .
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~ T . ~ ~ � r Y ~ -r ,
~ i ~ l f -'r ' i"~ . w
B ~ , t ~ ~ � t ~ ~ -
* ~ , x t ~ : ' t
~~r ~~i' ,l�~a'~.~~.. ~ ~~'1 .
~ ~ ti! l 1 ;
~C~ .~!'~,~'~~'~'i~'F'~_~SFC~~r. ~ 1 t ~ i '
' ` ~ ~ ' ,'j : � `G , ;
tY;~c.t;,~.~� r~:,:~~'!~~y ..1
: ~ 1 ; = " ~a; , {
~s ; a ` ~
.
; f ~ '
i
. (a, r E,` ~ _ t+
_ i ~ _ j, ~
~r... ,
::.j.
_ ,
_ , : _
f~~~.~'~,~_ r.. .~i { F_ _ :i . -
~e~ , , , , ;
~ ~
' %~,~~~:ii ibi .
Figure 32. Spectrograms of Pulsed-Whistle Signals* '
*It is assumed"that the whistle signal bears an emotional or individual-
recognition load, while the pulsed component serves to establish the
address channel of communications for directed transmission of any
whis tle s ignal .
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no less important for the locational perception of the targets as well,
since above all, it eliminates fatigue and the nervous centers' habituation
to uniform echosignals (Bel'kovich, 1972),
When using acoustic pulses, the dolphin is forced to tune oitt the pulsed
interference that is constantly present under natural conditions. The
spectrograms shown above are "cluttered" with the noise of surf, the
sounds of mussels, slzrimps, etc., which have a cunstant spectrum in
- time. This constancy of the spectrum is obviously one of the reliable
- criteria for singling out the signal, because the difference in the
spectrums and the modulated nature of the dolphins' sounding signals
make it possible to distinguish them easily from the interference.
It is thought that the principal and primary biological function of the -
pulsed signals is echolocation, which ensures the dolphins' orientation
in the aquatic environment, where its visual analysis is excluded at great
distances, but considerable possibilities arE afforded for the development
and, use of an acoustic system of monitoring (evaluation and analysis) of -
the environmental habitat and, accordingly, adequate organization of
behavior in it (Bel'kovich, 1972, 1974). The dolphins' echolocation
(acoustic) analyzer may serve as an example of the adaptive development
of this sys tem for acoustic analysis of the environment and its use in
organizing behavior, -
= It is expedient to show the given functional-ethological specificity of
_ various types of series of pulsed signals in relation to certain situations
and the nature of the dolphins' behavior, For example, series of pulsed
signals are use~' for orientation during migrations and when maneuvering
during a search for fish. Most of ten they are related to the detection
of the catch, but particularly with its pursuit during the hunt. In this
case an increase in the frequency of the transmission of pulses indicates
that the dolphin has begun active pursuit of the catch (Bel'kovich,
Reznikov, 1971), which is usually noted only from a distance of about
0,5-0,3 meters (Fig. 30.2g), This increase in the pulsation is usually
c~mpleted at 1/f M250-800 pulses/sec, which is perceived by our ear as a
roar ("mewing" or "quack"), which is emitted at the moment of seizing the
catch, Sometimes this echolocation series, with acceleration of the pulse
repetition does not end with a roar, which indicates that the dolphin has
- missed the catch. Therefore, one may calculate the number of fish eaten
during the course of the hunt by the number of roars.
Uniformity and slowing down in the pulse repetition frequency in the _
= series (5-30 pulses/sec) corresponds to the orientation of the dolphins =
and the scanning direction finding (of the fish) in the ir~3.tia1 phases of
the hunt. Individual clicks are u~ually also used by dolphins as a
duty practice, and ensure orientation and search of fish. Booms are ~
distinguished by gre3ter power (wid th of the spectrum) and a relatively
long duration (up to 60 msec), ihe functional significance of the booms
_ is not yet completely clear, but in consideration of their physical
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characteristics and the use mainly during active devouring of the fish =
by a group of dolphins (64,2%) (Table 7), it may be assumed that they are
used to disorient and deafen the fish during the hunt,

- Conclusion
The auditory analysis made showed that the pulsed signals used by the
dolphins occur during transposition of the characteristic sound image
representa*ions, which are easily memorized and clearly recognized by
the operator, It is possible that due to this, they may play a certain -
_ informative role for a group of dolphins as well, by creating an acous tic
picture of a behavioral situation as a whole and the action of each of the
members of the group individually, � -
In an auditory analysis with transposition of the acoustic circumstances ,
(context) in various behavioral situations, it was also noted that a
combination of different sounds creates fully definite sound images
(pictures) of the situation: "construction," "individual shooting,"
"air combat," "motorcycle races," etc, A set of such sound pictures
~ obviously is also limited, just as a set of types of group behavior~
This makes it possible, from the acoustic circumstances (picture), to
_ recreate in the first approximation the behavioral pictures of what is
taking place in the water, and to compare them with visual ethological
observations, This comprehensive ethological-acoustic approach was used
for the first time,
= There is no need to prove that such information can be useful in orgarlizing
group actions of dolphins under coiditions of the natural habitat, _
- particularly since our observations of the dolphins' beha.vior indicate a~ '
_ high level of interaction and organization in the behavior of dolphins
in a group,
- The abundant representation of pulsed signals in the situations (about
- 90% of all the signa.ls) gives reason to assume that in the process of the
evolution of dolphins' communicative system, the informative surplus of
pulsed signals began to be used by the dolphins for communicative purposes,
in combination with tlie whistle-like signals. This signaling could ~on-
tribute to more efficient intragroup organization and to the development
of the complex forms of behavior which ~ae Qbserve .
BIBLIOGRAPHY
1, Akopian, A, I.; Burdin, V. I.; Zaytseva, K, A,; Morozov, V. P.; and
Sokevykh, Yu, A,~ 1973, "The Effect of Noise on the Repetition
Frequency of Dolphins' Echolocation Signals,",Abstract of a report
at the Eighth All-Union Acoustic Confer'nce, Moscow, Vo1 3, p 9,
;
66
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2. Ayrapet'yants, E. Sh, and Konstantinov, A, I., 1972, "Ekholokatsiya -
v prirode" [Echolocation Under Natural Conditions], Leningrad, Nauka,
- Zd ed,, 1974.
3. Baranov, G, L, and Kovalenko, V, P., 1975. "New Information on Sonar
Detection in Dolphins," PRIRODA~ No 11, 84-90,
4. Bel'kovich, V. M., 1972. "Hearing and Location," Chapter 10 in the -
book, "Kity i del'finy" [4Jhales and Dolphinsj, Moscow, Nauka,
5, Bel'kovicr., V, M., 1974. "Cetacean Orientation," "Itogi nauki"
_ [Results of ScienceJ; Moscow, VINITI AN SSSR, Vol 6, 190-210.
6, Bel' kovich, t~ _:-i, ; Andreyev, F, V, ; Vronskaya, S, D,; and Cherdantsev,
A. I., 1975. "A S tudy of the Behavior of Bottlenose Dolphins Under
Natural Conditions," in the collection, "Morskiye mlekopitayushchiye"
[Marine Mammals], Naukova dumka, Pt 1, 24-25,
7. Bel'kovich, V, M, and Dubrovskiy, N, A., 1976. "Sensornyye osnovy
oriyentatsii kitoobraznykh" [Sensory Bases of the Orientation of
Cetaceans], Leningrad, Nauka.
~7 .
8. Vel'min, V. A, and Dubrovskiy, N, A,~ 1975. "Acoustic Analysis of
Pulsed Sounds by Dolphins," DAN SSSR~ Vol 225, No 2, 229-232,
9. Dreyer, Dzh, and Evans, U., 1969, "Communication Among Cetaceans,"
"Morskaya bioakustika" [Marine Bioacoustics], Leningrad, Sudostroyeniye,
10, Dubrovskiy, N, A,, 1975. "Echolocation in Cetaceans," "Rumb"
[Bearings],
11. Zlatoustova, L. V, and Nizova, A, B,, 1971. "An Experiment in
Acoustic Analysis of Certain Whistles of Bottlenose Dolphins," in the
book, "Morfologiya i ekologiya morskikh mlekopitayushchikh" [Morphology
and Ekology of Marine Ma.mmals], Moscow, Nauka.
12, Kaznadzey, V. V.; Kreychi, S, A,; and Khakhalkina, E. N,, 1976.
- ''Types of Communicative Signals of Dolphins and the Nature of Their -
Organization," AKUST, ZHUR,, Vo1 22, No 6.
13. Markov, V. I,; Tarchevskaya, V. A.; and Ostrovskaya, V. M., 1974.
"Organization of Acoustic Signals Among Black Sea Bottlenose Dolphins," -
in the collection, "Morfologiya, fiziologi.ya i akustika morskikh
mlekopitayushchikh" [Morphology, Physiology and Acoustics of Marine -
Ma.mmals], Moscow, Nauka, _
- 14. Markov, V, I., and Ostrovskaya, V, M,, 1975. "On the Combinatory
_ Nature of Whis t1e Signals in Bottlenose Dolphins," in the collection,
"Morskiye mlekopitayushchiye," PC 1, Kiev, Naukova dumka, 205-207. -
67
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15. Markov, V. I,, and Ostrovskaya, V. M,, 1975. "The Recognition Signal
_ in Dolphins," in the collection, "Morskiye mlekopitayushchiye," Pt 1,
Kiev, Naukova dumka,
= 16. Reznik, A, M., and Chupakov, A~ G., 1975. "The Structure of Speech
_ Signals of the Bottlenose Dolphin," BIONIKA, No 9, Kiev, Naukova
dumka ,
17. Romanenko, Ye. V., 1964, "The Sonar Detection Ability of Dolphins -
(A Survey)," AKUSTICHESKIY ZHURNAL~ Vol 10, No 4, 385-397. -
18, Titov, A, A,, and Tomilin, A, G,, 1970, "The Acoustic Activity of
the White-Sided Dolphin and the Porpoise in Various Situations,"
BIONIKA~ No 4, 88-94, Kiev, Naukov~: dumka,
19. Titov, A, A,, and Yurkevich, L, I,, 1971, "Physical Characteris tics
of Nonlocational Sounds in Black Sea Dolphins," BTONIKA~ No S, 57-62,
Kiev,
20, Titov, A, A,; Tomilin, A, G.; Baryshnikov, N. S,; Yurkevich, L. I.;
and Lekomtsev, V, M., 1971, "Communication-Emotional Signals in
Black Sea Dolphins," BIONIKA, No 5, Kiev, 67-72,
2Z. Khakhalkina, E, A,; Kaznadzey, V, V,; Kreychi, S, A,; and Nikolenko, `
~ G. V., 1975, "Emotional Signals in the Communicative Sys tem of
Dolphins," in the collectiori, "Ntorskiye mlekopitayushchiye," Pt 2,
Kiev, Naukova dumka~
22. Busnel, R. G., 1967. (Ed.), "Animal Sonar Systems. Biology and _
_ Bionics," Vols 1, 2, Frascati, Italy.
23, Busnel, R, G,, and Dziedzic, A,, 1966, "Acoustic Signals of the
Pilot Whale Globicephala melaena and of the Porpoise Delphinus delphis
and Phocoena phocoena," in, "Whales , Dolphins and Porpoises K. S,
Norris, (Ed,), Berkeley--Los Angeles, University of California.
24, Caldwell, M, C., and Caldwell, D, K., 1972, "Dolphins Communicate--
But They Don' t Talk," NAV. RES .
~
25. Caldwell, M, C., Caldwell, D. K.; and Hall, M, K., 1973. "The Ability
of the Atlantic Bottlenose Dolphin (Tursiops truncatus) To D iscriminate
Between and Potentially Identify to an Individual the Whistles of -
Another Species, the Common Dolphin (D, delphis)," CETOLOGY, 14,
1-7. .
26. Dierlcs, K. J.; Trochta, R. T,; and Ewans, W. E,, 1973. "Delphinid
Sonar: Measurement and Analysis," 3, ACOUST, SOC. AMER,, Vol 54,
No 1, 200-204. ~ .
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27, Dreher, J, J., and Evans, W. E,, 1964. "Cetacean Communication," in
"Marine Bioacous tics, Qxford-London-N,Y,
28. Evans, W. E., 1973, "Echlocation by Marine Delphinida and One
Species of Freshwater polphin,"
- 29. Kellog, W, N., 1961, "Porpoises and Sonar," Chicago Univ. Press,
30 . Lang, T, G., and Smith, H, A. P,, 1965 , "Communications Between
Dolphins in Separate Tanks by Means of an Electronic ~coustics Link,"
SCIENCE, 150, 1839-1844.
- 31 . Lilly, J, C., 1963 , "Dis tress Call of the Bottlenose Dolphin:
Stimuli and Evoked Behavioral Responses," SCIENCE, Vol 139, No 3550,
116-118 .
3'L. Lilly, J, C., 1967., "The Mind of the Dolphins," Doubleday, No 4.
33 . Norris , K , S , ; Evans , W . E , ; and Turner ~ R . N . , 1967 . "Echolocation
in an Atlantic Bottlenose Porpoise During Discrimination," in
"Animal Sonar Systems," Frascati, Italy, Vol 1, 409-437, -
34. Schevill, W. E., 1964. "Underwater Sounds of Cetaceans," "Marine
Bioacoustics," Pergamon Press, No 4, 307-316,
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UDC 534.88
- CHAR.ACTERISTICS OF THE DOLPHIN' SONAR FUNCTION ~
Moscow POVEDENIYE I BIOAKUSTIKA DEL'FINOV in Russian 1978 pp 104-].34
[Article by Ivanenko, Yu, V,, Ivanov, M. P,, Nonin, Yu, A., Chilingiris,
. V, I,]
[Text] This article gives a brief survey of our studies of the sonar
system of the Black Sea Bottlenose Dolphin (Tursiops truncatus), and also
formulates a number of hypotheses on the possible principles of the
functioning of certain elements of this system,
_ 1, As a locational click, the dolphin primarily uses one- or one-and-a-
half phase pulses with a duration of 7-100 msec, with a pulse dut;~ factor
somewhat greater than twice the time for transmission of the signal to
the object of location (Ivanenko, Ivanov, Telekhov, 1977).� .
All the existing physical models of the generation of location pulses of
the dolphin cannot explain the exis tence of the brief (tenths of a msec)
signals of a nonresonant nature, which we have recorded in various ex-
periments with the animals. To solve this problem we advanced the
hypothesis of the compression of the frequency modulation of the dolphin's
signals in the frontal protuberance (Nonin, 1977).
2. An analysis of histological specimens af the froatal protuberance
revealed the complexity and nonuniformity of its structure (Bel'kovich,
Nesterenko, 1971; Agarkov, Khomenko, Khadzhinskiy, 1974).
We will discuss tha frontal protuberance of the dolphin, which consists~
' of connective tissue and individual fat cells, as a two-component water-
like medium with varying values of density and compressibil.ity. By tnedium
we will mean the connective tissue, and by micro-nonunifortnities--the
fat cells .
When s tudying arbitrary, including wave, motions of the continuums, the
initial system oF equations consists of Euler equations, valid for the
ideal medium. In a one-dimensional case we have:
!
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dP ~ pv =0 ~ ~ 1) -
ar ax
~3v ~7v 1 dP
_ ~ v _ ~ ( 21 -
at ax P ax
P = P
(3)
where P is the density of the mixture of the two components, p is the
pressure and v is the velocity of the particles of the medium.
This system is complete and consists of a continuity equation (1), an
equation of motion (2) and an equation of state (3).
Passing through the frontal protuberance, the wave of pressure causes
a pulsation of the fat cells. Therefore, in the nonuniform medium under
discussion, Euler's equations should be supplemented with an equation for
the small oscillations of the cell. The movement of the surface of the
compressed cell may be described by Ra.yleigh's equation: -
RR + 3/2 R2 = ~ (p~_p~), ~4~
p0~
where R is the radius of the cell,~ 0 is the density of the medium, P
is the pressure inside the cell, ~ Q --the pressure in the medium. The
derivatives for time are designated by the points.
: When solving the system of equations (1) -(4) jointly, we arrive at a
variance ratio linking the speed of sound ) in the two-component model
of the frontal protuberance of the dolphin under discussion with the
frequency of the oscillations being transmitted (W ) and the relative
concentration of the fat cells ( n~/0
2
~1 Z ,
C~~CD (1 - nV~)2 �
2 '(s)
~ 1- W + ~vo ~
W 0 -
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Here ~p is the speed of sound in the tissue in the absence of fat cella
(~~/0 = Q); 43o is the resonance frequency of the cell, which according
to our estimates is a value in the order of 350-420 kHz.
From the ratio (5) it follows that with an increase in the concentration
ot the fa t cells ( nV~ the speed of sound will decrease ( Q); an
_ analogous phenomenon will be observed when ~J~ . The first fact '
indicates that in the dolphin's frontal protuberance there should be a
~ spatial dispersion of sound connected with the nonuniformity of its
' structure, and the second--that there is a frequency dispersion, with
the high-frequency harmonics being propagated more slowly tha.n the low-
frequency. The possibility of the dolphin's fatty cushion functioning as
a nonuniform coated retarding lens has been noted in works (Bel'kovich,
Nesterenko, 1971; Nesterenko, Rezvov, 1973), A mathematical examination
- of the acoustic properties of the dolphin's frontal protuberance reveals
a possible physical mechanism of an influence, through the nonuniformity
of the fatty cushion, on the passing sound wave, which lies in the
nonlinear and dispersional nature of the interaction of the wave and the
fat cells (Nonin, 1977).
Model experiments to det~rmine the speed of sound in specimens taken from
- various areas of the fatty cushion of the dolphin have shown that the
speed of sound in the center of the frontal protuberance is approximately
10% less than in the peripheral area (Norris, Harvey, 1974). In these -
experiments, however, it was assumed a fortiori tha.t the speed of sound
~aould not depend on the frequency and length of the transmission path of
_ the acoustic signal. Otherwise, the results should be different.
As was noted above, the existing physical models of the generation of the -
locational signal of the dolphin are unsatisfactory. Therefore, we made
an experimental attempt to ascertain the possibility of compression of
the frequency modulation of the signal in the frontal protuberance. The
experiment was conducted in a sound-anechoic tank measuring 290 X 60 X 50
cm, filled with sea water. The measured reflection factor of the acoustic
rubber used to deaden the sound did not exceed 0,1 in a frequency range ;
of 10-200 kHz,
- The dolphin's head was rigidly f ixed so that the distance from the emitter ~
introduced through the blow hole to the area of the tubular sacs up to the !
surface of the water was 20-25 cm. The pickup hydrophone was placed 15-20
cm from the tip of the rostrum on its axis. j
Used as the ocsillator was a sphere ~ 10 mm made from TsTS-19 piezoceramic, -
calibrated for the emission, and as the pickup hydroplane--a sphere ~ 7 mm
ma.de of the same material, ,
The si.gnal generator was a specially designed pulse frequency modulation ~
generator with the following parameters: ranges of change in the carrier ~
72
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frequency (f/r) 30-130 kHz; change in the deviation (Q ) from the value
of the carrier by 0-60 kHz according to the falling law, the duration of
the pulses could vary within the limits of 0,1-10 msec, the repetition
frequency--from 2 to 250 pu~s, and the nonlinearity of the output
signal, less than 3%.
The nonuniformity of the amplitude-frequency cha.racteristic of the entire
receiving-transmitting channel did not exceed 3 dB.
Figure 33 shoxs the results of the experiment, with the following values
of the parameters of the signal emitted: duration of the pulse 1 msec,
repetition frequency 40 Hz,~'~y = 120 kHz (Fig. 33 A). It turned out that
with the introduction of a negative frequency modulation, the signal being
transmitted~ in passing through the frontal pr~tuberance, breaks up into
two groups of pulses, and~ with an increase in the deviation, the duration
. ~f the groups themselves, as well as the components of the group of pulses,
decreases (Fig. 33 d-e). This fact ma.y be interpreted as a compression of
the emitted signal and--confirmation of the theoretical conclusions as to
- the existence in the frontal protuberar~ce of frequency dispersion.
~g)
e(f)
a (e)
~ .r..~ r ( d )
~ n ~
r. ' e ~ .
P:
. 9
, ' .
Figure 33. Cha.nge in the Structure of Signals Passing Through the Frontal
Protuberance, Depending on the Deviation ( a) of the Frequency
~ in kHz
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- rvn urrl~tru, ua~ vivLx
1~ should be nored that the results of the experiment depended on the
de~ree oL- freshness of- the specimens. (The experiment was conducted at
a di.Eferent time on three heads of bottlenose dolphins), After the head
had been in the refrigerator (at -3�C) for a certain time, the amplitude
oF the signal transmitted dropped, the compression disappeared and re-
verberation interference began to appear (defocusing occurred). Analogous
ctianges were also observed at the end of the experiment, if it lasted long
enough. These facts also indicate that the mecha.nism of interaction of the
sound field with the tissues of the frontal protuberance cannot be explained
merely from the standpoint of the nonuniformity of its structure (different
average density), The theoretical model suggested by us, even though it
may qualitatively explain the results of the experiment, for a quantitative
comparison requires detailed study of the physical properties of the -
dolphin's fatty cushion.
~ 3. In the behavioral experiments, conducted according to conditioned
reflex methodology, using multichannel recording of the sounding signals
(Ivanenko, Ivanov, Telekhov, 1977), ~ae studied the characteristics of the
directivity of the acoustic field of the dolphin during emission. The
resulrs of the experiment showed thaC the width of the characteris tics of
direcrivity for a level of 0.7, calculated according tfl the envelope of
the pulses, is 14� on the horizontal plane and 7~ on the vertical (Fig. 34).
The coefficient of concentration of the acoustic power is 550.
Along with the high directional properties, the dolphin's transmitting .
system makes it possible to form acoustic pulses, the temporal structure
of which depends on the direction of the propagation in space. If the
shortest and broadest-band signals (with the spec trum peak in the range
of 60-90 kHz) are transmitted in the direction of the rostrum (acoustic
axis), then with a deviation from the acoustic axis, the duration of the '
signals increases, and their spectrum, accordingly, is reduced in the
area of the lower frequencies.
A harmonic analysis of the integrated beam pattern, made on a BESM-6
electronic c~mputer, revealed that the nature of the formation of the
sound field for low and high frequencies was different in principle. If,
For the components of the spectrum in a range of 10-50 kHz, the directivity
is of a clearly marked double-lobe nature, with the minimum in the direction
of the acoustic axis up to (-10 ~-14) db, then in the area of frequencies
of 70 120 kHz a narrow emission.is observed. This fact makes it possible
to explain the changes in the spectral and temporal characteristics oi the
signals which are observed when they are recorded from different directions
(Ivanenko, Ivanov, Telekhov, 1977).
4. In the overall range of problems facing researchers of the dolphin's
echolocation system, a central place is occupied by the problem of the ,
animals' discrimination of the acoustic images, as well as by the problem
- of adaptation to a certain specific locational situation. '
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. g' -30' -20' -io' o lo' 10' 30' � e'
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f,e
~
~ s
~
. ~ .
~
~
_
R~ ~
~ ~ ~
~ i ~
' ~ ~
' ~ ~
~ ~ ~ .
' ~ ~
~ ~ ~
; ~ ~
~ ~ ,
~ ~ ~
~ i ~
~0' -f0' 0' 1D' ~0' !0' ~0' SO' f~~
Figure 34. Diagram of the D irectivity for the Emission, A--on the
horizontal, B--on the vertical plane
75
FOR OFFICIAL USE ONLY
APPROVED FOR RELEASE: 2007/02/08: CIA-RDP82-00850R000200020050-2

APPR~VED FOR RELEASE: 2007/02/08: CIA-RDP82-00850R000200020050-2
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