286

how is this relationship expressed in both ecological groups,

the aquatic and semi-aquatic mammals? In this scheme we have

attempted to examine simul:taneously the structure of a typical

rete mirabili of the dolphin, taking into account the possible

pathology, the structure of the mesenteric artery and of some

of the lymphatic nodes, representing in this case portions of

the lymphatic system.

MATERIALS AND . METHODS

We had four dolphin specimens available to us: three

common porpoises, Phocaena phocaena, and one bottlenose dol-

phin, Tursiops truncatus, from the Black Sea, and also four

fur seals, Callorhynus ursinus, two Steller's sea lions, Eume-

t0DiUS jubatus, and two ha:I:Lour seals, Phoca vitulina, obteined

on Robben Island in 1968.

Two of the investigated dolphins (one common porpoise

and the bottlenose dolphin) died from different diseases,

while the remaining two were dissected after an acute physio-

logical experiment.

Morphological and histological methods were used to

study sections of the intercostal, supraclavicular, axillary,

lumbar and iliac Portions of the retia mirabilia, sections of

the mesenteric artery, the mesenteric, axillary, pancreatic,

cervical and other lymph nodes. The sections were stained

with haematoxylin and eosin and resorcin-fuchsin.

The lymph nodes were measured in the fresh state; some

of them were weighed.

287 • 1- a

THE RETIA.MIRABILIA OF DOLPHINS

Common porpoises No. 2 and No. 3. The arterial net-

work in the intercostal region was investigated. The animals

were dissected after narcotization. The intercostal vascular

network is represented by a spongy tissue, composed of extreme-

ly coiled hollow vessels - the arterioles. Variations in the

diameter of these vessels are not great, from 200 to 1000 IL.

The lumen is gaping and filled with blood. The structure of

the walls of the vessels is typical of vessels of this category

(Figure 1). The internal layer is well developed, after this

comes a distinct thick-layered elastic membrane. In the tunica

media the circular smooth musculature is very dense and abun-

dant. It is bounded on the outside by a second elastic memb-

rane, which is also of considerable thickness and uninterrupted.

The tunica externa of the vessel is maximally reduced, it does 181

not contain collagen and is not firmly accreted to the surroun-

ding tissue. In this manner, the vessel is composed as it

were of a two-layered elastic tube, capable of withstanding

high pressure both from the inside as well as from the outside.

The tissue surrounding the retia mirabilia contains

lymphatic vess'els, sometimes covering the arterioles as it were

with a sheath. The veins form sometimes small sometimes wide,

very thin-walled sinuses. In places extravasations are evi-

dent in the dilated lymphatic capillaries and in the connective

tissue.

The erythrocytes which fill the arterioles are bi-

concave and sometimes crescent-shaped, which may indicate an-

aemia. In places arteritis obliterans was diagnosed, but no

infiltrates in the walls of the vessels were found.

288

p}!c. 1. RllyAectMa ceT6N 113 HaAx.110v11Tloif o6.nac•rf1 MopcKoi ► CIIIII[bil M 2

1-- etl^rpcttltstn 3.iacTUVCCI.aR oGoaolu<a; 2=-11apYacuast 3.7aCTH4CCFaA o1107o4Ha:.

, • :f - cpCAiIRA oûo, o,lha cocyAa; d- m1a+l( 14JTIMCCK11C cocyluIt.X8

Figure 1. Rete mirabile from the supraclavicular region.

of porpoise No. 2.

1 - inner elastic nnEmbrane;2 - outer elastic membrane;3 - tunica media of vessel;4. - lymphàtic vessels. X 8.

Next was studied the arterial "network" from the lum-

bar and iliac aorta (ventrally to this is situated the venous

"rete mirabile"). The general structure of the "network" is

similar to that in the thoracic cavity. All of the layers

are found in the walls of the arterioles. It seems, however,

that the elastic portions are here less strongly developed

and that the vessels are in a somewhat more collapsed state.

The cause of this is not cl.ear: of significance in this res-

pect may be the lowered pressure in the abdominal cavity, in

comparison with the thoracic cavity, the effect of the narcosis

on the autonomic centers, leading to spasms of the vessels,

or the proximity of nidi of helminth infestation.

289

In the lumbar "network" there appear sections, where

the walls of the vessels are • asophilic. From the tunica

interna, in the composition of which there are present muscle

fibres, cells are desquamated and fill the lumen of the vessel.

However alongside the described section there lie normal full-

value arterioles, typical of the retia mirabilia.

Common porpoise No. 1. The animal was dissected after

death, which occurred from an unknown cause. The thoracic

section of the retia mirabilia was traced from the intercostal

artery. The artery was not very elastic, the walls were friable

and the aorta itself was deformed. The rete mirabile starts

from the intercostal artery. Both of the elastic membranes

in the vessels of the network were damaged and the picture

resembled that of typical sclerosis. In the tunica media of

the arterioles there is clearly evident the large-mésh elastic

basis, while the continuity of the circular musculature is

infringed in places. Basophily of the walls and the occurrence

of collagen fibres in the tunica externa was noted.

Around the arterioles of the network there are clearly

expressed and very dilated lymphatic vessels. The veins here

are small and thin-walled. Within the thickness of the struc-

ture are encountered lymphoid follicles.

The bottlenose dolphin. This dolphin died of pneumo-

nia, which was confirmed by the dissection of the lungs. In

the arterial retia mirabilia the vessels lie in a very dense

mass. The diameter of the arterioles ranges within the same

limits as in the common porpoise. In their walls are represen-

ted all of the elements, including both of the elastic membranes.

290 • .1 •

Around the artery there are lymPhatic vessels, though these

are very narrow, and much fat tissue. The circular muscula-

ture in the tunica media of the arterioles is dense, abundant

and stains intensively with eosin. Collagen fibres were not

evident and, consequently, the vessel was in a state of full

tension and retained its elasticity. Apparently, the above-

described picture of arteriosclerosis was created by some other

pathological conditions.

The venous rete mirabile of dolphins. The venous net-

work is situated in the lumbar region ventrally to the aorta,

between the suprarenals. This structure is composed of small

calibre veins and venules. While the arterial network has the

form of a cushion of twisted vessels, the venous network is

composed of numerous anastomoses, braided into a lattice. The

vessels are sinuous, with compact walls. The normal layering

of the venous wall is here indistinctly expressed. Around the

veins are situated lymphatic vessels.

THE MESENTERIC ARTERY

It is known that the mesenteric artery of man has a

mainly circular arrangement of the musculature in its tunica

media. In the mammals there are also found bundles of longi-

tudinal smooth muscles. The longitudinal musculature in the

wall of the vessel provides for its peristaltic contractions,

giving rise to the so-called "peripheral hearts" (Zavarzin,

1939). In the animals studied by us the walls of this were

not uniformly constructed (Table 1, Figures 2-•).

*OTaeablIble peRKIIC no- :mum

480 •

32 40 30

162 280

400 720

foo 240 100 400

140

60

100

340

•180 180

Table 1. Thickness of the layers of the wall of the mesenteric artery, m.

Ta 6 auna 1

Tomnuna canea CTCHKII 6phiaceelmoil apTcpull,

291

Elements of treVer

'Harbour Stellerl I el cilnyq d .

sea-lion

Yur. Bottlenose mopcuow mwiuna

sel dolphin

A Burrpeminfi ujori .6e3 mycKyanyint

"Ilpopuribnan mycxyairrypa • B cpeRneti 0607otwe

C 11,upKyanpu an mycEya a- Typa 11 cpeaudi o6o-.710 , 1KC

D .1Iapywrian o6oacp.wa: . e naoTuan f pleaam

A - Inner layer without musculature

B - Longitudinal musculature in tunica media *Individual sparse fibres •

C - Circular musculature in tunica media D - Tunica externa:

e - dense f - loose

Bottlenose dolphin. The wall of the mesenteric artery

is compact. Its thickness in the bottlenose dolphin is about

620 f, which is about 6.5 times smaller than the outer diameter

of the artery. The tunica media is composed of circular mus-

culature, while the longitudinal musculature is very sparsely

interspersed in the form of individual fibres or bundles (Fi- 183

gure 2). The tunica externa has a thickness of 140m- in all.

There are few collagen fibres; the elastic part is clearly

expressed.

29 2

Puc, 2._CTeinca 6pbuKeet11lori apTepuu mopcKoil cmiltbut (Ns ,i 3):

Burrpotnyin actaouri; cpentinn o6otiogica .; 3 — tiapyxuiaii o6oaogKa.x8 .

Figure 2. Wall of the mesenteric artery of a common porpoise (No; 3)1

1 - tunica interna 2 - tunica media 3 -.tunica externa. X 8.

Seals. The overall thickness of the wall of the artery

is within the same limits as in the bottlenose dolphin; in

the fur seal and the harbour seal it is one sixth, and in

Steller's sea lion one fifth of the diameter of the artery.

Longitudinal musculature in these animals is found in all of

the layers of the wall of the vessel. This musculature may

lie in separate bundles in the center of the circular muscu-

lature (Figure 3), in a continuous layer along the periphery

Z93

4

^w.^^:^^1:::....r^a_.<à,^.w_.,... .^._....^.^.:.^^:.:,,.ks.;-.:.s^,.;M•.. . ^.x^^:;^=u:^

Pt+c. 3. CTCU!ca Gphl^+:ce'tnoii ap•rep!nt atopcr.oro xoin!:a (M, 3):

1- tutpsy.inpnaa ^q'cr.y^arypa cpeAnetl o6)o, ïo4^:n; 2 - npo;toat.ttanxycxy.nat)'ya cpcAned oGoao4xn; 3 - na wnan oGono4h3.?t8

Figure 3. Wall of the mesenteric artery of a fur seal (No. 3):

1- circular musculatùre.of tunica media,-2 -,longitudinal musculature of tunica media;3 - tunica externa. X 8.

of this or in both sections simultaneously, as it were "squee-

zing" the circular musculature and elastic base (Figure 4).

In those places where the longitudinal musculature is stronger

the elastic base is weak and barely evident.

The thickest layer of longitudinal musculature was

found in the Stelier's sea-lion, in which the ratio of its

thickness to the thickness of the circular musculature was

1811.

294

)'•

1„.

'''' • ''' '

Pitc. 4. C-reliKa 61) 1) clœytta: / —Bityrpeutinsi amogKa; 2— 3.rt acT imeci( a n NICNI Cipali a; 3— npop,om..- 11BSI MYCKYJ1TYP ii cpepmeit o6o.logEe; —Itirpxyosiptian mycKyazrry-

pa n cpeiweil o6o.runKe.X8

Figure 4 • Wall of the mesenteric artery of Steller's sea-lion:

1 - tunica interna; 2 - elastic membrane; • 3 - longitudinal musculature in tunica media; 4 - circular musculature in tunica media. X 8.

7 110. In the harbour seal this ratio was 3 110, while in the

fur seal it was 2 :10 (see Table 1).

it is possible that the tunica externa of the vessel

is also dependent on the strength of the contraction of the

longitudinal musculature. The thickness of the tunica externa

is'greatest in the Steller's sea-lion (only one tenth that of

the diameter of the artery, while in the harbour seal it is

295

one thirtieth the diameter, and in the fur seal - one twenty-

seventh). However, in the last two seals the tunica externa

is compact and is closely accreted to the tunica media. In

the Steller's sea-lion, however, the dense layer of the tunica 185

externa is not thick, in all being about 100 'A. in thickness;

over this there lies freely a thick and loose collagen layer,

which in some sections is 4 times as thick as the dense layer.

Probably, within this loose layer there may be accomplished

contractions of the vessel in the longitudinal direction.

LYMPHATIC NODES

The lymphatic nodes as a constituent part of the vas-

cular system are not homogeneous in the animals of the two

groups of animais under comparison:. A marked difference is

evident in the dimensions of the nodes of the dolphins and of 186

the seal (Table 2). For example, the packet of mesenteric

nodes of the dolphin is considerably smaller than that of any

of the seals. The weight of such a node in the dolphin is

170 - 200 gm, While in Steller's sea-lion it is 750 gm. In

the dolphin the nodes are dense and lie in the mesentry; in

the seals they are loose and enclosed in a sac, similarly to

the kidneys, ovaries and other organs.

Common porpoise No. 1. The cervical lymphatic nodes

were studied. In the nodes there were found abundant extra-

neous inclusions (possibly the eggs of helminths) about 12 r_ in diameter. The nodes were hyperemic. Plasmocytes were evi-

dent in the parenchyma, while in the sinuses there could be

seen typical reticular cells with processes, histiocytes with

14,3

15,8

• 6,1

11

13,8

10,6

7,8

139

200

147

158

71

107

103

203

195

120

163,5

114

12,0

I1x5

28x10

28x 9

I9x 8

10x 5

9x 3

700

500

170 — 200

2

12

6

Table 2.

Comparative characteristics of the mesenteric lymphatic nodes.

• T a6.neu.a 2

Cpasinnenbuan xapawrepic-nima me3enTernia.TIbHUX oP.AtcparviecKiix y3n0u

A .nniceie y3non e.11 ■ Iia

BHA rem .

Species

29 6

d % AJ11111C Keluutile C At TCAO

Bee ,riaKera.,

G 06wee 411C/10 yuou

1 rpeumalukciaii -TIO.ICIlb

2 •à.1çpcKoii . HOTIIK .bis2 3

3 MopcKoit 1(01I1K Ne 5

Cinirt •Nt, 1

.5 Jlapra .N'g I

6 .A(panima •

7 ..MapcKan c111111b11

A - Body length, cm B - Greatest leng:th of "packet" of nodes

C — cm d - % of body length

.

e - % of length of kidney

F - Weight of "packet", gin

G - Total number of nodes

1 - Harp seal • 2 - Fur seal No. 3 . 5 — Fur seal No. 5 4 - Steller's sea-lion No. 1 5 — Harbour seal No. 1 6 — Bottlenose dolphin 7 — Common porpoise .

large nuclei and cells with elongate rod-like nuclei. In the

trabeculae, infiltrated with cells, there were present collagen

and elastic fibres and musculature, which here was somewhat

weaker than in the other nodes.

297

Common porpoise No. 2. The lumbar lymphatic node

was studied. The structure of the node was not typical of

the mammals: the cortical substance was arranged along the

peri.phery of the node, however the follicles did not have a

spherical form but were rather thickened zig-zag shaped cords.

The capsule of the node is dense, over this is the muscular

mesenteric membrane, in which the node lies. The trabeculae

in the nodes are not wide,but they are dense and are composed

maxnly of muscle and elastic fibres. The sinuses of the node

are compzressed by the densely branching trabeculae, they are

completely choked up with histiocytes.. Possibly, these are 18?

modified reticular cells which have' rounded up and lost their

processes. In.this organ phagocytic properties of these cells

were not noted.

The nuclei of such cells of the node were light coloured.

The chromatin was disposed in a reticulum and fine clusters.

There were many large and medium-sized lymphocytes. There were

no reactive centers. In the sinuses there were found free

erythrocytes and eosinophilic leucocytes.

The mesenteric lymphatic nodes were studied. The whole

"packet" of nodes was 9 cm in length and 3 cm in width. The

nodes were irregular in form: in places the medullary layer

was found at the surface and was pierced'by a mass of blood-

carrying vessels. Spherical secondary nodules.were also not

found in these nodes. Trabeculation was strikingly expressed

in these nodes. The trabeculae branched so densely and so

finely that they occupied all of the free spaces in the node.

Thus, the sinuses of the node here also were not free, but were

298

CB111-11,11 J\re.). 2 Pm. 5. Me3eirrepnambnbtil enimtpanmecKiiit peg mopcnii LI,eirrpa.nbindi citnyc

I— cmiye; 2 — MHIZOTIILIC uillypbt.x8

Figure 5. Mesenteric lymph node of common porpoise No. 2. Central Sinus

- sinus; 2 - medullary cords. X 8.

filled with histiocytes and and a fibrous reticulum (Figure 5),

not at all similar to that which is formed by the reticular

syncitium in other animais. In rare places there could be

seen portions of the central sinus with an area of no more

than 70X 40}1. The histiocytes have a frothy cytoplasm and

extraneous inclusions. There were few true lymphocytes but

many eosinophils with strongly segmented nuclei.

Common porpoise No. 3. This dolphin, like No. 2 also,

was the freshest of our material and therefore the structure

of the nodes should be the most typical that can be obtained

under conditions of captivity.

299

The mesenteric lymphatic nodes were studied. The

cortical and medullary layers had the normal Proper relation-

ship, but spherical follicles were not found. The trabeculae

were dense and thick; in these there were found more smooth

muscles and elastic fibres, than were found in the preceding

animals. The sinuses were filled with a fibrous mass and

cells, which were similar to endothelial cells. All of the

arteries and veins are of the same caliber. Arteries witbout

a lumen, with a closed cavity. They resemble the arterial

sleeves of the spleen, which end blindly.

The pancreatic lymphatic node is complex: it is com-

posed of several nodes of regular form. Secondary nodules of

zig-zag form are present, but there are also spherical nodules

in which proliferation centers are evident.

The bottlenose dolphin. The overall structure of the

nodes is similar to that described above fvr the common por-

poise. A distinction is found in the finer branching of the

trabeculae and the richness of the musculo-elastic base in

these. The trabeculae surround the secondary nodules and cel-

lular cords. In cross-section the trabecula is composed of

several concentric layers of connective and musculo-elastic

tissue.

PinniPeds. The harbour seal. The meSenteric lymph

nodes form a "packet". The form of eàch node Is regular: the

spherical secondary nodules may be arranged in a crowded man-

ner in the surface portions of the node; in the center is an

extensive portion of medullary substance, The capsule of the

node is relatively thin. The trabeculae do not branch exten-

sively, they are thin, composed of connective and muscle tissue,

300

- with a moderate content of collagen fibres and a very

poor elasticity. Both the central sinus of the node as well

as all of the other sinuses are free of any fibrous elements.

The reticular cells form a weak syncitium within these sinuses.

There are many monocytes, as well as myelocytes and typical

plasma cells with a "cap-like" nucleus and strongly basophilic

cytoplasm. In some sinuses there is a mass of free erythro-

cytes.

The pancreatic lymph node was studied. The parietal

sheet of the capsule of the node is abundantly furnished with

vessels. This comprises the branched venous "rete mirabile"

of the seal, described earlier on other. organs (Sokolov, 1959;

Harrison and Tomlinson, 1956).

In the secondary nodules there can be seen the proli-

feration centers. In the medUllary cords there are many medium-

sized and large lymphocytes. Every secondary nodule on some

sections is enclosed by a trabecular membrane. Within the

sinuses there are found erythrocytes; - There are also cells

phagocytizing the erythrocytes. The latter fill the cytoplasm

of the cells.

Steller's sea-lion. The axillary lymphatic node is

of the regular form, but the secondary nodules may be somewhat

indistinct. The capsule of the node is very thick, in it, as

well as in the trabeculae, there are very many large collagen

fibres. In some places the collagen acquires basophilic pro-

perties. Individual collagen fibres may be encountered also

in the sinuses of the node. Elastic fibres are:distinCti.ni'. ,

the walls of the trabecular arteries.

301

Pile. 6. M.c3e1tTeponabilmit zumcparmecKnet y3e1 cnnytia Ueirrpaablibiri cuilyc

I — cimyc; 2 — mfaannbte tutlypbt; 3 —TpaGerzyma.X8

Figure 6. Mesenteric lymph node of Steller's sea-lion. Central sinus

1 - sinus; 2 - medullary cords; 3 - trabeculae. x8.

The thick and coarse-fibred trabeculae of the node

form large loops (Figure 6), as a result of which the Sinuses

are free for the lymph, both in the middle of the node as well

as in the region of the hilus. The cellular composition of

the sinuses is the saine.

The fur seal (adult male No. 3). The mesenteric lymph

nodes are of the regular form: the follicles lie around the

periphery. The form is spherical. The capsule is very thick,

but the trabeculae are not as thick as in the Steller's sea-

lion. They do not permeate the whole node and do not surround

the follicles. The collagen content in the trabeculae is more

moderate than in the trabeculae of the nodes of the Steller's

302

sea-lion. In the capsule of the node there were many sclero-

tized vessels. Fatty sections were found in the nodes. The

sinuses of the node are wide and free. Free lymphocytes and

eosinophilic leucocytes were noted.

CONCLUSIONS

To account for the structure of the retia mirabilia

E. Slijper (1962) put forward the proposal that in the thora-

cic cavity of the dolphin there is developed a high pressure,

which is considerably higher than that in the abdominal cavity

and other parts of the body. In addition, the arterial pres-

sure on the walls of the arterioles of the network from the

side of the heart is also great (Nagel, Morgane, McFarland and

Galliano, 1968). In such-a case the presence of the double

elastic membrane in the walls of the arterioles would be un-

derstandable. The poorly developed tunica externa of these

vessels and the exceptional paucity of collagen are, in all

probability, conditioned by the extreme lability of this struc-

ture.

The arterioles of the rete are surrounded by lymphatic

vessels; in the normal organ they are narrow, collapsed, while

in edema they are dilated. A similar picture is found in the

lungs of the dolphin, where the lymphatic vessels enclose, as

it were with a sheath, the pulmonary veins (Baudrimont, 1955),

which have a great capacity for contraction, since they carry

arterial blood and, possibly, regulate its flow. In this case

a heavy tunica externa of the vessel*is superfluous, while the

surrounding lymphatic vessels facilitate movement, playing the

mirabilia is, most probably,

animals are held in close and

in the end, to a deformatiun

303

role of a hydraulic shock-absorber. This shock-absorber may

also Protect the rete mirabile from the aspect of the intra-

thoracic pressure.

The fact that in one and the same animal the walls of

the vessels of the rete mirabile in the abdominal cavity are

less elastic than those of the thoracic cavity may, to a cer-

tain extent, demonstrate a difference in Pressure in the men-

tioned parts of the body. Besides the natural pathology

(pneumonia, arteritis, helminthoses etc.), the loss of elasti-

city by the vessels of the retia

a common phenomenon when aquatic

shallow enclosures. This leads,

of the vessels and to death.

The vessels of the venous rete mirabile, taken by us

from the lumbar region, in all probability are not typical.

They should be studied on more extensive material.

In seals under water the activity of the heart is

slowed down (Harrison and Tomlinson, 1962; Elsner, Franklin,

Citters and Kenney, 1966). By vasoconstriction and peristal-

tic contractions the peripheral vessels may set up an extra

pressure in these conditions. Among such vessels may be in-

cluded also the mesenteric artery, which possibly functions

even more intensively in this respect in the seals, which is

indicated by the abundant longitudinal musculature in its wall.

The active operation of this artery as a "peripheral heart"

organ (Zavarzin, 1939) should be expressed more clearly in

Steller's sea-lion than in the other seals and dolphins. This

morphological feature is probably connected with the peculia-

rities of the ecology of this species of seal.

304

• The mesenteric artery supplies both the intestine and

the lymph nodes which lie in the mesentry of the small intes-

tine. Undoubtedly there exists some sort of relationship bet-

ween the structure of the vessel and the organs which that

vessel supplies..

Slijper (1962) wrote that whales have an exceptionally

large number of very large lymph nodes. Rakhimov (1968) des-

cribed 7 groups of nodes in the dolphin. Not long ago a de- 191

tailed study was carried out of five lymph nodes in Black Sea

dolphins (Moskov, Schiwatschewa and Bonew, 1969).

In the Caspian seal Kurdyumov . (1962) noted a small

number of large lymph nodes. We found 18 groups of lymph nodes

in seals and a somewhat smaller number in dolphins. There

is no doubt however as to the difference in the size of the

nodes in the two groups of mammals (Table 2). On account of

what structural characteristics and tissues are the nodes of

seals enlarged in comparison to the nodes of dolphins? In

the first place, apparently, this is due to the free dilated

sinuses and lymphatic vessels, which is not found in the dol-

phins, where the sinuses are narrow and filled with densely-

fibrous tissue of the musculo-elastic type. In addition, the

nodes of the seals, and especially of Steller's sea-lion, are

exceptionally rich in connective tissue (in the form of large

loose collagen fibres), which is scarcely noted at all in the

dolphins. A third cause of the differences is the non-uniform

disposition and composition of the lymphoid tissue and its

disposition in the nodes. In the dolphins the nodes are most

frequently of an irregular structure, the cortical substance

305

does not form the usual spherical follicles but is concentra-

ted in the zig-zag shaped sections, which occupy various po-

sitions in the nodes, including also a central position. The

medullary portion of the nodes is considerably reduced.

There is little.typical lymphoid tissue in the form

of small lymphocytes within the nodes of dolphins, it is fre-

quently replaced by plasmocytes and eosinophils. Such a pic-

ture is observed both in dead animalsas well as in those that

have been specially narcotized. Histiocytes predominate in

the sinuses. Our observations are also confirmed by other

data ( Moskov, Schiwatschewa and Bonew, 1969).

The nodes of seals are richer in actual lymphoid tis-

sue, but here also there is a completely different picture from

that found in the nodes of the majority of terrestrzal mammals.

Within the sinuses of the lymph nodes of both dolphins

and seals there are frequently present free erythrocytes and

eosinophilic leucocytes. In the seals here there are found

macrophages with erythrocytes in their cytoplasm. Slijper

(1962) observed, under a microscope., the process of the break-

ing down of the erythrocytes in the lymph nodes.of the dolphin,

and in connection with this he proposed that the nodes partial-

ly take on the role of the spleen,in this process. However

the size of the spleen, according to our data, is larger 4n

those animals with larger lymph nodes. For example, the length

of the spleen in fur seal No. 3 was 4.3 cm, in the harbour seal

- 23 cm, in the bottlenose dolphin - 7.5 cm and only 2 cm in

the common porpoise.

306

The role of the mesenteric nodes in the deposition of

lymph has been mentioned more than once in the literature

(Bouviere and Valette, 1933; Horstmann, 1952; ghdanov, 1952;

Rusn'yak, Fel'di and Sabo, 1958). It is possible that in the

seals a large part of the lymph is deposited in the lymphatic

nodes, while the lymph nodes of dolphins, which have a great

contractine4 capacity in their trabecular base, may regulate

the movement of the lymph.

REFERENCES

1. Zhdanov D. A. The general anatomy and physiology of the

lymphatic system. Leningrad, Publ. Medgiz (State

Publishing House of Medical Literature), 1952.

2. Zavarzin A. A. A course in histology and microscopic

anatomy. 5th edition. Leningrad, 1939.

3. Ivanova E. I. The morphology of the lymphatic system of 192

the Russian desman. Trudy Khoperskogo gosudarstvennogo

zapovednika (Transactions of the Khoperskii State

Reservation), issue 4, 1970.

k. Kurdyumov N. A., The forms of the efferent lymphatic

pathways of certain animals in a comparative-anatomical

interpretation. Trudy Odesskogo meditsinskogo instituta

(Transactions of the Odessa Medical Institute), vol. 16,

1962.

5. Rakhimov Ya. A. The thoracic duct of mammals. Trudy

Tadzhikskogo meditsinskogo instituta (Transactions of

the Tadzik Medical Institute), vol. 94, 1968.

30?

6. Rusn'yak I., Fel'di M. and Sabo D. The physiology and

pathology of the lymphatic circulation. Budapest,

1957 (translation).

7. Sokolov A. S. Characteristics of the size and disposition

of some of the blood vessels in aquatic (seal) and

terrestrial (dog) mammals. Arkhiv anatomii, gistologii

émbriologii (Archives of Anatomy, Histology and Em-

bryology), 37, No. 12, 1959.

9. Baurdimont A. The structure of the pulmonary veins and

functional circulation of the lung of the common dol-

phin, Delphinus delphinus L.

13. Horstmann E. The functional structure of the mesenteric

lymphatic vessels.

15. Monro A. Comparison of the structure and physiology of

fish. Translated and annotated by J. G. Schneider.

Leipzig, 1787.

16. Moskov M., Schiwatschewa T. and Bonev St. Comparative

histological studies on the lymphatic nodes of mammals.

The lymphatic nodes of the dolphin.

19. Rouviere H and Valette G. The role of the lymph nodes

in the circulation of the lymph.

20. Slijper E. F. Comparative anatomy and systematics of

the Cetacea.

, 3 0 8

- JIHTEPATYPA

1 }Kaatioa ,IL. A. 06utan aturromuu H dnisno,norns: minulumotecuoil enure:Au. II., Mearna, 1952.

2 Sana patin A. A. Kypc rucToaoruit u muKpocuornmecxort auaTomiln. 113/t. 5-e. JI., 1939. . - • . 3 Ileanoaa E. PI. K mop(I)onorini allM(1)HTIPICCIZOÙ ClICTCNIbl pyecKori abtxyxc,in. .. • • lpy,i1A honepcicoro row,. aanoneAli., HUH. 4, 1970.

1.1.1(ypihomoa IL A. (1)0pm:A OE:1301511101x .7iiimdm -rmiecmix lire neKoTopmx >titiBoTimix u cpannuTeabilo-ana -rominiecKom ocoemeitint. Tpy,am OHCCCHCWO me,u.. lin- Ta , T. 16, 1962.

5 Paxu.moa 51. A. rpya,noil upoToK maeizonirraionnix. Tpy.alit Taa.uniNcKoro Nteil. WI-Ta, T. 94, 1968.

0 Perlb.r/K. H., cheizbau M.,.Ccr5o ,r7. (1)113nozonin Ii naToaormtemulioo6paine- muf. ByAmicwT, 1957 (iiepenoR). •

7 CoKome A. C. Oco6einiocTit Kaill6pa ii pacnpeaeâeium ItexoTopmx uponenoc-:lux cocyp,on y . no•rnimx (Tioaenb) il na3eminAx (co6amt) mneNoutrraiounix. Apxlin one.. i'lleT0.1. II Dm6pitomr., 37, ,N2 12, 1959.

• bi Bran K. .1■ I., Murdayglt H. V., Millen Jr. J. E., Robin E. D. Arterial constrictor res panse in a divin,,(Y Mammal. Science, vol. 152, no 3721, 1966.

9 Baurdinzotzt A. Structure des veines pulmonaires - et -circulation functionele du poumon du dauphin Common. Delphinus delphinus L. Bull. de friicrosccipie Apoliquce. 2 Serie. T. 5, no 5 et 6. 1955. . 1 0 Elsner R., Franklin D. E., Van Citters R, L, Kenney D, W, Cardiovascular Defense against Asphyxia. Science, Vol. 153, no 3739, 1966. 11 Harrison R. J., Tomlinson I. D. W. Observatibris on the venous system in . scitain Pinnipedia and Cetacea. Proc. Zool, Ces Soc. London, vol 126, 205, 1956. 12 Harrison R. J. and Tomlinson J. D. W. Anatomical .and physiological adap-Ohms in diving matnrnals. Viewpots in biology, 2..1963.

,

13 Horstmaruz E. Llber die funktionelle Struktur der mesenterialen Lymphgela–iCe.. .,, lorph. Jahrb. 91. 483, 1952. . . 1 e Irving L. Scholander P. F., Grinnel S, W, The. regulation of arterial blood pressuve in the seal during diving. Amer. Physiol. 135. 557, 1942,

, 1illonro A. Vergleichung des liaues und der Physiologie der Fische. Uebersegt FillC mit Bernerkungen versNien von Schneider, J. G. Leipzig. 1787. ,.....

, 10 Aloskort M., Schiwalschezea T., Bonev St. Vergleichshistologische Untersu- chimg der Lymphlmoten der Seg"er. Die Lymphlmoten des Delphins, Anat. Anzg.

' U.)j., N 1, 1969. . J. 7 Nagel E. Z., Morgane P. Z., McFarland W. L. Galliano R. E. Rete mirabile of Dolphin. Its pressure—damping effect on cerebral circulation. Sciense, vol. ,16k no 3844, 1968. lbh - Onzinaney F. D. The vascular Networks (Retia mirabilia) of the Fin—Whale (Balaenoptera physalus). Discovery Reports, 5, 1932. 19 Rouvicre H., Valette G. Role des ganglions lymphatiques dans la circulati- e de la lymphe. 13011. Acad. Med. 110. 189. 1933. '

, ZOSlijper E. F. Die Cetaceen vergleiehend-anatomisch und sYsternatisch. Ca-el, zoologica, VII. 1936. .

lijper E. F. Whaies, Hunchinson of London. 1962. . .. 22 Tyson E. Phocaena or the Anatomy of a Porpess, .dissected at Gresham Colledge. London. 1680. , . . .

309

UDC 599.5 A. S. Leontyuk

PECULIARITIES OF THE DEVELOPMENT OF THE BLOOD-VASCULAR 193

SYSTEM DURING THE EARLY EMBRYOGENESIS OF CETACEANS-

The aquatic mammals differ from the terrestrial mam-

mals by several characteristics, associated with the adapta-

tions to the aquatic medium and, above all, with the periodic

arrhythmic breathing and with the blood circulation. In the

blood-vascular system of the cetaceans there are known several

adaptations, which serve for providing the tissues with an

uninterrupted supply of oxygen. Many authors have noted the

considerable development and peculiarity of the vascular ple-

xuses in cetaceans, that are disposed from the base of the

skull along the thorax and form the so called "retia mirabilia".

Also characterized by having a considerable development in the

cetaceans is the subcutaneous network of blood vessels (B. A.

Zenkovich, 1938; A. G. Tomilin, 1946, 1951; V. E. Sokolov,

1958, 1963; V. M. Berkovich, 1961; Utrecht, 1958, and others),

while in the refion of the flippers there have been described

specific complex vessels (A. G. Tomilin, 1946-1962). To the

latter is alloted an important role in the processes of thermo-

regulation.

The adaptations of the cetacean embryos to supplying

the tissues with oxygen, in connection with the peculiarities

of the respiration and blood circulation of the mother, as was

noted by P. A. Korzhuev and N. N. Bulatova (1951), present an

"even greater puzzle". According to the data of these authors,

310

in cetacean embryos there is noted a high oxygen capacity

of the blood and a high concentration of haemoglobin, repre-

senting an adaptation to unfavourable conditions for embryo-

nic development. It is natural to assume that such adapta-

tions are not the only ones,and are reflected in se-

veral morphological charac -Éeristics of cetacean embryos. In

this connection we conducted studies of the vascular system

in cetacean embryos and directed our attention to adaptations

for ensuring a regular supply of oxygen to the tissues.

Serving as materials for these studies were complete

series of sagittal and transverse sections of cetacean embryos:

of the sperm whale-from 8.5 to 105 mm in length, of the

humpback whale - from 11.5 to 125 mm, and of the fin whale -

from 54 to 73 mm, stained by the Bielschowsky-Buke method, the

Nissl method and with haematoxylin-eosin. The value of silver

staining methods for revealing vascular networks has been con-

firmed on more than one occasion in the laboratories of V. V.

Kupriyanov, D. M. Golub, D. A. Zhdanov and others. The mate-

rial for the study was obtained thanks to the cooperation of

V. A. Zemskii, G. A. Budylenko and D. D. Tormosov, staff mem-

bers of the Atlantic Research Institute of Marine Fisheries

and Oceanography.

The results of the study show that the formation of

the vascular plexuses in cetaceans already occurs during very

early stages of embryonic development. At the same time the

vascular plexuses are formed in their most clearly expressed

form around the brain and spinal cord and around the spinal

ganglia. In a sperm whale embryo 14.5 mm in length, along the

periphery of the spinal cord there was developed a network of

311

•

01

thin-walled capillaries, filled with the formed elements of

the blood. Spreading out cranially, this network surrounds

the developing brain and initially is more clearly expressed

in the region of the posterior cranial fossa at the base of

the brain. The enrichment of the vascular networks proceeds

very intensively, and in the sperm whale embryo 16.2 mm in

length there appear at the base of the skull a considerable

number of large thin-walled veins, surrounded by a network of

finer vessels (Figure 1). Then the vascular system in the

cavity of the skull is constantly represented by very exten-

sive plexuses. At the same time there occurs an enlargement

of the diameter of the large venous trunks an an increase in

the number of small vessels in the network. In the older

group of sperm whale embryos (95 - 105 mm) there are found

extensive vascular plexuses in the cavity of the skull, that

are formed of both the large main veins as well as by the net-

work of small vessels of various diameters (Figure 2). The

vascular plexuses attain a considerable development in the

region of the base of the hypophysis, where they form the basis

of the sinus cavernosus. An intensive development of the vas-

cular networks in the region of the brain and spinal cord is

also characteristic of the embryos of the baleen whales. Thus,.

in a humpback whale embryo 95 mm in length the vascular plexus

around the brain was formed of thin-walled veins and arteries, 195-

located in all parts of the skull and and also extending out

onto the outer base of the skull through its openings.

A second zone of the extremely strong development of

the vascular plexuses is the region where the intervertebral

nodes and the initial sections of the spinal nerves are located.

312

pile. I. Cocy.ji+c'roe CII.Iereune 13 acnouauF nt 11epC(la 3a})o1LtIIla Kj•uiano-ra 16,2 srst ;iauuoil. i%l[ispocj>oro. 06- 2. or. 7

Figure 1. Vascular plexus at the base of the skull in a

sperm whale embryo 16.2 mm in length.

Photomicrogaph. Objective 2, acular 7.

Ÿ`.«^na.v..w.vv.«+..Fv.^'^:ii^+:'.l..t... .^. 7^ t•.'.L^:{ F^: ivi.. .:.117^:f13^^.i:.'.:'..^^t.:.:i: àF::i4r.J..:i.t.:.^

^^^ . . ^ ^" - ._. ...... ^.^ `.. . ^^'._ . . . ..... ^

Here there are formed extensive venous plexuses, surrounding

the spinal ganglia. These plexuses, which connect up with

the internal plexuses of the vertebral canal, are considerably

developed in all of the cetaceans (in both the toothed and the

baleen whales), and.are characteristic also of other mammals

and man. Thus, in a humpback whale embryo 95 mm in length

the venous plexuses are formed of blood vessels.of various

diameter, the branches of which penetrate even right into the

ganglia (Figure 3). In the sperm whale such plexuses in the

region of the spinal ganglia are found in an embryo 23 mm in

length, while later they become more and more extensive. A

characteristic feature of the blood vessels which form the

networks and plexuses in the region of the central nervous

system is the very small thickness of the vascular wall of

both thè arteries as well as the veins. This phenomenon,

which was noted in a dolphin embryo in the region of the

Pile. 2. COCYJUICTOC cnoe-re.ime B 0C110-

B8111111 neperia aapo.abuna KaWaJlOTa 97 AIM NI111101 .1, MinipmpoTo. 06. 2, OK. 5 V..

Figure 2. Vascular plexus at the base of the skull in a

. sperm whale embryo 97 mm in length. Photomicrograph.. Objective 2, ocular 5.

31 3

branching of the internal carotid and vertebral arteries by

E. S. Yakovleva (1951), is probably common to all of the

cetaceans and leads to a convergence of the conditions of

gaseous interchange in the arteries and veins.

Vascular networks and plexuses also develop along the 196

course of the large trunks of the cranial and spinal nerves,

where they are considerably expressed. The vascular plexuses

which accompany the spinal nerves are continuations of the

vascular plexuses of the vertebral canal. They are revealed,

for example, in a sperm whale embryo 65 mm in length, in the

form of a dense network surrounding the intercostal nerves.

A similar picture is found in a fin whale embryo 55 mm in

length. in the region of the trigeminal nerve and its branches

314

Par. 3, Cocy.aucToe cuaereane 13 n03a0•

110 ,1/10111 Eatia.ue II ao nepaci)epail rpy1-

111,1X cilainiomoaroabtx yanoe aapownia rop6aToro Rirra 95 .ADI Rau MIII:po-

4/0T0 . 06. 2, olz. 5

Figure 3. Vascular plexus in the spinal canal and along the

periphery of the thoracic spinal ganglia in a humpback whale embryo 95 mm in length. Photomicrograph. Objective 2, ocular 5.

in sperm whale embryos 97 and 105 mm in length therb is formed

an extensive network of vessels, entwining the trunks of the

nerves and giving rise to branches which penetrate into the

trunks. An extensive vascular network is developed also along

the course and within the trunk of the vagal nerves. The vas-

cular networks within the trunks of the vagal nerves are also

manifested in the embryos of the baleen whales: thus, in a

fin whale embryo 55 mm in length they were traced along the

course of the nerves over a considerable distance. The sympa-

thetic trunks are surrounded by developed vascular plexuses,

connected with the plexuses of the vertebral canal. In the

55 mm long fin whale embryo there are here formed wide-looped

• 315 à '

•

networks. The circum-sympathetic vascular plexuses are very

well expressed in the sperm whale embryos. Already in the

early stages of development of the sperm whale (in embryos

14.5 mm in length) along the course of the sympathetic trunk

there is found a network of blood vessels, having thin walls,

which surrounds the sympathetic trunk and its branches. Sub-

sequently, for example, in a 55 mm long sperm whale embryo

there develops around the sympathetic trunk an extensive net-

work of venous and arterial vessels. A similar vascular net-

work is also retained in the adult cetaceans. In the Atlantic

white-sided dolphin along the vertebral trunk there extend two

strands of vascular plexuses, within which are situated the

sympathetic trunks. In their structure these vessels resemble

the structure of vessels with endarteritis obliterans, when

the flow of blood is very considerably hindered (A. B. Khodos,

1967).

Thus, during the embryogenesis of cetaceans (of both

the toothed and the baleen whales) there is manifested in a

regular fashion the formation of the very strongly developed

and widely distributed vascular plexuses, formed by arterial

and venous vessels in the region of the brain and spinal cord,

of the cranial and spinal nerves, and of the sympathetic trunk.

We believe that such a localization of the vascular plexuses

is not fortuitous but is connected with the necessity for en-

suring the supply, in the first place - of oxygen, to the

centers of the nervous system and nerve trunks.

As has been noted by several investigators, in the

cetaceans the vascular networks and plexuses are also consider-

ably developed in other parts of the body. We directed

316

Pile. 4. Tonorpmpitsi DKerpaRapiLiianbiloro cocyaucToro cum:- realm y 3apo2t.butta Eatila.10Ta 55 .A1:11 ,/(,11111Ort:

/ cocyglictoc cluterelime: 2 — ani(Jaitiza JierKoro: 3-- 3a1uIEGINa Ka; 4-- gpoeemmilite cocyRbs 11 liPII7pa.141106 trverwe Tyaoainusa.

pegeutil cpe3. MeKpoilio -ro. 06. 2., OK. 5

Figure 4. ToPography of the extracardial vascular plexus in a sperm whale embryo 55 mm in lengths

1 - vascular plexus; 2 - lung rudiment; 3 - heart rudiment; 4 - blood vessels in ventral wall of body.

Transverse section. Photomicrograph. Objective 2, ocular 5.

cepn-floue-

attention to the richness of the vascular system, especially

of the venous part, primarily in the sub-pleural space of the

.thoracic cavity. L. B. Slavochinskaya (1965) considers that

the abundance of veins is explained by the level of metabolism

in the pleurae: the veins in the pleura participate in the

regulation of the metabolic processes in the thoracic cavity.

Under the parietal pleura in the region of ribs I - III and

the ventral part of the mediastinal pleura, commencing from

comparatively early stages (sperm whale embryos 34-35 mm in

length), there is formed a network of blood vessels, which

takes the form of an extensive plexus. In the region of the

dome of the pleura this plexus passes over onto the mediastinal

Pue. 5. Tonorpacimm 31:erpaRapirmanbito -

ro cocyRucToro cnacTeulta y aapoRbuua i:inuaaoTa 97 01,1(

3a1eam:a cepRu,a; 2— CT I30.1 Ariad)par-maabtioro neprui; 3— cocyttra

naenpm; 4 — cocyAbt, Ha rpyuoil noBep:otucTif Julad,parmbi. CarirrTam.nEtil cpe3

Mutcpoci)o-ro. 06. 2, OK. 5

317

Figure 5. Topography of the extracardial vàscular plexus

in a sperm whale embi'yo 97 mm in length:

1 - leart rudiment; 2 - trunk of diaphragmatic nerve; 3 - vessels of mediastinal pleura; L. - vessels on thoracic surface of diaphragm.

Sagittal section. Photomicrograph. Objective 2, ocular 5.

pleura and extends ventrally to the root of the lung to the

ventral wall of the body, connecting up ventrally with the

plexuses which are situated under the costal pleura (Figure 4).

Caudally a similar plexus extend down onto the diaphragm and

is distributed on its thoracic surface (Figure 5). As a re-

sult of this the ventral section of the thoracic cavity, in

which is situated the heart in the pericardial sac, is found

to be surrounded by the practiâally continuous extracardial

'vascular plexus, which is situated on the surface of the peri-

cardium. In its structure this network is composed of vessels

of two types: smaller iressels with a gradually forming wall

318

(of the arterial or venous type) which anastomose extensively

among themselves; the second type of vascular structures has

the form of thin-walled lacunae, characteristic of micro-

circulatory adaptations for increasing the capacity of the

vascular stream (V. V. Kupriyanov, 1969). In our preparations 198

these were found to be empty or only partially filled with

blood. Venous sinusoids of this type are considered in the

literature as a part of the cavernous-like formations (A. N. 199

Tikhomirov, 1967). Both types of vessels appear comparatively

early and can be distinctly found in embryos 55 and 65 mm in

length and larger. They are more markedly developed in embryos

97 - 105 mm in length (Figure 6). It should be noted that

the vascular network in the region of the parietal pleura is

distinguishable by its content of vessels of the arterial and venous

type. Characteristic of the mediastinal region, on the other

hand, is the presence of a large number of venous lacunae and

cavernous-like formations alongside the rich network of arterial

and venous vessels. It is interesting that this region in other

mammals is also distinguished by having a characteristic struc-

ture of the vascular system, which has been noted by several

authors (V. V. Kupriyanov, 1969; R. M. Petrova, 1967; L. B.

Slavochinskaya, 1967). Well expressed and widespread extra-

organic vascular plexuses are found in the region of the heart

and especially along the course of the large vessels which run

out of the heart, and further along the course of the aorta

and its branches (Figure 7). These pericardial and periarterial

venous plexuses anastomose extensively with the extracardial

vascular network in many sectors.

319

•

46

fin whale. IE

55 m771 in length we also find a fairly considerable network of 200

blood vessels, though this is more localized, primarily in the

dorso-cranial sectors of the thorax. This network is formed

mainly of arteries and veins. In the mediastinal pleura the

vascular plexus is considerably poorer and is represented by

a considerably less well expressed network of arteries and veins.

Pac, G. CTpyhTypa si.c•rpaxapaEEa.qi,uoïl cocY.aucToii ceriE sapoabi-wa KaA1aOTa 105 Arar ;1,-iuiEOïr.

I- cocy,ai+crbie nahyma; 2- CCTb TOIfKOCTC111i1.IX cocY;Zos. 91Eir.p04)0T0.06. 2, os. 5

Figure 6. The structure of the extracardial vascular net-.

work of a sperm whale embryo 105 mm in length:

1 - vascular lacunae;2 - network of thin-walled vessels.

Photorricrograph. Ubjective 2, ocular 5.

Thus, the disposition of the vascular plexuses in the

thoracic cavity of the sperm whale embryo is distinguished by

the marked peculiarity of the topography and the character of

the blood vessels. Within the thorax of the embryos of the

baleen whales (humpback whale, fin whale) the picture of the

disposition of the vascular networks differs markedly from

that described above. In the subpleural layer of a

I.

S)

Pic. 7. CocyglicToe clue:twine B o6Jtac - , Tit jyrn aopTbt 3apoluatia uall.laaoTa

97 it creitKa aoprm• 2 — cocymicroe cimere7-

Hue. Mulzpo(loTo. 06. 2, ow. 5 •

32 0

Figure 7. Vascular plexus in the region of the arch of the

aorta of a sperm whale embryo 97 mm in length: • 1 - wall of aorta;

2 - vascular plexus.

Photomicrograph. Objective 2, ocular 5.

Among the ecological characteristics of both the baleen

as well as the toothed whales, which have been noted by many

authors (A. G. Tomilin, S.E. Kleinenberg, V. M. Bel'kovich,

V. E. Sokolov and others), belongs the considerable develop-

ment of the network of subcutaneous vessels. The vascular

pattern can be seen distinctly through the integument of the

embryo, commencing from the earliest stages of development.

The architecture of the vascular pattern is especially well

marked out in the region of the head, the base of the flippers,

the base of the tail and the trunk. Only the development of

the pigment layer of the skin gradually conceals this vascular

pattern. With histological studies the development of vascular

networks is found at the base of the flippers and around the

• • 32 1

Pile. 8. lloRimmulan cocyRucTan cen, B 06.1111CTIL GOKOBal c-remui Ty.nounuut aapubmia KannknoTa 105 .11.11

MilKp04)0TO. 06. 2, ()K.' 5.

Figure 8. :Subcutaneous vascular network in the region of

the lateral wall of the trunk of a sperm whale

embryo 105 mm in length.

' Photomicrograph. Objective 2, ocular 5.

phalanges of the fingers, along the course of the bundles of

muscle fibres and onder the skin of the flippers (sperm whale

embryos 72 - 105 mm in length). Extensive vascular networks

are formed under the skin also in the muscle layers of the

trunk (Figure 8).

The data from our studies demonstrate the considerable

richness of the vascular system in cetacean embryos, especial- 201

ly of the toothed whales (sperm whale), manifested in the de-

velopment of vascular plexuses in various parts of the body,

mainly in the zones where vitally important organs are situated:

the central and peripheral nervous systems and the heart. In

this connection, naturally, the question arises as to the bio-

logical significance of such adaptations, their role in the

development of the embryo and, subsequently, in the functions

322

of the adult animals. The concept exists in the literature

that the vascular networks and plexuses in the body of ceta-

ceans fulfil an important, if not fundamental, role in supply-

ing the tissues with blood during diving, functioning as a

store of blood that is rich in oxygen and shortening the path-

way of the blood to the tissues (B. Howell, 1930; F. Ommaney,

1932; A. Krog, 1930; P. Schollander, 1940 - cited by S. E.

Kleinenberg, 1946).

Sharing this point of view, S. E. Kleinenberg empha-

sizes that during diving there is a decrease in the number and

strength of the cardiac contractions and, consequently, the

vascular store permits the blood reserve of oxygen to be more

economically expended and facilitates the work of the heart

during prolonged apnea. Our data permit one to consider that

such adaptations to a prolonged shut-down of external breathing

and the creation of stores of oxygen in the blood are formed

already during the early stages of embryonic development. It

should be borne in mind, however, that the marked development

of vascular plexuses in the region where the centers of nervous

regulation are situated is characteristic not only of the ce-

taceans but also of other mammals, including also man (A. S.

Ieontyuk, 1969), which are not characterized by having exten-

ded periods of apnea. Therefore it is possible to somewhat

expand the concept of the role of the vascular plexuses, which

are formed already in the embryos of the cetaceans and are

retained in the adult animals.

The character of the extra-organic, mainly venous,

plexuses in the region of the brain, spinal cord and peripheral

323

nerves is similar to those in tissues with an increased ex-

change (A. B. Miodos, 1967), since for obtaining the optimal

exchange the rate of the blood-flow should be reduced to a

minimum. It is probable that the developing vascular plexuses

are adapted in their structure primarily to the relative

hypoxia and moderate metabolic acidosis that is so character-

istic of developing embryos (I. A. Arshavskii, 1960; L. S.

Persianinov, 1967, and others). That this is actually the

case_is convincingly indicated by the data of several.authors

on the effect of hypoxia on the structure of the vascular

system (V,. I. Voitkevich, 1958'; E. N. Domontovich, 1958; E.

Van-Lir and K1. Stiknei, 1967; V. V. Kupriyanov, 1969, and

others ), which is manifested in an increased vascularization

of the tissues, a hypertrophy of existing vessels and a growth

of newly formed vessels. The resulting hyperemia leads to an

increased diffusion of oxygen, food substances and metabolites.

The changing conditions of the regional circulation of blood

favour a more complete assimilation of the oxygen in the blood

and the excretion of metabolic products, since the diffusion

of the oxygen from the blood is related to the area of the

diffusion and the partial pressure of oxygen in the venous

part of the vascular flow (A. I. Shik, 1966). V. V. Kupriyanov

(1969) emphasizes that the length of the venous pathways,,_the

diversity of their directions and the extensive roundabout

connections predetermine the pattern of the microcirculatory

system and, in a decisive manner, influence the hemodynamics

in the organs. Convincing data on the considerable reconstruc-

tion of the intra-organic vascular stream in cases of hypoxia

324.

under high mountain conditions were presented by Ya. A. Rakhi-

mov and L. E. Etingen and co-authots (1968).

Thus, the extra-organic network of blood vessels, main-

ly of venous•vessels with a retarded blood flow, which appears

in the early stages of development of cetaceans may be consi-

dered as an adaptation for supplying blood to the tissues in

the specific conditions of the intra-uterine development of

the foetus. In this connection the development of the special

vascular structures during the embryogenesis of the cetaceans

becomes understandable, and these may be considered not only

as storage structures in places with a maximal requirement for

oxygen but also as adaptations for the more complete assimila-

tion of the oxygen in the blood, and for the excretion of meta-

bolites in the zone of the disposition of the vitally impor-

tant organs and systems - the nervous system and the heart.

Taking into consideration the conditions of the life of the

toothed whales and the baleen whales, the more clearly expres-

sed development of the vascular plexuses in the toothed whales,

as compared to the baleen whales, is understandable as corres-

ponding to their ecology. It is evident that the adaptive

characteristics of the vascular system, which are set down in 203

early embryogenesis, retain their significance and also, pro-

bably, acquire new properties in the adult animals.

J111TEPATYPA a 1 Apumacutiii H. A. Onsno.Tiorun RpoBoo6pamemin BO nnyTpuprpotillom nepli- .

0A,e. M., 1960. . Beizbuoauct 13. M, 0 (1)11311 11CCKOrl Tepioperyantuni y 6e.aymi. Tp. conem. lixTit-

o onniecxoil KONHICCIIII AH CCCP, inn 12, 1961. •

Ban-Jl up 3., Cruuneii /(4. FlinoNcuil. M., 1967. • . Botiruearm B. H. 13.rinsinue xponinecKoro uncoopo,altoro r000.aantio na Epo-

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-rem pycciwii nbixyxo.nit. B vii.: «1op4loaorittiecuile oco6ounocTii BOP,111,1X mneHoini-Tammilx». M., «Haymi», 1964. • 8 Haanoaa E. II. Honoe e COCT051111111 «npecuoii CCTII» ,u,epuBwroplibix mexamis-

MOB y nenoTopmx rionyBoailux H■ 11130THIJIX. LI,olulaa..1,1 AH CCCP, T. 172, D1,111. 2, 1967. . . •

%. Ii &MOO(' E. II. OCO6e1/110CTII COCY,IIICT0ii CHCTC:111>1 131U1: 11/1X 14.1C1:01111T111011111X.

I Bcecomslioe cc-mental-me 110 liaplenum mopcmtx M.1C101111TD10111.11X. M., 1969.

1 9 Kneiinen6è pe C. E. M•neKonwraimune Lleplioro H 1-‘30eCKOTO Itropeii. M., 1956. 1 Kopmyea H. A., Bynaroaa H. H. Oco6ennocTil HbrikITC.11b110r1 (llyilmilin Bponit

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•Nicara II cnitimomosronidx Y:30011 D 9M61/110TCHC3C. B ii i.: <1P1111111C1/11:111.119 11 penac- By.impitaanitn nnyTpenitux opranon meToRom opralionencini». Muncx. 1969. • •

14: Ilepcum".,,... JI. C. Actl,,‘,iin ruioa,r; mi nonopozz -..leinioro. -M.. 1967.

15 Reptant! C. B., COK0.108 A. C., Tomumin A. r. Mopilm-cliyumutolia.-ibilbie oco-CiennocTu apTepliii it Ben Ima131111KOB .H.eabiliinion B C1151311 C .101011101111C11. 1V BCCCO-

103110C C011ellt HO nayii. mopcmix 1\1.1/e1:01111THIO111,11X. M., 1969. . 16 Herpona P. M. K nonpocy o TonorpatImil II minepBaumi me.anacTintaabunii Tuienphi. MaTepliambi OrleT11. liompop. Munemoro me.a,uncTwryTa aa 1963 r. MIII1CK,

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18 cAaaotimicKast .11. B. HCKOTOplaIC 13031M1CTIlble oco6entiocTil mitupotuipBymiTop-ublx erpyKTyp riplicTeito ■ mon rocupbt ttemneBa. B KII.: 4:Mop(1.fflorin-tecmie oco6eil-11 oon minwoulipBy.nmuni». M., 1967. • 19 Coicoma A. C. Ocori -ennocTii Kaoli6pa II pacime,rtenemin IICKOTOpbIX Bponeitoc- MAX COCYJI.OB y D0111,1X (T10.11C.111.1) H nasemnbix (CO6D1:11) NI.IHKOMITH101.11.11X. Apxnn anaTomun, rucTo.nornu u 3m6piumormi, T. 37, II1,111. 12, 1939. .

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■ ••

I 331

UDC 599.5 599.745

P. A. Korzhuev

THE PROBLEM OF THE OXYGEN RESERVES IN THE ORGANISM 205

OF AQUATIC MAMMALS

One of the most weakly elaborated problems of the bio-

logy of aquatic mammals is the problem of respiration, or more

precisely, the problem of the shutting-off of the external res-

piration during the period of diving. Up to the present time

in the literature there are present contradictory data on the

time spent by the animals under water. While the time of the

shutting-off of external respiration during diving in the diving

birds is measured in minutes, then for the aquatic mammals this

period is measured in tens of minutes, and for the large whales

- even in hours (Korzhuev, 196 )4). According to Pastukhov's

data (1969), the Baikal seal may spend more than an hour under

the water. It is true that this record duration of stay under

water, equaling one hour and eight minutes for one of the test

seals, was observed by the author only once, and later these

seals shut-off their breathing for only 5 - 8 minutes.

The author of the present paper haopened to be a wit-

ness to an experiment, conducted on a young seal in the Northern

Caspian in September of 1965, when after a forced stay under

water of 7 minutes the seal had a very lamentable appearance.

Experiments on the shutting-off of breathing in the Black Sea

dolphins, the white-sided dolphin and the nyalix±hm bottlenose

dolphin (Korzhuev and Bulatova, 1952), when the animals were

on dry land, showed that the white-sided dolphin calmly endured

such an experiment for only two minutes, while the bottlenose

332

dolphin endured this for four minutes. After the expiration

of this period there commenced a very violent reaction, accom-

panied by vomiting, which was.indicative of a very obvious

disturbance of the organism. It may be assumed that, if these

experiments had been conducted in water, the duration of the

experiment may possibly have been increased by a factor of two

or three, but at any rate for the white-sided dolphin this

would not be more than 6 minutes, while for the bottlenose

dolphin it would not exceed 12 minutes. Visual observations

indicate periods of about this order.

Apparently the periods of shutting-off breathing for

the pinnipeds are more prolonged than for the small cetaceans,

though these also measure 15 - 20 minutes (Mordvinov, 1969).

Set somewhat apart is one of the Antarctic species of seals,

the Weddell's seal, which in one experiment dived to a depth

of 600 m and stayed under water for 43 minutes and 20 seconds.

As to the representatives of the large whales, the du-

ration of the stay under water of which is measured in hours

(sperm whale - up to 75 minutes, bottlenose whale - up to two

hours, Irving, 1939), these data are purely visual and there-

fore require confirmation. Apparently, the problem of the

prolonged diving of aquatic mammals will obtain an effective

resolution only when a thorough quantitative determination is

made of those reserves of oxygen which the organism of these

animals has at its disposal.

It is scarcely possible to consider as convincing the

arguments of those investigators who consider that the duration

of diving is related to the capability of the animals to change

over to the anaerobic type of respiration during the period of

333

i diving. It seems to us that the basic condition affecting the

duration of diving and, by the same token also, the cutting-off

of breathing for some period or other is the presence of oxy-

gen reserves and their rational utilization.

However the problem of the oxygen reserves in aquatic

mammals has been very weakly elaborated. On this subject there

are more assumptions than precise facts. As an illustration

of this, one may point to the studies of Scholander (194•0) and

Robinson (1939)s who determined the oxygen reserves in cetaceans

and pinnipeds, for which determinations they assumed that the

probable amount of blood in the organism of the dolphin, for

example, equalled 15.0% of the body weight or that the quanti-

tative characteristics of the body musculature could be taken

as being equal to 35.0% of the body weight, although these

values were not precisely determined.

In this connection it-may be noted only that for the

terrestrial animals also the problem of the oxygen reserves is

one that has not been worked out at all. Only the attitude that

in these animals these reserves are negigible may.be considered

as indisputable.

This fact is often obscured, since during the process

of the evàlution of life on our planet the problem of the re-

serves of various vitally important substances has been resolved

very successfully. In different stages of the ontogeny of

various groups of animals there are formed reserves of proteins,

fats, carbohydrates and vitamins: it is known that the egg

cells of various animals (fishes, amphibians, reptiles, birds,

insects) contain the entire assortment of substances necessary

for normal development; for the period of hibernation

L

3314 •

heterothermal animals lay down reserves of fat, attaining up

to 35.0% of the body weight, while in the aquatic mammals the

fat reserves may be as high as half of the body weight.

However oxygen, which is one of the most important

elements, without which the organism cannot manage to exist,

is practically not stored at all by the organism. Terrestrial

mammals, including man, may cut-off their breathing for only

one to two minutes. This circumstance is quite incomprehensible

since oxygen reserves could protect the organism from many un-

desirable and sometimes also catastrophic consequences.

Apparently there exist some very weighty reasons which

prevent the setting up of considerable oxygen reserves within

the animal organism; the small 'group of aquatic mammals and

birds forms a certain exception to this rule.

To decypher the specific mechanisms, determining the

possibilities by which animals may shut-off their breathing

for some period or other, is a problem which still awaits its

solution. All the same, with respect to the terrestrial verte-

brate animals it may be said that one of the most fundamental

causes, preventing the setting up of oxygen reserves, is to be

found in the properties of oxygen itself.

The fact is, as was shown by Professor A. L. Chizhev-

skii (1960), that it is not a matter of indifference as to

what air we breath. Up to the present time it has been con-

sidered that for respiration it is important that the air should

have in its composition a certain amount of oxygen, nitrogen

and carbon dioxide; however not only the chemical composition

of the air turns out to be important but also the physical

335

state of its molecules and, in the first place, of the oxygen.

Among the factors determining this physical state, the first

place belongs to electricity: the saturation of the air with

electrical charges and the degree of its electrostatic charging,

as a consequence of which the oxygen molecules become ionized.

Particularly important for respiration are molecules of oxygen

having a negative electric charge. If the molecules of the air

are not ionized, then animals rapidly die in such an atmosphere,

in spite of the presence of oxygen molecules in the respired

air. These investigations of A. L. Chizhevskii shed light on

the causes which determine the absence of any considerable re-

serves of oxygen within the organism of animals.

A characteristic feature of the ionized oxygen mole-

cules is the fact that they •exist for a very short period of

time, of the order of ten minutes or a little more, after which

the charge is lost and the oxygen reverts into the inactive

form. This circumstance, apparently, is the fundamental cause

which deprives the animal organism of the possibility of storing

the ionized oxygen, which has a negative charge. The negative-

ly charged oxygen ion possesses several characteristics, such

as, for example, the presence of nine electrons (as in fluorine)

and a changed spin of its nucleus. 4parently these physical

changes, occurring at the level of the electrons and of the

nucleus, play a decisive role in the evolution of the animal

world, as a consequence of which also there is found the absence

of considerable reserves of oxygen even in the higher represen-

tatives of the vertebrate animals.

336

This is a very fundamental circumstance, which has not

been adequately considered by biologists. Nevertheless the

properties of the basic chemical elements, which play an im-

portant role in the life of animals, determine the character

of their evolution. Practically all of the characteristic

features of the organs of aeration and of the haemodynamics

are determined by the properties of oxygen.

As to the aquatic mammals, it seems that here there

occurs a deviation from this principle. It cannot be ruled out

that in the case of the marine mammals there may be some kind

of supplementary adaptations, directed towards overcoming these

properties of oxygen, as a consequence of which there may be

set up within the organism oxygen reserves that are more power-

ful than is characteristic of the terrestrial mammals. However,

until precise data are obtained on the actual periods of the

cutting-off of breathing in the large cetaceans and pinniPeds

in their natural habitats, there are no groundsfor speaking

of deviations from this principle. •

The experimental data that are available at the pres-

ent time permit one to say only that a real- basis for the pro-

longed cutting-off of breathing is the presence of certain oxy-

gen reserves, i.e. the retention of the aerobic type of respi-

ration. And this means that without a precise knowledge of the

oxygen reserves it is impossible to defenitively resolve the

question of the dtiratIon of diving.

UnÉortunately there are very few precise data available

on the amount of oxygen that is found within the organism of

animals. In Table 1 are presented data an the oxygen reserves 1W)

337

Table 1. Oxygen reserveè in the organism of man, certain

mammals and birds, cm3 (Robinson, 1939; Scholander, 1940; Korzhuev, 1964)

'Ta6onua 1

Pe3epnbi nitcnopoAa n oprannme qemonena, IICKOTOpbtX. wienounTaimunx M I1TMU, Cat 3

(PoGlincou 1939; Ilionann,ep, 1940; Kopniyen, 1964)

1 2 3

5 6 7

Kucao-

Beer° pona

"Tea- mtcao- Ha 1 KZ

Blut 13cc, K2 JlerKite Kponb albunnia ttenatt p„,a. Ben

Species We igit Lungs Blood Muscles *"a""h

' kg A. B. U.'

linunam 70000 300000 1400000 1400000 250000 3350 48,0

ByThIJJK0110C 1400 6000 45000 54000 4000 109 78,0

Reni.,(Inui 19 250 600 180 60 1,09 57,0

Moment, . 29 500 1100 '270 100 1,52 52,4

TIOJICIlb 70 . 545 2055 2530 245 5,37 77,0

1-1enoneic 70 900 1160 335 245 2,64 38,0

flinirnun 5,6 120 90 40 20 0,27 48,0

A. - Inter-tissue fluid B. - Total oxygen & 1 C. - Oxygen per 1 kg -body weight, cm3

1 - Fin .whale 2 - Bottlenose whale . 3 - Dolphin 4 - Seal 5 - Seal 6 - Man

Penguin

in the organism of certain representatives of the mammals and

birds. For these the structures of the organism that contain

oxygen are the organs of rspiration - the lungs; the blood,

as a liquid tissue, contains physically dissolved oxygen, as

does also the inter-tissue fluid. However these structures

have a relatively small part of the oxygen. The main portion 209

of this is contained in the form of chemically bound oxygen

jal the respiratory pigments of the blood and muscles in the

338

form of oxyhaemoglobin of the blood and muscles. At least,

in the large cetaceans the portion of oxygen contained in the

organs of aeration and tissue fluids in the form of physically

dissolved oxygen comes to about 10 - 15%, while the portion

of oxygen bound by the haemoglobin of the blood and muscles

comprises about 84% in the fin whale, about 90% in the bottle-

nose whale, from 85 to 90% in seals, and about 70% in the

dolphin. In man the respiratory pigments bind about 55% of

the total oxygen that is present in the organism. It should

be borne in mind that, in spite of the relatively large amount

of oxygen that is bound by the haemoglobin of the blood and

the muscles, in man, for example, the tissues utilize a com-

paratively small amount of this., since the venous blood of man

contains much unutilized oxygen (about 12 - 14 percent by vo-

lume). It is true that such a character of the utilization

of oxygen has its advantages, since the pressure gradient of

oxygen in the capillaries of the body is very high, which

favours a more effective supply of this to the tissues.

In this respect the cetaceans and pinnipeds occupy a

special place, since they utilize the oxygen of the arterial

blood more completely, so that in the venous blood there re-

mains a very small amount of this, of the order of one to two

percent.

Thus, a characteristic feature of the adaptation of

the cetaceans and the pinnipeds is their higher capability

for removing practically all of the oxygen from the blood,

in contrast to the terrestrial mammals.

339

0 If it is considered that the aquatic mammals find

themselves under conditions that approximate to a state of

weightlessness and, consequently, expend practically no energy

for supporting the weight of their own body, then it becomes

understandable that the energy losses in these are considerably

smaller than in their terrestrial relatives. It is very

strange that this circumstance has been completely ignored

in comparisons of the characteristic features of the biology

of aquatic and terrestrial mammals, since in actuality it is

very essential.

Some investigators turn their attention only to the

fact that aquatic mammals may shut-off their breathing for

more prolonged periods, as compared to the terrestrial mammals;

however the rest is accepted as being quite adequate to ter-

restrial conditions. It is considered that the energetics of

the organism are completely the same in both terrestrial as

well as aquatic conditions. However it is precisely in this

that there is a basic difference between the aquatic and ter-

restrial mammals.

Of course, a very important form of.adaptation is the

lowering of sensitivity of the respiratory center to the accu-

mulation of carbon dioxide in the blood. The behaviour of 210

the vascular system in the aquatic mammals at the time of

diving (the redistribution of the blood supply.to various

parts of the body) is completely different from that in the

terrestrial animals. It seems to us, however, that the cause

of this is not the necessity of ensuring a supply of oxygen

only to the vitally important organs (heart, brain) at the

34•0

expense of other parts of the body, which might seemingly be.

deprived of the necessary amounts of oxygen, at the same time

changing to an anaerobic type of respiration, but something

quite different.

From our point of view, the crux of.the matter lies

in the fact that this redistribution of the blood supply

during the period of diving occurs only because in the aquatic

conditions the requirement for oxygen is smaller than under

terrestrial conditions.

The correctness of this assumption finds confirmation

in the fact that the aquatic mammals, in'contrast to the ter-

restrial mammals, possess many times greater reserves of haemo-

globi.n, localized both in the blood and in the muscles of the

body. It is quite obvious that the large amounts of haemo-

globin are the main condition of the creation of the oxygen

reserves during the period of diving. Evidently the expendi-

ture of these reserves is more economical than is the case

under terrestrial conditions, where energy is expended not

only for locomotion but also for supporting the weight of the

body itself.

In other words, life under terrestrial conditions is

linked with the necessity of surmounting the forces of the

earth's attraction, or the forces of.gravitation, and this

surmounting presupposes losses of considerable amounts of ener-

gy, while under aquatic conditions, as a consequence of the

high density of the wate.r, the effect of the gravitational

forces is practically eliminated, although the gravitational

field is retained in the water, as on dry land. The data pre-

sented in Table 1 confirm the stated positions.

341

It may be thought that, if the cetaceans and pinnipeds,

in the process of their adaptation to the aquatic conditions

of existence, had changed over to a gill-type of respiration,

i.e. if they could obtain the oxygen directly from their sur-

rounding medium, as is the case in the fishes, then they would

not have such large reserves of haemoglobin and, by the same

token, of oxygen.

The fishes, as primarily aquatic animals, are charac-

terized by having a very small amount of blood in the organism

and a small amount of haemoglobin per unit weight of the body.

Thus, for example, in fishes on average there are about two

grams of haemoglobin per kilogram of body weight, while in dia-

dromous fishes, which accomplish extensive migrations to their

spawning sites, generally against currents, the degree of pro-

visioning of the organism with haemoglobin comprises about 3

grams per kilogram of body weight. It is true that in addition

to this there is present in these a small amount of physically

dissolved oxygen, contained in the blood and perivisceral

fluids, while in some fish species there is present a small

amount of oxygen that is bound by the muscle haemoglobin.

However, these additional amounts scarcely change the total

figures named above.

If one compares with these figures the degree of pro-

visioning of the organism with haemoglobin in the dolphins

and seals, then it is found that in the white-sided dolphin

this value is approximately equal to 25 g of haemoglobin per

kilogram of body weight, while in the seal it attains a level

of 40 g. This is almost 15 times greater than in fishes.

34,2

o These huge reserves of haemoglobin, required by cetaceans and

pinnipeds for binding oxygen for the periods of diving, would

be superfluous with the gill-type of respiration.

However, since in nature there is found the principle

of the irreversibility of evolution, first discovered and

formulated by the French paleontologist Louis Dollo in 1893,

the change to a gill-type of respiration is impossible. The

cetaceans and pinnipeds are forced to employ the lung-type

of respiration, with all of the consequences arising from

this. This means that the entire evolution of the aquatic

mammals was directed, in the first place, to providing for

one of the most basic functions, th function of respiration.

From this derive: the change in the rhythm of respiration,

the lowering of the sensitivity of the respiratory center to

the accumulation of carbon dioxide in the blood, the redistri-

bution of the blood-supply to the organs during the Period of

diving, and the marked stimulation of the synthesis of hà.emo-

globin not only in the bone marrow but also in the muscles of

the body. And all of this occurred so that it would be pos-

sible to do without breathing during the period of diving.

The aerial type of respiration turned out to be the only

thread which ties in the cetaceans, the group which has moved

furthest in its evolution to the aquatic way of life, with the

terrestrial animals.

34'3

JII'ITE.PATJ'PA

1 Koporcyeo 17. A. remor7o611n. CpaB11nTC7bnan 6noXiotilfl n 4p113110:lorltR.

•allayxa», 1964.2 Kop.mycc+ 17. A., Eqnaroea H, H. jlblaaTCAbnan fiynxunn xpoBn Ae.ab^nnoB

7py7b1 f1nccruryTa mop(Ilo;lorlut :i<liBOTnbix n;,t. Ceuepuoua AH CCCP, Bbin. 6,1952.3 hlopoaur{o(i 10. E. )JA11Ti':1bilOCTb npc6binannn uop, nolIoi! xacnlli'ICxnx Ttoae-

'ueïl. TC311cbt Aor,7a;tOB 4-ro IiceColo3noro coBentannfi no Diol)cxnSr nLlexonllTalO-7I nAt , Ka•aninlnrpan, 1969.

^^ 17(lC7IfX06 B. /^. Hei:OTOpblc Pe3y7hTaTbi na6.nto;tennH nap. 6a}(xaJ1bCt<oÏ1 nep-

n01i B yC.70BI15i\ 31<Cliepll\1e11Ta. Te3ncbl Aoi:.,laltos 4-ro I3Uc0i03110ro co8enjaili(llmo niopci<nm maicl<onirraioil.tu>t. 1Kaaltnnlirpaa„ 1969.5 yra;ccacs:uü A. Jl. Aapouonnc)nxaunsi B napo;tnoM xo3Ni'tcTBe. Ai., I'ocn.naiina-

A T, 1960;^ Harrison S. Experiments with diving Seals Nature. 188, 1960.`z Irving Z. Respiration in diving A'lammals Physiol, Rev., 19, 1939.tj Robinson D. 'Hie muscle hemoglobin of Seals as an Otygen Store in diving.

:Science, 90, 1939.g• Scholander P. Experimental Investigation. on the respiratory function in di-

-ving niaimnals and Birds. Hvalradets Skrifter; no 22, Oslo. 1940.

REFERENCES

1. Korzhuev P. A. Haemoglobin. Comparative biochemistry

and physiology. Moscow, Publ. "Nauka", 1964.

2. Korzhuev P. A. and BulatovaN. N. The respiratory function

of the blo.o.d of dolphins. Trudy Instituta morf ologii

zhivotnykh im. Severtsova AN SSSR (Transactions of the

Severtsov Institute of Animal Priorphology of the USSR

Academy of Sciences), no. 6., 1952.

3. Mordvinov Yu. E<. The duration of the stay under water of

the Caspian seals. Reports of Proceedings of the 4-th

All-Union Conference on marine Mammals, ' Kaliningrad,

1969.

4. Pastukhov V. D. Some results of observations on the Baikal

seal under experimental conditions. Reports of Pro-

ceedings of the II•th All-Union Conference on Marine

yammals, Kaliningrad, 1969.

5. Chizhevskii A. L. Aeroionification in the national economy.

T,loscow, Gosplanizdat (State Publishing house for Litera-

ture on Planning), 1960.

UDC 599.537

T. N. Glazova

344

QUANTITATIVE EVALUATION OF THE HAEMATOPOIETIC FUNCTION 212

OF THE SKELETON OF THE BLACK SEA DOLPHINS

It is considered that in the terrestrial vertebrates

the function of the skeleton consists of supporting the body

in a particular position, and also ofstrengthening and retain-

ing the musculature. In the aquatic vertebrates, according

to the opinion of E. Slijper (1962), the function of the ske-

leton that consists of supporting the body does not have a

great significance, since in the water the body is acted on

by an upward pushing force, which alleviates this function.

gle)

The importance of the latter circumstance is also emphasized

by P. A. Korzhuev (1963), who notes that the organism in the

water is as it were suspended and therefore its energy is ex-

pended only for locomotion, while under terrestrial conditions

the animal must expend considerable energy not only for loco-

motion but also for supporting the.weight of its body.

In both the aquatic and terrestrial conditions one of

the basic functions of the skeleton is the haematopoietic

function. From this point of view there is a special interest

in the study of the weight characteristics of the skeleton and

bone marrow of adult dolphins, as animals which shut-off their

breathing for prolonged periods, and also in the study of dol-

phin embryos, which are subjected to difficulties in acquiring

oxygen not only in particular stages of their development but

also at the time of the shutting-off of breathing by the mother.

345

. In the literature there exists a fairly considerable

amount of data on the weight of the skeleton of the large

cetaceans (Khar'kov, 19•0; lérochkov, 1953; sleptsov, 1961;

Bjarnason, 1954). These data are not precise and at times

are greatly exaggerated in connection with a lack of a thor-

ough cleaning of the skeleton. Probably, more precise data

on the weight of the skeleton of the large cetaceans are to

be found in E. Slijper's (1962) study. Characteristics of

the weight of the skeleton of representatives of the small

cetaceans have been presented in the studies of P. A. Korzhuev

and co-authors (1964, 1967). In these studies there are data

on the weight of the bone marrow in the adult cetaceans and

their foetuses, and also on the. weight of the skeleton of the

latter.

The aim of the present study was to show up the weight

relationships of the skeleton and bone marrow of the developing

embryos and sexually mature Black Sea dolphins.

There were studied embryos of different ages and adult

individuals of the bottlenose dolphin (Tursiops truncatus Mouf),

of the common dolPhin (DelPhinus delphis L.) and of the common

porpoise (Phocaena phocaena L.). The skeleton and bone marrow

was examined in 7 embryos of the . bottlenose dolphin, with a

weight of from 184 g to 16.5 kg and an age of from 4-5 to

11-12 months*; 4 adult individuals e . weighing from 30 to 67 kg,

and 10 embryos of the common dolphin weighing from 2.65 to 7.00

kg and aged 8-10 months; 11 adult individuals of the common

porpoise weighing from 13 to 47 kg and 11 embryos weighing from

* The age of the embryos was determined with an accuracy of to 0.5-1 months on the basis of the length of the body.

346

884 g to 1.996 kg and an age of 5 - 8 months, and also one

embryo weighing 3.07 kg aged 9-10 months.

For obtaining the weight characteristics of the ske-

leton, after the weighing of the whole body there were removed

the muscles, tendons, brain and spinal cord, and the cleaned

skeleton was weighed in sections. The volume of the bone mar-

row was determined by a method based on Archimedes' principle

(Petrov, 1952). For calculating the weight of the bone marrow

there was employed the specific weight of this in only the

sexually mature dolphins, in which it was possible to find

individual sections of cylindrical form in the ribs.

Knowing the specific weight and volume of the bone

marrow, its weight was calculated as a percentage of the body

weight, and the percentage relationships in the different

sections of the skeleton were also calculated.

According to the literature data (E. Slijper, 1962),

in the large crustaceans, such as the sel whale, fin whale and

blue whale, the weight of the skeleton comprises corres .ponding-

ly 13, 16 and 17% of the body weight. In the Black Sea dol-

phins, according to the data of P. A. Korzhuev et al. (1964),

the weight of the skeleton was half as great.

On thorough treatment of the skeleton it was found

that its weight relative to the weight of the body comprised,

on average, 6.5 in the common dolphin and 5.2% in the common

porpoise (Tables 1 and 2). In the Quantitative characteriza-

tion of the skeleton of the two species of dolphins there were

not found any statistically significant sex differences.

347

The axial skeleton takes up the main part of the ske-

leton of the dolphins. Thus, this comprises about 90% of the

weight in the skeleton of the common dolphin, while the ske-

leton of the limbs comprises only 10%. The relationship of

the parts of th skeleton does not remain constant: with in-

creasing weight and length of the dolphin (for example, of the

common porpoise) there is found a decrease in the relative

weight of the skull of from 32.6 to 24.0% of the total weight

of the skeleton, and an increase in the weight of the verteb-

ral column of from 46.0 to 53.7A (Table 2). It la interesting

that the relative weight of the ribs and sternum remains at .

almost the saine level.

While the fraction taken up by the anterior girdle and

free fore limbs in the common porpoise and the common dolphin

is 9 - 10, rin-i2ze the fraction falling to the posterior limbs,

which in dolphins are reduced to two short narrow bones lying

within the muscles, is measured in tenths of a percentage point.

The bones of cetaceans are filled with bone marrow. •

E. Slijper (1962) notes that the skeleton of young animals

contains red bone marrow but that with age there occurs a

replacement of this fraction by the yellow bone marrow.

According to our data, all of the studied dolphin in- 216

dividuals, including also the sexually mature individuals,

possessed only the red fraction of the bone marrow, i.e. the

haematopoietic fraction. The weight of the bone marrow in

the adult dolphins comprises about 2A of the body weight

(Tables 3 and 4). No sex differences were found in the weight

of the bone marrow in the dolphins. The bone marrow is loca-

lized identically in both species of dolphins. The greatest

iength Te2a,

noa •

Sexi cm oo fe" ut. r o

COI.umn

ljeee4 hind

rrepeliniii I 3azilux rore gepena skull

rleepand ,rpyzintst

sternum

44,8

44,1

48,7

eJ

46,4±1,2

10,7

9,8

10.5

10,5

10,4±0,2

6,9 6.9

6,2

6.7

6,5 ± 0,2

32,1

28.9

26,5

27.7

28,8±1,2

12,3

15.6

11,5

13,5

13,2 -±0,9

0,1

0,2

0,1

0,3

0.1-±-0,05

Table 1. The relative weight of sections of the skeleton of adult

, IsD common dolphins, as % of the total weight of the skeleton.

• Ta6au u, a 1

OTHOCHTeJ11,11b1ii BCC OT,Ile.10B CKCJ1CTa B2, p0C.11bIX 06bIKHOBC1111b1X Ae.11:411110B,, % OT e5tuero BCCa cliezeTa 1

TO jULliqadY re.:a, K

W e ght , kg

A • Bc CMC.1 CT2,

Cé CT Beca Ter: a

e"-Ig ul Dec o -r.z.c...:oa cxezeTa.

30,2

41,8

58,3

65,3

7-1-S7C 48,9 ± 7,95

camitaf 128

cameuM 157

camxaf 167.5

cameuM 174

camKa 155,0±10,2 11 CaMeLt f and m.

f - female m - male

A. - Weight of skeleton,:as % of body weight.

1 Translator's not. Sic. Should:be %, rather than kg. t, ;

c5D

6 .1 a 2

• 1: t, , Kg 71-regs--- 1(01! eq:10CTS: •

B .

Bee xpsuna. Ca Ben

body kelet

gepon. skull

Pe6q, ribs stenue

noni cro.163

column fore

nepea.. 'hind.

31.11111X

%04‘, ody

06Temi nec

wjUht, kg

Body length

/Limn rem,

CM

A. Bee cue- ;ICU. !'4 Or Ben

Tela

no.1

Sex

6.3

5,1

• 5,4

• 4,9

• 7,9

5,9 .

5,0

4,9.

4,6

5,2±0,2

0,5

0,3

0,3

0.2

0.8

0,2

0,1 0,2

0,2

0.2

6,2±0,05

7,7

5,5

6,3

3,5

10,5

4,1 9 ,5

3,5

3,8

3,4

4,2

31,2

.31,0

28,9

24,7

32,6

29,8 97,8

27,5

24,2

25,9

24,0

27,5±0,9

11,5

12,1

10,5

11,7

12,0

12,6 13,1 13,7 13,1

12,8

•12,2

12,4±0,3

re . 7,

Table 2. _

I • The relative weight of sections of the skeleton of adult

porpoises, as % of the total weight of the skeleton. common 0.:nocwreobt-rmil sec OT,Ze.,103 cxeneTa e3pocJ1bm mopcmux cBHHe, % OT eGulero Zeca cue-ieTà.

22,00

23,00

26,40

32,00

13,10*

37,45**

38,20"

40,00**

40,00 • •

40,00**

47,00**

xfSx_

34,61±2,54

canut male

» » '

emua . ema_

,»

canieül 129±3

0,9 46,8 9,3 •-.. 0.2

1,2 46,3 8,9 0,5

1.0 49, 9 9,5 - 0.8

1,2 52.9 8,8 0,8

1,2 46,0 ' 8.0 0,1

47.8 8,2 0,2

1,5 49.3 8,1 0,2

1,7 48,3 8,5 0,3

1,6 52,7 8,3 0.2

1,6 50,4 9,0 « 0,3

1,6 53,7 8,6 -

1,3±0,1 1 49,8±0,81 8,7±0,2

114

116

120

127

92 e 133

129

136

136

• 137

138

•

H • , I '

.camica • . m and f • 1lenonono3pezae ocoCh nim cravierinecgoil o6pa60-1- Ne ne

"' Camiza Gepemenue. ymIlThinaeach. •

0,4±-0,1

* Sexually immature individual, not taken into account in.the statistical treatment.

** Pregnant female

A. - Weight of skeleton, as % of body weight.

B. - Weight of cartilage, as % of weightof:

1 Translator's note. Sic. Should be %, rather than kg.

350

amount.ofi this is concentrated in the vertebral columnr ave-

raging 56.7 and 61.6;-) of the total weight of the bone marrow.

The minimal amount of bone marrow is found in the pos-

terior limbs of sexually immature animals. In a young common

porpoise weighing 13.1 kg the pelvic bones contained 0.2%

(Table 1I.). In the rudiments of the posterior limbs of the

sexually mature forms there was no bone marrow. In the skull

it is mainly localized in the occipital bone and is completely

absent in the lower jaw. The average relative weight of the

bone marrow in the skull comprises 20.6 and 15.5%. in both

species of dolphins about 12% is contained in the ribs and

sternum, and also in the bones of the fore limbs.

Before passing on to the description of the weight

characteristics of the skeleton and bone marrow in the embryos

of the Black Sea dolphins, in comparison with the adult forms,

we will give a short account of the regularities revealed for

each species of a particular age.

The greatest period of pre-natal development was co-

vered in the bottlenose dolphin, the embryos of which were re-

presented, on the one hand, by 3-4 month old individuals and,

on the other, by 11-12 month old individuals, i.e. just prior

to birth.

The relative weight of the skeleton of the embryos of

the bottlenose increases, in step with their growth, from 6.75f

in the 3--4. month old embryo to 10.4.0% of the body weight in

the foetus just before birth (Table 5). A positive correlation

is marked between the weight of the skeleton, on the one hand,

and the size of the foe-tus, on the otheri the relative weight

e.,

Z.!

weight of lug,gegpl,akeleton, kg1 andvertebral

peuep H noutouonoro Sen sternum cden

iore hina nepexunx 32AHHX

Length ezima Tua,

of fhdy, cm

A • Cennurt Bec HOCTHOTO

meBra. OT BCC2 Te.7a

30,2

41,8

58,3 65,3

x±Sx 48,9 ±7,95

camn f

cameu,

camxa f cameu,-M

camn CaMeg

f and m

128

157

167,5

174

156,6±10,2

2,0

1,7

1,9

2,2

1,9± 0,1

23,0 14,4

22,5

22,6

20,6±2,1

7,5

16,7

8,2

9,3

10,4±2,1

57,6

M,2

55,5

55,4

56,7 ± 0,7

11,8

10,3

PJ 12,3

12,0±0,7

0,1

0,3 0,2

0,4

0,2 ±0,05

Total body 06Lumg sec

Tem,gz weight, kg

Tion Sex

Table 3. The relative weight of the bone marrow.of adults of the common dolphin, as % of total weight of the bone marrow.

T a 6.1 1,111, a 3

OTHOCHTeablIber BeC KOCTHOr0 mo3ra B3poc.nux O6bIKHOBeliniX aeJ114}11013, % OT 0611.1,CTO Beca KOCTII0f0 mo3ra

f - female m - male

A. - Total weight of bone marrow, as % of body weight.

1 Translator's note. Sic. Should be % of total weight of bone marrow.

Table 4.

The relative weight of the bone marrow of adults of the common porpoise

,: in various sections of the skeleton, as % of total weight of bome 1,1ar1; o^r.

OTHOCSiTCRbHSIH BeC K4CTHoI'0 N.03ra B3110C.?6!Y "f0PCKfiY CBHY,C11 B .^.i137K4!Ib!Y OTAC:SaY CKe.i!eTa, clp CT OG::(efo E^.Ccl.

KOCTHOI'0 M03ra

Total ^ l` ^ iese on, icl 1:OCTt:01'0 bl03:'3 D Lne CrC.boGy I, g th 06ut::i! Bec CC

l^Tl }^SOGutafi sec AnHKa xocrnoro ver-tebralTeTa. c:e ^L,Jra, ?^ I•

c

n03no1roRr:orO xOP.e.liucrel!Te7a aB MOI

o^ ^j^d OT BCC2 TC7a ^CDCIIa pp6ea t rp}-ai+:crp1^e I n 1.nweigh^kg ;jex cm Y1 skull 1 _.^-bs sternum nepC;lc:, 321:131s

22,00

23,00

26,40

32,00

13,10*

37,45

38,20

40,00

40,00

40,00

47,00 **

x±Sx34,61-!-2,64

caaie!1male»

»

ii

casseu HcamKamale

114

116

120

127

92

133

1129

136

136

137

138

129.±2,9 2,0±0,1

and îenl le

13,8

16,6

17,9

13.6

•18,3

18,4

17,7

15,1

13,8

14,4

14,0

15.5.t:0,6

7,5

10,6

8.0

8,1.

8,3

12.3

10,0

8,811,09.2

9,5 ±0,5

1,2

1,9

1,2

1,7

1,3

1,9

2,3

2,8

2,5

2,5

2,0

65,5

59,1

61.0

65,5

62,3

56,7

55,9

60,3

65,3

G0,7

62,8

12,0

11,9

12,0

11,1

9,7

10,8

I1,1

11,6

9,6

11,5

12,0

2,0-?-0,2- 61,6±1,0 1 11,3--+-0,2

0,,8

" Heronoeo3peRas oc06b npit cTaracTKUCCxoi oGpaGoTne R O yti11Tai83naca.Camxa Gepevexaa.

^ Sexually immature individ1.zal,. not considered in the statistical

1

treatment.

Pregnant female

Translator's note. aJc.. Should be ô e

t;

353

of the skeleton increases with the increase in the weight and

length of the embryo. With the growth of the embryo, the re-

lative weight of the skull, which comprises a considerable

fraction of the total weight of the skeleton of the embryos

of the bottlenose dolphins, decreases from 44% in the 4-5 month

foetus to 31% in the 11-12 month foetus, i.e. just before birth.

The largest part of the skeleton consists of the vertebral

column, the relative weight of which increases from 38 to 53%.

The data characterizing the relative weight of the ribs with

the sternum, and also of the girdle and free anterior limbs,

did not display a change with age. It was found that these

parts of the skeleton were about the same in the bottlenose

dolphin embryos of different ages.

The relative amount of *cartilage decreases with the

growth of the embryo. This decrease is manifested most strik-

ingly when the weight of the cartilage is expressed as a per-

centage of the weight of the skeleton. While in the 4-5 em-

bryo weighing 4 kg 55% of the skeleton consists of cartilage,

just before birth in an embryo weighing 15.0 kg the cartilage

fraction comprises only 29%, which values are correspondingly 219

equivalent to 4.2 and 2.8% of the body weight (Table 5).

The bone marrow of the embryos of the bottlenose dol-

phin consists of the active red fraction. In step with the

growth, the total weight of the bone marrow in the skeleton of

the embryos of the bottlenose dolphiruincreases in both abso-

lute and relative terms: the weight of the bone marrow of the

4-5 month embryo comprises 1.7%, while in the embryo in the

last stages of intrauterine development it comprises 4.6% of

the body weight (Table 6).

• 351i-

Table 5 .The relative weight of sections of the skeleton

of embryos of the bottlenose dolphin, as % of

the total weight of the skeleton.

Tafiauua 5

UTIfOCfi7CllbHbiti üCC OrJleJ1OD CKCAC7'a 11a0t,t01f at^aANiib!,

% OT OÛIi(CCO i3CCa CHCJICTa

0,184

4,0

7,0

11,0

12,0

15,0

16,5

B

n o.1

C

AnnuaTe.7a,ru

caNtha^' 21,5

caa+ica^ 65

cahtica^ 95

ca^icqT^l 96

ca^ma 110

camica 120

ca^seü 114

DB03-pacT,MCCR-

1ICü

3--4

4-5

5-6

5-6

11-12

oxoao12

11-12

IiFe Bec X(v^Ita, çdcKC.le- OT BecaTa,

oT d,necÏ. a

a I cxe-TCAa nCTa

6,8

7,3

9,1

9 ,8

10,4

9,8

10,2

4,2

4,3

3,2

3,52,8

3,2

55,2

47,5

31,4

33,8

28,8

31,4

G Bec oTncnon Cr.e^1cr2, ICI,

^YV ^ x nctttcape6ep u utfovauo:co- u rte-ll

yepcna rpy. ru cro- Pcawlx

yllmw n6a Aoue't-ROCTeII

43,7

38,040,036,530,835,0

7,5

6,3

8,9

6,7

8,1

8,7

37,5

49,1

44,2

49,2

52,8

47,5

7,5

6,6

7,0

7,5

8,38,8

A. - Total weight of body, kg LL

B. - Sex

C. - Length of body, cm

D. - Age, months

E. - Weight of skeleton, as % of body weight

F. - Weight of cartilage, as % of weight of:a - bodyb - skeleton

G. - Weight of sections of the skeleton, kg1w - skullx - ribs and sternumy - vertebral columnz - pectoral girdle and fore limbs.

f - femalem - male

1 Translator's note. Sic. Should be %.

• • 355

Table 6,

The relative weight of the bone marrow of embryos

of the bottlenose dolphin in various sections of the skeleton, as % of the total weight of the bone marrow.

Ta6eltua

OTBOCETCJIMIblii uec Koc -ruoro moara nnoRon ati)a,auubt . paaaugubtx ax cxe.neTa, °,10 ov etimero ileca uoc -ruoro moara

A B C D E F Bec ICOCTII0r0 110311 la CIZeileTe. IC2

06tguii 06tuull oec w

BCC R:ulna Bo3pac-r. ROCTI1010

1C1)Y z

lumen H il

. Ten, 110.1 TC./13. ,,,,,,e, moan, ?k; 1103001104- nepeRinix CAt 01 tieca geperia rry _ 110F0 KOI1C.1110C- X2

Tena juintà CT0.1i511 TCii

•

0,184 caMIa 21,5 3-4 _ - - _ -

4,0 CaNIKa 65 4-5 1,7 54,0 9,0 30,3 6,7

7,0 camK‘ 95 5-6 2,4 42,7 4,4 44,3 8,7

11,0 camet 96 5-6 . - - - _ • _

12,0 ca.mx 110 11--12 3,5 31,8 5,7 54,9 7,7

15,5 UMK 120 OKW10 12 3,5 26,4 9,3 - 53,4 - 10,8

16,5 eaiei.fl 114 11-12 4,6 26,1 4,2 58,9 10,8

A. - Total weight of body, kg • B. - Sex

C. - Langth of body, cm

D. - Age, months E. - Total weight of bone marrow, as % of

body weight

F. - Weight of bone marrow in the skeleton, kg i w - skull x - ribs and sternum y - vertebral column z - pectoral girdle and fore limbs

f - female m - male

1 Translator's note. Sic. Should be %.

Bec cKene-Ta, qc

OT ueca Tena

42,7

40,1

38,2

40,0

35,1

37,7

35,1

35,4

28,3

MA

2,65

.3,00

:3,68

• 4,20

4,90

.5,30

.5,60

.6,00

.6,30

7,00

I) BoapacT, uecnoen

13ec xpnuta, 9t, OT neca

al Pce_ Tel a I n„a

(; Bec „tenon cKeneTa,

in pe6ép 110X0. H 1104H0-

`lePena rpy- ro cron- A1111b1 6a

A

lice Teaa,

he

noca II ne-peanux

.xouen-IIOCTeii

non ji.1{I u a Tena,

C.14

8,1

7,1

8,5

7,3

8.6 9,5

9,2

8,2

7,9 I

4,0

3,6

4,6

2,9

4,0

2,8

2,6

3,4

4,5

3,0

43,1

42,9

47,5

33,8

41,4

27,3

26,7

41,2

40,1

34,5

7-8

7-8

7-8

8-9

8-9

oxono 10

9-10

9-10

10

9-10

9,4

8,5

9,6

8,5

9,5

10,2

9,5

8,3

11,2

8,7

5,8

4,7

5,2

5,9 6,0

5,4

5,5

6,4

4,9

7,1

43,5

42,1 .

48,0

46,9. 50,3

47,4

54,4

49,0

58,6

50,0

ca mica ca men ,cameu.

ca men camxa

nmu

c a Wca CO mica camu

camen

66

67

72

79 • 81

84

83

81

90

84

• 356

o Table 7. The relative weight'of sections of the skeleton of embryos of the common dolphin, as % of the total weight of the skeleton

Ta6.-iiin.a 7

OTHOCIITOJIblibla Bee OTACJI011 cKe.nera naogon oeibncoonennoro.p,e,nbcfmna, % or o6inero neca cKeae -ra

A. - Total weight of body, kg

B. - Sex C. - Length of body, cm

D. - Age, months

E. - Weight of skeleton, as % of body weight

F. - Weight of cartilage, as % of weight of: a - body b - skeleton

G. - Wei&ht of sections of skeleton, kg1

w - skull x - ribs and sternum y - vertebral column z pectoral girdle and anterior limbs.

1 Translator's note.. Sic. Should be %.

357

By the 4-5th month of intra-uterine life about 50% of 220

all of the bone marrow of the bottlenose dolphin embryos is

concentrated in the bones of the skull, but towards the end

of intra-uterine development the greater part of the bone mar-

row, comprising about 60%, is localized in the vertebral column.

The weight of the skeleton of the embryos of the corn-

mon dolphin in the second half of their intra-uterine develop-

ment comprises, on average, 9. 14% of the body weight. As the

embryo grows there is found the same pattern of change of the

relative weight of the different parts of the skeleton and of

the cartilage, as occurs in the embryos of the bottlenose dol-

phin. Thus, with the growth of the embryo there can be traced

a tendency towards a decrease in the fraction of the cartilage

in the skeleton: from 4.6 to 2.8% of the body weight or from

47.5 to 26.7% of the weight of the skeleton (Table 7). In a

manner similar to that in the bottlenose dolphin embryos, there

is manifested a pattern of change with age of the growth of

the skull, the weight of which decreases from 46 to 28% of the

weight of the whole seleton with the increase in the size of

the embryo. In contrast to the development of the skull, in

step with the growth of the embryo there occurs an increase

in the weight of the bones of the vertebral column from 42 to

59% of the weight of the skeleton. Age fluctuations of the

remaining bones of the skeleton are insignificant. The weight

of the bOne marrow of the embryos comprises, on average, 3%

of the body weight (Table 8). On examining the data on the

distribution of the bone marrow in the different sections of

the skeleton of the embryos there is found a tendency towards

358

decreasing the amount of this in the bones of the skull (from

55 to 25% of the total weight of the bone marrow) and an in-

crease in the vertebral column (from 32 to 61%). A similar

pattern is found in the embryos of the bottlenose dolphin.

Unfortunately, the absence of material on embryos in the first

months of intra-uterine development, when there occurs the

formation of the bone marrow as a center of the synthesis of

haemoglobin, does not give us the opportunity of characterizing

this. However in the second half of the embryonic development

there is found only a gradual build-up in the mass of the hae-

matopoietic tissue, which to a considerable degree depends on

the individual characteristics of the developing organism.

On the whole, the distribution of the bone marrow in the ske-

leton of the embryos proceeds in such a manner that the largest

amount of this is localized in the vertebral column - about

50%, then in the skull - about 35%, in the bones of the fore

limbs - about 9%, and in the ribs and sternum - about 6% of

the weight of all of the bone marrow.

In the 5 - 8 month embryos of the common porpoise the

average weight of the skeleton, of which half is composed of

cartilage (53% of the weight of the skeleton), is equal to

8.4% of the body weight (Table 9). We note that the weight

of the cartilage in the skeleton of the adult common porpoises

comprises, on average, only 4.4% of the weight of the skeleton.

In the embryos of this age the relative weight of the skull

and of the vertebral column is about the same ( •2 and 44% cor-

respondingly). In the embryo just prior to birth the weight

of the skull skeleton decreases to 31%, while the weight of

the vertebral column, conversely, increases to 54% of the

Table 8.

A B C D E F Bee uocutoro T•1031":1 B cizene-re. et

06uutii 06nut6 BCC w X Y z

BCC RIIIBB licc e.n.r , KOCTUOCO pe6ep notica,

reia, lloa Te.na, ,„'",,,„„ M03 18, 5, 6 ii uo3uoitog- nepen„,,

cm OT ucca gepetta rpy_ Itur0 Kolictuloc-

IC2 re.qa Alum croA6a TCil ,

2,65 cana' 66 7-8 3,2 40,2 5,5 43,1 11,1

3,00 cameun 67 • 7-8 2,2 54,6 3,9 32,3 9,3

3,68 camearl 72 7-8 • 2,9 40,4 6,5 46,5 6,6

4,20 eamen; 72 8-9 2,5 33,5 7,8 54,0 4,6

4,90 calm 81 8-9 2,7 ' 33,9 5,1 50,1 11,0 .I.

5,30 eantKa 84 omviô 10 3,8 28,1 4,7 57,2 10,0

5,60 cMKak 83 9-10 3,9 26,3 6,2 61,2 6,3

(6,00 camxa 81 9--10 2,5 34,4 6,0 , 47,3 12,3

.6,30 camKa 90 10 3,6 27,5 6,2 52,4 13,9

7,00 caMe4 M 9-10 2,7 27,9 7,9 54,3 ' 9,8

`.7

359

The relative weight of the bone marrow of embryos of the common dolphin.in in various sections of the skeleton, as % of the total weight of the bone marrow.

T a 6 a 11, a 8

OTHOCIITC.,11,111:Iii nee nocTnoro mo3ra I1J10)1,0B o6humonennoro fleabcbitila

13 pa31114111AX oT,ncmax cy.eaera R% or o6atero Beca Eocruoro mo3ra

A. - Total weight of body, kg B. - Sex

C. - Length of body, cm

D. - Age, months

E. - Total weirht of bone marrow, as % of body weight

F. - Weight of bone marrow in the skeleton, kg 1 w - skull x - ribs and sternum

• y - vertebral column z - pectoral girdle and fore limbs

f - female m - male

* about 10

■ .

1 Translator's note. Sic. Should be %.

36 0

Table 9.

The relative weight of parts of the skeleton of embryos of the common porpoise, as % of the total weight of the skeleton.

Ta6aaaa 9

OrnocwreabBbtil Bec qacTeii caeaeTa n.noa,oa mopcaoà % OT o6lus ero Beca CKOJICTa

OT BCCa

G Bee oraeAoe cmeaeTa, E?

A 13.- C P03- BeeEeKe- F Bec xpetue, °6

06ani6 Bec " jltaxita pacT, Aera, % ir w x 1.,,

z Ron,' tocren

Tun, IC nOn TC33, NICCA- OT seca a: b ge pena pe6ep . rpyanub: n° ,3,n„,f1 " fore h.inci c.n Ilea re.la

Tula cKe.iera nepeainix samtex

0,884 ca n a 42 5-6 • 8,1 4,3 53,0 46,7 5,2 0,7 41,8 5,6 -

1,290. camen 48 5-6 8,6 4,6 53,5 42,7 7,4 0,6 42,0 7,2 0,1•

1,392 cameui 49 0_7l 4,7 53,2 41,3 5,3 0,9 45,0 7,5 0,1

1,485 camBa 46 . 5-6 '.: . 8,4 ' 50,9 42,8 5,1 0,7 45,0 6,4 -

1,508 camBa . 48 04-.-7. » 8,5 4,5 53,4 37,7 5,7 0,6 49,0 7,0 0,1

1,528 cameul 48 6-7 8,1 4,3 53,7 43,8 5,2 0,5 43,0 6,9 0,2

1,64 7 ." Camaa: 47 6-7 8,4 4,4 52,5 43,7 7,0 0,7 41,1 7,4 0,1

1,652 camel 50 6-7 9,0 4,5 50,4 42,2 5,6 0.7 43,9 7,5 0,1

1,833 ca meal 1 51 6-7 7,8 4,1 52,8 42,2 5,6 0,7 •44,6 6,8 0,1

1,847 ca /qua 1 52 7 7,7 4,9 54,6 40,7 7,6 0,9 42,4 8,4 0,1

1,996 camel!" 57 7-8 9,5 5,1 5,30 37,9 5,9 . 0,4 48,3 7,4 0,1

um . ramm; 70 9-10 9,9 - - 31,4 5.9 0,4 54,5 7,9 -

A. - Total weight of body, kg B. - Sex

G. - Length of body, cm D. - Age, months

E. - Weight of skeleton, % of body weight F. - Weight of cartilage, % of weight of:

a - body b skeleton

G. - Weight of sections of the skeleton, kg v - skull w - ribs X - sternum y - vertebral column • z - limbs

f - female m - male

é,, Table 10.

rioa lu ma

1103- p a cT,

mecm-uen

5-6

5-6

6-7

5-6

3-7 6-7

6-7

6-7

6-7

7

7-8

2:3

2,7

2,9

2,6

2,7 . 2,4

2,6

2:7

.2,4

.2,6

3,2

59,2 •

54,0

52,9

55,9

5-1,8

55,3

54,7

.54,7

56;7

50,4

57,1

8,6

10,8

9,3

4,6

8,1

9,1

10,0

6,9

8,4

11,1

6,7

0;9

0,3

• 0,4

0,2

0,02

0,3 0,5

• 1,4

0,3

0,4

0,3

26,1

26,9

29,6

29;1

31,5-

27,3

26,7

29,4

27,1

28,3

29,0

6,0

7,0

7,7

10,3

7,8

8,0

7,2

7,2

9,7

6,6

361

The relative weight of the red bone . marrow of the embryos of the common porpoise in . various sections of the skeleton, as % of the total meight of the bone marrow.

Ta6ouga M •

Bec xpacooro KOCTI-10r0 mo3ra 11.'1011,0B mopcBori C1311111,11.

Il pamipuou oraeaax cBeacTa, 0/0 OT oGalero Beca Bocruoro m031a

.06utues nee

. re.na, KO

061mill nec 10C 011010

moara,% OT

neca Toila

V •I gepena I peep

HY1110- H04,101-0

cioa6a

uonAnocre

J. 11111: laanunx

•

F Bec Knerioro Moira H cnenere,

X Wan-

caria " 42

cameo. 48

cameo .49

camaa 46

caMlV! 48

cameo 48

ca man 47

camen, 50

CaMen 51

calm 52

cameo 57

A. - Total weight of body, kg B. - Sex

C. - Langth of body, cm D. - Age, months

E. - Total weight of bane marrow, as.% of body weight

F. - Weight of bone marrow In the. skeleton, kg v - skull w - ribs X - sternum y - vertebral column z - limbs

1 - fore, 2 - hind

f - female m - male

0,884

1,290

1,392

1,485

1,508

1,528

1,647

1,652

1,833

1,847

1,996

0,2

0,1

0,1

0,2

0,1 0,3

0,3

0,1

0,3

I■J

„ , 362

weight of the skeleton. The relative weight of-the ribs and

sternum of the S- 8 month embryos comprises about 7% of the

weight of the skeleton, approximately equally divided between 223

the bones of the girdle and of the free fore limbs. The

smallest fraction in the skeleton is taken up by the rudimen-

tary bones of the hind limbs (averaging 0.1% of the weight of

the skeleton).

The amount of the bone marrow in the embryos ranges

from 2.3 to 3.2% of the body weight (Table 10). With the

growth of the embryo the fraction of the bone marrow in the

skeleton becomes gradually greater, probably mainly on account

of the development of haematopoietic tissue in. the vertebral

column, where there occurs a gradual displacement of the car-

tilage by the bone tissue. In the embryos the main part of

the bone marrow is localized in the bones of the skull (about

60%). Half as much of the bone marrow is contained in the

bones of the vertebral column. On average, 8% of the bone

marrow is contained in the ribs; in the sternum, represented

by a cartilaginous plate with a few centres of ossification,

in which there is also found haematopoi.etic tissue, is located

0.4% of the total bone marrow. In the skeleton of the girdle

and free fore limbs of the embryos there is concentrated about

8% of the bone marrow, while in the pelvic bones there is found

about 0.2% of its total weight.

In generalizing the materials on the weight of the

skeleton and of the bone marrow in the various bones of the

dolphins there are manifested several common patterns. During

the process of the intra-uterine development of the embryo

1 363

•

•

the cartilaginous skeleton is reulaced by the bony skeleton.

The process of the replacement of the cartilage by the bone

occurs especially intensively in the second half of the intra-

uterine development. While during the first phase of develop-

ment the skull occupies the largest part of the skeleton of

the embryo, during the following stages it is the bones of the

vertebral column which become dominant. The bones of the skull

are almost definitively formed towards the time of birth. Thus,

the weight of the skull of the embryos and adults of the white-

sided dolPhin is the same (approximately 28% of the weight of

the skeleton). In the vertebral column the cartilage is almost

completely displaced only in the sexually mature individuals.

With the development of the embryo the amount of bone marrow

increases in both absolute and relative terms, and at the same

time its relative amount decreases in the skull, while it in-

creases in the vertebral column.

According to the data of Z. I. Brodovskaya (1964), in

dolphins during the embryonic period there is found an inten-

sive haematogenesis in the sternum and the bones of the flip-

pers, which continues on also in the adult animals. According

to our data, during the urocess of the development of the em-

bryos there does not occur any marked increase in the relative

weight of the bone marrow in the flippers and sternum (see

Tables 6, 8 and 9), although in the adult animals the amount

of bone marrow in these sections of the skeleton is somewhat

increased (see Tables j and 4). The fluctuations which are

present in the weight of the bone marrow of these structures

in the sexually mature individuals probably, in the main, re-

flect individual variability.

364,

The haematopoiotic canabilities of newly born dolphins

may be judged only by indirect data. Thus, in the adult com-

mon porpoise the amount of bone marrow comprises 2.0%, while

in the 6 - 8 month old embryo this is higher - 2.7% of the

body weight. In the adults of the common dolphin the weight

of the bone marrow comprises 1.9%, while in their 9 - 10 month

old embryos (almost just prior to birth) the bone marrow com-

prises 3.0f of the body weight. Proceeding from these data

it may be concluded that in the newly born dolphins the amount

of bome marrow is one and a half to two times greater than in

the adult individuals. Between the weight of the skeleton of

the newly born and adult dolphins there is found this same re-

lationship. The study of the skeleton and bone marrow of

terrestrial mammals showed that the relative weight of the

skeleton and bone marrow of the newly born individuals is also

approximately twice as great as the corresponding values of

the adult animals (P. A. Korzhuev, 1964; A. N. Evstratova,

1965, and others ) .

During the period of intra-uterine development the em-

bryo finds itself in conditions of a relative physiological.,

hypoxia, the reaction to which is an intensification of haemato-

genesis on account of the increase zn the mass of the bone

marrow and also of its productivity. In the aquatic mammals

during the embryonic period the intensification of haemato-

genesis obviously proceeds to a greater degree along the path

of an increase in the productivity of the bone marrow, while

in the terrestrial mammals this occurs along the path of an

increase in its mass.

365

A basic feature of the adult Black Sea dolphins, as

compared to non-diving mammals that lead an active way of life,

is the smaller weight of their skeleton and bone marrow. Thus,

while the weight of the skeleton of the dolphins is as much

as 6.5% and the weight of the bone marrow - 2% of the body

weight, in the dog (P. A. Korzhuev et al., 1968) these values

are correspondingly equal to 10.5 and 3.3%, and in the reindeer,

according to the data of P. A. Korzhuev and I. S. Nikol'skaya

(1960), correspondingly 10.9 and 4.8% of the body weight.

In spite of the decreased weight of the skeleton and

bone marrow of the Black Sea dolphins, their haematopoietic

Productivity is fairly high, which may be judged by the supply

of haemoglobin to the organism. The haemoglobin supply of the

common dolphin (13 g haemoglobin per 1 kg body weight) is al-

most twice as great as the haemoglobin supply - to the organism

of the guinea pig (Cava Porcellus), which, according to our

data, has a bone marrow weight equalling the weight of the

bone marrow of the common dolohin, and a little e:reater than

the haemoglobin supply to the organism of the dog (11.9 g per

1 kg body weight).

The noted features of the quantitative characteristics

of the main haematopoietic organ of the Black Sea dolphins

are related to the shutting-off of the breathing for some

period or other during diving, and also to the life in a high

density medium, in which, in contrast to the terrestrial en-

vironment, the load on the body is evenly distributed.

366

,ili'iT1:I'ATJ'PA

1 Btm•poroua A. M. C)<e.ie•r, i<oc•rln,til aloar n relo11oa3 n Onrorenc3C y cntnlcïl.

jAol<.nattl,I TCXA, nbIn. 100, 1965.2 Ko/n.Icyca 17. A. J.[t,i\aTe:nwnasl (I)ylrt<IU151 xponll 11 cheseT n03n011091IL1X .[CnnoT-

tII,IX. «ycuexil coupenlelntail 611oaorlut», T. 39, nhln. 2, 1955.3 KopqrcYco 1%. A. IÏpOÛACAIa iieliOCo\toCTn C'l'O4IQI 3I)CI1}1A 3CdI1I03I q)I13IfOA0rnI1.

a. A^I., 1963.a^1,n'naüuolntan If troc^tllttecl<asI ^icAwnn[»^I Koparcyclr H. A., Gnaoûnnoc•.cr JT. B., L•c;crponoocz C. H., Moc7epuTOen 3. M.

OnI,IT xo:u111ecTnennoro onvcae.-Ienna rentor.1o6nr111 n t<ocTnoro I,f03ra y o6[,nttlo-nen110ro lICpuoatopCl:oro 1ze:1141tna. C6. «Mopc[<ne nl,nel<onnTatolunC». Al., «Hayi<a»,

1 3 J 4.5 Kop,;cl/ea 17. A., Tnc1306n 7'. H. O I<Onuvecrncnuaïl xapa[<Tepncrni<e hponn _II

hpouernopnt,Ix oprauon vclnlomOpcr.Iix 1eal,(j)iiiioa. «)Kypnaa 3UoalOlUfOnlloÎ[ 6tto-xur1 mt II c;)ust[onorml:>, T. 3, nwu. 2, 1967.

b Kop^u/ecr TT. A., I nasoaa 7'. H., Arnr:puncr:an 0.• 1•I., Kttnorti3apo6a Al. P.KonWIec•rlieunoe If xaaecTneunoe pacnpetenerlne I<ocrnoro mO3ra y 133poC.antx co-6ai<. c,i(ocmnmecr.asl 6noaor.In if nle,awtnna», jx^ 2, 1968.

rf Kopxc//eu 11. A., tlunoAr,cnasl H. C. KO.nïi1IecTno IrocTnoro uo3ra y cenepnoro

oriclln. RAI] CC'CP, T. 131, nl,nt. 1, 1960.8 h4po1Inoa K. A. I3econoïi if xnatnme.cl<uïI cocTan OTae:Ibnl,ix IIaCTCif Teaa B FIC-

xoroPLIX opranax (j)nnna.za. Tpynl>i ,T•41110, nl,tn. 25, 1953.9 CAeIU{oti M. M. PeayA[,Ta•rhI n3neuliluannst xpytnn,tX It ,Ie,Ii<nX HIlT0O6pa3nl,ls,

Ao61,niaenn,ix na Ila.a>>ucu 13ocTOI<e. Tpyttht 1 tAIïI^ All CCCP, nlrr. 34, 1961.

10 anpor nc< If, H. nlaTepna.tl,[ x neconoa)y If xnntityecr.oN)y cocTany xlITOn. Tpya),T-BIIi4PO, r,I,In. 15, 1940.11 Bjarltnsorl T. Some weight measurement of whales. 1\Torsk. Hvalfan;;sf.Tidende, no i, 1954.12 Slijpel• B. J. Whales. 1-flltchinson of London, 1962.

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367

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termination of the haemoglobin and bone marrow in the

common Black Sea dolphin. Symp. "Marine mammals".

Moscow, Publ. "rfauka", 1964.

5. Korzhuev P. A. and Glazova T. N. The quantitative charac-

teristics of the blood and of the blood-forming organs

of the Black Sea dolphins. "Zhurnal évolyutsionnoi

biokhimii j fiziologii" ("Journal of Evolutionary Bio-

chemistry and physiology"), vol. 3, No. 2, 1967.

6. Korzhuev P. A., 01azova T. N., Alyakrinskaya O. I. and

Kalandarova M. P. The quantitative and qualitative

distribution of the bone marrow in adult dogs.

"Kosmicheskaya biologiya meditsina" ("Space Biology

and Medicine"), No. 2, 1968.

7. Korzhuev P. A. and Nikol'skaya I. S. The quantity of bone

marrow in the reindeer. Doklady AN SSSR (Reports of

the USSR Academy of Sciences), vol. 13•, No. 1, 1960.

8. Mrochkov K. A. The weight and chemical composition of

different parts of the body in certain organs of the

fin whale. Trudy VNIRO (Transactions of the All-Union

Research Institute of Marine Fisheries and Oceanography),

No. 25, 1953.

Sleptsov M. M. Results of weighings of large and small

cetaceans, caught in the Far East. Trudy Instituta

morfologii zhivotnykh AN SSSR (Transactions of the In-

stitute of Animal Morphology of the USSR Academy of

Sciences), No. 34, 1961.

368

10. Khar'kov I. I. Information on the weight and chemical

composition of whales. Trudy VNIRO (Transactions of

the All-Union Research Institute of Marine Fisheries

and Oceanography), No , 15, 1940.

369

UDC 599.537

G. N. Solntseva

THE COMPARATIVE-ANATOMICAL AND HISTOLOGICAL CHARACTERISTICS 226

. OF THE STRUCTURE OF THE EXTERNAL AND INTERNAL EAR OF

CERTAIN DOLPHINS

The specific characteristics of the way of life of the

cetaceans and those data which we possess, concerning the un-

usually wide frequency stectrum utilized by these animals for

echo-location and communication purposes, makes the study of

their organs of hearing a very pressing question.

Up to the present time many experimental étudies have

been conducted on the capabilities of the cetaceans for per-

ceiving sounds, but insufficient attention has been paid to

the fine morphological structure of their organs of hearing.

For the elucidation of the functional possibilities of the or-

gan of hearing of the cetaceans, the detailed examination of

the morphology and histology of this organ presents consider-

able interest.

The organ of hearing of the terrestrial mammals has

been studied in detail by both Soviet as well as foreign in-

vestigators (Vinnikov and Titova, 1961). Some data are avail-

able on the anatomy and histology of the outer, middle and in-

ner ear of cetaceans. However the histological structure of

Corti's organ has not been studied in detail.

370

MATERIALS AND METHODS

The investigations were carried out on 5 white-sided

dolphins and 5 bottlenose dolphins.

The external and internal ear was studied histologi-

cally and anatomically. The material was fixed in 10:, forma-

lin and embedded in celloidin according to the usual method.

The bullae tympani were collected immediately after the death

of the animal and were fixed in Vitmaak's fixative for 3 weeks,

they were then decalcified and embedded in celloidin. Section

thickness 15e. Staining with haematoxylin-eosin.

EXTERNAL AUDITORY ATUS

It is known that many investigators (Hunter, 1787;

Yurie, 1873; Zille, 1915; jamada, 1953; Fraser and Purves,

1960; Reysenbach de Haan, 1957; Purves, 1955; Purves and

Utrecht, 1964), concerned themselves with the study of the

structure of the external auditory meatus in both the Mystaco-

ceti (Balaenoptera acutorostrata, B. musculus, B. physalus,

Megaptera nodosa) as well as in the Odontoceti (JlobicePhalus 227

melas,l'hyseter catodon, Phocaena Phocaena l Tursiops truncatus).

We have not found any studies in the Russian literature on the

structure of the external auditory meatus in the white-sided

dolphin and the bottlenose dolphin.

The external auditory meatus in the cetaceans starts

with a very small opening *behind the eyes. Further on it is

represented by a winding duct. Its inner end, which abuts on-

to the middle ear, is tightly closed by the tympanic membrane-

ligament, which separates the outer car from the middle ear.

' I •.

,5•7;

371

Pitc. 1. Flapywilmil cnyxonofi npoxo,]. (re,maToKenn1I11-3o3mi, 10x7)

Figure 1. The external auditory meatus (haematoxylin-eosin,

10 x 7).

The whole of the external auditory meatus was divided

into 5 sequential sectors (from the periphery to the center),

the preparations of which were serially prepared.

In sector 1 (Figure 1) the external auditory meatus

in the white-sided dolphin and the bottlenose dolphin is rep-

resented in the form of a very narrow slit. In sector 2 the

dimensions and form of the external auditory meatus change

markedly. The thickness of the wall increases considerably,

and at the saine time the auditory meatus itself is completely

obliterated. The lumen of the auditory meatus appears at the

very end of this sector (Figures 2 and 3). In sector 3 the

lumen of the external auditory meatus acquires an oval form

with protrusions (Figure 4). The wall of the auditory duct,

372

te) Plic. 2. II yliacTol: liztpyKlioro cayxotioro npoxcua

2a — coemiiiirresibuo.mannoe aapacTanne npoKcinta.-ibitoil macTit cnyxotsoro npoxoica, 26— I1J1OT1t.ft ilempopm.leiman coefimmt-

. Teabnayt TIUIIJb, 2o— numc:tuible nytint

Figure 2. Sector II of the external auditory meatus. 2a - connective-tissue obliteration of the pro-

ximal part of the auditory duct, 2b - dense unformed connective tissue, 2v - muscle bundles.

in comparison with the preceding sector, becomes thinner. In 228

sector 4 the auditory meatus has an oval form with a consider-

ably enlarged lumen, in comparison with the preceding sector

(Figure 5). The thickness of the walls of the auditory duct

becomes uniform. And, finally, in sector 5 the externai audi-

tory meatus differs but little in its form from the preceding

sector, though its dimensions are enlarged (Figure 6).

373

Yric. 3. Konüciuoil o1;tca lI YIlacrxa napyNCnoro c.,yronororipoxo^a

^2 - rIOflE.'1Ci'.1'C rlnpryCT^ n r^}rXf^Rp}r P.nO?_nnn 30 - CC.

KPCS1iLIC i4CAC;4bir OICj)^'R^2110111llC C7)'\01301Î no.`: (U

Figure 3. Terminal portion of sector II of the external

auditory meatus.

3a - appearance of lumen'in auditory meatus,3b - secretory glands, surrounding auditory duct.

In the preparations of sector 1 the external auditory

meatus is surrounded by numerous dermal papillae, between which,

within the epidermal cells,melanin is located in the form of

grains and granules.

In sectors 2, 3, 11' and 5 the inner wall of the auditory 229

meatus is surrounded in an annular manner by a layer of dense

connective tissue. In sectors 2 and 3 between the bundles of

connective tissue are situatedc•protein glands and individual

fat cells.

374

O

1) 11c. 4. III rtacToK Hapymoro cnyxonoro npoxon,a 4a — onaehmi (Impta cmyxocoro npoxpAa c nambucc3ut-

IIbMfl 13b11151411118111151M11, 46 — ccl<peTitte "Acme3b1

• Figure 4. Sector III of the external auditory meatus.

4a - oval form of auditory duct with finger-like protrusions,

4h - secretory glands.

In the dense connective tissue there are found almost

undifferentiated cells: fibroblasts and histiocytes. States

of mitosis and amitosis of the colis were found. lying adja-

cent to the layer of dense connective tissue are muscle bundles

(m. zygomatico auricularis and occipiti auricularis), which

extend along the auditory duct and have a varying thickness.

Between the individual muscle bundles and also between the

cartilage and the inner wall of the auditory duct is situated

loose connective tissue, including bundles of collagen fibres •

and seDarate elastic fibres.

375

Plic. '5. IV ytiacox napywnoro cayxoisoro npoxma 5a— oBamillast (popma c.uxonoro iipoxma, 56 111)0-

'11,1C EJICTIZI1*

Figure 5. Sector IV of the external auditory meatus.

5a - oval form of the auditory meatus, 5b - fat cells.

lb)

Directly around the auditory duct there are situated

many blood vessels and blood lacunae, filled with blood.

Commencing from sector 3, cartilage adjoins the exter-

na1 auditory meatus. Initially it appears on one side of the

auditory duct. Then it lies adjacent to the two narrower

sides of the auditory duct, in the form of two caps. Further

on there is traced the fusion of the two separate cartilages

of the preceding section, and already in sector 5 the cartilage

acquires a horseshoe-shaped form, surrounding the auditory

duct on 3 sides. The cartilage forms a tube which joins up

376

Piic. G. YttacTOK ü8P}'iKItOCO C.9)'XO6G1'O itj?OXOi(£IGa - Kpoael!octtvtït cocv^, (:ô - Kponel!octtaa n<u<ylla, Gc--

itAOTltitSi IICO(^lOj?1\lAClil!RSI COC;IUIIIITC^b1i8A TIdBlIb, Gi.-9:Ic2CTiI-

LICCKIIFi XpIIll(

`r'igure 6. Sector of external auditory meatus.

6a blood vessel,6b ^ blood lacuna,6v - dense unformed connective tissue,6g - elastic cartilage.

into the bulla tympani. By comparing the dimensions of the

external auditory ducts of the two species of dolphins under

consideration, by individual sectors, it can be ascertained

that these vary only insignificantly.

Thus, the external auditory meatus runs like a pigmen-

ted tube surrounded by fibrous tissue. After passing through

the fat layer the auditory duct enters a cone-shaped mass of

fibrous tissue, which is considered as being connected with

377 • the auricular muscles and the distal end of the auricular

cartilage (Purves and Utrecht, 196 )4 ).

INNER EAR

On investigating the inner ear of the dolphins it was

noted that, macromorphologically, the cochlea of the dolphins

was distinguishable from that of terrestrial mammals (for

example, of the guinea pig).

The cochlea of the dolphins is flat and completes 1.5

spiral turns, while in the guinea pig it is pyramidal and com-

pletes 4.5 turns (Figure 7).

As in the terrestrial mammals (Ya. A. Vinnikov and

L. K. Titova, 1961), the cochlear canal is divided by Reissner's

membrane into the two scalae, the upper one of which passes

to the apex of the cochlea, while the lower scala, running

from the apex, terminates at the round window.

The upper scala is divided by Reissner's membrane into

two unequal cavities, the smaller of which is the scala media.

Within the scala media on the vestibular surface of the basi-

lar membrane is located the organ of Corti. From the inside

closely abutting to the organ of Corti is the spiral lip, which

passes into the internal spiral notch. The spiral lip is formed

of flat epithelium, which in the dolphins, as in terrestrial

mammals, is arranged in Parallel rows. The cells of the epi-

thelium are large, with . large nuclei, and are disPosed along

the radius of the cochlea.

The cells of the spiral notch pass into the supporting

elements for the inner hair cells. At the point of termination

of the internal spiral notch are disposed the border cells,

which form two rows.

o 378

'

7.1".

Pue. 7. Buenumil uu,u, ,n,em41ma-6e:usGotwu _ ---

Figure 7. External appearance of the cochlea of the white-

sided dolphin.

The receptor elements of the organ of Corti are the

outer (NVK) (Figure 8) and inner hair cells (VVK) (Figure 9).

These cells differ in their structure. The inner hair cells

are arranged in a single row and have a jug-shaped form. The

large nucleus is situated at the base of the cell. They are

situated alonp,side the apices of the outer pillar cells.

In contrast to these, the outer hair cells are arranged in 234

three rows and have a cylindrical form. At their base is lo-

cated the round nucleus, which has a granular structure. The

cytoplasm is granulated.

379

8. kopTueu oprall. Ilapymmc BOAOCKOBMC E.-1C7Elf (remaToiccunuu-3031111,90x7 )

Figure 8. The organ of Corti. .Calter hair cells (haema-

toxylin - eosin, 90 x 7).

The supporting cells are represented by the outer and

inner pillar cells of the organ of Corti, which are located

on the basilar membrane. Between the supporting pillar cells

is located the tunnel and space of- Nun.

There can be clearly traced the inner border cells,

beyond which follows one row of the inner phalangeal cells,

which are situated between two inner hair cells, separating

one from another. Between the inner pillar cells and the bor-

der cells, in the cavity, are arranged the inner hair cells.

Lying adjacent to the inner pillar cells are the cells of

Deiters. With their bases they lie on the basilar membrane

and have a polygonal form with a large nucleus, situated in

the basal part of the cell.

; .

1 t.

38 0

Pile. 9. 1(opTucut oprau. Buyipennue no.nocKoable (teMaTOliC11:11111-

3631111, 90x7)

Figure 9. The organ of Corti. Inner hair cells (haema-

toxylin - eosin, 90 x 7).

• Next follow the cells of Hensen, which lie very closely

adjacent to the cells of Deiters. The cells of Hensen are

situated with their basal ends on the basilar membrane. The

apices of the cells are directed towards the outer hair cells.

They are large, with large nuclei. Beyond the cells of Hensen

are situated the cells of Claudius, which are represented by

short polygonal cells with very distinct intercellular borders.

Their cytoplasm contains a nucleus with nucleoli. The basilar

membrane, on which lie the elements of the organ of Corti,

is clearly traceable. Distinctly evident are the parallelly

arranged fibres, which are located at a uniform distance from

one another.

381

Thus, the analysis of these data makes it possible

to assume that, in contrast to the clearly observed macro-

morphological differences, there were not found any notable

histological differences in the structure of the cochlea in

the dolphins, in comparison with terrestrial mammals (guinea

pig).

CONCLUSIONS

In the study of the external auditory meatus we attempted

to elucidate the question of its obliteration in a particular

sector. Some investigators consider that, along its entire

length, the auditory duct in the toothed whales runs in the

form of an open tube (Fraser and Purves, 1960; Reysenbach de

Haan, 1957), while the studies of other authors indicate that

the external auditory meatus is a tube with a lumen that is

obliterated in some sector (Jamada, 1953).

Our data confirm the data of Jamada. The results of

the anatomical and histological studies showed that in the

white-sided dolphin and the bottlenose dolphin at a distance

of two centimeters from the surface of the skin there occurs

an obliteration of the auditory duct with connective tissue,

and this comprises approximately one twentieth of its length

(see Figure 2).

The obliteration of the auditory duct is of important

significance in the treatment of the mechanism of sound con-

duction. The comolete obliteration in the dolphins cannot

worsen the conduction of the signals to the middle ear in the

water medium and serves, mainly, to preVent water entering the

ear.

v.e^e..ea ve°^eaTRANSLATION 3185 5 of 7

F,a^e,°, 5e^ °^ea,==^°a SERIES NO.(S)aean^e^^°e °id.s°N^°° iame,

r

r

382

The f-Luznel--shaped a.uditory duct of the dolphins, which

is completely isolated from the medium, confirms the position

that they make use of another (as compared to the terrestrial

mammals) pathway for conducting the acoustic signals to the

middle ear.

On account of the "acoustà.c transparency" of the fat

layer of the hypodermis, the signals from the water penetrate,

with minimal losses, through the integument and, reaching the

cavity of the auditory duct, they move along this to the middle

ear.

As to the studies of the inner ear, there are all

grounds for assuming that in its anatomical structure the

cochlea of dolphins is well adapted for the perception of

ultra-sounds. This is indicated by the presence of the lower

loop of the cochlea, since, according to the existing electro-

physiological studies conducted on terrestrial mammals (Altukhov,

1911.9= Andreev, 1911.0, Bekesy, 1951H, Davis, 19,47, and others),

the shorter fibres lying at the base of the cochlea perceive

the high frequencies, while the longer fibres located in the

apex of the cochlea perceive the low f.requenciès.'

JJJ xTLPATYPA

• 1/tns•Yaoo l'. B. Yc,nonnopcrj),^c)iTOpllafl ;tes,TC.m>rioçTh co6axrt ripn srccncpuMc;r-Tü.'twnUT\r noupelaeutnl nepncI)epumCC).oro oTijensr c:Tysol;oro aua.a113aTOpa. «IIpUU,T_

tOIrlIr0JL flK)'CTIIKII», T. 1, 1949.2 flxopeea R. A. O q acoropUx uor,t,ix Aanrn>ix, xapar.Tepnsyloutnx ttesiTWhrrocTt,

nnyt;onoro aua.mtsa'ropa. 7-c coueutsulne no npo6.,,i. rs),tcru. nepnn. +ZesIT., nocn. na-1`u)'ru IJsrnnon<r: Tcsncw uor..q<)AOn. M.-,I1., 19,10. -

3 Bwttuu:oa Sl. A., Tlrrocra 31. K. Koprtrco opt'an. 1•13;>,-rso AH CCCP, M.-,r[_

l ir^31.I' Bckest/ V. Some F.Icctro - Mechanical Propertic^s of the organ of Coi ti.

Aun. or Olol., Rhinol. a. L.aryng., vol. 13, no 2, 1954.5 Davis 11. Biophysics- and physiology of the inner car. Physiol. Rcv., vol.

3, _ 1957.

6 Fraser F. C. and Purves P. E. flearing in 'Cetaceans, Bull , of the htitis-li. mus. (nat. hist.), Zoo logi, vol. 7, No I. 1960.

Huntcr J. Observations on the Strukture .and Oe.mnorny of Whales. Phil. Trans. 77. 1787. 8 Jantada. M. Contribution to the Anatomy of the Organ of Hearing of Wha-

les. Sei Re.p. Whales Research Inst. No 8, Japan. 1953. 9 Lilli J. C. Cetacea. Brit Antarctic. <Terra Nova». Expedition 1910. Nat.

IIist. Report. Zool. 1. no 3, 1915. 10 Marie J. On the Organisation of the Cealing Whale Globicephaius melas. Trans. Zoul. Soc., 8, 1873. _

1 1 Purves P. E. The Wax Plug in the external auditory meatus of the Mysti-ceti. Discovery. Reports 7,7, 1955. 12 Parues P. E. and van Utrecht W. Z. The anatomy and function of the ear of the bottlenosed dolphin Tursiops truncatus «Beaufortia» 9, no 111, 1964. 13 Reysettbach de Haan F. W. Hearing in Whales. Acta Oto— Laryngologica, vol. 137, 1957.

383

REFERENCES

1. Altukhov • G. V. Conditioned reflex activity of the dog

with experimental injury to the peripheral section of

the auditory analyzer. "Problemy fiziologicheskoi

akustiki" (Problems of i)hysiological acoustics), vol. 1,

1949 ;

2. Andreev L. A. Some new data, characterizing the activity

of the auditory analyzer. 7-e soveshchanie po problemam

vysshoi nervnoi deyatel'nosti, posvyashchanno pamyati

Pavlova. Tezisy dokladov (7th conference on problems

. of higher nervous activity, dedicated to the memory of

Pavlov. Reports of Proceedings). Moscow - Leningrad,

1940.

3. Vinnikov Ya. A. and Titiva L. K. The organ of Corti.

Publ. USSR Academy of Sciences, Moscow - Leningrad,

1961.

384.

UDC • 99.74.5

G. M. Kosygin and V. N. Gol'tsev

DATA ON THE MORPHOLOGY AND ECOLOGY OF THE HARBOUR SEAL 238

OF THE TATAR STRAIT

The study of the pinnipeds in the Tatar Strait was

initiated by S. V. Dorofeev (1935), who established that the

harbour seal, ringed seal, bearded seal and ribbon seal are

encountered here in the spring. The predominant species in

the Strait is the harbour seal and it comprises about 80% of

the catch of the hunting of the coastal inhabitants. The re-

maining species are not very numerous. The approximate periods

of whelping of the harbour seal, ringed seal and bearded seal,

and the period of moulting in the harbour seal were establtshed.

Later, P. G. Nikulin (1935) and E. A. Tikhomirov (1956)

placed the Tatar Strait in the category of unpromising regions

for commercial hunting.

These conclusions were made at a time when the seals

were caught for the sake of obtaining the fat. Now, however,

considerable attention is given to procuring fur skins. There

-f ore our attitude to the Tatar Strait as a commercially ex-

ploitable region should be re-examined. In an aero-visual

survey of seals, which was conducted by the Pacific ucean

Research Institute of Fisheries and Oceanography (TINRO) in

1968 (Fedoseev and others, 1968), a fairly large accumulation

of harbour seals was noted in the Tatar Strait. The study of

the harbour seal population that is living here, which has not

been affected by ship-based sealing operations, is of both pure-

ly scientific as well as applied interest.

385

Foin March 14 until April 7 of 1969 co-workers of the

TINR0 on the sealing ship "Sanzar" carried out field studies

in the Tatar Strait. Included in the purpose of the cruise

was the study of the distribution and biology of the seals,

and also the elucidation of the feasibility of commercial

exploitation of the animals. The exploratory transects covered

the region of the strait from 48 to 500 lat. N. The collected

materials were treated by the methods accepted at the TINRO

(Smirnov, 1934; Rokitskii, 1961; Chapskii 1963, 1967; Yablo-

kov, 1963, 1966; Tikhomirov and Klevezal', 1964; Sokolov et

al., 1966). Some of the materials on the harbour seal of the

Sea of Okhotsk were passed on by us to A. V. Yablokov. In •

addition to the authors, A. I. Corshkova participated in the

collection and analysis of the materials.

Many authors have devoted studies to the description

of the harbour seal, and yet the overall morphological charac-

terization of this seal is far from being complete. What has

been said applies to the same degree also to the harbour seal

population, living in the Tatar Strait.

Coloration of the hair coat. Earlier

(Kosygin and Tikhomirov, 1969) in describing the harbour seal

or mottled seal (Phoca larp-ha) its coloration was grouped into 239

4 types: light-coloured, slight mottling, very large mottling

and typical.

In the 64 harbour seals that were examined from the

Tatar Strait, the typical coloration predominated (6 0%); stan-

ding in second place was the slight mottling (33); 6 animals

had the very large mottling, and 1 was lightly coloured. The

colouration of the harbour seals is similar to that described

386

grw

by S. P. Naumov and N. A. Smirnov (1935), S. I. Ognev (1935)

and K. K. Chapskii (1963, 1966) on the basis of individuals

from other parts of the species range. Among all of the white-

coats of the harbour seal that were found in the Tatar Strait

only two pups were encountered that had the smoky coloration,

characteristic of the young of this species of seal from the

Bay of Peter the Great.

Length and weight of body. The average

length of the body (z0 ) of the harbour seal, living in the Sea

of Okhotsk, is 165.3 cm in the males and 151.6 cm in the fe-

males (Pikharev, 1941). There are available measurements of.Z0 on

the harbour seal from the Tatar Strait on individual animals

(Gagichko and Surzhin, 1935) from an autumn catch, which have

not been included with our data, because of the small numbers.

According to our data, the seals of this species from the Ta-

tar Strait are larger than those from the Sea of Okhotsk: the

body length (k) of the males and of the females was, corres-

pondingly, 169.0 and 167.7 cm. On the basis of this feature

they differ little from the harbour seal of the Bay of Peter very.

the Great (Sea of Japan) and are slgnificantly larger than the

females of the Bering Sea, Kuril and Sea of Okhotsk populations

(Table 1). There are also significant differences between the

males. The average length of the body of the whitecoats is

101 - 103 cm. On the basis of the total weight of the body .

(Table 2) and the weight of the carcass without the skin and

the fat ( sculp ) the sexually mature harbour seals from

the Tatar Strait are smaller than the individuals from the Bay

of Peter the Great. The weight of the whitecoats is the same

387

as in the Bering Sea. The weight of the sculp of the

harbour seals of the Tatar Strait ranges from 24 to 43 kg,

averaging 36.5 kg, in the adult males, and from 22 - 56 kg,

with an average of 35.1 kg, in the females. The relative .*

weight of the sculp comprised 31.40% (averaging 37%) in

the males and 29-49% (averaging 39.6%) in the females. In

the harbour seals from the Bay of Peter the Great during the

whelping 1-)eriod this equalled 43.3% in the males and 43.2% in

the females.

Meristic features. Many authors (Yablokov,

1963; Yablokov and Klevezal', 1964; Yablokov, 1966; Chapskii,

1967; Sokolov et al., 1968; Kosygin and Tikhomirov, 1969)

employ the number of vibrissae, of tracheal rings and of ribs

as a morphological feature.

The labial vibrissae of the harbour seal of the Tatar

Strait are seated in 7 - 8 rows. The same number of these

were counted in the harbour seal from the Sea of Okhotsk (Yab-

lokov, in litt.). The number of vibrissae in the males ranges

from 37 to 46, with an average of 43.2 ± 0.43; in the females

- from 37 to 50, with an average of 42.7 ± 0.44. In compari-

son with the harbour seal of the Bay of Peter the Great, no

great differences were found in their total numbers; on ave-

rage, there were somewhat fewer vibrissae in both the males

(t = 0.88) as well as the females (t = 0.36) of the former

population.

In two females (a whitecoat and a sexually mature ani- 242

mal) there was found a null row, in which there were from 1 to

vibrissae on each side of the snout. Such an occurrence has

* Tramslator's note. Sic. Presumably this should be 31 - 40%.

Pem

Rea:ion nøI

Sex

5,65 9,25

7,11 4, 9 1

7,99 6.43

-0,38

-0,03

+0,18

B3poce c

C2Me/t- camK3 .114 nz Urn

94-111 87-114

84-112 89-102

80-108 78-109

99-108 83-110

Toropc},:trik

3a.niel fleTpa Be./moro

5ep1Ir000 mope

Males CaMIZIpf: tj-3= +2,12

it--4= +2,60 Et-= +1,7O

Females Cammt t t -2,- 0,39

t 1-z +5,81 t_= +5,81

ti_4=±7.67

* Zammie A. H. Benustua. " ,11,anliNe 3. A. Tmomilpona.

* Data of A. N. Belkin ** Data of E. A. Tikhomirov

C CO CO

Table 1.

The body length of sexually mature individuals (z0v ) and

of whitetoats (ze. ) of the harbour seal from different regionsea6xxua 1 crn • ji(amia Te.na no.riosoapembtx oco6eii (Zcv) 6e.nbRo8 (Zc) oaprx paalibm paionon, c.e. •

7

9 7

60 47

4 4

lirn

Tata Strait

Bay df Peter the iGreat

BerinÉ Sea

Darili Islands* KYPubcKtrd ocTpona*

MO-

camen111 cammaf

corneure cammor

. ca eel camof

Ca MC11.-72, comNa.l.

comen.111 camIcaf

• B b X 1-t:

101,2±-1,91 102,7-±3,41

95.8 ± 2.37 M,0±1,61

95,9 ±-1,03 94,1 ±-0,93

103,0 103,7

16 146-173 158,0±-1.68 6,73 35 145-179 157,7-1-13,2 7,81

3 152-168 159,3 10 144-173 155,5±2,78 8,80

12 128-170 148,5±-4,14 14,33 13 123-156 141,2±2.52 9,09

14 134-163 150,0±2,58 9,66 6 135-154 1 45,5±0,89 2,19

19 135-171 153,3±2.18 9,49 16 134-155 142,1 ±-1,52 6,07

t male4 .

cam"a femare

+0,14

-

+1,51

+0,46

+4,10 Sea o'f OkhotgkUol;oe Pe"

m - male f - female

r

I t tLïat^

C3tiC1l-male -Q I Ca)tlSa femalé

Tatar^ Strait

Dây of Petertn_e Great-uril; Islands:<

,B-,^er in Sea

Table 2.Total body weight of the sexually mature individuals and

whitecoats of the harbour seal, caught during the period of ïlt^liiû^i^

in different regions, kg.Q611(L1H IIeC Tc:la iloJI0Q0317CRb1X 0^0Ûei! A^JeJIbISOS :

IaprY.^ J1,O6bITbiX II IIe17I10,^ ll.leHK41 B p23^^^ ^alt OI 1X+ hZ

3a:cIIB 1-leTQâBe:Illxoro

Kypl :zbcKi[eocTpo13a

BepII11rOüo Mope

canlltb 9canlKilf 7

ca1I11i, 4calIr.uf 4

cauub>ni 3Gca>IKII 34

13-30 19.4± 1,7811-31 22,0 =2,7G

8,0-27.0 12,9-_` 1,969,4-25,0 14,7--2,19

15,F-29.5 23,017,0-32.0 27,0

5,8-3:3.8 2 1.9 -!-1,257,3_.3:},7 22,2-!-1,12

5.047,3-1

5,$9 -O,6G 3 IOCi-125 t i3,3

5,80 10 80- 150 11•1,0--6,11

- .- 13 62-113 90,3! 3,99

7,53 - 0,19I 226,51, 16

iIpl1111Ujt HH031)aC9'C OT pomucuusl u 111^ioicl'110 .to oAtloro xecsuta.

Young- from birth to an age ofabou^ one month.

M - malesf - femal.es

t7-147I

55.4 -5,4866,0 - 3,2350-100

males :Ca1fIJ,b1: ES-3= `i' 1,ô'1

ts_,=+2,26

7 60-92 75,2+4,r9

19,3

14,4 +2,4112,7

25,7 +3,0412,9

'i'emales :Cfl1IKH: tI_z=-3,75

fi_a=+2,51t1_.=+5,50

t

,o

390

not -oreviously been noted in the harbour seal, either in the

Bering Sea (Chaoskii, 1967) or in the Bay of Peter the Great

(Kosygin and Tikhomirov, 1969). D)reover, vibrissae were found

in one sexually mature female on the lower jaw. They were

distributed as in the bearded seal (Kosygin, 1968).

In the harbour seals of both the Tatar Strait and the

Bay of Peter the Great there are 1 - 2 nasal vibrissae and 3 - 6

ocular vibrissae.

Between the males of the harbour seal from the Tatar

Straight and from the Bay of Peter the Great, in the average

numbers of labial vibrissae, there is found a significant dif-

ference in the first row (t = 2.5) and a highly significant

differenc (t = 3.33) in the foùrth row. In the remaining

Cases no siîrnificant differences were noted, either in the

total number of vibrissae or in the number of these in the dif-

ferent groups in the populations examined.

The trachea of the harbour seal from the Tatar Strait

has no significant differences in its structure from that which

has already been described by A. Si Sokolov et al. (1968).

In 6 out of 28 individuals (21.4%) the rings situated around

the bifurcation had a left overlap. In the harbour seals

caught in the bay of Peter the Great (26 head), tracheae of a

similar structure were present in 31A of the examined indivi-

duals.

59 - 76 rings were counted in the tracheae: in the

males the average was 67.5 ± 1.17, while in the females this

was 63.7 ± 1.07 rings (Table 3). The differences betweeen

these were significant (t = +2.48). As is evident from Table 3,

l'eeffls ii Um 1 M *m

16

16

02-76

61-71

64-76

61-75 65--73

+2,48

+1,18

+1,33

391

Table 3. Total number of tracheal rings in males and females of the harbour seal from different regions.

Tat-Sauna 3

()Stec nuantiecTuo Ko.nett, Tpaxeu camnors n canon naprif 13 pammix pafiouon

1

Camuu

n I 11m I M-•.m

P.U.LuS Cnmnu p.ca...t..e

Patiu Region

Tatar Strait TaTapcnnii upo- Jilin

Bay of Peter 3a.mil nona t e Great BumEon) Sea of ÛkhOtSPXOTenoe mope (according to (no 516.1oKo-Yablokov, 1. 066PY, 196.6; m in litt, 1969,5 1u' 1969) Kuril Islands' 30ToiTi .p" boepi<'1" e • Bering Sea Bepunr000 m ope

13

10

24

23 8

69,4 ± 0,54

69,5 ±0,74 68,5±:0,96 /1_3=1-1,12

h-4=-1,08 ti-5=-0,66

67,5± 1,17

65,9 ±0,82

98

11 7

«59-71

61-70

60-75

66-75 36-77

71,2±0,83 72,7 ±1,42 t1-2=-0,33 /1_3=-3;18 h-4=-5,73 /1-5=-5,17 •

63,7±1.01

64,1±0,64

68,0 ±0,84

—1,88 —2,46 •

the degree of the differences in this feature between the

harbour seal of the Tatar Strait and the other populations

of this species is not uniform: differences were not found

between the males, though, apparently, they are real between

the females from the Kuril Islands, the Bering Sea and the Sea

of Okhotsk.

A. V. Yablokov (in litt) made a count of the sternal

and and asternal ribs In 16 males and females of the harbour

seal from the Sea of Okhotsk and found a variation in their

number.

In the 23 harbour seals from the Tatar Strait that we

examine d no deviat ions were noted in the .number of ribs : each

had 10 sternal and 5 asternal ribs .

Some data on other internal organs .

The investigation of the internal organs of this spe cies is a

392

continuation of similar studies, which were started on pinni-

peds in 1960 (Scheffer, 1960; Kleinenberg et al., 1965; So-

kolov et al., 1966; Kosygin et al., 1969; Sokolov et al.,

1969).

In a description of the superficial structure of the

lungs of the harbour seal from the Bering Sea, attention was

drawn to the fact that in addition to the so-called typical

form (a not very deep, clearly evident notch on the acute edge)

there are alo found variations, in particular: among the 27

individuals examined in one individual the lungs had a lobed

structure (Sokolov et al., 1966).

A similar phenomenon has also been noted in other po-

pulations. Thus, among 31 specimens which were caught by us

in 1968 in the Bay of Peter the Great, in 8 the lungs had an

atypical form: in one animal there were deep notches and

grooves on the lungs, while in the other 7 there were no notches

of any kind at all.

Of the 80 examined harbour seals from the Tatar Strait,

24 individuals had atypical lungs:. 8 were entire and 16 with

deep notches. Some of the lungs had two notches with deep

grooves running from these, dividing the lungs into almost in-

dependent lobes (Figure 1). •

Either both of the lungs or one of them were atypical. 244

Both the right and left lunes, to an approximately equal deg-

ree, were noted as having a superficially different configura-

tion.

Gagichko and Surzhina (1935) had data on the weight

of the internal organs of the harbour seal from the Tatar Strait

393

Plic. 1. uacTo1•a ncrpevaeNtocrn pas.an11ttoilaropaum acrlnts .iaprn n nonYanunnx:

R- Sepunrona etopsr, 1 ; - 3anuna Ilerpa Bentnco-

100 [

cc ô a S C a e c

A. B % `%

o' UuUlY^

ea.d)opma aernnx:

a- 71muenan; G- a-rtmisynas[ nonblsaran; c- arn-tfanan ücawoasaccnsnsan.

Figure 1. The frequency of occurrence of the different forms

of lungs in harbour seal populations from:

A - Bering Sea; B - Bay of Peter the Great;V - Tatar Straight.

Form of lungs :a - typical; b - atypical lobular; c- atypical

41entire^

but, because of the fragmentary nature of these data, the oc-

casional lack of information on the sex of the animal and also

because this catch was made during the autumn period, these

data, unfortunately, could not be utilized. Our materials in-

dicate that in the adult males, as compared to the females,

the average weightsof the heart, liver, spleen and diaphragm

are somewhat greater and equal, correspondingly, for the former

0.56 ± 0.04; 2.63 ± 0.21; 0.28 ± 0.02; 0.54 ± 0.23 kg, while

for the latter the values are 0.52 ^ 0.006; 1.79 ± 0.07;

0.24• ± 0.02; O.1I-8 ± 0.02 kg. In this case the differences

in the weight of the liver are at a highly significant level

(t = 3.81). The index of this organ is also greater in the

males (Figure 2). In the females there is found a comparatively

394.

60 30 0 30 60C4b/ . Ca^A U,

mmes feriaPile. 2. Bec unyTpetitntx opraxou naprll

906b1TLIX B P8311bIX p8ÎIO11i1X.

060311 a q e fi il n:I- Tarapcrnrt npo.-Iun; 2- lieptrllrono nlopc; 3-3anun 17rrpa Be.vtroro; A- Kypu.il.ctulc orrpona

A- talmevlnlh: 6- tte•tculi; li - nérl<ttc; r- ri<cay;tor,;t( - cepAUe; F. -- ceneaet,ha; )n -- nuachparNla; 3-

noer.a; ff - noa 1;e:ryJw4ltast ;ece.nc3a

Figure 2. The weight of the internal organs of the harbour

seals caught in di.-fferent regions.

Legend

1-Tatar Stra i:t ; 2'Bering Sea; 3 - Bay ofPeter the Great; 4• - Kuril islands.

A - intestine; B - liver; V - lungs;. G - stomach;D - heart; F- spleen; Zh- diaphragm; Z - kid-ney; I - pancreas#

high relative weight of the lungs (22.6 ). Differences bet-

ween males and females in the cardiac and diaphragmal indices

were negligible (in the males and females these were corres-

pondingly equal to 8-5 and 8.6%, and 8.0 and 7.9%).

The fact should be emphasized that the relative weight

of the lungs and heart of the animals from the Tatar Strait

is somewhat I.ower than of those from the Bering Sea. Thus,

in the Bering Sea females the index of the lungs equals 2^a•.8%,

of the heart - 11.1%, of the diaphragm - 8.0%. In general, 21I•5

the relative weight of most organs of the females exceeds that

of the males.

395

The noted characteristics of the internal organs of the har-

bour seal depend on the living conditions, which are specific

to each region (Sokolov et al., 1969).

Craniometric characteristics are pre-

sented in Table 4. The index of the greatest length of the

skull, calculated from the body length (7 ) comprises 13.7

and 13.6%, respectively, in the males and females. Thus, the

sex differences in the leneth of the skull are insignificant.

In this feature the skull of females of the harbour seal from

the Tatar Strait is, on average, somewhat longer than in those

from the Bay of Peter the Great (t = +0.80), the Kuril Islands

(t = +0.73) and the Bering Sea - 207.8 mm (according to the

latest data of Chapskii, 1967).

Notable differences between males and females are pre-

sent only in the rostral width and in the length of the nasal

bones. The absolute length of the nasal bones and the right

upper row of molar teeth in the females from the Tatar Strait

is a little greater than in the females from the Bay of Peter

the Great.

In the skull of the females from the former region the

value of the zygomatic (greatest) width is somewhat smaller

(t = -2.26). This same difference is also noted in the rela-

tive dimensions. Thus the index of this feature, calculated

from the greatest length, on average is equal to 58.0% for the

individuals from the Tatar Strait and 61.4% - for the Bay of

Peter the Great.

The dimensions of the longitudinal diameter of the al-

veoli of the right upper canine show differences that are

close to significant (t = -2.57).

396

For a more complete characterization of the harbour

seal of the Tatar Strait it would be desirable to compare its

craniometric data with those of the neighbouring population,

from the Okhotsk Sea. Unfortunately, we do not have any of

our own skull materials on the Okhotsk Sea harbour seal, while

in the published studies these are either too scarce (Allen,

1902; Ognev, 1935) or have been presented without subdivision

according to sex and region of catch (Naumov and Smirnov, 1935).

Thus, the analysis of this material shows that, on the

basis of the morphological features presented, there are fea-

tures of similarity and noticeable differences in the harbour

seal which lives in the Tatar Strait, in comparison with indi-

viduals from other parts of the species range.

Food objects. Out of 76 harbour seals, aeed

1 to 41 years, that were caught, food remains were contained

in the stomachs of 27 animals. 20 of these animals were caught

on the ice and 7 afloat. In most of the seals the food was

strongly digested; in some stomachs there were retained only

parts of the skeletons and otoliths of fishes, and beaks of

cephalopod molluscs.

The food base of the harbour seal in the Tatar Strait

during March-April consisted of fishes, mainly the Korean cod.

These were found in 25 stomachs (93%). These data confirm

the conclusions of previous authors (Ognev, 1935; Chapskii,

1963; Kenyon, 1965, and others), that fishes play an impor- 247

tant role in the food- Of this seal.

In second place were cephalopod molluscs, found in 10

stomachs (37%), and last were crustaceans, in 3 stomachs*.

* The species affinity of the food objects was determined by L. I. Semenenko.

115

Greatest length

Greatest_zygoma- I1C CIT

Kastoid width

Rostral Width

Length of nasal bones i

Width of nasal bones

Main dimensions of the skull of sexually mature harbour 2 seals from "different regions, mm. . ' Tanza. 4

OcK0eubte pa3Mepb: nepena no.noB03pedimx aapr H3 pa3nb1x parionoti, itAt

Longitudinal dia-mêter of alveo-lus of ilipper can

Length of rostra' portion

Length of upper row of molgr eeth

.1.0.-1jd..I. ›D tel. Ci...L C. May u_L .L-- GUL t,11. U.L c.")..1.,

Tarapcmiet npoatm 3a:tun lima lienmcoro riptt3Harc 110.1 t

Feature sex n i un 1 Al+m 1 a n 1 Uni ' I .41-i:m I e

11aitgonEnuag es.,7u1!a cameo 8 210--225 215.5+1,77 5.01 • 3 205-230 215,9 - - caM!Ç 23 199-226 212,7:4:1,30 6,24 8 196-232 209,4±3,92 11,10 +0,80

Halifianwag criyzoBasi came 8 118-134 126,1 ±1,57 4,45 3 129-141 134,5 - - ampinra camNz 23 116-131 123,5+0,93 4,45 6 123-141 130,0±2,73 6,69 -2,26

Macrounast III npuna caMeLifl 6 117-129 • 123,7±1.70 4,27 3 120-133 125,8 - - ca..+I1 22 113-12G 120, 4 ±- 0,70 3,56 6 117-123 122,3±1,71 4,18 -1.02

PocTparibnam Lunpuna came! 8 37-43 39,4-±-0,65 1.85 3 39-45 42,7 - - cam; 23 32-42 37,7+0,47 2,24 11 35-47 40,4+1,45 3,56 -1,77

11 .1 i 1 f 1 a HOCOBIDIX Komi*/ CI MCTI 8 50-60 5C,6±-1,19 3.38 3 50-59 55,0 - - camK 20 46-59 52,6 ± 0.83 3,73 9 48-58 52,5±0,58 1,75 +0,05

U-iltptilla 1100013b!X KOCTeel camel 8 11-17 14,1 ±0,65 .1,58 3 15-16 15,6 - - CtIME 20 12-15 13.5+0,22 0,97 10 . 13-16 14,3+0,31 1.00 +1,97

/Luna pocTpaohnofi tiac .rn ' ca m e ! 8 103-109 106,3±0.88 2,50 2 101-108 104,5 - - cammz 21 97-115 105,2±0,93 4,28 6 100-119 107,8±2,89 6,85 -0,86

il.inlla Bcpxnero pH,g,a KOpeH- caeu 8 45-48 46.1±0,33 0.96 3 43-49 46.0 - -

111,1X 3y6OB ' • CZIMKt 21 44-50 46,8 ± 0,47 2,17 11 45,9+0,43 1,43 +1,25

11p00abHuii zHameTp aohne- camel 8 10--13 11,8±0.35 0,99 3 11-12 11,6 - - p.ibt Bepxnero Kohnia ca NTE. 21 • 10-13 11,2+0,20 0,92 11 11--14 12,0±0,22 0,72 -2,57 me •

r, camel's- caNika = -1-1j7 ea Nten -CaMxa = +1,97 cameu - cam = +1,79 ca camKa +2,12 male-female

camell - carom +0,88> /7 ca melt - ea mma = +0,86 /9 amen - camiza = +1,21 /9 ea nieu - camKa = +1,50

m male f - female,

398

The cephalopod mulluscs most frequently found Ln the

stomach contents were octoouses, lt is characteristic that

molluscs predominated in the stomachs of sexually mature ani-

mais. This characteristic of the feeding of the harbour seal

has also been observed in the Bering Sea (Gol'tsev, 1969).

Sand and small stones were found only in those stomachs

in which there were contained some benthic animals, which had

been the objects of feeding. It was obvious that these had

been ingested by the harbour seal while hunting for its prey,

as is the case among other species of pinnipeds (Ggnev, 1935;

Kind, 196 )4; Mohr, 1963 and others).

The maximal weight of food found in the stomach (3.27

kg) was discovered in a seven-year old female, containing the

remains of 19 specimens of Korean cod. A weight of food simi-

lar to that found in this seal is not a frequent phenomenon.

S. P. Naumov (1933) reported the finding of 80 specimens of

navaga (saffron cod) in the stomach of a harbour seal caught

on the Shantarskiye Islands. Unfortunately, their weight was

not given.

Distribution. In that part of the Tatar Strait

which was investigated by us, the harbour seal predominated

in encounters of pinnipeds. Females, which had whelped, •with

their young were mainly concentrated 20 - 30 miles from the

continental coast off Sovetskaya Gavan' (Figure 3), and here

there could sometimes be seen up to 77 whitecoats per day from

on board the ship.

Large patches of sexually immature harbour seals were

found in the northernmost part of the region of the studies,

L

399

• not far from the Sakhalin coast. The harbour seals were found

on both flat as well as on hummocky ice.

During the period of observation it was noted that

individual harbour seals whiçh were participating in the repro-

duction were, for the most pà.rt, situated close to the sea

edge of the ice, while the sexually immature and maturing ani-

r;:als kept themselves deep in the ice fields, at times at a con-

szderable distance from the edge.

Some representation of the age structure of the popu-

lation of harbour seals in the Tatar Strait may be obtained

from an analysis of the small series of our samples (76 speci-

mens). Since the hunting was conducted primarily on the whel-

ping patches, in the catch there was a predominance of adult

n.n3.::als and yoinig: of the previous year. There were few sexual-

ly immature animals, aged 1-^^years, and therefore the age

structure of our catch may to some extent characterize only

the part of the herd of bearing age.

1^1•The main part of the catch consisted of seals 5

years old (81%). The remaining 19% comprised animals aged from

15 to 11.1 years. The average age of this part of the herd was

11.8 years.'tTe will note that among the 611. harbour seals that 21I•8

were caught in 1,062 in the Bering Sea, no animals older than

23 years were found (Tikhomzr.ov, 1969). In the Ckhotsk Sea

and Bering Sea populations of harbour seals the average age

is, correspondingly, 11.5 and 9.9 years. The relatively high

average age of the harbour seal population of the Tatar Strait

indicates that it, like the two former populations, has been

little affected by commercial hunting.

4,1 /'i2°

■717 r

U25) IY • 41:,‘ ei3?M7elo r'l Q-ciirc 2

dd •41-9-)./ çt ri

-

A lE CO

r. CO CK

•\- 17 cis\ S *Or S 1

V -

.y Sovetskae Gavadee J rnEro

ç 0[3E1' CK ALI to Ugleh

fiA ri 8v-

5 13 "7r- G

ikoo

Puc. 3. BeTpmemoub Tionenefi n Ta -rapcKom npo:ntne 13

piton C 20 Malrfa no 6 anpeon 1969 r. 060311w-wino-I: A — anpra; B — oaxi.a8; /3 — cuily , t; RphulaT im.

R nun Irreoe — nc -rneveno 110.1011Cii, B 3,IaMelI aTC.1C — Ii TOM qucne 15C.ThKOB. chnrypbt. 1l3o6panienume cunounwil ounncii, — BCTIIC ■10CMOCTt, C 20 no 26 mapTa; npepunucTalt annun — C 27 maivra no 8 anpean; npepbt-

Blic-ran annnsi c TOMBOil C 3 no 6 anpe.in .

Figure 3. The frequency of occurrence of seals in the Tatar

Strait in the period from March 20th to Avril 6th

1969.

Legend: A - harbour seal; B - bearded seal; V - Steller's sea-lion; G - ribbon seal.

In the numerator - seals encountered, in the de-nominator - whitecoats included in above number. Figures, illustrated by a continuous line - en-counters from 1,.arch 20 to 26; interruped line - from iv_arch 27 to Avril 8; interrupted line with dots - from April 3 to 6.

14.01

o

o

249 In the habitats of the harbour seal in the Tatar

Strait there also live other species of pinniPeds. For example,

Steller's sea-lion is encountered here (Naumov, 1933; ügnev,

1935). The former author considered the encounters with the

Steller's sea-lions to be fortuitous.

We have noted considerable accumulations of Steller's

sea-lion. In groups of up to 15 head and singly, these seals

were found near the edge of the ice close to the Sakhalin coast.

Bulls and young animals predominated in these encounters. In

all, we registered 160 Steller's sea-lions. In addition to

these, there were encountered individual,speciplqns of the . .

bearded seal and the ribbon seal.(see

Among the factors determining the concentration of the

harbour seal in particular sections of the Strait, the main

ones, apparently, are the ice conditions and the feeding con-

ditions. According to an oral communication from G. A. Fedo-

seev, A. V. Evzerov and G. P. Kovalev, co-workers from the

T1NRO, who carried out an aerial survey of the seals, on April

1st 1969 in the Tatar Strait the white hummocky ice, suitable

for the whelpinz and resting of the harbour seal, was distri-

buted northwards from the region of our studies for a distance

of 60 miles, to the-latitude of the settlement of Aleksandovsk,

further on up to the narrowest part of the strait there was

slush, and no seals were registered on this. According to our

observations, the southern border of the ice extended not far

from the settlement of Uglegorsk. Here was noted the southern-

most point of encounters with the pinnipeds.

4.02

Under the action of the currents and winds the ice

fields gradually were displaced and extensive stretches of

open water were formed, at times near the continental coast

and at times near the island of Sakhalin. In accordance with

this, the animals were encountered on the edge of the ice

either close to the continent or near the island.

According to V. L. Andreev (1965), in February - April

in the Tatar Strait there occur spawning aggregations of the

Korean cod. These aggregations of the fish shift from place

to place. Probably, following these the harbour seals also

move from one part of the gulf to another.

Periods of' whelping and moulting.

During the period from Llarch 20Y -to April 2 among the encountered

pups there were whitecoats aged 3 - 5 - 10 days. Consequently

the whelping commenced in the second third of 1:.arch, which

once again confirms the conclusions made previously by S. V.

Dorofeev (1935) and G. P. Nikulin (1935).

With the aim of elucidating the conditions under which

the whelPinp.' of the harbour seals occurs, we conducted obser-

vations on the temPerature of the air and of the surface layer

of the water. In the Tatar Strait the temperature of the air

ranges from -2 to -9 ° C, while that of the water ranges from

-1 to 2 °C. It was found that during the period of the whelping

of the harbour seal in various parts of its range (Tatar Strait,

Bay of Peter the Great, Bering Sea) there are no great differ-

ences between the air and water temPeratures.

In many of the harbour seals that were caught in the

first third of April the ovaries already had mature follicles,

1403

which indicates that the animals are ready for ovulation and

the next Pregnancy.

At the end of March - beginning of Aoril the hair coat

of the snotted seals was robust. Among the sexually immature

animals there were found individuals with loosened hair, and

rarely - with portions of bare skin. The first moulting in-

dividuals of the younu harbour seals (ragged-jackets) were

encountered on April 1st.

Recommendations for the commercial

exploitation. It is known that at the present time the

sealing flotilla of the UMRZF* is paying considerable attention

to the procurement of fur pelts of seals. Thus, in the Sea

. of Gkhotsk in recent years the vessels of the UMRZF have been

catching about 6000 harbour seals, among other species of seals.

Because of the severe ice conditions during the period of the

wheloinu of the seals the whitecoats are caught in small num-

bers.

The volume of the procurement of fur pelts may be in-

creased by catches of harbour seals in the Tatar Strait. Ac-

cording to the data from the aerial survey, which was conducted

by the TINRO, about 9000 harbour seals were registered here.

In the past in the Tatar Strait, in the Bay of Tyk there were

caught about 800 head of this species in a year (Gagichko, 1931;

Gagichko ans Surzhin, 1935).

On the basis of the results of the aerial survey, of

the literature sources and the materials obtained during the

cruise of the sealing ship "Sanzar", we recommend the opening

up of a new region for commercial sealing: the Tatar Strait.

*Translator's note. I was unable to find the meaning of this abbreviation.

4o1,.,

Until further information is obtained on the harbour seal,

we propose to establish the quota for its catch for the first

year of the sealing operations at 1000 head.

As has been shown by the experience of the work on

the sealing ship "Sanzar", in years with favourable ice condi-

tions during the period of whelping, in the Tatar Strait it

may be possible to conduct the sealing operations for the

harbour seal both directly from the ship as well as from its

boats. Consequently, whitecoats and beaters may form a con-

siderable portion of the catch. It would obviously be advis able

to commence the sealing at the beginning of Pï.arch.

41J11•ITEPATY PA

1 15esrcux A. H. IIol.bin Bli-Z T IOr7i.!ICi C Kj'pnd!bCK11X oCTponOn -PI o aCinsularis

s^ n. RAt-I CCCP, T. 158, mm. 5, 19G^I.6e.vairt A. l1., Kocbrzurc 1'. il., 17axux. K. H. Hollble nlaTepua:!Lt 110 xapalcre-

I11,CT1!!CC OCTholl!IOrO TIO:IC!!fl.. CG. 4A'IQpCKIio DLRCI<Ol]!1TalO111nC». lIS., 1969.

^-.raau1tno C. !1. Ocetilnlïl lipoN!L1CCn naCTOUOrnX B manne Tbu:. <.PblGnoe xo-

3 tlcTno',Ua:!Lnc!ro I3ocTOxa», Ns l-2, 1931.^-•Fueuw:o C. 1I., Cypxcux C. H. Jlac•ronorue TaTapcsoro npo:111Qa il Amypc!coro

JnI9lana xaK np0\ILIIIS.iC!nloe cLlpbe. TPY;ibl BIiI•iPO, T. 3, 1935.

5^Zopo(1)ee(3 C. 13. A1aTepllaahl x llpO^ulCllosoii 611o1!0ril!t naCT0lfOr!tX B neceu-

lulïl :IC1[ollblü ucpno2 a TaTapexonl npo.nlluC. TpYnbl BIII1PO, T. 3. 1935.6 K,reüxeuüepa C. E., Slonohoa A. B,, K.lceesana r. A., Genbi:vauil B. ,ll.,

3TUx. B. A. Cnpauovuh!e nol<1saTC:m 110 s<;pal(Tel)IICTInCe ne!coTOpb!x ilacrollorl!x

1t KlITOOGpif311b1X. CG. «,^Iopcl:ueaL^er.ouuTa!ou.^ne». \•1., 1^1G5. •DlOpcjlo^or!i^!eCl:Oi{, ^apai:rep!!CT.!niC

^ KOCLL2uH% Ï . M. I'IC1:OTOpL1C DfaTCpna1!LI 110

n,1oRa :IasTal:a. 'fpy1L1 PIIIIPO, T. 65; 113B. "I•kII-IPO, T. 62, 1963.8 Kocnlzurt F. M., Coro loa A. C., Tu.romupoQ 0. A., LUtlcroo A. 17. Iloub!e

j(a11uL1E no nnTE:pbepY aaCT011ornS Cenepll0ïi 9aCTir Tiixoro oliealla. CG. «.^^Iopcr.ue

n! !clmnnTa!ontnc». Al., 1969.9 KOCb121tK r . M., Tuso.rupoa D. A. Jlapra 3an111la IIcTpa Bc 711K01'0. LICTBCpTOe

IiCec0103110C COnelünllnC no 113Y1ICI11!IO 1,101)Cli!IX 19CK01!IfTA1011(IIS. TC311C61^joxna-

Aou. M., 1969.10 !lay.stoa C. 17. Tlo.^!cllu CCCP. 1(oI-i3. i`2.-JL, 1933.

11 Hay.ston C. IL, CIIIIPH08 II. A. AlaTepitaabi no ClICTCMaTIIKU If reorpadmiecoomy pacupeite.neinuo Phocidae celleplion Tiixoro oocana. Tpyina BHPIPO, r. 3, 1935. 17 (knee C. 11..3Bepu CCCP ii npihneffluntx man, T. 3. MAL, 1935. 1.3 Iluxapeu I'. A. Tio.rienii ioru-aanannoii ‘IiICT11 Oxorcooro mopn. 113n. TI111P0,. T. 90, B:10,11.11110CTOK, 1941. 1,

POKIII{KIIII 17. ‘1, . Ocionna napilaimonna CTHTHCTI1KH T(.1151 6110aoron. 1961. -

Cieupnon H. A. 1.1ncTpyounn ,11;111 nomemix pa6or no 6iionoriiii cup:nun it siporpaiamid no inyileinuo npoiabic.nonoii cliaynia ii flp0Mblen0B. t. 1, ) 93l. J. CoK0.400 A. C. BCC0111.151 xapao -repitcruoa mianin opraiion ninuoeinin .macrono-nix. C6. «,\'Iopcolie ffleooriiiraiounie». M., 1969. 17 coso.um A. C. lieltOTOphIC 11T01- 11 H nepcneicrunm i(ciaoro-mopilianorinteconx IlccJIeJWnaliiIii THX0Olie.1111CKIIX Jiacronornx. C6. «Mopcone maeoonirraonnue».

re. C000ion A. C., Kochterm 1'. M., Tuxomupoe D. A. HeooTopide cneuenun o nece nny-rpeumix opranon siacronornx Bepunrona mops'. 143e. T141-1P0, T. 58. B.ria.au-BOCT01:, 1966. 19 C000..con A. C., .Kocbteun T. M., Illycroe A. II. Crpoeinie MerIiIIX 11 rpaxen y 6epiiiironomopconx ;lac -mot- 11x. T. 68. l'1311. T1.11-11)0, T. 62. M., 1968. 2 0 COK0.706 il. C., KOCb1el111 r. M., Ky31i11 A. E., Ilep.non A. C., Taxomupoci D. A., Illycron 17. Cripanotinbie 1101:a311TCAll no xapao-replicTitoe rmyrpeninix °prima Tilmonceancoux aac.roliorirx. C6. «I ■ lopeone ta.rieoonirraimitue». m„, 1969. 21 Tux0,1wpou .9. A. 0 TC1■111X nociipoilanoc.risa cenepo'riix000eznicoux T10:1C11Crl.

C6. «Alopcoini m.leoonirr a onmie». M., 1969. 22 Taxo.unpoa A„ Krietrearmb T. A. Meroulit oripeue.nemin 1303HaCra I101:0TO-

pux :1/ICT01101- 11X. B Ku.: eGiipeaesienue noapacrit nporaliteaonbtx ;lac-11moms: u pa- wionamblIOC IICH0.11,3011anite MOHC1:11>: M.11(.1:01111T11101101x>>. M., 1964. Z3 ganciaiii K. K. Onbrr nepecmoTpa cucTemia li manioc-molt rio.ieiieii noncemefi-cipa Pliouinae. Tp. T. 17, JI., 1955. 2 1 1. LInnentdi K. K. Orpiu Pinnipedia siacronorue. B ou.: «MneooluiTaionnie (Pay-

CCCP».1963. 25 ' ,/anckiili K. K. Mopqm.rioro-raocononintiecoan xapaorepucruoa Jinni

mush Tpy;(1,1 1 -11111P0, T. 21, Mypianco, 1967. • 2 0 /llyeron il. /1., f16.1oKoo A. B. Cpannivremaio-inopqm.imrunecoan xapaorepncrii-oa rpeimanRcooro n nomocaToro rio.rieneiÏ. TpyRta 1-I111IP 0 , 111111. 21, Alypmanco, 1967. 27 )76.fienwe A. 13. Heouropide ripe/inapirremaibie recreplia.nbt K mopilicaoriniecoort xapilKTCplICTIIKe rpeumailienoro T10J1C1111 15C.,10TO H Ppen.rianucooro raopefi. B c6.

liccneRon. pa6or 1962 T. (rpen.rialui,coliii T10,11C111, 11 xox.riati). Apxaureabco, 1963. 28 'M./Iowa A. B. 143mentinnocTb releKolurraionuix. M., 1966. • 29 516.aoKna A. B., Kiieneanizb r. A. Bn6puccia oirroo6pa3max u .nacronornx, ux pacnpe.c.ieii K', crpoenue 11 anal-feline. Mopipo.nornuecone oco6einiocrit nomibtx wieoomiraionnix. M., 1964. 30 Allen 1. A. History of North American Pinnipecls. U. S.- Geogr. S. Z. V, Ferrit. Publ., no .12, 1880. 31 Allen I. A. The Hair Seals (family Pilo' cidae) of the North Pacific ocean and Bering Sea. Bull. Amer. mus. nat.-- History, vol. 16, 1902. 32 Chapskiy K. K. Morphologie Systemaiique Differ'enciation Intra-specifique et Phylogenese du Sous-Genie Phoca sensu stricto Mammalia, t. 24, no 3, 19(30. 33 Doutl K. A review of the genus Phoca. Annals of the Carnegie Museum, vgl. 29, art 4, 1942. 3 14. Kenyon K. W. Food of harbor seals at Amchitka Island, Alacka. I. eMam-mol.», 46, no 1, 1965. 3 .D King 1. E. Seals of the World. Trustes of the British Museum (Natural History), London, 1964.

251

4.05

406

• 36 Mohr E. Ein neuer Westpazifischer Seelitind. Zoolog. Anz. Bd. 133. Nt 3/4. tiarnbur g, 1941.

• 37 -Mohr E. Dic Steine in Robbenrnagen. «Z. Situgetierlcunde», 28, no 3, 1963. . 3bike A4ohr E. Uber Phoea vitulina largha Pallas, 1811 .und Weissgeborene Seeh-

• niy.s. de . «Z. Siiiigetierkunde». Bd. 30. Hamburg. 1965. 3y Sell:4W 1/. B. Seals Sea Lions and Walruses. Stanford, 1958. LFO Scheiter V. B. Weights of organe- and glands in the Nortftprn Fur Seal. hammalia. vol. 24, 3, 1060.

•

REFERENCES

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2. Belkin A. N., Kosygin G. M. and Panin K. I. New materials

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3. Gae-iohko S. I. Autumn sealing in the Bay of Tyk. "Rybnoe

khozyaistvo Dal'nego Vostoka" ("The fisheries of the

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6. Kleinenberg S. E., Yablokov A. V., Klevezal' G. A., Bel'ko-

vich V. M. and tin V. A. Reference indices for the

characterization of certain pinnipeds and cetaceans.

Symp.: "Marine mammals". bloscow, 1965.

* Translator's note. "ostrovnoi tyulen" - literally "island seal", probably refers to the Kuril seal (Phoca stejnegeri)

407

7. Kosygin G. M. Some data on the morphological character- ,

istics of the foetus of the bearded seal. Trudy VNIRO,

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vol. 62, 1968.

8. Kosygin G. M., Sokolov A. S., Tikhomirov É. A. and Shustov A. P.

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Moscow, 1969.

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koscow, 1969.

10. Naumov S. P. Seals of the USSR. KOIZ (Vsesoyuznoe koope-

rativnoe izdatel'stvo - All-Union Cooperative Publishing

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11. Naumov S. P. and Smirnov N. A. Materials on the systema- 251

tics and geographical distribution of the Phocidae in

the northern part of the Pacific Ocean. Trudy VNIRO,

vol. 3, 1935.

12. Ognev S. I. The animals of the USSR and neighbouring

countries, vol. 3. Moscow - Leningrad, 1935.

13. Pikharev G. A. Seals of the south-western part of the

Sea of Okhotsk. Izvestiya TINRO, vol. 20, Vladivostok,

1941.

1•. Rokitskii P. F. Fundamentals of variational statistics

for biologists. Minsk, 1961.

• 408

15. Smirnov N. A. Instructions for field studies on the

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16. Sokolov A. S. The weight characteristics of the muscles

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17. Sokolov A. S. Some results and prospects of ecological

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19. Sokolov A. S., Kosygin G. M. and Shustov A. P. The struc-

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Pacific pinnipeds. Symp.: "Marine mammals". Moscow,

1969«

21. Tikhomirov El. A. Topics on the reproduction of the seals

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cow, 1969.

409

22. Tild-iomirov É. A. and Klevezal' G. A. Iiiethods for deter-

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1964.

23. Chapskii K. K. An attempt at a revision of the systema-

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24. Chapskii K. K. The order Pinnipedia, the pinnipeds.

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Leningrad, 1963-

25- Chapskii K. K. Morphological and taxonomical character-

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PIhlRG (Transactions of the M. Knipovich Polar Research

Institute of D':arine Fisheries and Oceanography), vol.

21, Murmansk, 1967.

26. Shustov A. P. and Yablokov A. V. The comparative morpho-

logical characteristics of-the harp and ribbon seals.

Trudy PIIVRC, issue 21, Murmansk, 1967..

27. Yablokov A. V. Some preliminary data on the morphologi-

cal characteristics of the harp seal of the White and

Greenland Seas. in symp.' of reseach studies 1962

(harp seal and hooded seal). Arkhangelsk, 1963-

28. Yablokov A. V. Variability in mammals. Moscow, 1966.

29. Yablokov A. V. and Klevezal' G. A. The vibrissae of

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and significance. Morphological characteristics of

aquat i c mammals. 141o s c ow , 1964.

4.10

32. Chapskiy K. K. The systematic morphology, intra-specific

differentiation and phylogeny of the subgenus Phoca

sensu strictu. Mammalia, vol. 24, no. 3, 1960.

36. Mohr E. A new seal species of the West Pacific.

37. Mohr E. Stones in the stomachs of seals.

38. Mohr E. Phoca vitulina largha Pallas, 1811, and seals

that are white at birth.

4, 11

UDC 599.745.2

V. A. Bychkov

THE SEASONAL VARIATION IN THE DENSITY OF THE FUR IN 253

THE HAIR BUNDLES OF THREE-YEAR-OLD BACHELOR FUR SEAIS

The three-year-old males of the fur seal, as compared

to other age groups, possess the most valuable fur and comprise

the main group that is exploited for commercial purposes.

Therefore, the study of the seasonal variation in the density

of the fur in the hair bundles of this particular age group

of fur seals is of considerable importance for determining

the optimal periods of the kill.

Studies which were carried out on the Pribilof Islands

showed that in all of the fur seal‘males (including also the

three-year-old males) the density of the fur in the bundles

remains almost unchanged during the course of the winter,

spring and summer (i.e. for the greater part of the annual

cycle of their life) and only in the autumn, when the new

hairs grow more rapidly, does the total number of the underfur

hairs in the bundle increase (Scheffer and Johnson, 1963).

Similar studies have not been conducted on the fur

seal population which reproduces on Robben Island. In connec-

tion with this, the present study is intended to elucidate the

characteristics of the variation in the density of the under-

fur hairs in the bundles on the body of the three-year-olds

in relation to the Periods of their kill e -on the basis of

which refinements may be made in the optimal periods for the

sealing operations on the fur seals on Robben Island.

412

The materials for the study of the seasonal variation

in the density of the fur in the bundles of the three-year-olds

were.collected on Robben Island from June to November during

1959 - 1961. There were collected 120 samples of pelt from

24 tagged bachelors of the local population of fur seals. The

samples, which were 2 - 3 sq. cm in size, were taken from the

mane, back, lumbar region, sides and belly (Figure 1), and

were fixed in a 10% solution of formalin. Histological prepa-

rations were prepared from each sample specimen in the form

of series of sections crosswise to the root part of the hair

bundle. The sections were prepared on a freezing microtome,

stained with haematoxylin and eosin, stuck onto a microscope

slide, embedded in gelatine-glycerin and covered with a cover

slip, according to the methods given in the textbook by G. I.

Roskin and L. B. Levinson (1957). The thickness of these

sections was 15 - 30 r . The evaluation of the density of the

fur was carried out under the microscope (with a magnification

of a X7 ocular and X8 objective) in the root portion of the

hair bundle at a depth of 1.0 - 1.5 mm (Figure 2). For each

sample there was determined the density of the fur in 25 homo-

geneous hair bundles. The hairs in each bundle were counted

three times and the average was taken as the "datum" for set-

ting up the variational series. In grouping these "data", the

class interval (i) equalled 5. The arithmetic mean (x), the

standarddeviaticn"cr50' thestandarderr°rofthemean' ) ' (mx

the confidence limits of the mean (5-c.± t 3m) and the confidence

limits of the series in question (51± teR) were calculated, to

one decimal place, by the methods recommended by N. A. Plokhin-

skii (1961).

4,13

rscv; `ls•^ ff':^^'',5;^°

.1^

^^^ ^:.\

^`^ `^.̂1

pile. 1. CxenlaTn1lnoe n3o6paxtellne Tynln xo^QcT»xa

co cnlnlm. (TollhaMll y1:a3al1U1 VtIaCTiQ[ 113flT![A npoû)

Figure 1. Schematic illustration of the body of a bachelor

from the.dorsal side. (The dots indicate the sec-

tions where samples were taken)

The results of the determinations of the density of

the fur in the bundles of the three-year-old bachelors are

presented in Tables 1 - 5, from which it can be seen that the

density of the fur varied in each month and on all parts of

the body of the bachelors. The amount of fur on the central

portions of the skin (mane, back sides) proved to be greater

than on the peripheral portions (belly, lumbar region). It

can also be noted that three-year-olds with the least amount

255

414

Pile. 2. 06milii isue, nonocanoro nyrwa na nonepemilom epe- • se iia ray6iiiie 1,0-1,5 .4t.e. boBepxtiocTil izom:11

– -- •

Figure 2. General appearance of a hair bundle in cross

• section at a depth of 1.0 - 1.5 mm from the surface of the skin. •

of fur in the bundles were most frequently found in August -

September. Furthermore, it was established that neither in

October nor in November were there animals among the examined

three-year-olds, in which the numerical values of the modal

classes of the density of the fur in the bundles would exceed

the corresponding indices in June - July.

Even in a comparison of the relative indices of the

density of the fur in the bundles on different parts oe the

body, only in one case was it noted that, in the fur on the

29

95

27

26

28

27

30

23

18

29

20

18

91

26

94

27

25

22

24

16

28

24

15

November

T a 6 el 1111a

Koantiecruo nyxoubtx no:toc n nytuce Ha rime Tpex..nemux xo.nonnicoa n opcnoro 1i:canna

Date of kill .riaTa AO6b14II 511Cp31

of animal

, Modal moe.mei

K111CC

class

Maxi-- Mean Mruzcit- CPung

mym

MUI

Confidence limits JLoiaeiiaJth'I hie

rpaumm

cr7

Mini upi MUM

June 24 moun 1960 r.

24 ». »

29 » »

29 » »

July 2 1110:1fl 1959 r.

10 » »

july 24 1110A51 1960 r.

24 » »

August 10 aurycra 1959 r.

• 11

• 11

• 11

September4ce 6 MrS urn_pst r.

. »

18 » >> •

ShtembeD2 ceirra6pn 1959 r.

Otober

>>

» »

8 non6pn 1959 r.

10 » »

10 » >>

10 » >>

33-37

33-37

38-42

33-37

33-37

38-42

38-42

33-37

28-32

28-32

28-32

28--39

23-27

23-27

33-37

28-32

38-42

33-37

28-32

33-37

33-37

38-42

33--37

28-32

44 36,8 -1- 0,65

46 36,6+1,22

38,4±1,05

49 35,4.10,84

44 36,4+0,81

44 362±0,98

47 37,010,94

45 34,6+0,93

38 28,4:11,02

43 28,6-10,72

43 32,4+1,27

38 30,2 ± 0,96

33 25,2±0,77

37 27,0+0,70

42 33,2-10,79

41 32,4±0,94

47 37,9+0,99

39 32,4:10,80

49 33,0-1.1,09

47 36,2±-0,90

44 35,4-.11,29

48 36,2-11,10

43 36,0±0,89

34 28,6+0,80

31,1-43,5

. 23,2-50,0

27,4-49,4

26,5-44,3

27,9-44,9

26,1-46,3

27,1--46,9

24,3--44,a

17,7-39,1

21,0-36,2

19,0-45,3

20,1-40,3.

15,1-33,3.

19,6-34,4

24,9-41,5.

22,5-42,3.

27,4-48,4

24,0-40,3

21,6-44,4

26,8-45,3

21,8-49,0

24,7-47,T

26,6-45,4

20,2-37,0

21 onrn61)11 1959 r.

23

23

26

256

Table 1.

The numbers of fur hairs in a bundle on the mane of three:7..year7ol.d._ ba.chelors__of... the.. seal.

Da

o

4ne

Jialy

Jiu1ly

ALg.

•

1

Sept.

Oct.

Nov.

1

T a .1 n, a 2

K0,111 1 1CCTLIO nyxonbix »mac U arum Ita mine TpexacTnnx. X0.110CTIIKOIS mopcicoro hOTJIKfl

, ,.;9nIictence ce of kill tini-odalbraxi mean biteptil eamthte Munn- Mo.n.i mud rt Ma melt- ,-Peallsig rpa Itil am

.1-1.a ra A061•1`111 -I Lie P`i mym Kaacc mym :.7÷ III:,:- x -F t1 a— C> animal mura clasi._., mum - x

24 Inomi 1960 r. 27 33-37 45 35,8 ± 0,88 26,6-45,0

24 >> » 26 38-42 47 37,4±1,10 25,8-49,0

29 >> » 31 38-42 47 40,0+0,85 31,1-48,9

29 » » 29 33-37 42 35,8 ± 0,71 25,4-43,2

2 inOnst 1959 r. 32 38-42 47 40,8±0,84 32,0-49,6

10 » » 29 33-37 45 36,4±-1,04 25,4-46,4

24 moan 1960 r. 32 38--42 46 36,8 -±- 0,93 27,1-46,5

24 >> » 24 28-32 46 33,8±0,91 24,4-43,2

10 anrycTa 1959 r. 2-1 33-37 - 43 33,8±1,14 21,9-45,7

11 >> » 27 33-37 49 35,2:9,66 28,3-42,1

11 >> » 20 28-32 45 31,2±1,17 . 18,9-43,5

11 » » 19 28-32 35 29,0-9,89 19,7-38,3

4 ceni Am 1958 r. 20 28-32 38 29,2-2:0,88 20,0-38,4

8 » » 23 28-32 38 30,2±0,87 21,1-39,3

18 >> » 25 . 28-32 41 31,6+0,84 22,8-40,4

22 ceiln 6p g 1959 r. 20 28-32 39 31.0±-1,06 19,9-742,1

21 oKrepn 1959 r. 24 33-37 45 35,6±1,14 24,6-46,4

23 » - >> 26 33-37 46 36,2±0,86 27,2-45,2

23 » » 20 33-37 42 34,4±0,95 24,4-44,4

26 » » 29 38-42 46 . 38,6±0,75 31,0-46,2

8 non6pn 1959 r. 14 33-37 49 3Q,2±1,63 19,0-53,4

10 >> » 20 33-37 43 32,6+1,20 20,0-43,2

10 » » 19 38-42 45 36,61:1,16 24,4-48,8

10 » » 25 28-32 36 31,2+0,66 24,3-38,1

limits

17 3aK. 2354

Table 2.

95T.

The numbers of fur hairs in a bun.dre on

_the_back of ..three-.year-old bachelors, of

the fur seal.

•

;e of kill Mini ModàlLicx..:- Mean ,11-eZWOW(5-

iiaTa Rowni 3„pg M\1,11,,Itl- M(Ui.a.,:litcll,lii M;I,K,Ce lfl- _q1) . 111,151 rpallIllthl X 1". Mx . Jt' ÷ t a-

- 1 .1.

r: animal mura Class In'15.1M

24 11101151 1960 r. 27 33-37 42 35,4±0,84 26,6-44,2

24 » » 25 33-37 46 34,2± 1,05 23,3-45,1

29 » » 25 38-42 46 37,0±1,28 23,6-50,4

29 » » 25 33-37 40 32,4±0,90 23,0-41,8

2 mo.9 a .1959 r. 31 33-37 43 36,6±068 29,5-43,7

10 » » 29 33-37 44 35,0+0,75 27,2-42,8

24 11102151 1960 r. 23 33-37 40 33,6±0,94 23,7-43,5

24 >> » 26 33-37 45 35,0 -±- 0,94 25,2-44,8

10 aBrycra 1959 r. 19 23-27 35 25,2+0,96 15,1-35,3

11 » » 25. 33-37 42 33,0±0,5 26,1-L-41,9

11 » » 17 28-32 38 30,0 -I= 1,09 18,5-41,5

11 » »' 16 23-27 35 26,2±1,06 15,1-37,3 .

4 ccurepn 1958 r. 26 28--32 41 30,8±0,73 23,0-38,6 -

8 » » 24 28-32 38 31,4±0,72 25.9-38,9

18 » » 24 33-37 42 32,8±0,99 22,4-43,8

22 ccurn6p1 1959 r 19 23-27 40 27,2±1,06 16,0-38,4

21 own 6p a 1959 r. 24 28-32 39 30,4±0,85 21,6-39,2 '

23 » » 24 38-42 43 36,6±1,05 25,6-47,7 .

23. » » 26 33-37 47 37,2±1,05 26,2-48,2

26 » , » 22 33-37 43 33,4 -± 1,1 21,6-45,2. .

8 1Io116pu 1959 r. 12 38-42 47 40,4±1,70 22,5-58,3

10 » » 11 38-42 44 35,4±1,51 19,5-41,3

10 » » 28 33-37 42 36,0+0,69 28,8-43,2

10 » » 23 28-32 35 29,4±0,65 22,6-36,2

limits Da

o

June

July

July

August

Sept.

Sept. Oct.

NOV •

Ta 6 muua 3

KOJ111 ,1CCTII0 uyxonox Bo.noc a urixe im 11051C11111e

TOCXJICT1111X XOTIOCTOKOB MO )cuoro KOTIUt

• 258

Table 3. The numbers of fur hairs in a bundle on

the lumbar region of three-year-old

bachelors of the fur seal.

^^

llate of kill›TA AOGL.^ut sne^a

oi^ animalI

19G0 r.June 24 1110119

»

29 » »

24 » »

Juiy 2 1110nA 19â9 r.

10 » »

1Jua.y24 }nonsl 1900 r.

24 » »

ug . 10 anrycTa 19,-)5) r.

il » »

11 » »

11 » »

1Co7114CCT}t0 nyx06L1X rnaac n}ivuce }la GOKY

TPCXTiCTI}I;X X0:lOCï}:KOC A10[iC1C}1X KOTHKOü

Sept • 4 ceuT}IGps} 1958 r

.1Sept. 22 celmiGpsl 195J r.

21 oxTS}ûpn 1959 r.

23 » »

23 » »

26 » »

Nov • 8 noslGps} 1959 r.

10 » »

10 » »

10 » »

17*

22

1,.odaliGio;ta.it,utatl

kT3cc

class

Pâxi,üat:cu-

mf \(

muni

33-37

33-37

38-42

33-37

38-42

38-42

38-42

33-37

23-27

33-37

28-32

28-32

28-32

28-32

23-27

28-32

33-37

33-37

33-37

38-42

43-47

33-37

38-42

28-32

4G

!^-.e anCIte.lnst st

.r±ni.C

36,0.-L 1,0G

34,0=0,98

37,G-!-1,10

34,2±0,92

37,2 ±0,75

38,8:L0,^02

3G,8z:E 1,09

32,8:h0,89

2G,1± 1,01

35,4±0,G9

31,0-±-1,02

29,1-±- 0,8G

30,4±0,93

30,8 ±0,84

27,0±0,97

31,G± 1,09

33,2±0,9G

37,2±0,70

35,0-1-0,94

38,2±0,93

42,0.±:1,06

30,G:L 1,39

37,2 ±0,7G

31,0 ± 0,75

TaG.-1 nüa 4

Cunfi-d-91rce l imit sAonepnTeat,tiwe

rpcwnma .

.l _fl ..•

24,9-47,1

23,7---44,3

2G,1-49,1

24,6-43,8

29,3-45,1

30,2-47,4.

25,3-48,3

23,4-42,Z'

15,5-37,3

28,2-42,G

20,3-41,7

20,1-38,1

20,7-40,1

22,0-39,G

26,8-37,2

20,1-43,1

23,2-43,2

29,7-44,7

25,1-44,9

28,4-48,0

30,8-53,2

15,9-45,5

29,2-45,2

23,1-38,9

259

Table 4.The numbers of fur hairs'in a bundle on

threé'year-^old bachelors of

the fur seal.

23

20

419 •

24

25

24

22

18

23

21

25

13

19

14

18

18

20

21

18

11

21

24

17

11

G

18

15

;

Sept.

Sept.

T a .1 n 1.t a 5

KOJIIPIeCITO nyxonmx BOJIOC i nruce na 61noxe . TpexaeTnnx. XOJIOCTIII(011 mopcicoro »Tun

Date of kill AaTa ao6lagn »elm

of animal,

• Modal Di .ace mym class mun

Jean ni7

-C-ontidence limits rffl .u. Munn-

mym

MUM

June 24 'Ilona 1960 r.

24 » »

29 » »

29 » »

July 2 mail 1959 r.

10 »

July 24 nio,nst 1960 r.

24 » »

August 11 » »

11 » »

11 » »

4 con -15161m 1958 r.

8

18

22 cenuGpa 1959 r

ÙCtObel^ 21 cuanGpn 1959 r.

23 »

23 >>

26 .»

8 nodipn 159 r.

10 »

10 »

10 .»•

33-37

33-37

33-37

28-32

33-37

28-39

28-32

28-32

18-22

23-27

28-32

23-27

23-27

23-27

28-32

23-27

18-22

33—r

33-37

33-37

33-37

18-22

33-37

18-22

32,3+0,89

31,8±0,89

34,6:1=0,93

30,2+0,99

34,2-±0,84

31,81=0,99

29,6±0,84

32,0 ± 0,80

21,2±0,M

25,8±0,67

27,4 1=0,90

26,6 0,89

24,0+0,69

24,8 ±0,77

27,8±0,81

25,0+0,80

22,0±0,97-

31,6:1:0,89

34,8+0,87

31,4±.-1,28

34,8+1,78

21,8±1,41

34,2±-0,84

20,2+0,72

22,9-41,7

22,4-41,2

24,9-44,3

19,8-40,6

25,4-43,0

21,4-42,2

20,8-38,4

23,6-10,4

12,4-30,0

18,8-32,8

18,0-36;8

17,3-35,9

my-31,3

17,1-32,5

19,0-36,6

16,6-33,4

11,8-32,2

22,3-40,9

25,7-43,9

17,9-44,9

15,9-53,7

6,9-36,7

25,4-43,0

22,6-27,8

39

40

42

39

39

40

38

39

33

3 9

33

33

31

36

37

34

30

38

43

44

40

30

39

26

10 anrycia 1959 r.

NOV

260

Table 5. The numbers of fur hairs in a bundle on

the belly . of three-year-old bachelors of

the fur seal.

Mane

Back

Lumbar region

Si de

Belly

420

4 0 -

f- - -...

30 - 0. c \ , 2 5 - •

40- Cp 1- - -i■ -71- - ... 3 5

- - --. ■̀. '

'., 30- - C

25 - .

40 -

35 - f- - --f. , , 4- - --i 30 -

• ' 1- - -}- tC 0 25 - C

it 0 -

k 35 - I- - -I- r̀) 30- ,0 25 -

0 , 'ç 35 - 4 f- -- i,, , ck 3o—

-f - 1 . z `() 25-

i , . 1 , c ' t) e> V/ V// V/// IX , X X/

>> k U 44 e C e U bf

1. 2 Months 1- parts Pan. 3. Cpe,queB3nemennaa (x:=1:_t atn4 2- amount

ceaonnoro pa3noo6pa3n1 rycToni nyxa B nyincax na 'rule XOSIOCTBKOB 3 OCT

Figure 3. Weighted mean (5I± t3m) of the seasonal variat4n

in density of the fur in the bundles on the body

of 3-year old bachelors.

side of an animal, caught on the 8th of November 1959, the

value of the mode was found to be higher than in animals that

were killed in June - July. The analysis of the individual 261

mean values and confidence limits of the variation in the den:.

sity of the fur in the bundles of the three-year-olds also

indicates the small probability of finding in uctober - November

animals on the body of which there would be found bundles

containing more fur hairs than in June - July.

4.21

The above described characteristics of the change in

the density of the fur in the bundles of three-year-olds is

manifested even more obviously in ,a comparison of the weighted

mean values of this feature (Figure 3). Fromt}ae figure pre-

sented it can be seen that over the entire body-of the bache-

lors the numbers of hairs in August - Septembur are lower

than in the other periods (this•differencé is found to be

statistically significant), while in the comparison of these

same indices in the animals that were killed in June - July

with the animals killed in October - November'only a small 262

difference was detected between these, and moreover this was

not statistically significant.

On the basis of the data obtained, it is not possible

to agr.ee with that interpretation of the seasonal variation,

of the density of the fur in the bundles in bachelors, which

was given by Scheffer and Johnson (1963). The scheme that

was indicated by them is, apparently, atypical. In our opi-

nion, it is more reasonable to propose that the seasonal

changes in the density of the fur in the bundles proceed ac-

cording to the following scheme: as a result of the massive

falling out of the fur hairs of the previous year's generation

their numbers in the bundles on the body of the bachelors de-

creases markedly in August - September. Then, during October

- November, as the fur hairs of the new generation grow up,

the density of the fur in the hair bundles again increases,

but on the average it does not exceed that amount which is

found on the bachelors at the beginning of their appearance

on the hauling grounds in June - July. Taking into account

422

the fact that at the moment of the departure of the bachelors

from the hauling grounds the majority of them have not com-

pletely moulted, the possibility cannot be excluded that the

growth of some portion of the new fur hairs will continue

during the marine period of their life, as a consequence of

which the total number of fur hairs in a bundle will build up

during the winter.

Thus, from the data presented above, it may be conclu-

ded that in August - September the quantity of fur hairs in

the bundles on the body of the three-year-olds is, as a rule,

lower than during other periods of the annual cycle of their

life. However, the violent falling out of the fur from the

bundles in August - September, in its turn, leads to the si-

tuation that the total density of the fur hairs on the skin

of the three-year-olds at this time also decreases. Conse-

quently, under all conditions August - September cannot be.

recommended for the sealing operations on fur seals on Robben

Island.

j1I-ITEPATYPA

. 1 POCKI1H I'. II. .aesancon II. E. Min:pocizontmeexasi Tex a. M., 1957. • 2 1-1.7oxunci;aû il. A. Bname-rplist. 143,-t-uo CO CCCP, Honocemtl)m, 1901. 3 &holier V. B. and Johnson A. AI. MGR in the Northern fin seal. U. S. Dept. Jnter., Fish. Serv., 450, Washington D. C. 1963.

REFEREN CES

1. Roskin G. I. and Levinson L. B. Macroscopic technique.

Moscow. 1957.

2. Plokhinskii N. A. Biometrics. Publ. Siberian Branch

of the USSR Academy of Sciences, Novosibirsk, 1961.

4,23

UDC 599.711•5.2

G. A. Nesterov

THE EFFECT OF THE MUSCLE RELAXANT DITILINE ON THE ORGANTSIvI _ 263.

OF FUR SEALS

In 1963 on the Komandorskiye Islands there were set

up the first tests on the use of the curare-like preparations,

ditiline and diplacâ:n, in the slaughtering of fur seals. Di-

tiline was found to be the most suitable for these purposes.

The effect of the preparation commences after 30-II•0 seconds.

At the moment that the fur seals became completely immobile

they were exsanguinated with a knife.

This method of slaughtering, in comparison with that

existing at the presemt time (killing with a club), has seve-

ral advantages. It is more humane and permits selective

slaughtering to be carried out. The meat and meat products

are suitable for food. It is more economical, since it elimi-

nates bruisings on the pelts. In a slaughter of 10 000 fur

seals the profit may comprise II-5-50 thousand rubels. All of

this permits this method of slaughtering to be recommended for

introduction.

In the selection of methods for the pre-slaughter

stunning of the animals, an important criterion is the possi-

bility of retaining the cardiac activity in the animals, which

is necessary for a better exsanguination. The latter is also

important in the sealing operations on the fur seals, and there-

fore we set ourselves the aim of determining the effect of

ditiline on the organism of the fur seals.

1-12k

From the literature sources it is known that during

the narcotization of animals with non-lethal doses of ditiline

there are not noted any drastically expressed changes in the

blood circulation and respiration (Zhulenko, 19631 Zhulenko

and Korneva, 1965).

Similar studies were carried out by us with fur seals.

As a result of this work there were determined the mean effec-

tive, optimal and lethal doses of ditiline. The effect of the

preparation on the cardiovascular and respiratory systems and

on the body temperature was studied, and at the same time the

consequences of the effect of ditiline on these animals were

revealed.

METHODS OF STUDY

The investigations on the immobilization of the fur

seals with the muscle relaxant were carried out on animals .

which were kept in cages. The animals were first weighed,

after which 5% ttnd 10% solutions of ditiline in various doses

were administered intramuscularly. In the experimental ani-

mals electrocardiograms were taken, the respiration was mea-

sured and the body temperature was measured every five minutes. 264

The electrocardiogram was recorded from 3 standard points of

contact at a current voltage of 1 mv. The changes in the bio-

electrical potentials of the heart under the influence of di-

tiline were registered from the fore and hind flippers on pho-

tographic film with the aid of a portable EKP-60 electrocardio-

graph (Figure 1). The respiration was measured only in animals

which were not held down. The body temperature was measured

rectally.

425

Puc. 1. Dpucoe.nunenne wiewrpo,],on K

Te.ly KoTru.:a npu perucTpanuu cTau- AapTubix celnciummii— I, II, III

Figure 1. The attachment of the electrodes to the body of a fur seal for recording from the standard con-tact points - I, II, III. •

DESCRIPTION AND RESULTS OF EXPERIMENTS

For the purpose of determining the mean effectiveness,

the optimal and lethal doses for the fur seals,there were

tested the following amounts of the preparation: 0.25; 0.5;

0.75; 1.0; 1.25; 1.5; 1.75 and 2.0 mg per 1 kg weight of

the animal. Each dose was tested on 6 male fur seals, aged

2 - 4 years.

The results of the tests indicate that 0.25 mg/kg shows

no effect. A weakly expressed narcotization sets in at doses

of 0.5 and 0.75 mg/kg, within which range was found the mean

effective dose. The optimal immobilizing dose, which brought

about a clearly expressed immobilization of the animal in all

5

ilereTBIle 2111TWOUTA

1.r10 2,00_ Effect of ditiline

He WICITIIWIO Did not ensue Flammwm Ensued' aemumAmmuLethal out corne

0

5

1

9

6

426

of the tests without having a lethal outcome, is'found to be

between 1.0 and 1.25 mg/kg. A dose of 1.5 mg/kg causes a

stoppage of breathing (apnea) in individual animals. Death

always ensued after the administration of 2.0 mg/kg. This is

the absolute lethal dose (Table 1).

Table 1. 265

The effect of the muscle relaxant ditiline on the

organism of fur seals. - -

Ta6.nia 1

Ram-lime mbunequcro peaaucapTa perumpia na opranuam mopcium KOTHKOB

•

The mg/kg

oe 0.50 035

6 4 1

.1triK2

1,00 1,21

0 0

6 6

0 0

o • • 200

• tj

•

s O ()Up - • • e

,r-i too

CI) o

H so

Dos ,a

e me/kg 030`,

£1 75 1,0 1,25 1,5- 2,0

Pitc. 2. linteuenue nyabca y mopci:m Horuzon OT Be:111 ,11M 1 ,1 ;1, 031,1 :11IT 11,11111 a

,,,,, . . • • 0 20 4* /3 .70 0 12 30 0 20 10 30 0 20 43 30 0 40 10 30

apv4sr, A41.0r

Time, min.

Figure 2. Change in the pulse of fur seals depending on

the size of the dose of ditiline.

427

With the aid of the electrocardiograms it was shown

that in optimal doses the °reparation slows down the cardiac

activity, which later returns to the original level (Figure

2). The duration of the intervals in the cardiac cycle is

directly dependent on the dose (Figure 3). The initial part

of the ventricular complex (GPS) is subject to the least

changes. On emergence of the experimental animal from the

state of adynamia, the duration of the intervals in the cardiac

cycle returns to the initial state.

The pulmonary respiration in the narcotized animals

is changed to a greater degree, the greater the amount of the

muscle relaxant reaching the blood. With the administration

of mean effective doses there is an insignificant depression

of respiration, while optimal doses impair gaseous exchange

and may even evoke short-term arrest of respiration (figure 4).

The mean effective dose caused an increase in the body

temperature of the fur seals of 0.6° C; the optimal dose evoked

a temperature increase of 1.0 - 1.5 ° C, while the absolute le-

thal dose evoked a temperature increase of 3 - 4°0 (Figure

With the restoration of the capacity for movement, the body

temperature gradually attains the initial level.

Our investigations showed that the muscle relaxant

affects the fur seals for 15 - 30 minutes, after which the

'animals recover their locomotory function. With the aim of

elucidating the influence of the preparation on the fur seals

on the days following the injection, observations were carried

out on the animals for 1 - 4 days in a cage, after which they

were released onto the rookery. Before release, the fur seals

5 ).

* State of animal Normal

Incomplete immobilization .1••••• ••■•••

Dose mg/kg ,a0 3 a mr/icr

0,25 0,75 1,0 1.5 2,0

428

Dose m5/kg 1. 11* Ilo çja,mr

1.4. 075 fO 1,25 15 j 2.0 j

,14 eft/Mite 1,1 hen rr711110/1

0 01- * Odceioe Hue .xeugammoro ; 1 i

....

–.— tlenomine 0.lu.:ev&vaie i , (1) – C(,` T. ----- if OpMailbmae ■

c•-:---,-- - IlOntioe ocre3gikeyrtdomue i'à i i

.1)

• it i

e02/ 1 . J i f\iN . .1.A\ .. .,.....4 .• 1

E-IÇ:Q eif . • i I I I I i I

P pa.eun ernmem p Fegassi 11/2/5/ P Pcese rerrtus£ piee:greerneee p e7Gias1 fasitgRe 11 fri M epeoebt intervals

-Pm. a. FI31eiteitire npoRomKtrre.ibilocTit unTepnamon 11 cepitetinom unK.ie n 31313IICMIOCTII OT ne.-nnunim no3b1 munanna

11>

Figure .3. The change in the duration of the intervals in the cardiac cycle depending on the quantity of the dose of ditiline.

-1 11•■ •••••■•• # Complete immobilization

V4 •

-P • 70

1`

60 ‘.0 •

0 E 90

0 • te. PO r0 PA •

. 12

"

05 wa 20255101520255 ffl 1520e 5 i0 a 20 e 5 weee epemg 17170X0,YraeH5IR Orkena MUH '

Duration of test, min Pnc. 4. II3menenne •n,bixannn y mopcimx KOT111:00 110/1,

B:111511111CM

Figure I. The change in the respiration of fur seals under the influence of ditiline.

41Ï

Iiose mg/kg joaa. ht r nr.

0.75 1,0 1.25

429

17 5 10 1520 5 10 15 20 51015 20250 5 10 15 20 2539 gpemn -np0x0.7.yeemag onbuncr,

Duration of test, min. Pile. 5. alliSInue airrnaima Ha Temlieparyp ■,. -re-

mopciaix KOTIIKOB

Figure 5. The effect of ditiline on the body temperature of fur seals.

were tagged by clipping the guard hairs on the head and front

flippers. The clipped areas varied in their form and position 267

on the body of the fur seals. During the course of the entire

daylight period constant observations were conducted on the

fur seals. 14 out of 23 experimental animals (60.8%) were

again found on the ilauling out grounds. They came out onto

the shore and were caught in some of the sealing drives. In

their behaviour these fur seals did not differ in any way from

the remaining animals. Those tagged fur seals that were not

found were apparently in the sea or in such places in the

rookery where it was extremely difficult to catch sight of

them, since the percentage of encounters with control animals,

to which no ditiline was administered, was about the same.

From this we conclude that, when administered in optimal doses,

the muscle relaxant ditiline does not produce harmful conse-

quences to the organism of fur seals.

o 0 0

Wti 39

-P D W% P.37

P-1°-

(1.) -p 3g

1:1 0 - IA

14.30

CONCLUSIONS

The mean effective dose of ditiline for fur seals aged

2 - 4 years lies between 0.5 and 0.75 mg/kg; the optimal dose

- between 1.0 and 1.25 mg/kg, while the absolute lethal dose

is 2.0 mg/kg.

The mean effective and optimal doses, at the start of

the action of the preparation, slow down the cardiac activity

in the fur seals, which later returns to the initial level.

The more ditiline is administered, the more marked is

the dePression of the pulmonary respiration in the narcotized

animals.

In the fur seals the ditiline gave rise to a short-term

increase in body temperature.

No harmful consequences of the effect of ditiline on

the fur seals were observed after the administration of mean

effective and optimal doses of this muscle relaxant to these

animals.

1a-31,d ,L

.rII-rrr:I1,aTS•IIA1%Kymexr:o B. II..IIpe.ay6oiinoe o6cs;LBn;xnuauuc :r.nBO•cuMtX cj)apatar.o;tortt1tc-

ct;uMnt BeutecTaaMut. 1MaTepea.n111 Aox.TIanoB III nay"tnoïr xotujtepenqnn BCTCpnltap-rto•cannT,1.pnoro q)atq•.1r,Te•ra 1\1 11IBLNIII, 196.3.2)I(y,•:exr o B. H., I^oprrecaa B. H. O6c:32Btur:nBarnte Bo.,ncoB n;,tewe;[cü 11,11T11-

nom. ^<I3ercptntap1151:^, \ë 11,'19û5.3 Hecae'poc I'. A. OntAT rtpuNtettenct» ;Urrit.iima na so•rm:oBO.m npoNnAc.-te. I43B.

TI•lI-IPO, r. 58, 1966.

REFER.ENGES

1. Zhulenko V. N. Pre-slaughtering immobilization of animals

with pharmacological substances. Materialy.dokladov

III nauchnoi konferentsii veterinarno-sanitarno fakul-

teta nSTIMMP (Reports of Ilird scientific conference

of the veterinary-sanitary faculty of the IGSoscow Techno-

logical Institute of the ^`Ieat and Dairy Industry), 1063-

2. Zhulenko V. N. and Korneeva V. I. Narcotization of wolves

and bears with ditiline. "Veterinariya" ("Veterinary

Xledi.cine" ), No. 11, 1965-

3. Nesterov G. A. A test of the application of ditiline in

fur sealing operations. Izvestiya TINRO (Proceedings

of the Paci;Cic Ocean Research Institute of Fisheries

and Oceanography), vol. 58, 1966.

4.32

UDC 599.745

V. N. Mineev

QUESTIONS OF CONSERVATION AND REGULATION OF THE STOCKS 269

OF MARINE MAMMALS

In recent times questions concerning the rational ex-

ploitation of the stocks of marine mammals have acquired a

particularly important significance, since these stocks in

different basins have either been undermined or are found in

a stressed state.

The Ministry of Fisheries of the USSR is Paying con-

siderable attention to the rational utilization of the stocks

of marine animals and considerable efforts are being undertaken,

especially in recent years, in this direction.

THE FAR-EASTERN BASIN

The hunting of the sea-otter has been banned everywhere,

which has had a verY favourable influence on the re-establish-

ment of its stocks. The numbers of this species at the present

time in the waters of the Soviet Union comprise 5.5 - 6 thou-

sand head. Ma recent years measures have been undertaken to

strengthen the conservation of the sea-otter; throughout the

course of the whole summer period vessels of the Sakhalin Fi-

sheries Service and the Kamchatka Fisheries Service patrol the

region off the Kuril Islands and Mednyy Island; in 1967 there

were organized two permanent posts for the conservation of sea-

otters on the Kuril Islands, on the islands of Paramushir and

Urup, while in 1968 a third post was set up on the island of

Iturup.

• • 433

o

o

For the further increase in the population of this

very valuable marine animal the ban on its hunting should be

maintained. It is necessary to intensify the work on the

acclimatization of the sea-otter on Bering Island and in other

regions, which used to be its habitats.

Recently commercial organizations are proposing the

introduction of a prophylactic culling of the sea-otters. In

the opinion of Far Eastern scientists, a commercial culling

operation of a so-called "selective" cull afloat is Premature.

At the present time it is sensible to restrict ourselves to

collecting the pelts of sea-otters which have died as a result

of the natural mortality and to carry out further tests on

culling.

Sealing operations on the fur seals are being conducted

on the Komandorskiye Islands and Robben island.

The measures for the conservation and regulation of

the sealing operations on the fur seals on the on-shore rooke-

ries favour the growth in their population numbers.

After the signing in 1957 of the Provisional Convention

on the conservation of fur seals in the northern part of the

Pacific Ocean, the numbers of these in the Far East increased 270

by almost 2.5 times and attained a level of 400 - 420 thousand

head (Komandorskiye Islands - 200; Robben Island - 180 - 200

thousand head).

The dynamics of the catches from 1961 appear as follows:

in 1961 there were killed 12 thousand head, in 1962 - 13.5 thou-

sand, in 1963 - 1•.7 thousand, in 1964 - 18.9 thousand, in

1965 - about 20 thousand, in 1966 - 18.9 thousand, in 1967 -

4•3y., ,.

17.7 thousand, in 1968 - 15.0 thousand and in 1969 - 15.6

thousand head. It should be noted that from 1966 the fur seal

kill began to be reduced in connection with the decrease in

the numbers of the commercially exploited bachelors. The

causes of this decrease still remain unclear, in spite of the

deliberations of a special conference and the organization in

1969 of a trip of a commission to the Kuril Islands ..

It may be supposed that one of the causes of the de-

crease in the numbers of bachelors in the rookeries was the

overestimate of the catch limits, which were established in

previous years, and also the prolongation of the sealing ope-'

rations after August 1st and the frequent driving-off of these

animals from the bachelor hauling grounds.

In recent years several measures have been undertaken

for the further strengthening of the conservation of the fur

sealsi a 30-mile prohibited zone has been established around

the Kuril Islands and Robben Island. It has been resolved to-

kill bachelor males that are no younger than 3-4• years and

of a size not less than 108 and no more than 150 cm. Drives

of fur seals are authorized only from bachelor hauling grounds

with not fewer than 100 head of the commercially exploitable

animals present on the hauling ground.

The population numpers of Steller's sea-lion have

attained a level of 11•0 - 60 thousand head, but these are vir-

tually not being commercially exploited. Thus, "lial'ryba"n°

has cought the following numbers of Steller's sea-lion in dif-

ferent years : in 1964. - 329, in 1965, - 14-6, in 1966 - 40, in

196? - Va., in 1968 -1I•20 head, while about 3000 head could be

caught each year.

i` ":llal'ryba" - Far Fast Trust of Fishing Industry. Enterprises.

4,35

in connection with the reduction in the stocks of the

true seals in the Far East, a limitation of their catches

was initiated from 1967:

In 1967 limits were established for the ringed seal

(11-5-0 thousand head) in the Sea of Okhotsk; the ribbon seal

(5.0 thousand head) in the Bering Sea;

in 1968 for the ringed seal (32.0 thousand head), the

ribbon seal (5.0 thousand head), the harbour seal (11.7 thou-

sand head), the bearded seal - ban in the Sea of Okhotsk; in

the Bering Sea: for the ringed seal (1.0 thousand head), the

ribbon seal (5.0 thousand head), the harbour seal (3.0 thou-

sand head), the bearded seal (1.0 thousand head);

in 1969 for the ringed seal (32.0 thousand head), the

ribbon seql (5.0 thousand head), the harbour seal (11.7 thou-

sand head), the bearded seal - ban in the Sea of Okhotsk; in

the Bering Sea: for the ringed seal (26.0 thousand head), the

ribbon seal (5.0 thousand head), the harbour seal (5.2 thousand

head) and the bearded seal (2.0 thousand head).

From 1970 the sealing has been subdivided into coastal

(sovkhoz*), state shipborne and for the needs of the local

native vovulation.

We consider it advisable.to. switch the hunting opera-

tions for the seals from the spring to the autumn, since during

the autumn sealing all of the seal pelts may be utilized as

high-quality fur raw material. It is necessary that scientific 271

organizations, within a short period of time, give their propo-

sals on the localities and periods for the sealing operations

and on the rational exploitation of the pinniped stocks.

* sovkhoz - state farm.

L36

In connection with the small numbers of the "antur"

of which there are 4.0 thousand head on the Komandorskiye

Islands and about 2.0 thousand head on the Kuril Islands,

there has been a ban on its hunting from 1970.

With the aim of conserving and re-establishing the

stock of the Pacific Ocean walrus, the numbers of which are

estimated at 53 thousand head, the catch limit will be main-

tained at 1.0 thousand head for the needs of the native popu-

lation of Chukotka with a complete ban on shipborne hunting.

In addition, there have been introduced several measures

to strengthen the conservation of the walrus on the hauling

grounds. Every year during the hauling out period of the wal-

ruses an inspector will be present on Wrangel Island.

Commercial whaling for the beluga is not developed in

the waters of the Far East. The numbers of the beluga are not

known. From 1970 the catch of older individuals has been autho-

rized, with the exception of belugas with suckling young.

Scientists should devote their attention to the study

of this animal and should recommend methods of organizing its

exploitation to the industry.

THE NORTHERN COMMERCIAL BASIN

The main regions of the commercial operations are the

White and Barents Seas. From 1965 shipborne sealing for the

harp seal in the White Sea was banned for five years, the

catch of whitecoats did not exceed 20 thousand head.

In 1969 the period of the ban expired but the re-estab-

lishment of the stocks is proceeding extremely slowly and

therefore the limit for 1970 was set at 23 thousand head with a comolete ban on killing the females.

437

o

o

In 1969 for the first time there were carried out ob-

servations on the Norwegian sealing operations in the Barents

Sea. No infringements were found on the part of the Norwe-

gians.

From 1970, in connection with the unsatisfactory state

of the stocks, bans were established agaist sealing for the

bearded seal in the White Sea and the grey seal in the White

and Barents Seas.

At the present time sealing operations for the ringed

seal are not restricted. The SevPINRU (Northern PINRO) and

PINK) (The M. Knipovich Polar Research Institute of Marine

Fisheries and Oceanography) should, in the very near future,

study the state of the stocks of marine mammals in the White,

Barents and Kara Seas, and provide conclusions on their ratio-

nal exploitation.

In recent years the Jan Mayen and Newfoundland stocks

of the harp seal have not been exploited by the Soviet sealing

industry.

CASPIAN BASIN 272

In recent years the hunting for the adult Caspian seal

has been banned, with the exception of the hunting for unmoulted

whitecoats and greycoats during the period from January 25th

to March 15th on the ice in the Northern Caspian within the

limits of the quota. For 1970 the catch limit was set at 60

thousand head.

4.38

THE BAIKAL BASIN

The numbers of the Baikal sea at the present time

comprise 33 thousand head, which permits a yearly catch of

2.5 thousand head. The catch of moulted individuals WO tO 1

year of age ("serka", "kumatkana"*) is authorized during the

period from April 25th to the termination of the sledge sealing

by means of rifles of 5.6 mm caliber with fortified cartridges

and by means of fixed capron** nets under the ice, in the re-

gion to the north of the line which joins the 01'khanskie Vo-

rota to Cape Tonkii, except for the three kilometer zone around

the Ushkan'iye Islands.

In recent years measures have been undertaken to inten-

sify the conservation of the Baikal seal, the staff of inspec- (-)

tors has been increased and additional transportation has been

alloted.

THE BALTIC AND LADOGA BASINS

In the Baltic Sea a limit of 2.5 thousand head has been

set for sealing operations.

In Lake Ladoga a catch of the Ladoga seal in the amount

of 500 head has been authorized.

In accordance with the recommendations of the 3rd All-

Union Conference on Marine Mammals, the "Glavrybvod" (The Main

Directorate for Fish-breeding and Protection of Fishes) devised

Translator's notes. * "serka","kumatkana" - names of older stages of the seal pups. For the harp seal, "serka" would be translated by "beater".

**"capron" - a synthetic fibre

439

the "Provisional regulations for the conservation and exploi-

tation of marine animals for Soviet ships, organizations and

citizens", which were confirmed by the Ministry of Fisheries

of the USSR and came into force from January 1st 1970. After

these have been in force for 1 - 2 years it willbecome clear

as to how good they are and then the question will be decided

on the promulgation of permanent regulations.

UDC 599.7 11.5.2 + 599.7/1,513

D. A. Mikeladze

KEEPING FUR SEALS AND CASPIAN SEAIS IN THE 273

BATUMI AQUARIUM

The study of the way of life, behaviour and many other

questions of the biology of marine mammals is made difficult

by the lack of opportunity for systematic prolonged observa-

tions on individual animals. The keeping of these animals in

an aquarium or oceanarium permits such observations to be made

and, in addition, provides an opportunity for studying such

questions as the dynamics of the growth, the change in weight

and other aspects during particular periods of the year and

in step with the increase in the age of the animal. However,

it should be borne in mind that life in captivity naturally

to some extent changes the instincts and habits of the animals.

In Japan and certain other countries there has already

been accumulated a fairly considerable experience on the keep-

ing in captivity of certain species of pinnipeds, including

also the fur seals. ln the Soviet Union such work has not

been carried out, with the exception of individual short-term

experiments on the purposeful keeping of fur seals in enclosures

and the more prolonged keeping of Caspian seals (Badamshin,

1959). The opportunity for organizing and conducting such

studies in our country has emerged with the bringing into ope-

ration of the marine aquarium in Batumi.

In Uctober of 1966 there were brought by airplane to

the Batumi aquarium three Caspian seals, which had been caught .

441.,,

on Zhemchuzhno Island in the Caspian Sea, but which died in

1`dovember of 1967. Immediately (in this same month) there

were again brought to the aquarium five Caspian seals, that

had been caught on the same island (Table 1). The animals

were transpor.ted by airplane, where they had been placed in

wooden crates without water.

Fur seals were delivered to the Batumi aquarium on

three occasions (Table 2). From Robben Island, located in the

southern part of the Sea of Okhotsk, in August of 1967 there

were brought, in the same way as the Caspian seals, by airplane

in wooden crates six fur seals, including one pup that had

been born this same year. The next batch, amounting to four

fur seals, was brought in September of 1968 also from Robben

Island. Finally, in August of 1969 from Bering Island (the

Isomandorskiye Islands in the Bering Sea) there was brought one

adult female, that was presumably pregnant.

One of the fur seals was an unweaned pup and the attempts

to artificially feed it were not successful. The simultaneous

death of the three Caspian seals and four fur seals in 1967

occurred from food poisoning. U:,0 to December 1969 three Cas- 275

pian seals had survived in the aquarium for 2 years, one fur

seal for more than two years, 3 fur seals for one year and

one female was delivered in August of 1969.

In the northern part of the Pacific Ocean the fur seals

accomplish regular seasonal migrations. Spending the winter

months in the open sea, in the spring, as the waters become

warmer, they move to the north and spend the summer months on

the on-shore rookeries, located on Robben Island and the Kuril,

i;ornandorskiye and Pribilof Islands. The water temperature in

w.n^air.u.r--......r-...r-...-..l.^.ua..iiw.a-L+._`^..1- a..^ .v._......

Table 1.Information on the delivery of -Caspian seals

TaG:i[Iüa ^

CBejkenNH 0 3aCO3C KacnBVlCKnx Tlo!leueïl

►IonSex

Bospacr

AgeCaalert male 2 ro;>,a

yearsCamKa f e Iria.l - 2 ronLa

CaTIKa f I ro;t

Caueültl

Caaleli m

CaNIKa f

Caa1Ka fCaatKa f

Table 2.

2 ro;ta

2 roTLa

I ro;i

1 roi

1 ro:2

Date of!la a,

a.nPuHO3a

devery

OKTS16pb 1966 r.UctoberOKTaGpb 1966 r.

C<KTRGpb 1966 r.

24 IIOAGpfl 1967 r.

24 IIoslGps[ 1967 r.

24 11os16pst 1967 r.

flovember24 11o1Gp5i 1967 r.

24 non6pi 1967 r.

I1puutcuauusiRema.rks

17orn6 8 noslGpa Died,1967 r.

TIorllGna 8 1Io^16psI1967 r. I

IlorltG:[a 8 ]loaGpx1967 r.

I'vov..

of of

it

ilorltG 7 1<Iasl 1969 r.,

I7orI16,na 30 slflBaps[1968 r.

Information on the delivery of fur seals T 11 it a 2

CBeJl,ennH 0 3IIB03e hSOpCK}Ig KOTIIKOB

I'vIe1g-ht Date of Time i^11on 1i03p3C2 Bec, sz Aaru pp1uzo3a 13(: hm nnryrn 1ïPn^^e iai na

Sex lL^e kg dellvery trans ^ y Ke^nar.ks.

Caatetx Il7

Camett M

Carleu 111

Ca;`nca f

Camnta f

Camuca f

Caali:a f

CamKa f

CanlKa fCameu-ill

CaAtKa,f

2 roaa

2 roaa

year:2 roZa

2 ro;la1 ro^L

III

Pup.3 ro;ia

2 ro;ua

I ro:

.1 roa

7 :IeT

year

25

25

23.

21

19

12

26

14

19.5

18,5

42

23 anrYcra ' 1967 r.

23 anrycTa 1967 r.

' august23 anr}•cTa 1967 r.

23 asrycTa 1967 r.

23 aurycra 1967 r.

23 aurycra 1967 r.

9 celn•nGpa 1968 r.

9 cexTSlGpa 1968 r.september

9 cenTStGpst 1968 r.

9 CCIITflGpR 1968 1'.

4 aBr\'CTa 1969 1'.

august

52 ^taca

52 =Iaca

hour52 yaca

52 yaca

52 llaca

52 'Jaca

131uac.

131 llàc.

131 llac.

131 nac.

53 tlaca

17or116 8 1I0aG- Died, iVOT:pH 1967 r.

To >Ice

To Ace

To >Ice

to

W_ayJan.

ifet

n

Ilorit6aa 31 il Sep"CCI1TflG1)H

1967 r.

-çr1lIIorIIGm 21 an " il-

pe,Ia 1969 r.

274

1143

• • -•■ • •

PIIC. 1. 1(0T111:11 13 aKBapityme

Figure 1. Fur seals in the aquarium.

() the regions of the winter habitats of the fur seals keeDs

within the limits of 1-2 to 9-10 oC, occasionally dropping to

0° C. The air temperature in these regions rarely exceeds

12-13°C. With the warming of the water above 14-15 °0 the fur

seals leave the wintering regions and move to the north (Panin

and Panina, 1968; •Arsen'ev, 1964).

The regions of the on-shore rookeries are characterized

by having cloudy weather with drizzling rains and an air tem-

perature ranging from 10 to 15 °0. On occasional sunny days

this may increase to 2000 and more, and then the fur seals

tend to spend the greater part of the day in the sea, where

the temperature of the water generally does not exceed 10-12 00.

In the Batumi aquarium the fur seals encountered com-

pletely different conditions (Figure 1). The water temperature

in the aquarium varied from 6.5 to 25.4°C, while the air tem-

perature varied from 2.5 to 30 0C. The differences in the

CC Ile :flom 1:0>iaa-nopcime wrpo-

BM

6a -rpm KO:Wall:10 1' -

CKIIC OCT pOLIZI

BwryNm

Komait- ',-topewlic ,errrpo-

im Gal-pm

21,2

MA 21,3

94,9

21,5

24,0

6,8

8,2

10,8

11,5

7,8

9,3

6,1

7,7

10,3

11,2

7,2

9,4

Table 3. Comparative data on the water and air temperatures, o C.

, T a 6 au u, a 3

CpanuuTenbume Rauume o TemnepaType nogm H noagyxa, °C

.A. • TeNusepa'Typa B • Temm,parypa ito:./tyxa

d ico.rie6anim von umecm

Year and month

Komatutop- CKIle

f ourpoca

(I KoacCiamist CCpcansm

BaTVMIL

mo June iliolib

July Mon' AUM

Sept . CeltTuCipb

190

June HIM,

July Miom,

18.0-23,5

23,2-2-1,6

22,7-25,6

22,9-25,1

18,0-23,4

23,2-24,6

5,4-8,2

7,6-9,2

9,4-12.6

9,9-13,5

5,8-8,2

6,7-9,8

19 .8 22.4

99,6

21,4

20,9

21,4

14,6-25,6

15,8-28,6

17 ,5-28,7

W1-29,0

17,0-30,6

17,6,-29,3

4,0-7,5

8,5-11,8

9,4-12,6

r)

A. - Water temperature B. - Air temperature

c - Average d - Ranges

e - Batumi f Komandorskiye Islands

4‘)

temperature conditions in the aquarium and in the region of

the Komandorskiye Islands are shown in Table 3.

As can be seen from the table, the hydrometeorological

conditions of the habitats of the fur seals in the region of

the Komandorskiye Islands and in the Batumi aquarium differ

considerably. So abrupt and rapid a change in the conditions 276

of the habitat may lead to certain changes in the organism of

the animais, but the investigation of these questions is a

task for further studie .s.

All that has been said also applies, to a somewhat

lesser degree, to the Caspian seals.

445

With their placement in the aquarium, there also

occurred a considerable change in the food objects of the

animals. In the northern part of the Pacifie Ocean within the

stomachs of the fur seals there have been found the remains

of 45 species of fishes and 8 species of squids. The main

food objects in different regions are the Korean cod, the sand

lance, greenlings, lanternfishes, herring and certain squids

(Panina, 196 )). Under natural conditions the Caspian seals

feed on gobies, "kilka", herring, silversides and gammarids

(Badamshin, 1948, 1959). In the aquarium both the fur seals

and the Caspian seals were fed on the Black Sea scad, and more

rarely on the picarel and the Black Sea anchovy. Predominantly

defrosted fish was provided, more rarely live fish (Figure 2).

Under natural conditions in both the fur seals and the

Caspian seals the weight of the food consumed per day ranged

from 5 to 15; of the weight of the animal itself, though in

this case the regularity of the feeding could not be observed

(Boitsov, 1934; Dorofeev, 1964), since at times the animals

cannot obtain food for themselves over the course of relatively

long periods, while at other times the food is plentiful.

In the aquarium the animals were Provided a regular

feeding twice a day, and in this case, as our investigations

showed, the amount of food consumed ranged within the limits

of 10 - 15% of the body weight. In i:ndividual cases over the

course of two to three days the amount of food consumed was

as much as 20-26% of the weight of the animal, but then the

demand for food dropped markedly. It is possible that with

the higher caloric value food, obtained by the animals in the

aquarium, the feeding rate decreased somewhat.

ena mom Date of delivery

Mm

.S'ex

23 anryna 1967 r. 9 ceuTsiGim 1968 r. ;Se Di 9 ce11r96p1 1968 r. 9 ceuTsulpn 1968 r. 4 anrye -ra. 1968 r. Aug.

31,5 31,5

. 21.5 23,5

- 42,0

Pile. 2. lopm.leuue Kuriu:ors

Figure 2. Feeding the fur seals.

Table 4.

Change in weight of the animals.

T a 6 u a 4

Ilamenenue Lulea ,',,, nOTHMX

4.46

B.Decnaunannyme,ta

13e,nc„,1,1,P," uc,t Aug. 12 ownwpsi j 3 .11IapTa I 12 anrycrn

e2 196S r. 1069 r. 1969 r. A.

AlopcKue Korutai Fur seals Ca meuM 25.0 36,0 28.0 1 Ca mat. 26.0 - 25,0 25,0 Cameum 18.5 18,0 17,0 CamRaf 19,5 19,0 18,5 Ca mizaf 42,0 ' — —

Kacludicraie 710.7efill Caspian seals " 24 11096p9 1967 r. :NOV • Caetti 20,0 I 23.0 22,0 29,0

24 no516ps 1967 r. C9MNaIj 16,0 19,0 19.5 . 25,5 . 24 non6pH 1967 r. Cana1 15,0 !« 1,0 18,5 24,5

3FrIplimegaaue. B nepeol c nena6pn 196S no main. 1939 ro.u. a Asuno-rnbte n ann'apnyme-ne GLUM oi.:c.,cnegenbi z.ocTa -romnum E.0.111 ,1eCTI1ONt KOpM a.

* Note. During the period frOm December 1968 to March 1969 the animals in the aquarium were not provided with an adequate amount of food.

A. - Weight at delivery, kg B. - Weight in aquarium, kg

m - male f - female

F I. I.

44?

o In Table 4 is shown the change in weight of the fur

seals and Caspian seals during the period of their stay in

the aquarium.

A task for further studies is the detailed investiga- 278

tion of the feeding of the animals under the holding conditions

in the aquarium and the influence of this factor on the growth

and development of the organism.

In the Batumi aquarium constant observations are con-

ducted on the behaviour of the animals which are being kept,

and some Preliminary results of these observations may be re-

ported.

The fur seals easily and rapidly become accustomed to

people, especially to those who constantly attend them. They

quickly begin to recognize those who feed them and quietly sub-

mit themselves to these people. Fur seal No. 1, an aborigine

of the Batumi aquarium (the only survivor of the first batch,

who is now already 4 years old) and the adult female do not

• resist when their mouth is opened to examine the state of the

teeth.

The Caspian seals have also become accustomed to people,

they calmly take food from the hand but do not allow themselves

to be touched.

Observations show that the marked change in the living

conditions, and apparently primarily of the temperature regime,

has exerted a considerable influence on the periods of moulting.

Thus, in fur seal No. 1 in 1969 the moult commenced on May 28th,

approximately one month or even earlier than is usual under

the natural conditions. The so-called third stage of the moult.

448

o was noted at the beginning of July, while on the islands this

stage is generally observed in the first half of August (les-

terov, 1968; Bychkov, 1964). In the Caspian seals also the

moult commenced at an earlier period than in the Caspian Sea.

Of great interest is the elucidation of the question

as to the influence of the life under the aquarium conditions

on the quality of the fur pelt of the animals, especially of

the fur seals.

The work with the pinniPeds, which are being kept in

the Batumi aquarium, is only beginning. Included among future

tasks are comprehensive studies on the effect of the stay in

artificial conditions on the vital activity, growth and deve-

lopment of the organism.

011TEPATYPA

1 Apcenbee B. A. 0 cmeninnantitt nom:amuït mopcirttx KOTI1E011. Mopcirrte 10111- lonlamblicro Boctrora. TI1HPO —BHIIPO. M., 1964. 2 Baan.utittut B. Il. 0 nirrounit racintiicKoro Toaenn. Tpybt Boaro-racrinficuoft

cTanuilli, T. 10, ACTPaXallb. 1948. 3 Baaottinin B. II. Onbur co.:tepwatinst racinnicroro Tionenn n neuone. C6opinur

cieeri no 61(o.loritit. Tpy.arr KnenlIFIPO. fliuneupomii3nar, M., 1959. 44. Batioa JI. B. KOTIIIMBOC X0331 fiCTBO. M., 1934.

Bb1lltiO8 B. il. MaTeptianbi no antimite moperux ROTIIIZOB na ocTpoce TIOciellbeM.

opciie KO Bocrora. THI-IPO — BHIIPO, M., 1964.

6 ,z/opooepa C. B. Ceriepirbie mopcirne. ro-rurit (CaIlorllinus ursinus). kt.operile ro-rinat ii,a,unero Bocrora. THHP0— B1-11•IPO. M., 1964 .

7 Ilecrepoo I'. A. MaT.epilaabi o cporax .11111bE.II EOTILKÔB Ha KomatuopcKux ocTpouax. ilacTotiorne ceucpnoii mac-rit Tiixoro oKeatia. M., 1968. •

8 IlaRtma r. K. Flirt- aune No-rin:ori u StnolicRom mope. MopcKire Eo•riikti ,aa.tibue-Boctrinia. T'IWO-B(1'1P°, M., 196-1.

9 Hamm K. II.. [lamina F. K. ',.-/Konorlui ii murpannir OTI1KOB il alimile-necett-unii tiepno,k 1.1 griouch:em ,flacTottot ne ceueptioii tiacTit Tri xoro

1 IIHPO—B1-11•11)0. M., 1968.

449

REFERENCES

1. Arsen'ev V. A. The mixing of populations of fur seals.

Fur seals of the Far East. TINRU - VNIRU (Pacific

Ccean Institute of Fisheries and Gceanography - All-

Union Research Institute of Marine Fisheries and Oceano-

graphy). Moscow, 1964.

2. Badamshin V. I. The feeding of the Caspian seal. Trudy

volgO-kaspiiskoi nauchnoi rybokhozyaistvennoi stantsii

(Transactions of the Volga-Caspian Fisheries Research

Station), vol. 10, Astrakhan'. 1948.

3. Badamshin V. I. An attempt at keeping the Caspian seal

in captivity. Sbornik statei po biologii. Trudy

KaspNIRO (Collection of papers on biology. Transactions

of the Caspian Research institute of Fisheries and

Oceanography). Pishchepromizdat (State Pulishing House

for Literature on the Food Industry), Moscow, 1959.

4, Boitsov L. V. Fur seal_management. Moscow, 1934.

5. Bychkov V. A. 'Information on the moulting of fur seals

on Robben Island. Fur seals of the Far East. TINRU -

VNIRO, Moscow, 1964.

6. Dorofeev S. V. Northern fur seals (Callorhinus ursinus).

Fur seals of the Far East. TINRO VNIRU. Moscow, 1964.

7. Nesterov G. A. Information on the periods of the moulting

of the fur seals on the Komandorskiye Islands. The

pinnipeds of the northern part of the Pacific Ocean.

Moscow, 1968.

8. Panina G. K. The feeding of the fur seals in the Sea of

Japan. Fur seals of the Far-East. TINRU - VNIRG,

Moscow, 1964.

9. Panin K. I. and Panina G. K. The ecology and migrations

of fur seals during the winter and spring period in

the Sea of Japan. The pinnipeds of the northern part

of the Pacific Ocean. TIivRO - VNIRO. Moscow, 1968.

45 1

Une 599.745.3

V. D. Pastukhov

THE FEEDING OF THE BAIKAL SEAL 280

UNDER EXPERIMENTAL CONDITIONS

The study of the feeding of the seal under natural

conditions presents certain difficulties. In connection with

the fact that the animals feed mainly during the crepuscular

periods, while, as a rule, they are caught during the day,

the stomachs of the animals are found to be empty. However

in the large (rectal) portion of the intestine there is always

contained a greater or lesser amount of otoliths, which are

very convenient to use for determining the species composition

of the fishes. Moreover, in the sorting of the otolith samples,

removed from the intestine of the seals, a striking feature

was that the otoliths that belong to one and the same fish

species differ markedly from one another in their dimensions.

This suggested to us the idea that the fishes consumed varied

in age, size and weight. From this, naturally, it seemed

possible to establish the data on these fishes on the basis

of their otoliths. However for this there had to be answered

other, no less complex questions: could the otoliths be diges-

ted, how long were they retained and what was the degree of

their state of preservation during their passage through the

gastro-intestinal tract of the seal. All of these points, as

well as a whole series of other questions, could be resolved

only in an experiment.

•-• 452

With these aims I during the period from September -

November 1967 to April - June 1968 there were held in large

tanks (1.2 m3 ; 2.3 m3 ; 7.4 m3 ) three Baikal seals: Ushkan

(aged 1.5 years), Vega (of the same age) and an adult female

Manyunya (more than 10 years old). In the 7.4 m3 tank there

was constant running water, while in the two other tanks the

water was completely changed once a day. The water tempera-

ture in the tanks was close to the mean annual temperature of

the surface waters of Lake Baikal, about 5 0 0.

The animals were fed with fishes, which commonly occur

in their food: the Baikal oil-fish, and pelagic and benthic .

gobies. However, while the Baikal oil-fish predominates in

the natural rations, in the conditions of the experiment the

seals were fed primarily on pelagic gobies. Mainly fresh,

recently caught fish were provided.

For . determining the periods of retention of the undi-

gested remains in the digestive tract, marked food was manu-

factured. Coloured . taas of three different types were employed:

plastic platelets (25 x 6 x 1 mm); finely cut up pieces of

this same Plastic (2 x 2 x 1 mm), which are close in their

dimensions to the otoliths of fishes, and pieces (30 x 10 mn)

of thin rubber (Table 1).

The plastic platelets were retained for a particularly 281

extended period of time. The rubber tags emerged three times

as rapidly as the former. The small pieces of plastic appeared

in the tanks, on average, 1.5 times more rapidly than the rub-

ber tags. Evidently, as a result of the working of the stomach

and the Peristalsis of the intestine, small undigested particles

are expelled more rapidly from the digestive system of animals.

1 • - ■

o 47 8

3. Lilly D.* Man and dolphin. Moscow, "Mysl" ("Thought"),

1965.

L. Rei K.** The flight of three whales. "Amerika" ("America"),

No. 87, 1963.

5. Sokolov V. E. et al., An attempt at studying the charac-

teristics of the swimmïng'of dolphins in a water channel.

Tezisy dokladov 4-go Vsesoyuznogo soveshchaniya -co

izucheniyu morskikh mlekopitayushchikh (Reports of

Proceedings of the kth All-Union Conference on the

Study of 1,arine Mammals). Kaliningrad, 1969.

6. Tomilin A. G. A story of a blind sperm whale. Moscow,

"Iauka" ("Science"), 1965.

Translator's notes. * sic. Should be e. C. Lilly.

** This is a direct transliteration of the Russian uhonetic transliteration. •Probably I... Ray.

4'79L

LDC 599-537

D. A. Yorozov

TRANSPORTATION OF LIVE DOLPHINS 296

The transportation of live dolphins is one of the im-

portant questions in the overall.problem of the maintenance

of these animals in captivity. Dependant on the quality of

the transportation are the health of the animals and the dura-

tion of their life in captivity., and consequently also the

opportunity for conducting variôus studies and experiments.

Within a single experimental station the animals are moved from

the summer sea pens to the winter ;:)ools with warmed water.

Durin`s the course of the cleaning of the pools and in the ?pre-

para.tion of various experiments it is necessary to move the

dolphins from one from one building to another. A mastering

of the technique of transportation of dolphins permits one to

transport experimental animals from one specialized station

to another, to employ trained dolphins in spectacle shows and

to solve problems of an applied character (searches for under-

water objects and securing diving work).

In connection with the expansion of experimental stu-

dies it seems useful to provide a generalization of our expe-

rience concer.ninx?, the transportation of dolphins, which has

been accumulated in conjunction with F. A. Leontovich, V. A.

Kalganov, V. V. T^.',,anukhov, V. V. Belyaev and others.

Cot.ltmencin.!^, this work in 1965 and having available very

limited information on achievements abroad, we were forced to

make and test devices that we had constructed ourselves and

to try out various methods of transportation. 'îdotwithstanding

the diversity of the means of transportation which have been

utilized for this purpose (motor boats, launches, medium Black

Sea seiners, dry-freight transporters, special and passanger

shi.los, several types of motor vehicles, helicopters etc.),

at the present time we may single out only three basic methods:

towing in the sea, placing in a limited volume of water and

transportation without water.

Particularly unfavourable to the dolphins are the con-

ditions of transportation "without water". Under natural. çon-

ditions these animals are found in.a state that is close to

equi_librium and analogous to weightlessness. The force of

buoyancy, evenly applied to the whole body of the dolphin,

.neut,l"ali7.es the force of the weight, and at the same time only

the hydrostatic pressure acts on the internal organs. During

deep dives under the influence of the hydrostatic pressure,

as a consequence of the great motility of the thorax, the vo-

lume of the lungs in the dolphin may decrease tenfold for a

short period (V. V. Babenko and D. A. Prorozov, 1968). During

transportation without water the lungs and other internal or-

gans are compressed for an extended period under the action

of the weight of the animal. Dolphins are unaccustomed to the

drastic limitation of space and mobility, and to the isolation

of animals from one another.

During transportation the following undesirable effects

and traumas may arise.,

1. Compression of the internal organs, which makes

breathing and the work of the heart more difficult.

2Q?

k81

2. Injury to develouinq embryos and premature births.

3. Dislocation or soraining of the joints of the

pectoral flippers.

• . Fechanical injuries by the transporting apparatus:

cuts, bruises, scratches, abrasions and tears to the edges of

the flippers, leading to considerable losses of blood.

5. Temporary loss of motility of the locomotory mus-

culature.

6. Drying of the eyes and Portions of the skin, heat

and sun burns.

7. Overheating of the organism and heat stroke.

8. Entry of water into the blowhole during inhalation.

9. The impossibility of raising the dolphin to the

surface of the water for the normal exchange of air in the

lungs, as a result of entanglement in nets or the dolphin being

under other animals.

10. Stress with a lethal result.

The main inconvenience in the transportation of the

Azov doluhins (common porpoises) is caused by their habit of

being constantly in motion. The active movement of the tail

makes every stage of the transportation of these animals more

difficult. They are inconvenient to carry by hand or in a

stretcher, they frequently push one another down under the water,

they acquire traumas and, conseauently, suffer more from loss

of blood. The specific. difficulties in the transportation of

the bottlenose dolphins arise as a result of the great weiPht

of the adult animals, which in some cases may be as high as

300 kg.

4.82

We will pass on to a description of the procedures

and contrivances which assist in averting the enumerated

troubles. Entanglement of the dolphin in the net may occur

during the catch1ng, when the ourse seine is hauled in.

This is most likely to occur in those places where the net

gathers into folds or is pushed-in by the current into an

overhanging wall. Much more rarely the dolphins will ram

perpendicularly hanging nets. Having gotten caught in the

mesh by its beak and attempting to raise itself to the surface

of the water by the shortest path, the dolphin takes up a

vertical position in the water and in doing this it tangles

its jaws still more deeply in the net. Sometimes at a depth

of only one or two meters from the surface of the sea there

arises a threat of suffocation and drowning of the dolphin.

Therefore, several of those who are participating in the trans-

portation get dressed beforehand in their diving suits, fins

and masks, and, in case of necessity, jump into the water.

Tt is generally possible to free the animal from the net with-

out an aqualung on the diver, but at the saine time watching

the divers from the ship are one or two aqualung divers, who

are ready to quickly come to assist*.

* In working with nets the prototype model "Ukrainy-3" proved itself in an excellent manner; this is a new aqualung by the designer A. I. Gnamm. Special arches (bulges) prevent nets from hooking onto this aqualung. The AVM-1-1',1 aqualung often hooks onto a net with its charging nozzle, minimal Pressuraindi-cator, manometer and shut-off valve. I..ornover, the minimal pressure indicator and other metallic comoonents on the left shoulder strao may injure the delicate skin of the dolphins. For working- in nets one may remove from the AV1.-1-1,-. aqualung the charging nozzle and all of its fittings. A charging Point end-cap cov'érs the ooening of the high pressure nozzle near the shut- off valve. A supplementary arch Prevents the nets from hooking onto the valve.

14.83

NC. 1. Renhcbulibi B «noxrounemiom» Henoese

Figure 1. Dolphins in "submerged" net.

Une of the procedures for protecting the bottlenose

dolphin from injury by the net is the placement of a diver

between the dolphin and the net. The head of the animal is

placed on the shoulder of the man, the net is kept away from

the tail by using one's legs, while the hands protect the

Pectoral flippers from getting tangled in the mesh of the net.

The water suits "Sadko" and "Super-Calypso" without the weighted

belts have a Positive buoyancy and this permits one to lie on

the surface of the water without effort. The animals are

calmed by careful stroking. We did not have a single case of

a dolphin biting while it was being caught, although fairly

frequently it was necessary to grasp them by the jaws with

one:s hands aad even to remove the net from the teeth. The

ideal procedure for protecting the doluhins in a partially

484

hauled in purse seine would be to position divers beside each

dolphin, although under the conditions of mass capture this

is generally not possible (Figure 1).

The ship which transports the dolPhins does not ap-

proach right up to the nurse seine net, and the animals are

towed to it in special cages. The frame of the cage is welded

of duralumin tubing o in such a way that water cannot get into

the internal cavities of the tubes. To increase its buoyancy

and stability, there may be attached to the cage additional

foam-plastic floats. Over the frame of the cage is stretched

fine-mesh capron material, if possible - made of thick threads.

The dolphin is moved out of the purse seine through

the open end side of the net (Figure 2). After one or two

dolphins are placed Ln the cage, it is towed by a row-boat or

small motor boat to the transporting vessel. The Position of

the animals in the cage during the towing is controlled by

swimmers (Figure 3), Particularly restless dolphins are

placed in the cage with a man, whose job it is to support the

head of the animal during the towing, to avoid entanglement

in the net.

The raising of the animal from the water onto the

deck of the transporting ship is.the most difficult and

dangerous, to the animals, operation. Norris (1966) warned

against attempts to raise the dolphins from the water by the

tail, since in this case the blowhole is under the water, the

breathing is impeded and the animals often drown. Pulling out

the animals by hand into the small boat generally leads to

traumatization of the animals against the hard fixtures on the

side of the boat, the ribs and other parts of the boat. The

* Translator's note. sic. However the actual figure 3 is not found in the oa2er.

!".

14.85

Pm. 2. lipzcupoma meril C iteithepincamit

Figure 2. Towing the cage with dolphins.

best method up to the present time remains the method of .

raising the dolphins from the water on a stretcher, usine a

derrick. On special ships the cage with the animal may be

pulled up a slip or through an open ramp.

In Figure /1.*is illustrated the most successful design

for a transporting cage, in which the end walls fold'down on

hinges for the entry of the dolohin into the cage. Other

designs, which do not have the folding-down walls, have to be 300

sunk down on one side in order to, as it were, scoop up the

animal, but in the confinement of the partly hauled in net

this is very inconvenient. The latter design permits one to

take up the dolphin without bringing the cage into the net 301

but placing only the folded-down wall over the top line. The

second foldine wall Permits one to brine the dolphin out of

the cage head first. The convexly arched side wall struts

and the upper connecting tubes have a form which decreases

the -Drobability of the animal beinz injured by them. The skids

*Translator s note. Figure 3.

sic. Description, however, refers to

^^ --^--^--^`C.^ ^-^ ^ ',- ^.,^•^ ^. ^.-^.

^^`^^^^^

Puc. 3. TpaucnopTUan K:ICTb IIOaO}f }iOIICTpYIitlltN

4

figure 3. A tr. ansporting cage of a new design.

^` {1--w ^..+^^•.r ^ ^ ^

Pue. 4. TpattcnopTUpoa:.a Ae1b^titta ma r."t ► ccttpYto-uicPi pame

Figure II.. mrans?porting a dolphin on a hydroplaning frame.

^protect the ;e s roi:l be i-n^^ de.r^a:ed when it i s '?u11ed. u^^ on

shore or u-o a;,li,,) onto the cleck and raise uA), above the Ç1ip-

wa^i or -round, the bottom of the cage on which the dolphin is

1.yinÛ. It is best to make the bottom of rubberized ca^^ron.

4-87

it should have small openings for the drainage of water, so

that when the cage is lifted the excess water is not lifted

with it. The most convenient variant has a removable bottom.

In this case the bottom has a separate framework with 4 - 6

ri 11^ bolts for the attachmient of the slings of the derrick.

The si?e of the botton of the cage, 3.5 x 0.8 m, permits

loading of three-meter Black 'S)ea bottlenose dolphins. The

corresi)ondence of the dimensions of the removable bottom of

the cage with the dimensions of the tubs used for transporta-

tion i)ermi-bs one to place the dolphin into the tub on this

same bottom, avoiding superfluous transfers. A reserve supply

of interchangeable botto,ms considerably exuedites the work of

loading and unloading. A cage of this design may be employed

for the temporary isolation of dolphins in a pen or pool.

For the towing of dol^hins there can be proposed a

hydroplaning frame design (Figure j'`) which was elaborated by

us. The outside dimensions of the frame (3.5 x 0.8 m) permit

it to be placed into a transporting tub. While the removable

bottom of the cage should have a negative buoyancy, the front

and side tubes of this frame should hold it on the surface,

while supporting, a dol-phin. After the securement of the ani-

mal with. straps to the canvas panel the posterior edge of the

frame sinks down. This is achieved by the fact that the

posterior tube fills up with water through its openings and

does not 'serve as a float. Additional .trim at the stern may

be obtained by installing foam-plastic floats in the front

-part of the frame. Gn account of the trim there I s created

the initial angle of attack (see Figure 21,

Translator's notes. * and *,-. sic. * - Figure 1I•. ** - second

f igure in Figure 4.

488

of the towinir, the frame emerges to hydroplane along the sur-

face. This considerably decreases the towing force required.

A distinguishing feature of the towing on the hydroplaning

frame is the securement of the dolphin with straps and the

hydrodynamic lifting of the blowhole above the water. in

securing the dolphin partiCular attention should be Paid to

tiFrhtly fastening down the caudal flukes to the canvas panel,

this prevents the possibility of the animal rolling over onto

its side. The trimming of the frame at the stern helps to

lift the blowhole of the dolphin out of the water even when

the frame is not moving, which is of particular value when it

is installed in the transporting tub. For the greater comfort

of the animal openings are cut In the canvas of the frame for

the Pectoral flippers.

The frame may be utilized as a cradle for ensuring the 302

immobility of a dolphin during the course of a medical exami-

nation of the animal or for experimental purposes.

The described ways of towing may be used in bringing

the dolPhins from the purse seine to the transporting ship,

from the transporting ship to the pen (through a sPecial pas-

sage in the wall of the pen), in moving the animals from one

section of a holding Pen to another or in moving the dolphins

along channels, connecting several pools.

The second method of transporting dolphins, conveyance

in a limited amount of water, was introduced into Practice in

the USSR by F. A. Leontovich*. Aqualung divers were first used

* This information was published by F. A. Leontovich in the re-port "A test of underwater filmirur of dolPhins" at the 3rd ple-nary session of the Section of Underwater Studies of the uceano-graphic Commission of the USSR Academy of Sciences in January of 1966.

489

for removing dolphins from a purse seine on the 9th of July

1965. They held the dolphin and brought up under it in the

water a special stretcher, on which the animals were raised

from the water, transfered and suspended in containers with

water. Used as the containers were the canvas deck tubs on

a metallic frame, of the type that is found aboard the Black

Sea refrigerator ships and seiners. F. A. Leontovich noted

that dolphins suffer considerably from all reloadings and

transfers, and recommended that a large number of stretchers

be employed, so that each animal is only once placed onto and

once removed from the stretcher during the entire journey.

This principle of "straight from the sea onto the suspension

device and to the place of holding without transfers" was set

at the basis of the further imProvement of the Procedures for

the transportation of dolphins.

F. A. Leontovich's observations showed that the dol-

phins which were placed in an individual canvas tub, 2.0 x 0.5

x 0.6 m in size, began to struugle and manifested nervous be-

haviour, while in a transparent aquarium of perspex of the same

size they behaved much more calmly. This was taken into con-

sideration in the design of the sPecial tubs for the transpor-

tation of the dolphins. Before these tubs were made there

were used canvas brine tubs, stretched out on ropes in the hold

of the ship or in the back of a GAZ-69 A truck, and canvas

tubs on wooden end-panel frames. There were also used ply-

wood boxes, lined with porolon*, within which was inserted an

inner lining of polyvinyl chloride film. Lesides these, there

have also been utilized rectangular steel boxes, welded of thin

sheet metal, inflatable rubber boats etc.

* Translator's note. "porolon" - a foam plastic.

4.90

John Lilly (1962) prepared boxes according to the di-

mensions of the animals and attemrted to decrease the volume

of water in these to a minimum. The tubs were sha-.)ed out of

thin plexiszlass in the form of the body of a dolphin, while

the strength and rigidity of the design was ensured by plywood

ribs. In order to lessen -Che spillage of water en route s the 303

box was covered with a lid, with a large slot for the dorsal

fin. In this case there was utilized the principle of simul-

taneous loading of the tub together with the animal and water,

though such a tub was suitable only for a dolphin of a Parti-

cular size.

Cur tubs were designed for the transportation of

various suecies of dolphins. They had a rectangular form,

which permitted, depending on the size of the animals, the

transportation in one tub of one or two bottlenose dolphins,

or 1.11) to five white sided dolphins, or up to nine Azov common

porpoises simultaneously. The most suitable type proved to

be welded tubs of duralumin, which were much lighter than

steel tubs, more durable than plywood tubs, and did not pro-

duce such scratches and abrasions to the delicate skin of the

dolphins as did the canvas tubs. The internal dimensions of

the transporting tubs should be no less than 3.5 x 0.8 x 0.8 m.

In the anterior part of the tub are mounted three rectangular

windows of plexiglass.

In the preparation of the tubs Particular attention

should be Paid to the method of securing the windows. In the

first design the six-sided heads of the bolts which projected

inside the tub and the edges of the tub at the window were the

cause, in several cases, of traumas to the dolphins that were

4•91

being transported. The second design eliminated this defect

by the placement of the bolts on the outside and by the selec-

tion of the bevelling on the plexiglass. ^:;e note that from

the outside the plexiglass should be clamyped down with nuts

through a steel strip which borders the window along its peri--

meter. This ensures a more even distribution of the load on

the fastenings. The hydraulic impacts which arose during

rolling and pitching caused cracks in the windows and loss of

water, if ordinary washers were placed under the nuts instead

of such a strip.

Edging the upper edges of the tub with 7cm diameter

tubing considerably decreases the probability of traumas during

the loading of the dolphin into the tub. The bottom of the

tub was provided with skids as a convenience for moving it,

while on the lower part of the sides of the tub there were

fixed handrails, which could be used for carrÿing the empty

tub by hand or for raising it mechanically. The presence of

drainage plugs considerably facilitates keeping the tub clean

and allows for the possibility of changing the water level in

it. It should be borne in mind that painting the tub with red

lead and the subsequent trans°cortation of an animal in standing

water in such a tub may lead to the poisoning and death of the

animal.

Aboard ship the presence of running water in the tubs

is provided for by making use of sea water from the fire mains.

on motor vehicles, loss of water from the tubs occurs on star-

ting, durin^; turns and. on brakin:; and also when moving over

an uneven road. Strips of porolon, placed on the surface of

1-1.9 2

the water, decrease the splashing. En route, fresh water may

be added: the movement of dolphins into rivers, sometimes

for hundreds of kilometers upstream from the mouth of the ri-

ver, indicates that brief sojourns in fresh water do not harm

them. The susPension of the dolphin in the tub on a stretcher 304

prevents injury of the animal against the walls. In this man-

ner, in 1968 and 1969, there were transported by motor vehicle

four bottlenose dolphins and four white-sided dolphins over a

distance of 250 km. Without a stretcher it is fairly difficult

to restrain an animal on an uneven road § though enthusiasts

were able to deliver dolphins without injury, at the cost of

a five-hour stay in the water together with the animal. The

main thing in this situation is the timely raising, of the del-

Phin by the jaw, so that its blowhole is above the water.

Thanks to the experiments of A. G. Tomilin (1951) it

became known that the loss of heat from the organism of the

dolPhin to the surrounding medium occurs through the flippers*.

If the flippers are not moistened with water for a prolonged

period, then, because of the smaller heat capacity of air,

there may occur overheating of the organism and. death of the

dolphin from heat shock. Drying of the eyes may cause blind-

ness. Some authors recommend wrapping the dolphins in wet

woolen blankets or in sheets. This method has its disadvan-

tages: The fabric does not adhere closely to all parts of the

skin and it hides dried-out portions from the observer. Such

coverings, however, protect the skin from burns by the direct

rays of the sun.

* Translator's note. The Russian word "plavnikin refers not only to the flippers but also to the dorsal fin and tail flukes.

11.93

Spending a prolonged time without movement during trans-

portation may cause a temoorary numbness of the locomotory mus-

culature in dol_ohins. Therefore, when it is let out into the

pen or pool the dolphin sometimes may not be able to raise it-

self to the surface to exchanP - e the air in its lungs. At such

times it is desirable to have an aqualung diver in attendance.

Further studies of the physiology and characteristics

of behaviour of dolphins under experimental conditions will

permit improvements to be made in the methods of transporting

these animals.

JIHTF.PATYPA

1 EaGengo B. B., It lowrion 1.1euoToplite (1)U31ItiecK1Ie 3aKonomepnoe-rn npn - Reaml)nnolu. C6. eMexann3Mht nepetunt:KeHlin H op uieuîa 1H III 1 UROT111,1X».

dlayKorsa Aymt;a», knell, 1968. 2 TO.U11.111H A. F. IleantitiKu oprauta TepNtopery.rnunni laiT006p8311b1X. «PbI6110e

X03grICTDO», 12 u 1950. Tomimivi. A. r. O Tepmoperymmun y Kirr006pa3twx. «Flpnpoaa», N2 6. 1951. I.illi J. C. Man and dolphin. no 4, 1061. 5 whales dolphins and porpoises. Ed. by Norris K. S. Berkeley and Los;

Angeles. Syrnpos. 1965.

REFERENCES

1. Babenko V. V. and ïorozov D. A. Some Physical regulari-

ties in the diving of dolphins. Symp.: "Mechanisms

of locomotion and orientation of animals". Publ.

"Kaukova dumka", Kiev, 1968.

2. Tomilin A. G. The flippers, organs of thermoregulation of

cetaceans. "Rybnoe khozyaistvo" ("Fish L.anagement"),

12, 1950.

3. Tomilin A. G. Thermoreq.ulation in cetaceans. "Priroda"

("nature"), no. 6, 1951.

494

UDC 599.74.5.3

K. K. Chapskii

THE SYSTEMATIC RANK AND SUPRAGENERIC DIFFERENTIATION 305

OF THE 8-INCISOR SEALS

Already in a Paper published ten years ago (Chapskii,

1961) the author argued his views on the subfamily structure

of the ohocids, the main criterion for the systematic subdivi-

sion of which was taken to be the incisor formula. Un this

basis was confirmed the classical suprageneric structure of

the seals in the form of the three subfamilies: the Phocinae,

I,onachinae and Cystophorinae, which had been accepted three

years previously by Scheffer (1958) in his review of the seals

of the world. It may be recalled that it was namely this sys-

tematics of the phocids which were adhered to by the Predomi-

nant majority of mammalogists, commencing with those who laid

the foundations of the systematics of the pinnipeds: Gray,

Gill and others. Many other prominent zoologists of our time

have gone even further and have foilnd it possible to divide

the phocids into four subfamilies (Kellog, 1922; Hay, 1930;

Simpson, 1945; 2rechkop, 1955; Ellerman and Morrison-Scott,

1957, and others).

However the arguments presented for such a division

were, in large Part, very limited and inaccurate.

This "fra=entation" of the family began in the last

quarter of the last century and, aPparently, pertains to Gray

(1874), who, after including the 8-incisor seals in a single

subfamily or tribe for many years, decidedto divide these,

1405

separatinz from the monk seals the southern 8-incisor seals

and including these in the tribe (essentially, from the taxo-

nomic aspect, equated to a subfamily) Jtenorhynchina, equiva-

lent to the Lobodoninae. In the final analysis, even though

this did not improve the systematics as a whole (since in the

process of the rearrangemelit of the genera the monk seal was

placed together with the 10-incisor seals), it focused the

attention of mammalogists on the taxonomie heterogeneity of

the 8-incisor seals and, in particular, on the specific cha-

racteristics of their Antarctic representatives.

THE INCONSISTENCY IN THE TAXONOMIC INTERPRETATION OF THE

8-INCISOR SEALS

The differing views on the systematic position of the

seals that have in common a single incisor formula are, un-•

doubtedly, conditioned by the insufficient clarification of

the elements of similarity and difference (primarily of the

craniological features) between the subtropical and Antarctic

components of this group of phocids.

r'ven in contemoorary reviews it is difficult to find

indications of any distinct osteological differences between

the seals of the genus r._onachus, on the one hand, and the

whole aggregate of the 8-incisor seals from the Southern hemi-

sphere, on the other. This situation may be illustrated by

several sPecific examples.

Almost fifty years after the systematics of the phocids

were enriched, following Gray 's initiative, by the new name

of a distinct subfamily of the southern 8-incisor seals,

o

4.96

Kellog (1922) was forced to state that it was difficult to

find any Precise differences between the Antarctic seals

(Lobodoninae) and the monk seal. Scheffer (ibid.) also aureed

with this, oddityg that, in the Presence of the already existing

subfamily of 8-incisor seals of the Sothern hemisphere, the

seY)aration of a distinct subfamily for the seals of the genus

Uonachus seems to be unsubstantiated.

Not without interest in this connection is the state-

ment by King (1966) that the intergeneric skull differences

in the seals were in general so great that it was difficult

to find any features that were common to the grouP. It is

impossible not to ouree with this statement. Unfortunately,

however, from this King arrived .at extremely different conclu-

sions in another direction.

THE NEWEST ATTEMPT AT A RADICAL RECONSTRUCTION OF THE

SUBFAMILY STRUCTURE OF THE PHOCIDS

In King's opinion, which was expressed in the previous-

ly mentioned paper, the mammalogists who created and perfected

the systematics of the Dhocids had, as it were, "gone along

the line of least resistance" in making use of the number of

incisors as a criterion of the subfamily for precisely the

reason that this is a very convenient.feature, although, from

her point of view, this is by no means sufficiently reliable.

In her own studies, directed towards elucidating the genetic

relationships between the various groups of phocids and, con-

sequently, to the creation of a more rational systematics of

1407

the latter, she completely abandoned the incisor criterion,

considering it as being too variable.

She preferred to place completely new features at the

basis of her reconstituted systematic scheme:

1. The size of the posterior lacerate foramen (fora-

men lacerum).

2. The partially visible, from above, mastoid (os

mastoideum).

3. The ventral part of the petrous bone (os petrosum)

which projects from the lacerate foramen.

It is already known what followed from this. The re-

constructed systematic scheme was found to be reduced to two

subfamilies, which were quite sufficient, as it seemed to King,

to include all of the seals. In one of these, the subfamily

of the northern phocids (Phocinae), she included the 10-incisor

seals and the hooded seal, in the other (Monachinae) - all of

the seals of the Southern hemisphere, including also the ele-

phant seal. King admits that in these subdivisions it is dif-

ficult to clearly Place: firstly, the bearded seal (genus

Erignathus), in which the posterior lacerate foramen is abbre-

viated, and secondly, the monk seal itself (genus Monachus),

since in this two of the basic features (the visibility of the

edge of the mastoid in dorsal view and the distinctly projec-

ting petrous bone in the lacerate foramen) do not correspond

to the requirements of affinity to the subfamily Monachinae

in its new interpretation. Acknowledging these deviations,

King found a way out of a difficult situation by considering

that both of these genera of seals were transitional forms,

k98

ikï

linking the subfamilies of the northern and the southern seals.

Strictly speaking, there are no grounds for disputing this

thesis in general, even if only for the reason that the Antarc-

tic 8-incisor seals undoubtedly belong to one and the same

genetic line with the monk seal, from the ancestral form of

which they also originated. Likewise, there is also no doubt

that the contemporary monk seal (from the ancestor of which

there commenced the divergence which led to the appearance of

the whole group of genera of 8-incisor seals of the Southern

hemisphere) is itself a derivative of the more primitive 10-

incisor seals. •

Probably there is nobody who doubts also the fact of

the relatively more primary and primitive character of the nu-

merous morphological features that are characteristic of the

seals of the genus Erignathus, which aspect has already been

discussed by Winge (1924, 1941), Naumov and Smirnov (1936),

and Chapskii (1955). Nevertheless, it by no means follows

from this that the seals of the Sothern hemisphere could have

originated directly from the predecessor of the contemporary

bearded seal.

The term "transitional form" is a concept from the

phylogenetic arsenal, and by itself is essentially not con-

nected with systematics. Whatever this form is, the compo-

nents which it embraces are taxonomie categories, and these

should have a fully defined position, for which arguments

have been presented in the proper manner. However, in the

system which has been constructed by King on the new morpholo-

gical criteria the position of both the monk seal and of the

11-99

bearded seal is fairly precarious. This is a postulated posi-

tion, rather than one which corresponds to the formal require-

ments of systematics. In actual fact, the motives for inclu-

ding the first of the named species in the subfamily T+'ionachi-

nae (against which, of course, there can be no objections)

clearly will not pass criticism from the formal aspect, since

out of the three criteria of the subfamily, -proposed by King,

two do not fit. At the same time the only and, obviously, in

these circumstances, the decisive feature, the specific cha-

racter of the lacerate foramen, cannot play a similar role

in the second case, in establishing the systematic status of

the bearded seal.

',+hat else can explain these facts, if not the weak 308

aspects of the systematic scheme itself and the unreliability

of the arguments on which it is based? do references to the

transitional character of the species can rectify the situation

here.

The transitional position of any particular species,

genus, subfamily etc. may manifest itself only in those fea-

tures of the structure.which do not belon` to the diagnosis.

To rephrase this postulate, it may be formulated in another

ways the features which are distinguished by an inconstancy

of their manifestation cannot serve as diagnostic features.

Cbservance of these requirements is obligatory for all

of the links of the systematic scheme, particularly for cate-

gories of so hi^^h a rank as the subfamily. Deviations may be

encountered, but only in the form of extremely rare exceptions

to the rule in individual cases. In the system proposed by

500

King, however, the infringement of those few structural fea-

tures to which an exclusively diagnostic significance is as-

cribed seems to be the rule rather than the exceotion! What

then, under these conditions, is the advantage of the new sys-

tematic scheme and why should it be given preference?

SOIE CRITICAL REMARKS CONCERNING THE DIVISION OF THE PHOCIDS

INTO TWO SUBFAMILIES

All that has been said above (the inadequate diagnos-

tic value of the new features which were erected by King to

the position of the primary craniological criteria of the sub-

families, and, as a consequence of this, the presence of the

mentinned "transitional forms") already by itself very much

weakens King's position, which so decisively broke with the

classical systematics of this family of pinnipeds.

There are, however, also other serious grounds for not

agreeing with the new taxonomic structure.

In the first place, there calls attention to itself

the undervaluation of the dental criteria (the incisor formula),

the negative attitude towards which was not argued out in a

proper manner.

The main objections, however, arise from the results

of a special craniological study, undertaken by the author

with the aim of re-examining the taxonomic problem that has

arisen. Having checked.the mentioned new criteria of the sub-

familial integration and, at the same time, having reviewed

the craniological material on the phocids that is available

in the osteological de .00sitory of the Zoological Institute of .

,501

the USSR Academy of Sciences with the aim of revealing more

completely, as far as was possible, the similarities and dif-

ferences between the various groups which were being compared,

the author reached a firm conviction as to the stability of

the previous classical systematic scheme for the phocids

(Chapskii, 1969, in litt.). Without repeating here all of his

arguments, the author nevertheless considers that it would be

useful to present in a very brief form the main motives, on

the basis of which he disputes such radical changes, introduced

by King into the systematics of the phocids.

The objections may be reduced, basically, to the fdl-

lowing points.

1. The significance of the craniological elements of

similarity, which are in fact present in the hooded seal and

the 10-incisor seals, were exaggerated in King's study; at

the same time the cranial differences between these seals

were revealed by her very incompletely and were underrated.

îeanwhile, in fact the number of similar features in these

seals is only about half of the number of the features of

difference: there are about fifteen of the former and at

least twenty five of the latter. In this situation, the cate-

gorical judgement of King on the.inadequate diagnostic value

of the incisor criterion does not appear to be very convincing.

If however one takes into consideration certain additional dia-

,gnostic features, namely: a) the narrowed spindle-shaped bulge

of the mastoid part of the temporal complex (precisely that

portion which is visible beyond the lateral mastoid process

from dorsal view and which in cross section does not exceed

502

half of the length of the tympanicum); h) the direction of

the axis of this bulge, which is inclined markedly downwards

behind the mastoid process; v) the marked swelling of the

maxillary bones immediately in front of the orbits; g) the

well developed anterior palatine foramina, situated in groove-

like depressions, - then, together with the incisor formula,

the subfamily of the ten-incisor seals acquires a fairly good

craniological basis.

2. Speaking convincingly against the systematic bring-

ing together of the elephant seal and the .8-incisor (in-parti-

cular, the Antarctic) seals is the marked predominance of fea-

tures of difference between these over the features of simila-

rity. Strictly speaking, these seals are linked only by the

abbreviated lacerate foramen, while on the basis of almost all

of the remaining elements of the craniology they are differen-

tiated. If, however, there are scrupulously counted all of

the features by which the compared seals are in any way simi-

lar and these are set beside those features which differentiate

them, then, with such an approach, it is found that there are

almost three times as many features of difference as there are

features of similarity. In these conditions it is difficult

to speak of closer genetic ties between the seals which are

being compared.

3. en the other hand, the similarity between the

elephant seal and the hooded seal is far from being exhausted

by those few and, in Ring's opinion, unimportant features which

she mentions. LA actuality, the similarity between them is

much greater. With an attentive examination of the skulls,

é 503

it turns out that there are twice as many elements which bring

these two genera of seals together as there are elements by

which they are differentiated, although there are fairly many

(about 20) of the latter, since these are different genera.

Thus, if the dental formula is accepted as the criterion for

the systematic ,grouping of the genera Cystophora and Mirounga,

i.e. in this case the presence of six incisors (i = 2/1), then

in the seals which are united by this characteristic there are 310

found to be about 40 similar features, and only about 20 dis-

similar features. If however, on the contrary, one is guided

not by the incisor formula but by the relatively little exten-

ded lacerate foramen, and, consequently, one unites the ele-

phant seal with the southern 8-incisor seals, then the total

number of similar elements of the cranial structure in these

will be no greater than five, while at least a dozen features

of difference can be compiled. •

With such numerical ratios, one would think that it

would be difficult to insist on uniting the elephant seal in

one subfamily with the 8-incisor seals. Similarly, it is un-

natural to combine in the second subfamily the hooded seal

with the 10-incisor seals and, as a result, to liquidate the

third subfamily of the 6-incisor seals, the Cystophorinae.

To sum up, the truth in the taxonomic structure of the

phocids is found to be on the side of the classical systematics,

and as the leading criterion of this, at any rate for the con-

temporary members of the family, we may consider the dental

formula*.

* (This footnote is given on the next page of the'translatiOn.)

504

CRANIOLOGICAL SPECIFIC CHARACTERISTICS OF THE 8-INCISOR SEALS

Accepting the incisor formula as being sufficiently

weighty, from the taxonomic aspect, and giving preference to

this classical criterion in_subdividing the phocids into sub-

families, it is impossible not to see also how difficult it

is to find any other craniological features which would in the

same way unite all of the 8-incisor seals and would be speci-

fic only to these. This most important, if not the only, dia-

gnostic element could be expanded by taking into account cer-

tain additional features, namelys

1. The low positioning of the anterio-inferior edge

of the cheekbones (Figure 1).

2. The markedly more developed, than in other seals,

and further backwardly directed zygomatic process of the maxil-

lary bones (Figure 2).

3. The narrowed anterior angle, which is markedly

smaller than a right angle, of the lumen of the orbit in the

horizontal plane (Figure 2).

However all of these additional craniological charac-

teristics are not very expressive, are subject to considerable

variability, are not free of exceptions and may sometimes be

found also in other seals.

Still less diagnostic are several other structural fea-

tures, which, though characteristic of all of the members of

* (From previous page). The author is forced to make the stipulation with respect to

the contemporary phocids at the present time on the grounds that in the upper Tertiary ancestors of the monk seals (genus 1:.ono-therium) there were allegedly present not 2 but ) incisors in each half of the upper jaw (Kellogg, 1922; Boettger, 1951).

505

4

Pttc. 1. KonTypl.[ netsoït ct<ynoneïtI:oCTtt (n^loi^lttnt.) Ttone1[aAFOttI x^I

(8eflxBltït j111C.) il TlD:lettfl y9a^[ ^^I87CthlliCKÿ Rlo'soü(».)oc'fil, Q-

Otio9tte'IC1111S1: a rc noi!^'fe31[If911oC nrRepcrtfe. ee nepei;He-Ila;:fnuil tipafFq

CYCAIIC^RCj1Xl111Ïf uTj1UC70t: i^OC'l'ft,. ^ ^ .-

Figure 1. The contuurs P f the le ft zyg o:matiç boi:e (profile)

of the monk seal (upper figure) and the Weddel

seal (lower figure).

Ilegend: . a - suborbitai foramen, b- body of ^ygo=-matic bone., v-- an^erio-superior proçess of bone,g -'its anterio--inferior edge,

this subfamily without exception, are at the same time inhe-

rent also to the representatives of the subfarriily of the six-

incisor seals. JLmong -these bel-4g; the wide but not niarkedl,y 311

protruding bulge of the mas^oid, whieh is not direeted down-

wards behind the mas.wid proçess; the eonçave outïines of the

projection of the lateral contours of the yre-orb.i.tal part onto

a horizontal plane; the disappearing anterior -Palatine fora-

mina; developed pre-orbital proçesses; a wide inter-ocular

space; abbreviated lacerate foramina.

506

Puc. 2. CxeataTt['tecxoe nsoGpaxtenne ^tacreït vepena• cuepxy:

n-- itwtTypt,t aepcna naprn (npcAcTau11TcAH ABCHTnpC3qOUWX TtoIc-ncit); 6--- t(rparatcuT npmroü cTOporu.t ^tcp, na -1 ro:tcuH-n+onaxa: a- raaa=Hu4nan 4acTb npaEOit croponLt vepena Ttoncnn Y9AACnAa; a- To Ace,

Ttosctts[ Poccaq pcpl,nlncT6tMn JtitlntHa+tt oÜoaitaHctta ;tJnnta (no npoAoJtbftoti oeil 4C-

pena) chy.nonoro oTpocTr;a acascunns:pnoCt hocrn

Figure 2. Schematic depiction of parts of the skull from above s

a - the contours of the skull of the harbour seal(a representative of the 10-incisor seals);

b - fragment of right side of skull of the monk seal;v- orbital part of right side of skull of the

Wed.dell seal;g - the same, of the Ross seal

The interrupted lines indicate the length (alongthe longitudinal axis of the'skull) of the zygoma-tic process of the maxillary bone.

•

5 0 7

o CRANIOLOGICAL GROUNDS FOR THE TAXONOMIC DIFFERENTIATION

OF THE SUBFAMILY OF THE 8-INCISOR SEALS

The inadeouacy of the available characteristics of the

tribes, of their diagnostic features, makes it urgently neces-

sary to have a proper reviSion and more precise definition of

the craniological basis of the established taxonomic subdivi-

sions of the 8-incisor phocids, the P,Ionachinae.

Gray (1874) was probably the first to present certain 312

definite and fairly accurately grasped specific characteristics

of the bones of the skull (in particular, the fusion and elon-

gation of the nasals), which permitted one to more or lesS re-

liably identify the 8-incisor seals of the Southern hemisphere.

Subsequently to these was added one other feature, the elonga-

ted zygomatic processes of the maxillary bones.

AS a result of the most recent examination of these

diagnostic features and all measurements of the craniological

difference of the 8-incisor seals of the Southern hemisphere 313

(Lobodonini) from the subtropical representatives of the

Monachinae, there were found, on the one hand, certain weak

aspects of Gray's diagnosis, while on the other hand there were

revealed several new and more precise elements of difference*.

It was found, in particular, that even such a charac-

teristic feature of the southern 1Lonachinae as the fusion of

the nasal bones is not completely reliable, since divided nasal

bones are not infrequent in Weddell seals (and not only in the

* This work was carried out mainly on the same craniological material in the osteological section of the zoological insti-tute of the USSR Academy of sciences.

508 F

young); it is encountered in the crabeater seal, and a ten-

dency to this is also noted in the leopard seal.

In all of the subtropical monk seals the nasal bones,

of course, are always divided. The second feature, the elon-

gated frontal part ("auex") of the nasal bones, also sometimes

does not give the typical picture in the southern seals, and

at times manifests the opposite development also in the monk

seal. Reference has also been made to the variability in the

length of the zygomatic processes.

Thus, in spite of the characteristic nature of the

craniological features, which one cannot avoid taking into

account, they nevertheless cannot have a primary diagnostic

• sienificance.

Taking into consideration all that has been said, the

cranioloeical part of the expanded diagnoses of both tribes

of seals of the subfamily Monachinae may be represented in

the following form.

TRIBE MONACHINI

Zygomatic bone and profile with a fairly clearly out-

lined anterio-inferior edge, permitting the length of its body

to be measured (without the anteior articular process). The

inferior edee, from the same position, along its whole length

has the form of an arch, which is elevated to the greatest de-

gree in the middle part of the body of the bone (Figure 1).

The end of the anterio-superior process of the zygo-

matic bone extends far anteriorly and terminates almost over

the interior edge of the suborbital foramen. The latter is

509

Pile. 3. Cxehta c -rpeemist mae -rit 'wane erepopm mepena ertepe,mt. rft Ea3HHO cocyroomeulle NICH ■ :(y riestemcennem ripe,D,raa3nwiliere o -reepcTim (ypepeob 'compere e -rmetielt upephusirerell :tumid') H

iiilem eKyaorteii 1:0CTIL (opereem ee pep:mere Hpayi) 060311aulitur. m— TIO.ICI1b-M011ZIX, Il — MO)CI ■Orl neonapjk

Figure 3. Schematic diagram of Part of the right side of the

skull from the front. There is shown the relation-

ship between the position of the pre-orbital fora-

men (the level of which is marked by the double

interrupted line) and the position of the zygoma-

tic bone (the trough of its upper edge).

Legend: M:- monk seal, N - leopard seal.

situated no higher than the level of the greatest deflection

downwards of the upper edge of the body of the zygomatic bone

(Figures 1 and 3).

The nasal bones are not grOwn together with one ano-

ther; the frontal portion of these is generally not longer

than the remaining (anterior) portion (Figures 2b and 4).

The fontanelles in the reeion of the presphenoid are

huge and round. -

The tribe includes the three sPecies of monk seals of

the genus 1\.onachus 21em.

A A

510

Puc. 4. (1)0Inta CJIMAX 11000BIAX 1:0CTal TIOJICIICfi 10)uuoro nouywainut: Tumeun Pocca (A) u mopcuo-

ro Aconapua (5) orpeilliamu ii upephuulcumu Juouumu noicalana

noGnore (*nepuntimi») noconbrx Nocreii. 06038atieinun Jr Kont, 2I —Bepxite!te.moeritari,

n — H000111,10 nocrit _

Figure 4 . The form of the fused nasal bones of the seals of

the Southern hemisphere: the Ross seal (A) and the leopard seal (3).

The arrows and interrupted lines indicate the length of the frontal portion ("apex") of the nasal b-ones.

Legend: fr - frontal bone, m - maxillary bone, n - nasal bones.

TRIBE LOBODONINI*

Profile of the zygomatic bone without any angular bor-

der between the maxillary process•and the body of the bone;

* The 8-incisor seals of the Southern hemisphere were first grouped in a separate tribe, as far as may be judged now on the basis of the available literature sources, by Gray (1871, 1874), thouP:h the author made use of a preoccupied name, Steno-rhynchina, which was subsequently re .placed by another. Since the Presently acceoted name for this tribe is derived from the crenus Lobodon (not Lobodont), then from this it seems that the more correct derivative is namely Lobodini, by which name the tribe is here designated.

511

its inferior edge is either completely without -an arch-sha1ped

curvature, or with a barely expressed, shallow arch only in

the posterior half of the bone. The anterio-suuerior end of

the zygomatic bone reaches far short of the suborbital foramen,

terminating at the level of the latter or below this. The

zygomatic bones in general are set considerably lower and the

greatest bend downwards of their upper edge is situated below

the suborbital foramen (Figures 1 and 3).

The nasal bones, as a rule, are grown together, while

their frontal >?orti.o-n is considerably longer than the remaining

part (Figure 11• ) .

The paired fontanelles in the region of the presphenoid

are in the form of narrow slits, and if they are wider-then

nevertheless they have a distinct tapering in front and behind.

iIMTI•:FATYPA

1 Hatt.lroa C., C.ilupHoa H. MaTepuant>I no cnc•reaa7111;e It reorpacilm[ecr,omy pac-npe,jeneHluo Phocidae cenepxoïi vacTn Tnxoro olceaaa. TpyAtii Bcec01o311oro uay^tn.-ncc:l. III]-Ta aiopcsot•O p1,l611. X03-l;a It o[tea1Iotpa4-un. T. 111, 1936.2 Yancxuit K. K. Onrr nepecaloTpa cucTeiltit it ,Zo-[artlocTIncu Tio.neueit 110;1CedIC31-

crua Phocinac. TpyAbl 3oo:rornvecsoro lui-Ta AI-1 CCCP, T. XVII, 1955.3 yancl:nO K. K. Coupeateatuoe cocTO>:uue If 11pOGae\ibl CIICTC\taTnlU( AaCT0110171IX.

T. corieruanuïi i-lxruo.nornvlccuoü soafliccnli Ali CCCP, nhiu. 12, 1961.

A- yaucr:uù K. K. Bonpo:.bt Taxcouotauvecxoi't crpvlerypbt cenleilcrua n cneTC hpa-niloltorntICCKIiX Ila11u61X. LICTBepTOe BCCC0103IIOe C013enta113ie qo n3Vtie[inlo NtopCliliX

DtTieüOllHTalolllHX. Te311Ch1 j(Oli.7a;1,013. N1., '1969.

5 Boettger S. Notizer zur Verbreitung und iiber Vcrwaridtscha-ftbezichungenctcr nlonchsrobbe (Alonachus albivcntcr I3ocidaert). Zool. Anz. -Leipzig, Bd. 147,Heft 11-12. 1951.6 Gray J. E. Notes on seals (Phocidaè) and the changes in the form of the

lower jaw durinh growth. Ann. k1a5. Nat. Hist. ser. 4.1869.

7 Gray J. E. Supplement to the catalogue of seals and whales in the BritishMuseum. London , Brit. Pylus. 1871.8 Gray J. E. I-land-list of seal'morses, sea-lions, and sea-bears in the British

h•luseuni. London. 1874.9 Eliervuan. and Morrison-Scott. Cheslist of Palaearctic and Indian mammals.

London. 1951.10 Frechkop. Ordre, des P;nnipecies. Traite de Zoologie vol. 17. 1955.

11 Kellogg R. Pinnipeds from Miocene and Pleist, deposits of California. Univ.Calif. Pablic. Gcol. SCi, vol. 13, no .1, 1922.

12 King J. E. The monk seàls (venus Alonachus). Bull. of the British Museum(Nat. 1-list) Zoology. vol. 3, no 5 L. 1956.

51 2

13 King J. E. Relationships of the Hooded and Ekphant se.als (genera Cye-

• tophora and Mirounga) J. of Zoology. Proe. Zoolog. Society of London, vol

.11 18. part 4: 385-398. 1956. •

Sciwf jer V. Seals, sea lions and wali uses. Stanford California. 1958.

1.5 Simpson G. 'Hie principles of classification and a classification of Main- •

Ines. Bull, of the Amer. Mus. of Nat. Ilist. vol . 85. New York. 1915.

10 Weber M. Die Sàugetiere. I3d. 2, Unterordnung: Carnivore Pinnipedia, Jena.

Winge H. Pettedyrsleo.gter. Kobenhavn. 1924 lb Winge il. The interrelationships of the mammalian genera. vol. 11. Koben-

havn. 1941.

REFEREN CES

1. Naumov S. and Smirnov N. Information on the systematics

. and geographical distribution of the Phocidae of the

northern part of the Pacific Ocean. Trudy VNIRO

(Transactions of the All-Union Research Institute of

Marine Fisheries and Oceanography), Vol. III, 1936.

2. Chapskii K. K. An attempt at a revision of the systema-

tics and diagnostics the seals of the subfamily

Phocinae. Trudy Zoologicheskogo instituta AN SSSR

(Transactions of the Zoological Institute of the USSR

Academy of Sciences), 'Vol. XVII, 1955.

3. Chapskii K. K. The contemporary state of and problems in

the systematics of the pinnipeds. Trudy soveshchanii

Ikhtiologicheskoi komissii AN SSSR (Transactions of

conferences of the Ichthyological Commission of the

USSR Academy of Sciences), issue 12, 1961.

F. Chapskii K. K. Questions of the'taxonomic structure of

the family in the light of craniological data. kth

All-Union Conference on the Study of Marine Mammals.

Reports of Proceedings, Moscow, 1969.

513

5. Boettger S. Notes on the propagation and relationships

of the monk seal (Monachus albiventer Boddaert)

10. Èrechkop. The order of the pinnipeds.

16. Weber M. The mammals. * Vol. 2, Ilborder: Carnivora

Pinnipedia. Jena, -1928.

17. Winge H. Mammalian genera. Copenhagen, 1924.

o

•

\ • 514

A. A. Antonyuk

UDC 599 .7115

QUANTITATIVE CHANGES IN THE VERTEBRAE IN THE 3 1 7

VERTEBRAL.COLUMN OF PINNIPEDS

'WO

In considerations of the post-cranial skeleton of main-

mais it is customary to determine the overall vertebral formu-

la. To an equal degree this applies also to the representa-

tives of the order of the pinnipeds. Thus, Flower (1885) and

Turner (1912), on the basis of an examination of about 46 in-

dividuals of various species of the true and eared seals, de-

rived similar formulas for the whole order of the pinnipeds,

namely: C 7, Th 15, L 5, S Cd 8-15 *. An exception was

presented by the walrus, in which these authors noted 14 ver-

tebrae in the thoracic section and 6 in the lumbar section.

Many years later this was still accepted as the vertebral for-

mula and this conclusion was confirmed by King (1956). Two

years later Scheffer (1958) introduced certain corrections,

as a result of which the formula tOok on the form: C 7, Th 15,

L 5, S 3-4, Cd 10-12, with no exceptions being made. Still

later King (1964) corrected this once again: C 7, Th 15, L 5,

S 3, Cs 10-12, an exception to which still remains the walrus

with its 14 thoracic and 6 lumbar vertebrae.

This small review of the literature shows that there

is no agreement in the opinions on the numerical values of the

* C cervical section (pars cervicalis), Th - thoracic section (pars thoracicalis), L - lumbar section (pars lumbaris), S - sacral section (pars sacralis), Gd - caudal section (pars cau-dalis).

51 5

vertebral formula. In the present paper an attempt is made

to investigate the causes of this and, as far as possible, to

clarify the actual situation concerning the vertebral formula

in the pinniped order. With this aim the author again ana-

lyzed the fairly extensive.material which is kept in the col-

lections of the Zoological Institute of the USSR Academy of

Sciences. There were examined two hundred vertebral columns

of almost all of the species of this grouP of animals, of

various sex and and age.

The results are presented in the table, from which

it is evident that, on the basis of the character of the ver-

tebral formulae, there may be established at least seven groups

for the pinnipeds. The first group, which embraces the largest

number of cases (66.5;:; of all of the examined individuals),-is

characterized by the formula: C 7, Th 15, L 5, S 4, Cd 8-16.

In this group are included representatives of all three fami-

lies. In the second, less numerous.group (19 of the examined

individuals), the vertebral formula is: C 7, Th 15, L 5, S 3,

Cd 8-15. In this group the walrus family is completely lacking,

but for the rest the composition is identical to that of the

preceding group. In the third group (6%),with a vertebral

formula of C 7, Th 14, L 6, S 4, Cd 11-12, there are included

in all only two*Udobenus rosmarus and Erignathus barbatus.

The fourth group, which is just as small (5%), includes Histrio-

phoca fasciata, Phoca hisnida lado7ensis and Eumetonias jubatus.

Its vertebral formula is C 7, Th 16, L k, S 4, Cd 8-16. The

* Translator's note. Sic. - Presumably . - "... two species •.."

516

T1G.ttttta

Konliecrno 1103nouhors no oTjtenant no3nonomtitay npeJlcraoure.ncii orpsl,ka nacTOUornx

p11,qu I

Sueciesn I C Th I L I S I Cd

Pt1OCIDA E.Phoca hispida 25 7 15 5 4 12--16

» • 4 7 15 5 3+ 13» 2 7 15 5 3 14-15» 1 7 15 G 4 -

Phoca hispida lado-gensis 5 7 16 4 4 14--16

Phoca sibirica 15 .7 15 5 4 13.-15Phoca caspica . 26 7 15 5 4 13-15Pagophoca ;;roenlsn-

dica 7 7 15 5 4 12-15» 1 7 15 5 4 13

1-listriophoca fasciata 2 7 15 5 4 12» 1 7 15 G 4 --» 2 7 15 . 5 3+ 11» 2 7 16 4 4 12

Phoca Vitulina 18 7 15 5 4 10-12Halichacrus t{rylsus 6 7 15 5 4 11-14 '

» 1 7 15 5 3;- 12Erignathus barbatus 5 7 15 5 4 12-13

» 4 7 14 6 4 11-12» 1 7 16 4 4 15

Cystophora cristata 4 7 15 5 4 10» 2 •1 15 5 3+. 12

» 1 '1 15 5 3 14Monuchus monachus 2 '1 15 5 4 10Leptonychotes wedde- 2 7 15 5 3 --

11t 1 7- ,15 5 4» 2 7 15 5 2 -» 2 7 14 G 3 12-14

Hydrurga leptonyxLobodon carcinopha- 3 7 15 5 3 12-13

ga 2 7 15 5 3 -Mirounga leonina 5 7 15 5 3 11-12Omtnatophoca rossil 2 7 15 5 2 12-13

»

OT.-IRIIDAB8 7 15 5 3+ 8-10

Callorhinus ursinus 9 7 15 5 4 8-92 7 16 4 3+ 9-10

Eumetopias jubatus83

77

1516

44

4,4

8-1412

» 1 7 15 5 3+ -7 7' 15 5 • 3-{- 10-11

Zalophus californianus 4 7 15 5 4 11

OD013G,VIDAEOdobenus râsmarus 7 7 14 6 4 12

» 1 . 7 15 5 4

318

Table

HO d':

CT(!;1

ptits;L:e ^TI)It

xci :;occ>^

chclL 5.IiO

LG,I-qI

ycI-i.nCtul.s`t iT1Sil.'üCiJn

(1'

3t=C1CItciii^s+

M.I'to"

I

The number of vertebrae in different

sections of the -vertebral -column

representatives of the pinniped order.

517

remaining vertebral formulae are scarcely characteristic of

the pinnipeds and each of them is represented by two speci-

mens of some different species, which comprises 1% of the

total number of examined vertebral columns. Thus in a

Weddell seal (Leptonychotes weddelli) the vertebral formula

had the form: C 7, Th 15, L 5, S 2, Cd 11. In a fur seal

(Callorhinus ursinus) there was found a somewhat different

formula: C 7, Th 16, L 4, S 3, Cd 9-10. Finally, the formula

C 7, Th 14, L 6, S 3, Cd 12-14 is charaCteristic of the leo-

pard seal (Hydrurga leutonyx).

In all of the vertebral columns of pinnipeds which

were examined by us and also by other investigators (Allen,

1880; Flower, 1885; King, 1956, 1964, 1969; Scheffer, 1958)

the cervical section was invariably represented by 7 vertebrae.

In the thoracic and lumbar sections there are counted, in to-

tal, 20 trunk vertebrae, which is characteristic also of the

representatives of the order of carnivores. Unly in two cases:

in one ringed seal (Phoca hispida) and one ribbon seal (His-

triouhoca fasciata) were there 21 trunk vertebrae. Such cases

may be considered as atypical deviations from the norm.

In the thoracic section 15 vertebrae were found in 87%

of the examined vertebral columns. This number, evidently,

should be considered as being the most typical for the pinni-

peds. However, as can be seen from the summary table, there

are encountered individuals with 14 and 16 thoracic vertebrae

(6.5% in each case). A thoracic section with 14 vertebrae,

as has already been mentioned above, is typical of the walrus.

The second species in which there are 114 thoracic vertebrae

518

is the bearded seal (Erignathus barbatus) and moreover this

species was also characterized by having 15 and even 16 tho-

racic vertebrae. Such a variation of the vertebrae under con-

sideration possibly conforms, to some extent, with the con-

cepts of certain authors (Winge, 1941; ChaPskii, 1955) on

this species as being little specialized and primitive. It

is interesting to note that in the Ladoga ringed seal (Phoca

hisuida ladogensis) the thoracic section is formed exclusively

of 16 vertebrae. This feature distinguishes this seal not

only from the soecies Phoca hisPida but also from the whole

of the genus Pusa and, possibly, may prove to be a taxonomie

feature. Perhaps the same applies also, though this is less

convincing because of the very small number of individuals

examined, to one other species of true seal - the leopard seal,

the thoracic section of which is composed of 14 vertebrae.

In the eared seals a variation in the number of thora-

oie vertebrae, from 15 to 16, is observed only in the fur seal

(Callorhinus ursinus) and Steller's sea lion (EumetoPias juba-

tus).

The lumbar section in the pinnipeds characteristically 320

has 5 vertebrae, though more rarely there may be encountered

4 or 6. Since, as has been mentioned above, the number of the

trunk vertebrae in the pinnipeds is constant, the frequency

of occurrence of such cases is precisely the same as the nulliber

of encounters of 14 and 16 thoracic vertebrae. In view of the

fact that these values are correlated, all of the arguments

rresented above on the variability in the number of the thora-

oie vertebrae will be valid also for the lumbar vertebrae.

519

As is evident from the literature references, each

author considers that for the oinnipeds that number of sacral

vertebrae is characteristic, which seems to be in accordance

with the results of his own analysis of.the vertebral formula.

In summing up, the authors agree that for the true and eared

seals the typical sacrum is composed of both 4 and 3 vertebrae.

If, however, one again turns to the summary table, then it

can be noted that thm± all of the enumerated investigators

were to some degree right, for in the sacrum there are found

four, three and even two vertebrae. Most often (79%) the

sacrum is comuosed of four fused vertebrae, which, apParently,

may be considered as bein9,- typical of all of the eared and

true seals of the Northern hemiSPhere. Sometimes, however,

in these seals there may be found a sacral bone which is formed

of three fused vertebrae. With a mire detailed examination

of such cases it turns out that the sacrum, even if only mor-

phologically, is nevertheless formed of four vertebrae. Two

facts lead one to this conclusion. Firstly, the caudal epi-

physis of the third sacral vertebra and the cranial euiPhysis,

in this case, of the first caudal vertebra are considerably

flattened, their edges have lost the rounded outlines (figure

1), in contrast to all of the epi -ohyses of the bodies of the

vertebrae in the other sections (Figure 2). Secondly, the

transverse Processes (procesus transversus) of the sacral ver-

tebrae, which here form longitudinal ridges (pars laterales),

in the normal four-vertebral sacrum zradually become narrower

and thinner from the first vertebra to the fourth, merginF

completely with the body of the fourth and last vertebra in

520

Pm. 1. Kayanabilan TpexrIo3B0111:0130r0 kpe- . emit napril ii nepnbtri knocToeoii 110300110K (BIM. C

nekTpambuofi cTopakm, nopuko- Bbie komepa i10 -2B0IIK013 B kpecTu.e)

Figure 1. Caudal portion of three-vertebral sacrum of a

harbour seal and first caudal vertebra (view

from ventral side, the numerals indicate the

ordinal , number of the.vertebrae in the sacrum).

Pue. 2. 4e-rupexao3nokkoublii «TiopmaabkhuI» KpecTeu iaprui (aka c ecin-paibuoii cTopoubl)

Figure 2. Four-vertebral "normal" sacrum of a harbour

seal (view from ventral side). •

its caudal portion (see Figure 1*). In the case, however, of

the three-vertebral sacrum these ridges, on the contrary,

become slightly wider and thicker on the body of the third

di? vertebra, which is clearly not "ended", and only on the first

caudal vertebra do they become thinner and merge with its

body (see 2igure 2*). Such a "morphological incompleteness"

* Translator's note. Sic. Text references to figures obviously

5- 21

i

•n

of some of the three--ve.rtebral sacra considerably reduces,

if it does not practically stop, the freedom of movement of

the J_ast sacral and first caudal vertebrae relative to one

another (such cases are marked in the table by the + sign).

Consequently, the thought arises that such sacra function

like the four-vertebral sacra. Thus, the number of individuals

for which a sacrum composed of four vertebrae is typical be-

comes even 9-:reate-r. There are also encountered cases of a

clearly normal sacrum that is formed out of 'three or even two

vertebrae, representing a morphologi cal ly single sacral bone.

Such a sacrum is bas i ca:lly characteristic of all of the Imtarc-

tic seals, in which there is observed a tendency towards a

reduction in the nurnber of the constituent elements of the

sacrum. In -cne examined series there were also noted two cases,

xkxra* in one individual of the Caspian seal (Phoca caspica)

and one harbour seal (Phoca vitulina largha), in which the

sacrum was composed of five fused vertebrae. This phenomenon

represents a clear deviation from the norm and may be consi-

dered as ateratism s in both cases the first sacral vertebra

was separated from all of the others, while the junctions of

the last sacral vertebra with the first caudal vertebra bear

traces of a distinct ankylosis.

In the caudal section it is generally impossible to

establish any regular pattern of the variation in the number

of vertebrae. in one and the same species the number of ver- 322

tebrae may vary from ô to 12-16. Therefore it is scarcely

possible to utilize the number of vertebrae in this section

as a taxonomic feature of the species, subspecies or popula-

tion rank, at least within the order of the pinnipeds (Yablokov,

522

4i

1963). The only feature which may be noted is the abbrevi-

ated nature of the tail of the eared seals s the tail in

these is formed of a smaller number of vertebrae (from 8 to

].Ii•) than in the true seals, in which the caudal section is

composed of 10 - 16 vertebrae.

Summing up all of the information ,oresented above, the

general vertebral formula of the pinnipeds may be presented

in the following form o C 7, Th 1l1•-16, L 4•--6, S 2--1I•, Cd 8-16.

JI I•ITIà PATS' P A

I ya!u.cai!ï K. K. Onbrr nepecasarpa cisc.reta!,t nRnarnoc•ru!cti Tsoneneit ncsacensers-

c•r!!a Phocinae. TpynL! 3oonor!!vecror(, nneT!rryra AH CCCP, T. XVI1, 1055.2 Sld.ior:oa A. 13. i•ferozopLae npez!iap!rre.^t.!n,!e MaTepna.•!!,! K >sopcpo.nrm!ec!coï!

xapa!nrpt!c•rn!<c rhe!!;!a!!:1c!coro •r!o.aenst lie:!oro n l'pc+nna!tilcxoro mopeil. Ccï•nayt!.-ncc.n. pc16oT CerfllIllPO, Apza!n•c.i!,c!c.'1963.

Allen J. A. 1-iislmy North American Pitinilieds. Washinhton. 1880.^. Flotc^e; IV. 1/. Ost^^oln^y of the mai^^ur!alia. London. I885.

Kinr; J. E. The nsonk seals (hcnus dlonachns). 13u11. Brit. Mus. (Nat.I tit;, Zool. 3: 201--956, M56.

Ring J. G. Seals of the world. London. 196,1.King J. E. Some aspects of the n!!a;tomy 'of the Ross seal, Ommotophoca

r^si (Pinnihedia: Phocidae). British Antarctic Survey, Scientific`reporls, no, 63.London. 1969.8 Scüc(f.er 17. B. Seals, sea lions and walruses. London. 1958.C^ Turner W. 'flic marine mammals in the anatomical rnuscum of the Univer-

s(tv of Hdinburf*. London. 1912.10 ü%in{;r. II. The interrelatiouships of the mammalian l;enera. Vol. 11, Koben-havn, 1Jfl.

REFL,RE:IdCES

1. Chapskii K. K. An attempt at a revision of the systema-

-tics and diagnostics of the seals of the subfamily

Phocinae. Trudy Zoologicheskogo instituta AN SSSR

(Transactions of the Zoolo^ical Institute of the USSR

Academy of Sciences), vol. XVI1:, 1955-

523

2. Yablokov A. Y. Some preliminary data on the morpholo-

gical characteristics of the harp seal of the White

and Greenlad Seas. Sbornik nauchno-issledovatel'skikh

rabot SevPINR0 Uorthern Polar Research and Planning

Institute of Larine Fisheries and ceanogra.)hy),

Archangel, 1963.

524 .

UDC 599 ,7/15 .3

V. A. Zheglov and K. K. Chapskii

TEST OF AN AERIAL SURVEY OF THE RINGED SEAL AND THE GREY SEAL

AND OF THEIR HOLES IN THE GULFS OF THE BALTIC SEA

AND ON IAKE IADOGA

Over the course of many years in the northern and

eastern part of the Baltic the ringed seal (Pusa hisoida gm*)

and the grey seal (Halichoerus grypus F.) have served as

objects of the marine sealing industry; these seals have

been caught here at all periods of the year and without any

limitations. «Furthermore, in Sweden and Finland a bounty is

paid for catching the seals (Curry-Lindahl, 1965).

Every year there are caught by these countries in the

coastal waters about 15-16 thousand seals (Ropelewski, 1952;

Bergman, 1958).

it is natural that such destructive sealing has af-

fected the stocks of the seals.

In the territorial waters of the Soviet Union (in the

Gulf of Riga and the Gulf of Finland) the sealing is carried

out by sealers from fishery kolkhozs in Estonia and Latvia,

the annual catch of which does not exceed 1200 - 1500 head

(according to official data from the Estrybakkolkhozsoyuz**,

in which each 100 luy of oroduction from sealing ooerations is

considered as equivalent to 3 seals).

711rom 1970 a limit has been placed on the catch of Bal-

tic seals in the USSR.

Translator's notes. * Sic. Schreber is the species author. ** Acronym for Estonian fishery collectives union.

525

o

o

Up to the present time the representations of the

numbers of seals in the Baltic remain very indefinite. More-

over, there has been noted a marked reduction in their popu-

lation numbers.

Thus, according to Bergman (1958), "the stocks of the

ringed seal in the coastal waters of Finland have decreased

considerably from the twenties and thirties up to the late

fifties of this century. In the southern part of the Gulf of

Bothnia they are oarticularly small, in the middle part of the

Gulf of Finland and in the central portion of the Gulf of

Bothnia (Salkameri and Melenkuru) they are rather small, in

the shallow waters off the islands of Valgrund and Berge they

are not more than average, and in the eastern part of the

Gulf of Finland they are average, at the level of the 1920s and

1930s. In comparison with the 30s, even in the northern part

of the Gulf of Bothnia the population numbers of the grey seal

have decreased considerably. In the western part of the Gulf

of Finland there were considerably fewer grey seals in 194(7)-

1950 than there were in 1920; there were a few more of these

in the eastern part of the Gulf of Finland. The total number

of grey seals within their territorial waters was, on average,

considerably smaller than the number of ringed seals. Their

ratio, according to Gotberg's data, comprises 1 :3".

According to the tentative estimate of Sundstrem (1962),

the Baltic population of the grey seal at the end of the fif-

ties was about 10 thousand head. A marked lowering of its

population numbers was pointed out in the middle sixties al-

ready by Curry-Lindahl (1965). According to Smith (1966), the

stock of grey seals in the Baltic comprises 5 thousand head.

52 6

The only approximate indication of the population num-

bers of the Baltic ringed seal was found bills in a Finnish

fishermen's journal ("Kalamichen Viesti", no. 1, 1966), where

it was stated that each year after the sealing operations in

Finnish waters there remain about 20 thousand rineed seals.

There are very few operational data of any kind, on the basis

of which judgements can be made on the population numbers of

the two species of seals in the Gulf of Finland and the Gulf

of Riga.

It is even more difficult to judge the population

numbers of the Ladoga ringed seal, of which about 500 head

are cauet each year by hunters and fishermen.

K. K. Chapskii (1932) assumed the possibility that

about 20 thousand seals could be living in Lake Ladoga; in

the opinion of S. P.. Sokorin (1958) the population numbers of

the Ladoga seal were nor high, while A. I. Zubov (1965), re-

ferring to data from the State Research Institute of Lake

Fisheries (GWnIL.RKh), estimates the stocks of the Ladoga

seal at 5 - 10 thousand head.

In connection with the growinz demand of the sealing

industry there has come to a head the necessity for reorga-

nizing this on a scientific basis. A most important condition

for the rational utilization of its resources should be the

knowledrre of the population numbers and other questions re-

lated to this.

2rom 1969 the AtlantnIR() (Atlantic Research institute

of i:.arine Fisheries and ceano,9raphy) has been conductinE work

on the study of the seals in the eastern coastal region of the

527

Baltic Sea and Lake Ladoga, and therefore in arch-April of

1970 there was made the first attempt at an aerial survey of

these seals by workers of the Atlant .HIRu.

MATERIALS AND METHODS

As far back as the forties Dorofeev (1940) wrote about

the advisability of employing aviation for surveying marine

mammals.

af

•

In recent years Tikhomirov (1966), Fedoseev (1966),

Shustov (1969) and other investigators have conducted aerial

surveys and aero-visual counts of seals in the Far East.

In the Gulf of Riga and Gulf of Finland and on Lake

Ladoga no.special aerial surveys of seals were conducted up

to this time.

Conducting an aerial survey of the ringed seal . and the

grey seal in *these regions was accompanied by considerable

difficulties. UD to the present time there has been no metho-

dology worked out fàr the counting of these seals in the zone

of the landfast ice, although we know of the attemnt by Sviri-

dov (1954) and by the far-eastern investigators, who conducted

aero-visual counts of the ringed seal and other seals on broken

drift ice. At the same time, it is known that the ringed seal,

as a rule, does not form concentrated whelping and moulting

patches, similar to the patches of the harp seal, and therefore

we did not employ continuous aerial photo=aphy.

In elaborating the methodolopy for the aerial survey,

we made use of information obtained by Questioning local sealers

and the experience acquired in counting seals by the far-eastern

investigators and other authors.

a M + m C. V.

45 3

Table 1.

Some indices of the periods of retention of tags

in the digestive tract of the experimental animals,

hours.

Ta6auua 1

He KoTopme nourtaaTeas spemess aa,sepiass meTok s nnut,esapinemmom Tpatne no;tonbusbtx :seirtoTtmx, trac

miiTepu la Material

11.TacTmaconb1e MCTIZII 73,5±16,4 73,5 100 i'lastic tags KvottiYieceS oi rubber

59,0 25 ,cm stmatibt 22,64.2,7 13,4

111e,utut Kycotput macTmaccht 14,9±0,8 3,4 22,7 20 Small pieces of

plastic * nPume ■mum.. B CLIN3B C TCM, 4T0 cymcr Beim ,tx pa3.111411rf B 0;1licam1l!bn:

me-rot: cpc.au >1:1moTm ■ IX IIC TZlem Hue 11 imucaeti b! oGoCim,esume ;mumble.

*Note. In view of the fact that significant dif-ferences were not found in the retention times of tags of the same type. between the animals, generalized data are presented in the table.

The retention of the tags and, probably, Of the undi-

gested remains of the food also depends on the amount of food

consumed by the animals and on the characteristics of the

working of the digestive organs. Thus, the plastic platelets

that were given to Ushkan before a prolonged fast were found

after 55 days, while with Vega (in a similar case) they were

found after 20 days. In contrast, when Vega was fed abundantly

during the first third of February (the daily ration exceeded

4 kg) the tags often appeared in the tank 4 - 5 hours after

the ingestion of the food.

However it should be noted that the small pieces of

plastic, just like the otoliths, did not pass through the

gastro-intestinal tract simultaneously: on the average, about

20 t =3,1 AB

t =3,5

2,7 BB

454

80:, of these appeared after 10 hours, about 15 approximately

after one day, and 5 - two days after being given to the

animals. If the working of the digestive system of the ani-

mais under the natural conditions does not differ substantial-

ly from that in the test conditions, it may be assumed that

the otoliths, collected from the intestines of caught animals,

must belong to fishes that were caught by the seals within a

period of 2 - 3 days, although the main mass of the consumed

fishes had been digested less than half a day previously.

In spite of the fact that the animals were held in

small tanks, i.e. in almost immobile conditions, the assimila-

tion rate of the food was found to be very high. The faeces,

which are composed to a considerable degree of undigested or 282

poorly digested remains of fishes (bones, otoliths, crystalline

lenses of the eyes etc.), under a moderate feeding regime

(March - May) on the average had a weight of about 4 - 5% of

the value of the daily ration*. There were found differences

among the animals in the weights of the faeces, expressed as

percentages of the daily ration, though statistically (with

P 0.0) these were insignificant.

Because of an insufficiency of food, the animals did

not always obtain the necessary amount of fish. Moreover,

from October 10th 1967 till January 9th 1968 Vega did not feed

at ail, while Ushkan did not feed at all from January 13th

until March 3rd 1968. During 60 days of fasting Vegas weiet

fell by 10.6 kg (from 38.8 to 28.2 kg), by 29.7% of the initial

weight. (ver 50 days of an enforced fast the weight of Ushkan

dropped from 40 to 29.45 kg or by 26.4%, i.e. he lost 10.55 kg.

The mean daily weight losses comprised: 177 g for Vega and 211 g for Ushkan.

* Urinary wastes were not taken into account. -

455

o The two month fast of the seals in the described con-

ditions should not be considered as the limiting value.

Over 222 days (including the period of fasting) Ushkan

consumed 301.4 kg of fish, i.e. on the average he ate 1358 g

per day. From the initial (39 kg) to the last weighing he lost

3.6 kg or 9.8% of his initial weight. Over 154 days (also in-

cludine the period of fasting) Vega consumed 191.5 kg of fish,

i.e. on the average 1244 g per day. Her weight dropped by

7.95 kg (from 38.8 to 30.85 kg), i.e. by 20.5%.

Ilianyunya obtained food in some amount or other over

the course of almost the whole of the holding period. Never-

theless, the lack of food during the winter period also affected

her. While initially her weight was 65.8 kg, after 184 days

this was equal to 61.9 kg, i.e. she lost 3.9 kg or 6.3% of

the initial weight.

The animals reacted very sensitively to changes in the

size of the ration by changes in their own weight (Figure 1).

With a low daily norm over several days, the weight of the

animals began to fall, while with an increase in the norm the

weight began to increase. At the same time, in the changes

of the body weight with respect to the amount of food consumed

there was observed a certain asynchrony, a semi-antiphase re-

lationship of the maxima and minima. It is quite possible

that this time lag in the changes in weight in relation to the

size of the ration was caused by a differing assimilability

of the food.

In the conditions of the experiment, i.e. with the

markedly restricted movements of the animals, it was found to

be possible to approach the question of estimating the daily

66

621

56 4

2

CIII1 0

pa 401 • 1 36

3 '1 I 28i

L__

456

361

Pic. 1. 1,13menemie peca Teaa ileput MMICIMOCU IDT'IleM1 , 11111U

Cr(VMOrOpMWOlia

I — BCC Tesla (Ke); II— crrognmil palm. on (K2):

a — Dora, 6— Mainoim, o — YulKalt_

Figure 1. The change in body weight of the seals in relation to the size of the daily ration.

- body weight (kg); II - daily ration (kg);

a - Vega, b - lanyunya, v - Ushkan.

ration which, in energetic respects, is close to the value of

the standard (basal) metabolism. With this aim, in March -

May 1968, when the animals were provided with food to an

adequate degree, we attempted to select for each animal just

such a daily ration, i.e. to establish such a moderate feeding 283

regime in which the weight of the seals would remain more or

less stable (Table 2).

The animals were primarily fed with the yellowfin Bai- 284

kal sculpin*. Both species of Baikal oil-fish**, the longfin

Baikal sculpin*** and also the benthic gobies were given in

the form of a small supplement. Even under such conditions,

Translator's notes. * Cottocomephorus comephoroides or C. grewingki ** the "big golomanka" Comephorus baicalensis and the "little. golomanka" - C. dybowskii. *** cottocomeohorus inermis

11•5 .7

Table 2.

Some indices of the daily ration with a stabilized

weight of the experimental animals.

Ta6:inua 2

1-1eNororu:e noha3aTen11 cyTO11110r0 pacqirona npn craGn:11+3nponaHnont eecenoAont,lirthlx xcunoTnblx

[lapaM1terpt,t 26. 111-30. IV 21.11==2.1V

yacupclteicnttü s ï n e^aii^.rF,'^lrE'•ters J'mt<an Ilera

-of distribution Bec eC Body

34,29±0,10

34,0-34,70,24

0,7

6

33,13±0,09

33,2-33,8

0,23

0,7

7

M^E m

lim

IfC. V.

n

^teee c^ô Nozoo par;da^^^j^ ration,

2097-!:49,1

1900-2500282

13,4

36

1872:L 84,2

150-3000

533

28,4

41

12. 11t-14.V

r:dnvûnyaD1autbust

kg.

59,54 ±0,48

59,3-61,8

1,45

2.4

9

g

2559-54,91000-4500

429

16,7

63

when the energy losses for all possible movements were reduced

to a minimum, the animals required a fairly large quantity of

food. In percentages of the body weight the rations compr.ised :

for A:anyunya - 4.2fo, for Vega - 5.6% and for Ushkan - 6.1/.

There is no doubt that under natural conditions the ration is

at least 1.5 times greater than in the conditions of the test.

Just like with other mammals, on a per unit body weight

higherbasis the young animals required a.inaut of energy accumulated

in the food than did the adults.

The food coefficient changes with the age of the ani-

mals. ;t1hile for a body weight gain of 1 kg Vega required 23 kg

of yellowfin i3aikal sculpin, and Ushkan required 26 kg, the

adult female had to consume twice as much of this same fish or

511• kg .

weight,

45 8

The maximal uptake of food at one time comprised 2 kg.

Generally, however, the animals consumed not more than 1 kg

in one meal (within 1 - 5 minutes). When fed "ad libitum"

Manyunya consumed not more than 5.6 kg of pelagic gobies in a

day, Ushkan - 5.3 kg, and Vega - 4.5 kg.

In spite of the fact that the food was usually given

during the day (once every 2 )-i hours), the main mass of the

food was consumed in the evening (creuscular) period.

In March - April, during the period of the optimal

feeding regime of the animals, there were set up experiments

on the digestibility of the otoliths of the big and little

golomankas and also of the yellowfin Baikal sculpin, i.e. the

otoliths of those fishes which comprise the basis of the feed-

ing of the seal under natural conditions. The otoliths were

removed from freshly caught fish, measured under a binocular

along their long and short axes with a precision of to 0.05 mm,

fed to the animals, and later these same otoliths were picked

out of the faeces of the seals and again measured. Subsequent

comparison showed that the average values of the dimensions

of the otoliths of the control and experimental series differed

from one another at a high level of statistical significance

(Table 3). On average, for the three seals, the length of the

otoliths of the big golomanka decreased by 5% after passage

through the gastro-intestinal tract, of the little golomanka -

by 3.4% and of the yellowfin Baikal sculpin - by 4.3%; the

widths decreased, respectively, by 6.9, 4.2 and 5.7% from the

initial dimensions.

M + m C C. V. tAB

A

A

A

A , 13

A

A

11,7

11,5

459

Table 3. Changes in the dimensions of the otoliths of fishes with their passage through the gastro-intestinal tract of the experimental animals, mm.

T a 6 a It us a 3

Ilanicaeuan paudepoa oToasyron pb16' lipli i< ripoxo.:«aellau . . nepea ›Keay,r(oliao-icitaiegablil Tpaar noaonbrrahlx. ›auncrrumx, iLSt

--01011Lhs Parameters—of the di7st-1-1-bution Ilapameind pactipulenetinii

1 Inweimtm c'elm01

chal_g_eL.,g9r ies*

Bo/Lamm coAcmuitea Big golomanka

,annuaL

InpaaaW

RmtnaL

L

• IlInpunziVi

2,246:120,0095 0,288 12.8 919 „

. 2,134±0,0105 0,293 13,7 774

1,799+0,0005 0,197 10,9 919 ,,„ ek

1,675± 0,0077 0,215 12,8 774 '`"-'

MaAaa cOACMIJIlea Little golomanka

1,677:3-0,0036 0,161 9,6 9090 n i

1,620±0,0052 0,179 11,5 1237 ̀.'"

1,279i:0,0031 0,140 10,9 2090

1,224.±0,0015 0,151 12,6 1937 10,1 . . .

XCJITOlepbtAblit 6blitOle. Ye 1101f scu pin

1,883±0,0053 0,145 7,7 748

1,803:120,0044 0,177 9,8 1599

1,096±0,0025 0,069 (3,3 748 1,034:120,0043 › 0,191 18,8 1599

e A — J&O onma, 13 — ontaTa.

* A - before experiment, B - after experiment.

L - length W - width

The degree of digestion of the otoliths was found not

to be uniform in each of the three seals. The otoliths were

found to be somewhat more strongly digested in Manyunya, more

weakly in Vega. lt may be assumed that the intensity of the

digestion of the otoliths is under the control of several fac-

tors, being dependent on the time spent in the gastro-intesti-

nal tract, on the age of the animals, their nutritional state,

motility etc. It is a curious fact that in each of the animals

the otoliths of the big golomanka were most strongly digested, • 285

4•60

less so - of the yellowfin Baikal sculpin, and still less -

of the little golomanka. In this, one cannot but perceive a

species si)ecificity of the otoliths themselves. It is also

interesting that the otoliths decreased in size somewhat more

markedly along their short axis than along the long axis.

Apparently, the digestion of the otoliths depends, among other

factors, also on the surface that comes into.contact with the

gastric juice.

The digestibility of the otoliths, which on the whole

is very similar in the golomankas and in the yellowfin Baikal

sculpin, should nevertheless be evaluated as being insignifi-

cant. The otoliths which.have been in the digestive system

of the seals do not differ superficially in any way from those

which were removed from the fishes. By visual evaluation there

are also no differences in their optical structure: the annual

rings are distinguishable with the same degree of definition

as in the original otoliths.

Thus, it may be stated that the study of the feeding

of the seal on the basis of the otoliths is well-founded.

Lending itself to reconstruction by this method is not o•nly'

the species spectrum of the food but also the size, weight

and age of the fishes consumed by the seals. This last circum-.

stance is of considerable significance for acquiring an under-

standing of the inter-relationships which are established

between the representatives of the two final trophic levels

in the pelagial of Lake Baikal: the golomankas and goby fishes

and the seals.

UDC 599.53r

4.6 1

V. Nt. Hel'kovich and^V. S. Gurevich

QUESTIONS RELATING TO THE CATCHING AND LONG-TERM: HOLDJ:NG

OF DOLPHINS IN CAPTIVITY

The representatives of the suborder of the toothed

cetaceans, the dolphins, attract investigators of the most

diverse specialities. This is connected with the study of the

profound adaptive changes, brought about by the habitat, and

several biological characteristics which are of considerable

interest. Among these, primarily, belong: the methods of

orientation and navigation.in the ocean; the characteristics

of the hydrodynamics; the diving to great depths and for pro-

longed periods; the characteristics of the behaviour and

higher nervous activity.

The necessity for conducting experimental studies to

resolve these problems has forced us to seek ways of trans-

forming dolphins into experimental animals.

In this paper an attempt is made to generalize the

information available to us on the catching, transportation

and medical servicing of experimental dolphins abroad, and

there are presented some of our own studies from the period

1965 - 1969.

THE CATCH

There are no special scientific reports devoted to the

methods and procedure for catching and transporting live dol-

phins. The details of the methods employed are professional

286

4.6 2

secrets of the firms which are engaged in -providing the an!-

mais for aquaria and oceanaria (i'orris, 1966). However, the

scattered framentary data in scientific ouplications.of re

' cent years and our own experience permit us to present a ge-

neral picture. Catching the dolphins may be subdivided into

the catching of individual animals and the catching of groups

of animals.

Individual catching permits one to select the required

specimen from a school and decreases the mortality. A pioneer

of this method was M. Santini, who supplied dolphins to the

Florida "Marineland". In addition to him, such catching is

practiced by F. Brokato, the captain of the ship "Geronimo",

who supplies animais to this same oceanarium, and by Jaques

Ives Cousteau.

Generally the catchers on fast-moving vessels follow

the school until they are able to isolate and catch the marked

out dolphin by means of a throw net or a Special catching de-

vice. M. Santini jumps onto the dolphin in the water and holds

it until the launch approaches.

The catching of groups of dolphins is carried out, as 287

a rule, with the help of nets. In addition, with this method

there are employed vessels or motor boats as beaters, aerial

reconnaissance for finding accumulations of the animals and

also means of transporataion for moving the captured dolphins

to their destination. Such methods of catching were utilized

by K. Rei (1963) for capturing belugas, and by D. Gascin (1966)

for catching bottlenose dolphins.

1 1 ,63

PliC. 1. Re.111.41111b! B • aaomane. CdBlep Bp01131100,1iT no,acynnq

CCT11, a no,nnn pacTnrnnaior ce, trro61,1 ne c.no);;n:m TegenueNt

*Figure 1. Dolphins in "aloman" (purse seine). The seiner takes up the net, while the boats stretch it out so that it is not furled by the current.

R. Busnel (1966) worked out a special method for

catching dolphins with the aid of a powerful acoustic signal

which temporarily stuns the animal.

The existing method for catching dolphins in the Black

Sea was designed for catching groups of animals (Figure 1).

In spite of the fact that it is considered as a commercial

method, the sweep net catch is acknowledged as being the most

effective method up to the present time. The details and tech-

nique of the commercial fishing have been described in detail

in our literature by Tatarinov, Eremaev, Larti, Freiman,

Kleinenberg, Gubenko and others (according to Kleinenberg S. E.

et al., 1964.), and therefore we will not stop to consider this

in detail here.

When we commenced the experimental work with the dol-

phins, utilizing accumulated experience, we somewhat modified

464

the commercial purse seine fishing method, which turned out

to be fully justified. Thus, out of the recently caught dol-

phins (white-sided and bottlenose dolphins) only 24% of the

animals died at the time of the catch. Protective measures

were employed during the catching operations: care was taken

that the current would not furl the purse seine, while during

the hauling in of the net by the boats and the SChS*the net

was kept stretched; during the approach of the transport

vessel to the side of the SChS for the trans-shipping of the

animals between these there were set up spacing struts.

The captured animals were Pulled out of the water with 288

• the aid of stretchers, inflatable rubber mattresses, by suspen-

sion with a soft sling by the tail or simply by hand, depen-

ding on the species and weight of the animal. The main factor

in this operation is speed. The more rapidly the animals are

brought onto deck from the drawn up seine net, the lesser will

be the traumas and the less likely they are to get entangled

in the nets and to choke.

With the contemporary nethods of catching groups of

small dolphins (the common dolphin and the common porpoise)

the best reeults are obtained by the use of a fine-mesh kapron**

Purse seine net (of the scad purse seine type), since with this

the losses of the animals are reduced.

The best method for trans-shipping is the use of a

stretcher with a soft covering and an opening for the Pectoral

fins. The tans-shipping of the small species of dolphins

Translator's notes. * SChS Black Sea semer

** kapron - a synthetic fibre.

465

may be successfully carried out by hand; in transferring large

specimens, as an extreme measure, suspension by the tail in a

soft sling may be tolerated.

TRANSPORTATION

The transportation of dolphins at the Present time is

accomplished by various methods: by ships, airplanes and motor

vehicles.

Transportation of dolphins by sea was first accomplished

in the seventies of the last century, when two belugas were

brought to the Brighton Aquarium in England from Labrador and

Newfoundland in the hold of a ship. Both animals died (Slijper,

1962). In 1907 the New York Aquarium contained in its pools

a group of bottlenose dolphins, while in 1912 in transporting

a new batch of dolphins to this aquarium Taylor first employed

soft litters for their transportation. In recent years the

technique of transporting dolphins by sea has been considerably

improved, which permits them to be transported over fairly

great distances. However, the most successful transportation

is accomplished to those aquaria which are located on the

coast and may receive the dolphins 1 - 2 hours after they are

caueht.

Successful transportations by sea in recent years have

been accomplished by: Bel'kovich (1964), Kritzler (1949), Ray

(1963), Andersen and Dziedzic (1964) and Gascin (1966).

The curator of the ilapier oceanarium in New Zealand

considers that the transportation of dolphins by airplane, in

particular of the bottlenose dolphin, over considerable distances

4.66

in air without water over the course of a period of more than

two days apparently does not cause them any harm (Gascin, 1966).

In 1960 D. Lilly transported two bottlenose dolPhins

by airplane. The dolphins were placed in a special box of

perspex and Plywood, 2.8 x 0.6 m in size. Within the box, at

a distance of 17.5 cm from one another, there were installed

plywood partition-frames, strengthened with perspex. From the

town of St. Augustine (Florida) to the island of St. Thomas 289

the animais were conveyed aboard a cargo plane, the flight of

which was at an altitude of 2000 m and lasted 7 hours. During

the course of the flight the respiratory rhythm of the animals

was measured. The altitude does not have any effect on the

frequency of the respiration, if the dolphins are in water

"in a suspended state". K. Rei (1963) transported three belugas

from the south-western coast of Alaska to the New York ocean-

arium. The animais made the 5900 km journey on stretchers,

boats, a fishing vessel and then on a special B-17 airplane.

The animais were Placed in wooden boxes, partially filled with

water, the Walls and bottom of which were covered with a soft

plastic lining. All three belugas withstood well the 24-hour

non-stop flight. However, immediately after landing at New

York airport one of the belugas died.

Dudok van Heel (1966) successfully transuorted by air-

plane four bottlenose dolphins from the USA to Holland. He

indicated that for the successful transportation by air the

airplane should fly at an altitude of 1500 m. To protect them

from contusions, the animais were suspended in hammocks. The

flight lasted 16 hours and ended successfully.

467

In 1969 there was carried out a successful transporta-

tion of two male common porpoises aboard an IL-14 airplane from

the Crimea to rix)scow; the animals were delivered in a satis-

factory state (Sokolov et al., 1969).

Transportation by motor vehicle is employed mainly

over short distances: from the shore to the oceanarium or

from the oceanarium to the airport. The transportation is

accomplished in open vehicles, employing soft linings and maintain-

ing protective measures to prevent the action of the direct

rays of the sun on the integument of the animals.

It is difficult to say who first employed a truck for

the transportation of dolphins, but as early as 1948 Kritzler

transported two young male pilot whales (Globicephalus macro-

rhynchus), which had been stranded ashore in Florida to Farine-

land.

D. Lilly (1960) delivered by truck, from Marineland

to the airport in St. Augustine, two bottlenose dolphins over

a distance of approimately 30 km. W. Kellog (1966) conveyed

an adult female bottlenose dolphin to his laboratory in an open

truck. Among other investigators, who have used used the me-

thod of conveyina dolphins to the laboratory by motor vehicles,

should be mentioned Schevill and Lawrence (1956), M. Caldwell

and D. Caldwell (1963), Ray (1963), Andersen and Dziedzic

(1964), Heel (1966) and Sokolov (1969).

For the transportation of captured animals we utilized

a refrigerator ship (TKI1S), and moreover we employed several

variants, which may conventionally be divided into two methods:

"semi-aquatic" and "aquatic".

4, 68

The basic factor, necessary for the successful trans- 290

portation of dolphins, is the establishment of optimal condi-

tions for thermoregulation. A prolonged sojourn of the dol-

phins in air, especially under the action of the direct rays

of the sun, leads to a fairly rapid death of the animals. This

is explained not only by the low thermal conductivity of the

air, which causes overheating and heat stroke during the summer

period in literally a few hours, but also by a disturbance in

the activity of the cardio-vascular and respiratory systems.

The latter occurs as a consequence of the infringement of the

"practical weightlessness" on dry land and the compression of

the thorax by the weight of the animal, and also from the

marked shrivelling of the skin,, wrinkling and flaking-off of

fl the euidermis.

11, Thus, the transportation of dolphins out of water brings

to the forefront several requirements, on the correct fulfilment

of which deuends the success of the entire transportation. The

animals must be protected from the action of the direct rays

of the sun, care being taken that the skin is kept moist all

of the time; in addition, to avoid the formation of bed sores,

the dolphins should be placed on thick, fairly soft linings,

which may to some extent compensate the temporary loss of the

state of "suspension" by increasing the area of support of the

body. For this on the deck, shaded by awnings from the rays

of the sun, there is set up a soft bedding; of inflatable

rubber mattresses, boats or thick porolon*. Un these the ani-

mais are Placed individually. A sufficiently thick and soft

beddinP: will somewhat decrease the normal loads in the region

of the thorax and abdomen.

* Translator's note. "Porolon" - presumably an aerated plastic of the styrofoam type.

469

Pnc. 2. Reamjnin B 6opro13or1 II a mi e jia ramane. Ha crinne muna mapaeBan nanne,na

Figure 2. Dolphin on a hammock in a tub aboard ship.

The gauze coverlet is evident on the back of the animal.

The second method of transportation is the "aquatic"

method. For,this there is utilized a tub aboard the trans-

porting vessel, with running water, or special containers, in

which the dolphins are secured in hammocks (Figure 2). The

securing is absolutely necessary,even when there is only a

slight rolling and pitching, in order to minimize the trauma-

tism in general. In addition to this, the individual secure-

ment prevents the possibility of the occurrence of a panic

among the dolphins, during which they fight, ripping the skin

and inflicting serious injury to one another with their power-

ful tails. The securement also prevents, and this is probably

the most important factor, the entry of water into the blowhole.

In one standard shipboard tub, 180 x 250 cm in size,

there are accomodated, depending on their length, 1 - 3 bottle-

nose dolphins or 5 - 6 white-sided dolphins. The containers

are for individuals.

47 0

The elaborated method for transPorting the animals

on the special soft bedding and in the shipboard tubs on the

refrigerator ships (TKhS) were found to be suitable for main-

taining the health of the dolphins for 54 hours and for their

subsequent prolonged holding in captivity.

MEDICAL SERVICING

The problem of the medical servicing of dolphins, kept

in captivity, has attracted the attention of many investigators

(Brown, 1962; Brown and Norris, 1956; Brown and McInture,

1960; Ridgway, 1965; Ridgway and McCormick, 1967; Nagel

et al., 1964, and others).

Dolphins that are kept in captivity most frequently

suffer from skin diseases, which cause real distress and lad,

as a rule, to a lethal outcome (Russel, 1966). In addition

there have been found in them: erysipelatous inflammations,

stomach ulcers, jaundice, insults and infarcts of the myocar-

dium, diabetes, scurvy, inflammation of the lungs, and numerous

external and cavitary parasites (Ridgeway, 1955). One of the

most serious problems is the swallowing by the dolphins of

extraneous objects that fall into the Pools (balls, gloves,

toys). In recent years (Ridgeway, 1955, 1967; Nagel et al.,

1964) there have been elaborated methods for the catherization

and venous puncture of dolphins, of anaesthesia for serious

surgical oPerations, and standards for the application of

antibiotics, tranquilizers, drus and vitamins. In the Tobo

Aquarium (JaPan) there has been established a special hospital

for diseased fishes and dolphins, where they undergo a course

of treatment (Russel, 1.966 ). It has been established that

the medicines which help )eople also help dolphins, in par-

ticular tablets that aid the digestion of food and vitamins,

as a suwlenient to the food ration.

In the work with the dol1?hins we carry out a medico-

biological control on the state of their health (Figure 3).

The importance of this work is dictated among other factors

by the fact that commercial hunting of dolphins in the USSR

is banned and every catch of experimental animals is turned 292

into a complex problem, and therefore we consider that it is

more important to preserve the animals than to catch new ones.

Serving as objective symptoms that are indicative of

the state of health of the animals are : the rate of swimming

and diving; intercourse with other animals; degree of acti-

vity at feeding; character of respiration. A slowing in the

rate of swimialng, long periods spent on the surface of the

water and a reduction in activity at feeding are all patho-

logical symptoms, and in the presence of these the animal

should be examined.

The most important disease symptoms of the animal are:

A disturbance in the rhythm and intensity of the respiratory

act - a prolonged res-piratory act with a sluggish exhalation-

inhalation and the appearance of foaming from the blowhole at

the moment of exhalation; assistance given to any animal on

the part of the others; a yellow or grey colour of the faecal

niasses; a categorical refusal of food. In such cases it is

necessary to immediately examine the dolphin.

lv:ucilagenous discharges from the blowhole (of the spu-

tum type) and a putrescent odour of the expired air are

472

NC. 3. Chcpciutoft àle,locmoTp. CII1TI1e KaplutorpammiA c (1)1111a-eao6nuli

Figure 3. Routine medical examination. Obtaining a cardio-

gram from a white-sided doluhin.

indicative of a flooding of the' respiratory tracts with water

and the presence of inflammatory processes in the lungs. In

this case it is necessary to isolate the diseased animal from

the general group, placing it in a laboratory aquarium for 4 -

5 - days and treating it with antibiotics (penicillin or bicillin*

in the usual doses per 1 kg weight).

Liquid faeces of a yellow or grey colour are indicative

of diseases of the gastro-intestinal tract. In this case it

is necessary that the animal be fed only fresh fish, with which

there may be administered tablets of laevomycetin. Besides

this, use is made of antibiotics (biomycin, synthomycin*1 and 293

vitamins of the "B" group (in the usual doses per 1 kg body

weight). When there is a lack of aupetite and deterioration

of the state, it is advisable to isolate the dolphin for a pro-

longed period from the remaining stocke to avoid the spread of

Translator's notes. * bitsillin or bicillin benzathine penicillin G. ** synthomycin - synthetic streptomycin

473

the infection among the remaining animals, and to carry out

an intensive penicillin therapy.

In the case of the presence on some part of the body

or other of fungal diseases, ulcerations, bedsores and other

skin diseases, there were employed intramuscular injections

of antibiotics and administrations of polyvitamins in the food.

Simultaneously with this there was carried out a treatment of

the infected portions of skin with antiseptic solutions (iodine,

alcohol, potassium permanganate), with a subsequent drying in

air and smearing of the affected sites with sPecial dermato-

logical ointments. A depressive state of the animals may be

revealed only vith systematic observations The superficial

symptoms are: aggressiveness, swimming separately from the

group, loss of appetite and increased caution. All of these

may be evoked by a depressive state, if it proves on examina-

tion that other diseases are absent. In these cases aminazine

was employed.

The methods of medical control which were used in our

work proved themselves to be fully justified. For illustration

we present two examples when the treatment was effective and

operative.

en the 24th of June 1969 a young female bottlenose dol-

phin, which was living its second year in captivity, refused

its food. On the following two days she continued to ignore

the food, swam apart from the group on the surface of the

water, and did not react at all to man and the dolphins. She

was given intramuscular injections of bicillin, 600 thousand

units, and a comolex of the vitamins B 1 , 1-36 . and B121 in 2m1

v

v

aliquots. After 3 - la, hours she becan to move-mors actively

about the pool but continued to refuse the food. Over the

course of the next few c3.ays she made attempts to eat fish,

but without success. On June 30th a thourough medical exami-

nation was made, since the suspicion arose that the dolphin

had swallowed a small rubber ball. In 6 days the animal had

grown considerably thinner, the back was sunk in, the mucôsa

of the mouth cavity was inflazned and a distinct putrescent

odour emanated from this. A strut was placed into the mouth

and an attemnt was made to introduce some machine lubr.icating

oil into the stomach through a rubber tube. This was not

successful. However an inclination to vomiting was noted.

The cavity of the mouth was treated with penicillin and a

solution of potassium permanganate, Intramuscular injections

were macle of bicillin - 1 million 800 thousand units, and a

complex of vitamins (i3i and Bi2 ) in !I ml aliquots. Already

on the following day there were noted attempts, though it is

true that these were still fairly sluggish, by the dolphin to

play with a black rubber ring and with the other dolphins.

On July 3rd repeated injections were made of bicillin

(1 million 200 thousand units) and a complex of vitamins (B1

and B12 ) in 11, nil aliquots, while during the examination the

mouth cavity was aaain irrigated, although the state of the

mucosa was already satisfactory. Towards the evening of this

day the bottlenose dolphin fairly readily and energeticall.y

played with a ball and with rubber rings. Her movements be-

came more confident, energetic and pronounced. She readily

began to eat fish. She began to swim in ai.)air with a young

4.75

bottlenose dolphin, reacting in a lively fashion to the ap-

pearance of a man and making contact with him. During the

course of the following week she was given 3)olyvitamins and

oxytetracycline. This animal soon recovered completely.

In a white-sided dolDhin, which had been caught in May

of 1969, during a routine change on July 16th there was noted

typical loss of coordination of movement, which was expressed

by a rolline over onto the side and bumping into the walls of

the oen. After several convulsive movements of the head she

suddenly sank. We were able to quickly raise her from the

bottom, but she could not maintain herself afloat. It was

necessary to transfer her into a laboratory aquarium and to

secure her on a hammock, so that she would not choke. She was

given an intramuscular injection of bicillin (1 million 200

thousand units) and mellipromin (2m1), after which the water

in the aquarium was disinfected with a solution of potassium

permanganate, while the skin was treated with alcohol and

iodine solutions. The respiration of the animal had a rate

of 31 times in 5 minutes, the heart-beat frequency was 110

beats per minute. On July 19th we repeated the injection of

bicillin (600 thousand units) and of the vitamins B6 and B12

(4 ml of each). The frequency of respiration decreased con-

siderably, to 12 times in 5 minutes. The pulse - to 90 beats.

The irrigation was rel)eated with the disinfectant solutions

and the infected portions of the skin were smeared with un-

decine ointment.

On July 20th, after she had been kept in the laboratory

aquarium for 87 hours without food (refusing the food), she

4.7 6

was let out into the pool 1 here she began to swim actively,

and after 5 minutes to take scad greedily from the hand. Over

the course of a week we continued to give oxytetracycline and

polyvitamins with the food. This white-sided dolphin is still

alive at the present time.

The catching, transportation and relocations, and the

medical servicing during the course of long-term holding in

captivity are important constituent parts, on each of which

is dePendent the overall success of the work with the dolphins.

The tested methods for carrying out the catch and

transportation, taking into account the available practical

possibilities under the conditions in the Black Sea basin,

have shown that they may be successfully utilized in future

work.

The transoortation of the captured animals on a ref-

rigerator ship in containers with running sea water, with the

animais being secured in hammocks, was found to be the most

favourable form of transportation of dolphins by sea. However,

the extension of work with dolphins sets investigators the

task of elaborating and employing method of transportation on

dry land, which is a more complex problem.

The apPlication of therapeutic measures, utilized by

man, deserves by all means to be maintained and to be intro-

duced even more boldly into the practice of the routine work

with dolphins. However, in soite of the assistance rendered,

the mortality among dolphins during long-term holding in cap-

tivity remains high. Therefore there are needed further thor-

ough studies on the causes of the mortality of the animals and

on the elimination of these.

47?

Gur work does not pretend to provide a complete eluci-

dation of the problems that have been nai.sed. One of its aims

was to prepare the soil for the further development of the

above listed problems, connected with the long-term holding

of dolphins in caotivity.

JIIdTEPt1TYPfl

^ IJen^tiueu^t B. M. tt Op. 3ara;uta ot;cana. n4.. «,^io:to;last raap;ltts3», 1JG5.

2 Knetïxettôcp, C. E. u Op. BeayNa. M., 1964.

3 rlttr,.^tt %I. Lle:to1nex il r^c:n,c^slt1. \^I., «\Itac:tt,», 1965.

,lj• j'aii K. rloacr Tpc\ xtrrots. «>lntcp3n;ar, N, 87. 1963.

5 Cor;naoa B. E. u iJp.Ont,n• nccrte;totlaun;l ocoGewtuc^eü naat:<uulsl Aenli^snttols

L' rltAPos:asta^C. "fe3. Aotc:l. '1-ro licecoloSnoro cotSCnlaliltn Ilo u3)•9ctwto ntohclaix

DtA holtnl'a1o11UfX. Kaan1111111'1>F1;1, 1969.

.^ fjTo.suatttt A. I. Ilcrop,lsl caenoro saluaaoTa. 1\9.. «1Ia^r,a», 1t^G;s.

7 Andersen S., Dziedwic /1 Behavior patterns of cantiv^ Ilarbor porpoise(Paena ll,rocacna), 13u11. Inst. Occano..f^r. Monaco, vol. 63, no 116. 1961,

hro:on D. 1I. »Flic Hcalth problcros of \Valrus colves and retnarl.s on theirgeneral progress in captivity. cln[ernal. 7oo learboolc», 4. 1962.

9 Bro;crt 17. 11., .+Iiclrttttrè R.N. lleallh prublenls of caplive dolphins ,Incl

scals. 4. Amer. Velerin. Mcd. Assoc». vol. 137, no (à, 1960.

ti j.(Y3rowtt D. H.,.i\'orris K. S. Observations of captive and wild Cetacea.

«J. b•1aminal», vol. 37, no 3, 1956.Caldwell D., Brown D., Calcic;:cil M. ]nicr;cncric bchavior by a

iive Pacific Pilot Whalc. «Los i1n;;^elcs Conntv Alns,» no 70. 19G3.

M Gast:Lt D. E.i)olphil, ai 11tc ;^Iapier t^I lri ulancl. «\c\r• "l.ealancl Sei. lic^•.»,

lrurn^l•'M , no 2. 196G.1 D«dok vcrtt Ilcc! W. N. Observations in flizllit re-tctionti of Tursiops

S lil^oilcr ^.u^^^^ ,• ^l,tus (M) with sUllic sUf;'esttonS on 'tli5til planning. «Z.

no 5, 1966.1Il• Keliog Il?. H., Rice C. E. Vistlal Discrimination and Probl^m Solvinl; in aBottlenose Dolphin. «1Vhales, Dolph. a Porp.». 1966.15 Kritzlcr N:

The Pilot \V hale al c^•IarinclancL «1tlt. Ilislor}'», GG, tlo 7, •191J...

jb Nagel E. L., Morgane P. J., :1icl'arland N. L. Anesthesia .for the I3ottlenosellolphin (Tttrsiops irnncatus). «Scicnce», vol. 146, no 3G51. 1961.Nortis N. S. Practical problems. ^\^'halcs, Dolshins <t. Porpoises». 1966.

] Ridgujny S.H. Miclical care of marine Inammttls. «J. Amer. Veterin.:lssos»,

147, no 10, 196,3,19Riilgtonll S. Il, t1lcCorntic J.

G. Ancsthctization of porpoises for major sur-

Hospital. «\emr vol.Scicntisi», 21, no. 6. 19G6.gnvRtsscl>'JI Jal:an's I0iGsil 1

967.

21 Scltevill W. E., Lawrence D. rood-finriin; by a captive porpoise (Tursiopstruncattts). r.I3reviora», no. 63, 1966.22Slijper E. L. l^thales. «I-Itttcilinsun of I-ondon». 1962.

REF FREIN iCES

1. Bel'kovich V. M. et al., A riddle of the ocean. Il+,oscow,

"I^_olodaya. Ûvar• diya" ("The Young Guard"), 1065-

2. !,,Ieinenberg S. E. et al.,

("Science" ), 1965 .

The beluga. TY:oscow, "r'iauka"

52 8

The fairly constant maintenance of the holes by the

ringed seals (Chapskii, 1940) suggested to us the possibility

of counting the number of seals and of their holes as these

were encountered; therefore, in conducting the aerial survey,

we recorded encounters with both seals and their holes, situ-

ated both in direct proximity to a lying animal and also with

no animal present.

It is still difficult to make a completely definitive

judgement as to the number of holes that are made and main-

tained by different individuals of the ringed seal, though

in the opinion of Greve (1896) each ringed seal in the Baltic

Sea has from 5 to 8 breathing holes (according to Ropelewski,

1952).

With respect to the holes of the grey seal Holm (1921)

writes; "There occur cases, when in severe freezing weather

the exit holes of the seals not infrequently freeze over and

many animals are forced to leave these and accumulate at one

hole in numbers of 10 - 15 individuals".

As a result of the studies, conducted on the landfast

ice in the Gulf of Finland in Earch of 1970, by one of the

authors there was thoroughly investigated the region between

Malyi and Bol'shoi Tyuters Islands. In the investigated area

there were found 17 seals and 17 holes of various diameter

(from 3 ).l to 40 cm), visible from above. The ratio of their

upper diameter (on the outer surface of the ice) to the lower

diameter (the edge of the ice that is under the water) com-

prised from 0.57 to 0.63. it is Possible that the size of the

hole is directly deoendent on the size of the animal. The holes

were situated at a distance of from 1 - 1.5 to 3 km from one

another.

529

o

.e4

Besides the 17 separately situated holes, there were

found by us in one place three holes of various sizes situated

next to one another (Figure 1).

Taking into consideration the smaller diameter of two

of these on their upper edge (16 x 15 and 20 x 17 cm), in com-

oarison with the large hole which was found beside these (37 x

40 cm), the thickness of the ice in the place where these holes

were located (about 30 cm) and the tracks of small sized flip-

pers around them (around the large hole there were tracks of

an adult seal), we assume that these two holes were used by

a young animal (possibly born in this same or Ln the previous

year). Such groupines of holes, of 2, 3 or even 4 together,

were observed by us on more than one occasion also from the

airolane.

Further studies of the characteristic features of the

distribution of the seals, and of the disposition and arrange-

ment of their holes will allow us to explain this phenomenon

more specifically.

According to the observations of V. A. Zhelgov every

ringed seal on the landfast ice makes a hole for coming out

onto the surface in the freezim; over patches and cracks

("smorozi") between the ice floes that have been drifting from

the middle of the winter. Later the thickness of the ice on

these "smorozi" increases but it always is less than the thick-

ness of the main frozen together ice floes. In the Gulf of

Finland this difference comprised from 10 to 20 cm. The holes

which are maintained by the ringed seal in the snow drifts

amidst the ice hummocks are also located on "smorozi". As the

thickness of the ice on the "smorozi" increases, the ringed

530

Pile. I. Patnoao)Renue pa3.-uvj- Iforo pa3mepa 13 ()Rawl mec-re. Map .r.

(baueKiiii (Pow A. F. &ammo u B. .4. }Keziwea '

Figure 1. The disposition of holes of various size in ohe

location. March. Gulf of Finland.

Photograph by A. G. Esipenko and V. A. Zheglov.

seal constantly maintains the hole, for which even in the

middle of 1arch it must break through, once or twice a day,

the crust of ice which forms on it (up to 2 cm in a night).

In the course of the investigations on the ice it was

noted that a ringed seal which had been frightened away did

not appear at its hole until the next day. Cbservations in

the regions closest to the hole (in a radius of 3 - 4 km)

showed that the ringed seal did not appear within this distance

on the surface of the ice, even though the weather conditions

for its emer.cecrIce onto the ice were favourable (clear, light

wind, daytime temoerature of +1 to - 2°c),

It is possible that during this period it either made

use of its hiding places (in the hummocks and snow drifts

5 31

with an access hole from the water) or it moved away from this

hole to a distance greater than 3 - 4 km.

Ln the basis of the information presented we assume

that on the landfast ice each ringed seal, up to a certain time

(until the holes are broken up during the break-up of the ice

or due to other factors), makes use of one hole for emerging

onto the surface of the ice and several hiding places with an

access to the water, situated in the ice hummocks, where the

ringed seal lies during bad weather, utilizes these for re-

Plenishing its air reserves during feeding and gives birth to

its younF. The hole which the ringed seal uses for emergence

onto the surface of the ice is clearly visible from an air-

plane. The holes that are located in the hiding places cannot

be counted from an airplane.

During the aerial survey we noted that the break-up

of the ice occurs mainly along the sites of the "smorozi" and

hummockinF. This has also been noted by hydrometeorologists

(Lethodological instructions, 1959). In such cases, both the

holes which are concealed in the hiding places as well as the

open holes, used by the ringed seal for emergence onto the sur-

face of the ice during its break-up, in our opinion, are des-

troyed in anproximately the same ratio. The ringed seals,

whose holes have been destroyed, are apparently forced to make

use of the open patches of water or the holes of their "neigh-

bours".

Taking into consideration the aoproximate periods of

whelpine and moulting of the ringed seal and of the grey seal,

the count of the seals in the Gulfs of Finland and Riga and

532

on Lake Ladoga was conducted in two stages t first - on 1arch

9 - 11, and second - on April 3 - 7. On the basis of informa-

tion obtained from questioning the local sealers and our own

assumutions on the possible distribution of the seals in the

regions under study, the transects for the observation flight

in 1\.arch were set up ahead of time, while in Korn these were

set UP on the basis of the observations made during the first

flights and the direct observations made on the ice in March.

On detection of a region with a more concentrated dis-

tribution of the seals there were "set up" counting areas (see

the schematic maos of the aerial survey). The region was in-

tersected by several counting lines. This permitted us to

estimate more reliably the density of the hauling-out of the

seals in the given region and to determine the borders of their

hauling-out patches.

The flights were made on LI-2 and IL-14 airplanes at

an altitude of from 100 to 150 m, at an average speed of 200

km per hour, from 97.10 a.m. to 3-6 n.m.

The observations were carried out from the right and

left side from the blisters (transparent hemispheres, which

were mounted in place of the portholes in the navigator's com-

partment), which Permitted the observer to see not only to the

side but also to the front and underneath the airplane.

The data on the state of the ice cover was periodical-

ly recorded in the log-book using the generally accepted me-

thod for charting ice mans in accordance with the methodologi-

cal instructions for aerial observations on ice conditions at

sea.

533

Special attention was paid to determining the degree

of hummocking, the snow cover and the closeness of the ice.

Generally on the state of these factors depended the distri-

bution and concentration of the seals and their holes, the

numbers of which and the characteristics of their disPosition

were carefully recorded in the flight logs according to the

time of encounter.

According to our observations, it is best to make thé

simultaneous count of the seals and the holes t from one side,

over a counting strip not more than 150 m in width, although

the seals may be discerned also from the side at a distance

of 300 - 350 m from the airplane. At a greater distance from

the airplane the holes are poorly discernable and it is Pos-

sible to miss them. It is necessary to mention also the par-

ticularly cautious behaviour of the animals themselves in the

Baltic. In the Gulfs of Finland and Riga the ringed seals

escape into their holes 100 - 150 m ahead of the approach of

an LI-2 airplane to it and 200 - 250 m ahead of an IL-14 air-

plane, which forced the observer to look ahead, along the

flight path of the airPlane, from time to time.

The Ladoga seal behaves more calmly in the presence

of the ID-14 airplane, and escapes from it into the water at

a distance of 75-100 m.

It is possible that such differences In the behaviour

of the Ladoga seal and the ringed seal of the Baltic during

the sPring period depend on their physiological state or on

sonie other factors connected with living in bodies of water

with different hydroloe- ical characteristics.

5311-

Ne. 2. llpuuaïl, o6paa0uannLlft 113 upmlocnurn.ac,ta. Anpc:n,. Pnxcit;nit 3:1aua. ( CTpc.Rxa>lu yxa-3a11L1 1I)'1101 K0.7L98T6IX 11Cpn. Ot1nC?ff Iia 1111X

nca:nT ncpna)Asporjioro A. K. 111.voxoru:a

"?ilgure 2. La_nffast ice, formed of ice which has drifted in.

t^-pril. Gulf of Riga. (The arrows indicate holes

of the ringed seal. 13eside one of these lies a.

ses.l ).

Aerial pho-to^ra?)h by A. :•". Shlyundin.

a - hole of sealb - rinEred seal and hole

From the air the ringed seal is similar in form to a

comma (2ed.oseev, 1966). The holes of the seals in the Baltic

are very even dark c i rcles situated beside hummocks. From a

heit:ht of 9.00 - 1.J0 in around the holes there can be clearly

seen the snow that has been crushed down by the seals. The

ed.,es of the holes are not fro..f._: o^c^er., as in the holes of the

haryp seals (Figure 2).

It is almost impossible to de-termine the size of ani-

nials lying singly from an airplane. In April, when groups of

'inirl?a1s were found l^T^.n^ in directdirect ^^ro._ir:i-ty to one another,

t\re observed ?both small as well as lar^,-e seals. it is fairly

cai.ffi ct?l.t to discern awh.itet:oat from an airplane. ',.e no-ted

these only s..oains t ice of a background colour darker than them.

535

The whitecoats are distinzuishable from the adult animals by

their colour and smaller dimensions (about half the length of

the body beside a lying adult animal).

In :arch and J+.nril in the Gulfs of the i3altic Sea we •

noted 2 and 3 whitecoats.

The ,grey seal was visible as a dark-grey comma against 329

the background of the white and grey-white ice. It has a much

lighter colour and is larger than the ringed seal, and lies

more frequently beside cracks. It scarcely reacts at all to

the airplane, and if it does escape into the water, which oc-

curred only in two cases, it does this only right under the

airUane.

• The tentative count of the numbers of the rin,g;ed seal

and the grey seal in the regions under investigation was con-

ducted accordin.q: to the following schemes

1. Drawing the counting transect on a large-scale map

on the basis of the times and orientation points.

2. Transferring from the flight log onto the schema-

tic mao of the transect the information on the sites of finding

of the seals and their holes, and, as changes occurred, the

ice conditions.

3. Gutlining the contours of uniform ice conditions

(according to the types of ice formation), in which we utilized

our own observations and the data from the ice survey by the

hydro-meteorological service, which conducted its work simul-

taneously to us.

4. Calculating the areas of similar ice conditions.

5. Calculating the area of the counting strips for

each type of ice formation (the width of the total counting

5 36

strip from both sides of the airolane was 300 m), determining

the density of the distribution of the seals and their holes

on the counting strips and in the areas with more concentrated

haulinE-out Patches (see the schematic maps of the surveys and

the counting areas on these) for each SQ. km . for each cate-

gory of ice formation.

6, CalculatinP: the approximate numbers of seals and

holes over the area for each type of ice formation (separately

for each Gulf and for Lake Ladoga) by the method of extrapola-

tion of the obtained densities and their distribution.

RESULTS OF SURVEY

As a result of the data obtained by us from the aero-

visual observations, the ice types accepted by other investi-

gators - white, grey-white, grey etc. (Popov, 1966; Shustov,

1969, and other authors), did not permit us to characterize

the conditions and characteristics of the distribution of the

rinp:ed seal and the'exey seal in the Gulfs of the Baltic Sea

and on Lake Ladoga.

For the characterization of the habitat conditions and

the distribution of the seals and their holes we distinguished

several types of ice formations, which were characterized by

a particular density of the distribution. Under the concept

of "density" we imply the average number of seals and of their

holes Per one square kilometer.

1 9 Landfast ice of local origin is

characterized by_a thick ice cover with a thickness of the ice

of more than 30 - 40 cm (from our ice observations), with a

1

537

weak hummocking (up to strength 1), formed by a certain shif-

ting of the ice during the winter; its snow 'cover in the

middle of the winter has an index of up to 3. The grey seal

was not found by us in this type of ice formation; the rineed

seal and its holes are occasionally encou.ntered in hummocks

or beside these .

No seals or their holes were found at a.11 either from

the (airplane or during the observations on the ice in the

Gulfs of the Baltic Sea on landfast ice of local oriein which •

overlies depths of up to 10 - 15 m (see schematic maps of

a.erial survey ). At present it is difficult to explain this.

It is possible that this was due to the absence of the basic

food objects of the seals at small dePths during the winter

period or by the small number of hummoclçs, which are needec3.

'oy the seals for the construction of their hiding places.

Therefore we did not take these areas into• consideration in

calculating the areas of landfast ice of local origin.

2. Landfast ice, formed by the freezing

together of' white ice t.hat has drifted in e

with a degree of hummocking of strength 2 - Li, in our opinion,

i.s the most favourable for the construction of the hiding

places and for the 'maintenance of the holes by the ringed seal

(Figure 3).

At the beginning of January - February 1 during the

hummocking and free 'zing together of the 3_0e le ,thich is drifting

in the ,lfs, the ringed seal, in all probability, is distri-

buted alorrY the o,)e.n oaLtches of water Etmong the ice floes.

Subsequently, when these freeze tor;ether and ice is formed on

the open :.)atches, the seal here makes and mainta.ins its holes,

and in the hummocks - its hidinp_..; -31a.c.:es

- 5 38 • • I

•

. Pile. 3. Tonotinoe mecTo mytioK 1<0.711,‘11iTal

nepnbi an npituae 113 np11110C11010 J11,)1,8. MapT. (1)liocKaii 3a-

. .11113 nepeRuem niane 1:031b,IiITCH"1 nepubl). I'm.° B. A. )Keennea

5. Fir-Jure Typical site for the construction of holes by the

ringed seal on landfast ice derived from drift ice.

r;ulf of Finland. (ln the foreround is a

hole of the ringed seal).

Ehoteraph by V. A. Zheluov

3. Breaking-up landfast ice of frozen

together drift ice is formed mainly with the onset

of warm weather.

In the Gulf of Riga in 1970. this landfast ice began

to be formed from the beginning of Larch; in the Gulf of

2inland and on Lake Ladoga - from the beginning of April

(the closeness of the ice had an index of 8 - 10).

In Larch in the Gulf of Riga.the density of the distri- 331

bution of the seals on the broken up landfast ice (with an

- 9 index of closeness of the ice) attained a level of 0.43

head and the holes had a density of 0.41; in April these

values were only 0.01 head and 0.02 holes. This is probably

connected with the breaking up of the holes and the distribution

of the animals alonu the cracks and on ice formations of other tr e s.

539

In the Gulf of -7'inland on the broken up landfast ice

(closeness of the ice with an index of about 10) in April the

density of the seals was0.15 head, while the density of the

holes was 0. 3 7.

The smaller density of the holesin the Gulf of Riga,

in comparison with the Gulf of loinland, was obviously related

to the greater break-un of the ice, and therefore of the holes,

in this Gulf.

In the Gulf of Finland the opening up of the landfast

ice and its subsequent break-up occurs later than in the Gulf

of Riga, by 10 - 15 days (Atlas of ice conditions in the Bal-

tic Sea, 1960).

4, Ice fields of white and grey-white

ice are formed of frozen together driftine ice floes in

the central and western Part of the Gulf of Finland at the end

of January - beginning of i;'ebruary, and in the middle of the

Gulf of Riga in the middle to the end of February (Atlas of

ice conditions in the Baltic Sea, 1960).

It was noted by us that in . April the ringed seal and

its holes were encountered more frequently (density up to 0.17

head and 0.17 holes) on the ice formations of this type in the

Gulf of Finland than on the landfast ice of local origin.

We consider that the more broken up ice of this type

in the Gulf of Riga also exPlains the smaller density of the

seals and of their holes on this ice (correspondingly 0.02

and 0.13).

In karch Anril, as the ice fields break u), there

occurs a translocation of the seals onto firmer ice, where they

51a-0

distribute themselves along the cracks and oPen :patches of

water, and someti.mes also utilize the holes of the "old resi-

derits". Thus, according to our aerial observa-tions, neither

seals nor holes were found in April on the broken white and

grey-white ice in the Gulf of r'inland and the Gulf of Riga.

At the same time on the ice fields and broken up

landfast ice beside one hole or crack there lay 2 -- 5 seals,

which was noted by us on more than one occasion in April in

the Gulfs of Riga and Finland.

This is indicative of their concentration at the be-

^,.-innin` of A-pril on the firmer ice. In the Gulf of Riga the

fragmentation of the ice occurs in the direction of from south

to north of the -gulf, while in the Gulf of Finland this occurs

from the west towards the north and east (Atlas of ice condi-

tions in the ^,̂ alti.c Sea, 1960). In step with the fragmentation

of the ice the redistribution of the seals apparently occurs

in the same direction.

5. Broke,n-up white and grey-white i.ce,

in our opinion, is unfavourable, in I::arch and in April, for

the hauling out of the ringed seal and the Grey seal. The

areas with this type of ice in the Gulfs of the Baltic Sea

are small (see schematic maps of aerial survey).

In T"arch on ice formations of this type in the region 332

of the Irbenst.i? Strait we noted a relatively high density of

ringed seals, 0.33 head. We explain this as a result of the

dri ftinz of the breakinE--up ice fields through the Irbenskii

^,`),trait into the ouen sea, where the ring,ecl seal does not .)ass

with the ice, but as the broken u-p ice drifts out to the west

the seals concentrate in the :trbenskii Strait.

541

Pitc. 4. Cxema artuaytteTitoro mapulpyra ii aeaonoft o6cration1at I(1)1111éKOM 3a.ritute (mapT 1970 r.)

*. .

Figure 4. Schematic map of the aerial survey transect and ice conditions in the Gulf of Finland (P;arch,'1970).

A. - Counting area.

In Aril, when the edge of the broken-up landfast ice

and of the ice fields moves to the north, and the more broken-

uo grey-white ice is carried out into the open sea, the ringed

seal is no longer encountered here.

6. Light and very light floe grey

ice (area of floes from 40 to 200 sq. m) is formed at the

edge of the ice fields and as it were girdles them with a 5 -

10 km belt: in the Gulf of l'aga from the south, and Ln the

Gulf of Finland from the south-west (see schematic ma) of

aerial survey). The ar:eas of this are small, and in the sprin

it is rapidly broen up. The seals clearly avoid such ice and

were not found by us here.

0 A

®U

00

FED

5 9

®b

EE

it

12

Yc.ionul,te o6o3ua4enlln tc piic. 4-8

1. I]pün<lü ^secTUOro nponc^o.xa^ulsst2. Ilpuuait 113 npuiloalorn cmcp3uicroc5l :n,aa

3. B3.noataunt,tïi npnnai: 113 npniloCnoro ca+epsiue-

rocsi rli>^a4. ^Ielütui,ic iloast Ge-'()ro ti cepo 6e loro .at 1:i5. Iàntl,tïi 6c.1litü n ccpo Geni,li'i 10a6. l^p)'nuo- il nlear.eGnrt+ïi cepi,+ü tc [7. Bo;ta8. I pf111fII[6i 'Cn(IU6 ICdUCLI\ o61)a3oüfln1lÎt il CLIeT-

u[,is naoulaAeït

9. blilpnlp)'T 8nll,lY'IeT110. 13erpeaelnlasl na ',saplupçre xo ibllaT 1 isi nepna

il. IicTpet+euui,lii lui i,\taplnpvTe cep1,tü Tlo.leni,12. L'crpevellnan na asapulp^'Te :nnlca

6",

8 -LT:?`:cnd. to .E''iy^hres I• --

La.nd_C'ast ice of local ori"in.Landfast ice of drifted-in frozen to^,ether ice.Broken-up landfast ice of frozen together drift ice.Ice -.fields of white and Erey-white ice.3rol_en-uu white and gr ey-whi.te ice.I:iEl>.t and very 11.21->.t floe Erey ice.Water.,o_rd.ers of the types of ice formations and counting areas.Transect of aerial sur;Tey. vRinged seal encountered along transect.Grey seal encountered along transect.Hole encountered along transect.

.Un ice formations of other types (ice rind, pancake

ice) the seals were also not found. This is connected with

the fra^^;ilit,y and remoteness of such ice from the ice fields.

The distributions of the listed ice formations, the

encounters with seals and holes alon- the transects, and the

re4dons of their concentrations (counting areas) in the

5 4.3

Pm'. 5. Cxema aDilaytteTnoro maptupyra 06CTa110131:11 Cinalcizom 3amme (aripeab 1970 r.)

Figure 5. Schematic map of the aerial survey transect and

ice conditions in the Gulf of Finland (April, 1970).

investigated bodies of water are presented in the schematic

maps of the aerial survey (Figures 4 - 8).

DYNAMICS OF THE FREQUENCY OF OCCURRENCE OF THE SEALS AND

OF THEIR HOLES IN THE GULF OF FINLAND

In comparing the data on the density of the distribu-

tion of the encountered seals and their holes on the various

types of ice formations, we found:

In March the maximal density of hauling out of the

rinued seal occurred on the landfast ice of drifted-in ice

(up to 0.62 head). A smaller density occurred on the ice

fields of white ice, 0.17 head. The densities of the distri-

bution of the holes were correspondingly expressed as 0.4 and

544

A.- Counting area

Pue. 6. Cxema anuavtieruoro mapuipyra u :Icaonoff (De clauouu B PiGi■CHOM malice (maim. 19/0 r.)

Figure 6. SchematiC mao of the aerial Survey transect and

ice conditions in the Gulf of Riga (iiiarch, 1970).

0.17 holes. There was no broken-uo landfast ice in March.

In summing up, from the calculations, in March there were

about 5000 seals and 4500 holes.

Durine the course of the oeriod between the surveys

profound changes occurred over the whole ice area in the Gulf.

In April there was a decrease in the size of the areas

of nf continuous landfast ice of frozen together ice; these

were replaced by broken-up landfast ice and partially broken-up

A.- Counting area

54.5

•

Pnc. 7. èxema annaytieTnoro-mapniprra n .Icaonort. o6cTa- 110111C11 n PII)liCKOM 3asnme (anpc.lib 1970 r.)

Figure 7. SchematiC map of the aerial survey transect and ice conditions in the Gulf of Riga (April, 1970).

ice fields. The average density . of hauling-out of the ringed

seals on the broken-up landfast ice of frozen together drift

ice in April was 0.15 head, while the density of the holes was

0.37. The ringed seal was not found on the ice fields (the

size of the floes here had decreased to 0.3 - 0.7 sq. km .),

where there were still occasionally found traces, i.e. the holes,

of their prior habitation.

546

The absence of seals on this type of ice formation

is explained, apparently, by their migration to other regions.

According to our calculations, in April in the Gulf

of Finland there were present on the ice about 900 ringed

seals and about 3000 holes.

It is generally known that the ringed seal does not 336

carry out distant migrations, with the exception of slight

seasonal migrations, related to shifts in the edge of the ice

(Bergman, 1958; Chapskii, 1963), and therefore we consider

that the number of ringed seals calculated on the basis of the

encounters in A'uril does not reflect their actual numbers.

The decrease in the frequency of encounters with the

ringed seals on all of the types of ice formations in j1pril

is explained by us as being due to their redistribution onto 337

the edges of the broken up landfast ice, where they form denser

moulting patches, for the detection of which a large counting

area is necessary. Three such patches of ringed seals were

found by us in April in the region of the Islands of ï'ykh ^ya--

Ukhti, Vigrund and 11'ayak. The seals lay here in groups of

2 5 individuals beside one hole or beside an open patch of

water in direct proximity to one another. Groups of ringed

seals were situated at a distance of 0.5 -- 1.5 km from one

another.

The decrease in the total number of holes encountered

in April (according to our calculations there were up to 3000

of these) is related, in our opinion, to the widening of the

"smorozi" and the fragmentation of the hummocked sections on

the ice fields and broken-up landfast ice.

338

5 11.7

o

o Pue. 8. Cxenta annaynernoro maptupyra u sieRonoii o6-

cranonizu n ioclot o3epe (anpe,ib 1970 1.) _ . . . •

2igure 8. Schematic map of the aerial survey transect and

ice conditions on Lake Ladoga (April, 1970).

The data obtained on the distribution of the seals,

on the basis of the calculations for the entire area covered

by ice in the Gulf of Finland, permitted us to make a tenta-

tive estimate of their approximate numbers and to determine

the regions of their spring concentration. It was found that

the main part of the population of the ringed seal in the Gulf

of Finland at this time is concentrated in the eastern part

of the Gulf between the islands of Vaindlo and Seyskari.

548

IMF

o

The approximate number of the ringed seals, present

on the ice, is reckoned by us, on the basis of the encounters

in F.arch, at about 5000 head, with about 4500 holes. The en-

counter with one ig-ey seal in April was a natural occurrence,

since the area investig.ated at this time was 1.6 times lare_er

than in March, and these seals are very rarely encountered on

the ice during this period.

We consider that the number of ringed seals and of

their holes which was calculated by us on the basis of the

density is underestimated. In our case, during the survey

all of the record entries were made by the individual conduc-

ting the count, who had to interrupt his observations to do

this. It has been established experimentally by American in-

vestigators that the error in counting from an airplane, even.

in the case of ungulates on tundra, leads to an underestimate

of the results by up to 20;0 on average. It is therefore re-

commended that those making the count should transmit their

observations, without interrupting these, to a co-worker who

makes all of the record entries (Chervonnyi, 1969).

Takinginto consideration a similar miscalculation in

our work and the approximate percentage of occurrence of the

seals in the water (about 30,%0) duping the daytime in March,

it may be assumed that in the Gulf of Finland during this

period there were oresent about 8 thousand head of the ringed

seal. The calculated number of holes, on the basis of the

findings in b_arch (with a correction for omissions of 20A),

is about 5.3 thousand.

54 9

THE-GULF OF RIGA

The overall character of the distribution of the seals

and their holes according to the types of ice formation is

similar to their distribution in the Gulf of Finland (see

Figure 7).

The number of ringed seals calculated by us on the

basis of the frequency of their encounters'in April is also

smaller (in March - 3.1 thousand head, and in April only 0.12

thousand head). It is probable that the ringed seal in the

Gulf of Riga in April also quits the moving ice and concen-

trates on the edges of the firmer ice (the stationary and

broken-up landfast ice).

In April, in connection with the breaking-up of the

"smorozi" and hummocks, the number of holes decreased consi.L.

derably (according to the calculations there were 3.9 thousand

of these in March, while in April there were only 1.5 thousand

holes).

The number Of ringed seals obtained on the basis of

its encounters in April undoubtedly does not reflect the actual

stocks (as is the case in the Gulf of Finland), and therefore

we calculate the tentative Population numbers of the ringed

seal on the basis of its encounters in March.

In March and April on the breking-up landfast ice in

the Gulf of Riga there were encountered individual grey seals.

In April at the edEe of the ice fields of white and grey-white

ice (with a closeness index of 7-8, and a degree of hummocking

of 2-3) there was found by us a hauline-out patch of grey seals,

where a counting area was set up. The seals here were diffusely

550

distributed, with 1 - 2 beside one hole or edge of an open

uatch of water, at a distance of 0.3 - 1.5 km from one another.

According to our calculations (density 0.89 head), in

this area there were lying about 200 - 250 seals. Besides

this, individual grey seals were also encountered on the

breakine-uu landfast ice.

The finding of a single hauling-out uatch of grey

seals in April evidently cannot serve for evaluating their

population numbers in the Gulf. According to the data of

Sundstrem (1962), adult grey seals are attached to their own

reuions and migrate only in the summer. Therefore we assume

the possible oresence of such hauling-out patches in other

regions of the Gulf.

Un the basis of the results of the aerial survey,

which was carried out by us in D.arch - April, there were reck-

oned to be about 3 thousand ringed seals on the ice in the

Gulf of Riga. Taking into consideration that at the moment

of the count a part of the animals were in the water and an-

other part was omitted while making the record entries, and

introducing the appropriate corrections (tentatively 30% -in

the water, and 20% - for omissions), the auproximate numbers

of ringed seals in the Gulf of Riga were estimated at about

4.5 thousand head.

The calculated number of holes on the basis of the

encounters in 'earch was 3.9 thousand; with the correction

(20%) for omissions the numbers of these come to 4.68 thousand

holes.

551

16 LAKE UDOGA

A tentative picture concerning the places of the dis-

tribution of the Ladoga seal (on the basis of findings of

holes) over the expanse of the lake was provided by the first

test flight over this in T^:ar.ch. holes of the ringed seals

were occasionally found from 5-to 30 km from the eastern and

western shores of the lake.

At the beginning of April up to 20jo of the surface of

the lake remained covered with stationary ice, 60Jo of its re-

maining area was covered with breaking-up landfast ice (Figure

8). The snow cover on the breaking-up landfast ice had an in-

dex ranging from 1 to 2.

At this time on the ice there were found not only holes,

but also ringed seals beside the holes or open patches of water.

The Ladoga seal, like the ringed seal of the Baltic,

prefers landfast ice derived from drifted-in frozen-together

ice. In the northern part of the lake, up to the line of the

Valaam archipelago,.on the landfast ice with very sparse cracks

and hummocks the density of the hauling-out of the ringed

seals in April was 0.028 head, while the density of the holes

was 0.081-1•. The density of the hauling-out of the seals on the

breaking-up landfast ice derived from drifted-in ice was 0.19.

head, while the density of the, holes was 0.36.

The main part of the population of the Ladoga seal

during the syprinf; period was dispersed around the central part 311-0

of the lake, which part is the last to freeze up (in mild win-

ters - in Pebruary, and in severe winters - in January). it

is interesting that, according to the data of Kalesnik (1968),

these places are the basic regions for catching fish.

552

o The number of seals calculated on the basis of their

encounters (2200 head), in our opinion, does not reflect the

actual stocks, since the average yearly catch of these seals

of about 0.5 thousand head has been maintained for several

years without changing. Moreover, the small number of encoun-

ters with these seals during the survey in April may be ex-

plained by the fact that at this time there was cloudy and

rainy weather over Lake Ladoga.

The number of holes of the seals on Lake Ladoga, with

the correction for the inherent error . (20%), is about 5.1 thou-

sand. The partial breking-up of the landfast ice undoubtedly

influenced the numbei's of these, but at the present time we

cannot estimate any correction .factor for this. It can only

be assumed that the numbers of ringed seals in this case should

be greater than the calculated number of holes.

In the analysis of the data obtained in March and

April on the distribution of seals beside the holes and open

patches of water some interesting data were obtained.

In the Gulf of Finland in narch 85% of the seals were

found beside the holes, while in April - only 44%. In the

Gulf of Riga the corresponding value was 48% in Larch, while

in April the rined seals were encountered only beside open

patches of water.

In Lake LFidoga in April 79% Of the ringed seals were

found beside the holes and 21% beside open patches of water.

Un more than one occasion, while conducting the aerial survey,

we noted that on the approach of the airplane the ringed seal

escapes into the water not by way of the polynya but through

its hole, which is sometimes situated fairly close to the

.553

polynya (uo to 3 m). On this basis we judge that there is a

considerable attachment of the seal to its hole. The differing

percentage of the finding of the animals beside the holes or

beside the ooen patches of water in March and in April may be

explained by the availability of the holes, the number of which

decreases as the ice breaks up, or by a differing physiologi-

cal state of the animals, which compels the seals to make

greater use of open patches of water and cracks.

CONCLUSIONS

The comparison of the results of the aerial survey in

March and April showed that:

1. The ringed seal in March and April prefers landfast

ice derived from drift ice and ice fields.

In April it is concentrated on the edges of the breaking-

up landfast ice, where it forms scattered hauling-out patches

(possibly, moulting patches). The number of holes decreases

on the gradually breakin-up ice.

2. On the basis of the présence of whitecoats of the

ringed seal in the Gulfs of Riga and 2inland at the beginning

of March and in April there may be assumed to exist a certain

protraction in the periods of its reproduction.

3. The grey seal (in hauled7out groups and singly)

is found only on broken-up white ice.

I. Taking into consideration that this was the first

attempt to survey the ringed seal and grey seal on the landfast

ice in the conditions of the severe winter of 1969-1970, the

data obtained on the distribution and the numbers of the 'seals

5 54.

o should be considered as tentative , it is possible that the

spring concentrations and numbers of seals in the Gulfs of the

Baltic are not constant and depend on the climatic conditions

each year. In the future, after a thorouFfh study has been

conducted on the ice on the ecology and character of the dis-

tribution of the seals, it would be useful to conduct a repeated

aerial survey of these seals.

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'frak6w, 1952.

2F5. Suncl.str8rii K. The "re,y seals of the northern <ali,.^c .

"Sve r. natur. .jLrsbok, 1962, :lro j 53". Stockholm,

1962.

ri

559

TABLE OF CONTENTS 343

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Section 1. The ecology and distribution of cetaceans

B. A. Zenkovich. The fate of whales 7 4

V. 1. Krylov and L. P. Medvedev. Ubservations on cetaceans and pinnipeds during the 13th Soviet Antarctic Ext-)edition (SAE) 28 42 .

G. A. Klevezal' and D. D. Tormosov. The separation of local zroupins of sperm whales by the character of the dentine layers of the teeth 35

V. V. Zimushko. information on the reproduction of grey whales 44. 68

V. L. Yukhov. Some data on the feeding of sperm whales in the high latitudes of the Antarc- tic . . . 54 83

L. P. Medvedev. Information on the feeding of the beluga in the Dikson region 60 92

V. 1. Shevchenko. A contribution to the question of the origin of "white scars" on the body of whales 67 105

M. V. Ivashin. Cases of multiple pregnancies and foetal monsters in the baleen whales . . . 75 118

Section 11. The ecology, distribution and commercial exploitation of

seals, fur seals and sea otters

G. A. Fedoseev. The distribution and numbers of seals on whel -oing and moulting patches in the Sea of Ukhotsk 87 135

L. A. Popov. The causes and dimensions of the natural mortality of young harp seals durinsu the lactation period 100 159

• 54

560

*

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V. I. Krylov. The feeding of the Pacific walrus (Ldobaenus rosmarus divergens Ill.) . . . . 110 177

V. D. Pastukhov. Counting the young stock of the Baikal seal 117 190

V. L. Kogai. The state of the stock and the pro- gnosis of the feasible catch of fur seals on Robben Island 125 204

A. V. Zasosov. A method for determining the size of the fur seal oopulation and of the maximal possible stable catch quota 130 211

. Arsen'ev. The local distribution of fur seals in the Sea of Japan 138 226

F. G. Chelnokov. I:ovements of fur seals in the rookeries on Uednyy Island 151 240

V. E. Sokolov and N. S. Stepanov. An experiment on the radio tracking of fur seals 157 250

T. I. Chupakhina. The helminth fauna of the fur seals on Robben island 166 265

A. M. Nikolaev. 1211grations and local regrouoings of the Kuril sea otter 171 274

Section III. The morphology and:physiology of marine mammals and othet questions

E. I. Ivanova. The morphology of the vascular system of cetaceans and pinnipeds 179 285

A. S. Leontyuk. Peculiarities of the development of the blood-vascular system during the early embryogenesis of cetaceans 193 309

— P. A. Korzhuev. The problem of the oxygen réserves in the organism of aouatic mammals . . . . 205

V.

331

T. N. Glazova. Quantitative evaluation of the haematopoietic function of the skeleton of the Black Sea dolphins 212 344

G. N. Solntseva. The comparative-anatomical and histological characteristics of the structure of the external and Internal ear of certain

dolphins 226 369

and Caspian 273 440

Baikal seal

D. A. Morozov. Transportation of live dolphins . 296 479

K. K. Chapskii. generic seals .

The systematic rank and supra-differentiation of the 8-incisor

305 494

561

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G. M. Kosygin and V. N. Gol'tsev. Data on the morphology and ecology of the harbour seal of the Tatar Strait 238 384

V. A. Bychkov. The seasonal variation in the density of the fur in the hair bundles of three-year- old bachelor fur seals 253 411

G. A. Nesterov. The effect of the muscle relaxant ditiline on the organism of fur seals . . . 263 • 423

- V. N. Mineev. Questions of conservation and regula- tion of the stocks of marine mammals . . . 269 432

D. A. Eikeladze. Keeping fur seals seals in the Batumi Aquarium

V. D. Pastukhov. The feeding of the under experimental conditions 280 451

V. E. Bel'kovich and V. S. Gurevich. Questions re- lating to the catching and long-term holding of dolphins in captivity 286 461

A. A. Antonyuk. Quantitative changes in the ver- tebrae in the vertebral column of pinnipeds 317 514

V. A. Zheglov and K. K. ChaPskii. Test of an aerial survey of the ringed seal and of the grey seal and of their holes in the gulfs of the Baltic Sea and on Lake Ladoga . . • 323 524

4 562

RESEARCH ON MARINE MAMMALS

Transactions, issue XXXIX

7i'ditor Ê. S. Mil'chenko

Technical editor T. M. Andreeva

Artistic editor G. P. Burov Proof-reader E. I. Arenko

Delivered for type-setting 10th uctober 1972. Sent to press

19th 1,.ay 1972. Form .Ab uf paper 60 x 90 1/16. Conventional

printers' sheets 21.5. Registered publishers' sheets 21.18.

Edition 1000 copies. KU 04382. Larder 2354. Price 1 rouble, 60 cooecks.

Atlantic Research Institute of Fisheries and Oceanography.

Kaliningrad obl., ul. Dm. Donskogo, 5.

Printing-house of the "Kaliningradskaya pravda" publiShers.

Kaliningrad obi., ul. Karla Marksa, 18.

eee,74.

DUE DATE

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201-6503 . printed in USA