translation 3977

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ARCHIVE S
«lb
FISHERIES AND MARINE SERVICE
Translation Series No. 3977
The position of the longfin squid Loligo pealei Les. in the ecoystem
by A. N. Vovk
Original, title: Ilélozhenie dlinnoperogo kal'mara Loligo pealel Les. v ekosisteme
From: Tr. Atl. Nauchno Issled. Inst. Rybn. Khoz. Okeanogr. 58: 168-176, 1974
Translated by the Translation Bureau (WDP)
Multilingual Services Division
Department of the Secretary of State of Canada
Department of the Environment
Fisheries and Marine Service
Biological Station
St. John's, Nfld.
1977
13 pages typescript
Fdne141 39-T7
DEPARTMENT OF THE SECRETARY OF . STATE
SECRÉTARIAT D'ÉTAT
Freje
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ez.fm
-tezete
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DIVISION DES SERVICES
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TRANSLATED FROM - TRADUCTION CLE
INTO -- EN
Russian
_dnglish
AUTHOR - AUTEUR
A.N. Vovk
TITLE IN ENGLISH - TITRE ANGLAIS
The Position of the Longfin
bquid.Loligo
pealei Les , in the
.Zcosystem
TITLE IN FOREIGN LANGUAGE (TRANSLITERATE FOREIGN CHARACTERS)
TITRE EN LANGUE ÉTRANGERS (TRANSCRIRE EN CARACTERES ROMAINS)
Polozhenie dilinnoperogo kaltmara Qjjo pealei Les, v ekosistame.
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Trudy Atlant. Nauchno-issled. Inst. hyb. Khoz. Okeanogr.
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Transactions cif the Atlantic Scientific Research Institute of Fisheries
and Oceanography
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APR --1 1977
Trudy_ Atlant. Nauchno-issled. Inst. Ryb. Khoz. Okeanogr. (Transactions of the
Atlantic Scientific Research Institute of Fisheries and Oceanography), Vol. 58,
pp 176-184, 1974 (USSR)
UDC 794.5
The Position of the Longfin Squid
Loligo pealei Les, in the Ecosystem
By A. N. Vovk
Translator's note. Blank spaces in the first two paragraphs of this
translation correspond to illegible portions in the photocopy of the
Russian original.
Owing to their large numbers and wide occurrence, squid play an
enormous role in biological interrelations and represent one of the most
. important elements in the trophic structure of the sea and ocean biocoenoses
[1, 2, 6, 7,
established,
18/.
How great this role is, however, •has not yet been
investigators
analyze what significance
attach to squids, we find two different .
.
In some papers [1
and 10],we feel, the role of squid in biological productivity has been
exaggerated.
has determined
I. Akimushkin [1], for example, minimizes
the biomass of cephalopods as being no less than 2.2
billion tons, which represents about 6 tons per square kilometer of the
oceans of the world.
•
* Numbers in the right-hand margin indicate the corresponding pages in the
original.
UNEDITED r?, A 51.ATI ofq
Forirriz.--7m;.,".:". o.1 on:ly
TRAtViCii-0 NF.; ri
Information soulornorl SEC
5-25T (6/76)
/168*
2
Other authors [9] underate their value in food
L.A. Zenkevich [5] is probably right when he says that we still know very
little about the processes of secondary production
transformation of organic matter created in the upper
or about the
concerning
the amount and biological role of the fish population
cephalopods and other groups of ocean inhabitants.
get even general outlines of the interaction of the different
in the food chains helps to expand our knowledge of the biological structure
of the ocean.
We have attempted to present and to analyze what we feel are the
typical trophic relations of one of the important cephalopod species,
e
Loligo pe alei Les., and to determine its role and place in the ecosystem
of the shelf waters of thé northwest Atlantic (Fig. 1).
The material for our analysis included, on the one hand, data on the
feeding of squids, and on the other, [information on] fish and other animals
eaten by them (Table 1). The region of our research included the shelf and
continental slope from Cape Hatteras to Georges Bank. The available material
on the feeding of the squid and fish is of unequal value and hence enables us
to examine trophic interrelations only in general outline. Where we were
able to, we looked at the quantitative aspect of the interrelations.
The food items of the longfin squid include organisms on at least
three trophic levels:
a)
phytophages (copepods, euphauCiids, etc.);
b)
zooplanktophages (different groups of invertebrates, juvenile fish and
squids, planktophage fish and others);
c)
predators, (cods, porgies and squids) [4].
/171
27.
• •
26.mev _pbi5a m enue „a1),(u 28 olecgaelôo
21 oheobteld hi apitu. 14
oKapoban aKyna
5 drama
22\
171yHeal
otrUMOBCIR axyna
oKtimoôaft
17Kelg
25 ,; 17onocambe 1 I
c/Synop axyna
rnyHg4(
20 ceeeamuda
3 ewna-monorni
ilepited
3-3 Wyna-Ka
Fee
g . riumna_ _
I
iianacambtù petib 35.
Xapôecrruptuis â .
aCITANaR potea3?
lluHoewort 3
ry§.
ill >18cm
centeo
4 gmeç.z.timuvecicap
'Kpyznap ceilodo
(---"-1
Do s
1.f-elan:ye
.
Cicymapug 4b 0
i
•.-- '
e''
,, ,it'a.i.iman c 4tt:
m ''' • ,. ' Alanpenew of-wi
'eloY e , Cexbâ. 48.
78.!./774letee. 490
CazurnrneÉgier
affl y.c.,/,
77U"‘ILtHeU /.70/6 ;,)
U0 . .
PornomoeAtq
paxti 5;5
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PHoegoo,
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..-., !...-: ' -: i;:.
it it. „... ... ,..:,• e.,.„...,
ge",.:..110:nepor_à
, , c, _ , .: trà.nbmapa:
.:-:::1 ,'.,.- 1,1)1p ieci ;;;;., • i33sem
pilc. 1..›
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2! -,.-...,;.,...:-... r,..iwe cusialt; ..f,-• -; .-...., d c ,. f.: ., .--_, .
. . 1 ..C'Ct.il.P!Ible C1311311
-. . . .
.
• "-•
-
-
-
-
-
.
Fig. 1 - Trophic interrelations of thé longfin squid.
[illegible]
associations
[illegible]
associations
Key:
1.
2.
3.
4.
5.
6.
7.
8.
9.
i7
\
05 <zu_u_diai_res2
•
42.
Cfnewmall_w440
gongebee-xieterbeure
. rammapud.,
',.: .62
hoponeecean pei5a40
biepny3a . 43 d>
Ctreu5pue 6
gaeondyril
edy3b1
Koponuconepoiô nonama
2. flouwHonepold Kanbm'ap
a.
f.__.--Tayrnoaa 41.
--- ----1 /embipexnerrulacniag Naht5aea
4
67.
[17
ollepHblû MOPCKOil OlfgHb
j0 -1- Obellret1Hblà Clf77
î tea
7 Cûte a-
2. Cppe5pucnioniVeA
2;
Cenebecicnitou xee lu, reen
13 .
5e1bià xem
1
14.
ben_bià Hanum
15.
Irnackepù Hanum
30
&eyed
Gonatus [transcribed from original Russian";
longfin squid;
shortfin squid;
Atlantic herring;
[literally] round herring;
mackerel;
alewife;
jellyfish (medusa);
haddock
T
4
Figure 1 (Cont'd)
Key:
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
cod;
coalfish;
silver hake;
white hake [Latin name not available];
white hake [Urophycis tenuis]
red hake;
skate (ray);
thornback ray;
Levis skate [original Russian skat-levis; no other equivalent available];
stingray;
bonito;
white marlin;
bluefin tuna;
skipjack tuna;
bluefin tuna, tunny [distinction between 22. and 24. uncertain; 24 is
literally "black tune];
.
•
whales;
swordfish;
great black-backed gulls;
blue nurse shark;
cat (carpet) shark;
whale shark;
smooth hound [Mustelus mustelusl;
smooth hammerhead shark;
southern spiny dogfish [Squalus fernandinus];
black sea bass;
banded sea perch [Dules auriga];
harvestfish [direct transliteration from original Russian];
butterfish [Palinurichthys pringlei];
Cynoscyon [spelling uncertain; suitable equivalent not available;
suggested rendition only of original Russian Tsinastsion];
tautog (blackfish) [Tautoga onitisl;
pejerrey (kingfish) [Basilichthys];
tautog pautoga onitis] (American wrasse) [distinction between 39. and
41. uncertain];
fourspot flounder;
(common) hake;
stenotomus [transliterated directly from original Russian];
mackerel;
alewife;
Atlantic saury/skipper/billfish pcomberesox saurus];
herring;
anchovies;
lanternfishes;
sagittas [taken directly from original Russian sagitty];
fish larvae;
Stomatopoda;
Anomura;
•
crabs;
5
56. shrimp;
57. Hyperiidae [suitable equivalent not available; taken directly from
original Russian Giperiidy];
58. Polychaeta;
59. crabs;
60. euphausiids;
61. [illegible];
62. [illegible];
63. lobsters;
64. Ilysidacea;
65. crab larvae;
66. gammarids;
67. predatory copepods.
The squid in turn is one of the most important food items for predators
included in the third to fourth trophic levels. To these we may assign many
representatives of the cods, skates, tautogs (blackfishes), tunas, sharks
and other fish. In addition, we have often observed longfin squid feeding
on different species of seabirds. Vinogradov's observations [3] have also
confirmed this fact.
We have sometimes noted bottlenose whales (Hyperoodon)
in areas of accumulations of longfin squid. We know that
/172
Loligo eat small
dolphins, including harbour porpoiseâ Phocaena phocaena (oral communication
by K.N. Nesisa). Hence we have also included whales in the food chain. Thus
the longfin squid is in food interrelationships with animals on at least four
trophic levels. We shall now look at the nature of these interrelationships
on each of the levels.
The interrelationships of the longfin squid with other organisms on
the first three trophic levels involve its predatoriness and are primarily
vertical. The algae found in squid stomachs may be viewed as chance food
items; hence the relation of squids to plant organisms is most probably
casual.
The organisms on the first trophic level of consumed items* (copepods,
euphausiids and juvenile crustaceans) are the basic food items of small (2 to 10 cm)
*Translator's note. "Consumed items" is a suggested *rendition of the original
Russian noun konsument, for which no other more suitable equivalent is available.
6
Table 1
List of Squid Food Items
T a 6 .1 u it a
1
Ctiicoi *11110TIlbtX, turraiontuxcn Hanbmapanut
Latin. naine
Scyphozoa
Loligo pealei
PyccKoe nanamie
Author
Verrill, 1882 Vovic.,
Meayahl
13013K A. H., 1972; Verrill, 1882; Williams, 1909.;
Stevenson, 1934
jt.1111111011epblil liaahmap
11Iex illecebrostis •
OdontaspiS tricuspidatus
Ginglymostorna cirratum
Rhincodon typus
Mustelus griseus
Squalus açanthias
•
Sphyrna zygaena
•
. Raja egkintcria
KopoTxoriephiri xa.nbmap
rlecvalian axy.na .
. axy.na konpa
IÇurooan axyaa
Kyuba axyaa •
Axy.na-Karpan
Axy.na-mo.nor
Lliniiooamiii exaT
Raja erinacea
• R. laevis'
06bunionenmeil cxaT
Cxar-aeanc
•
XBOCTOK0.1
Cepe6pncmil xex.
Dasyatis centroura
Merluccius bilincaris
• /1*.rop
Russian name
Our data
Haum Àaiiiii;ie
Ilepomyrrep, 1970 •
erlalutter •
Our. date
Perlgutter •
Haunt
AalllibIO
Hepamyrrep, 1970
Pericoubirt orner• BAIPT
aA.T.naliT» 1065-1966 rr.;
Builorpaaoa, • CT),J10Ba,
1971.-1972 (ro..-konbie OTnOTbl)
M..albidus
1
• Benift xex
1
Perlmutter
Pomolobus mediocris
Homo.no6ye •
Gadus morhua
Tpeexa
Melanogrammus aeglefinus
Pollachius virens
flmuua
.
Cafma
•
Kpacithlfc na.num
•'
Pericoubie orgeibi BMPT
«Ar.naur» 1965-1966 rr., .
1966-1967 rr.
orter BMPT
«AT.nairr» 1965-1966 rr.;
flep.nmyrrep, 1970
Verrill, 1882; flaunt ;tauPble
our data
Urophycis tennis
U. chuss
-
31
I-rep:imp-rep, 1970; Binio-.
rpa.lon, 1970 (roitonort
cal er); pciiconbui
BMPT «Amur>, 1965—
1966 rr.
41
POPKOBLA OVWT
5
CPTP
«06.nopcx», 1968 r.
Bittiorpa,ion, 1970 (roan-
nun °vier)
170
Translator's Note. The second column of this table has not been translated,
as it supplies merely the Russian equivalents of the Latin
naines in the first column.
7
Table 1, colitinued
JlarnucHOC uaauannc
Latin name _
Paralichthys oblon6us
Sarda sarda
Katsuwonus. pclamis
Thunnus thvnnus
T. atlanticus
Makaira albidâ
Yiphias gladius
PYCCKOC 11a.4
Ilponoi,ccnne raF .1.
lie
Kussian name
LteTLlpeXns(TlulcraA 'xaat•
6ana
I7c:I a am;r,a
,IloaocaTl,til Tyttctl.
O6LI]u10l)cnlll,ül rVlteti
ticpublit Tyncu
Seaulil atapJnnt
A•teu-pwGa
Aorop
Au
^ thor
•ilep:IMyTTCp, 1970
erl.mutter
dïCplalyTTep, 1970
Dragovich, 1969
Dragovich, 1970
Banc, 1965
o0istrnrnnmxon, 19F0; Tlep.t.MyTTep, 1970; Verrill, 1882
7 .Onnnllllxon,
1970;
pHn,1970
P.oronotus triacanthus
Peprilus alepidotus
Rocctls saxatilis
Centropristçs striatus
Tàutoba onitis
Cynoscion sp:
Pomatomus sali.atrix'
Scomber scombrus
]1lcnticirrus amcricanus
Clupea harengus
Etrunieus teres,
Larus sp. p.
,
MacnsniaR pb16a
Kpyr:isax
`
To'
T1Qp.7xyTTCp, 1970
PexJ..mutter
tleprnbIit atopcxoit oxynb
3ycn,.Ilccuc, 1971 Zuev &
3ycn, I-icctlc, 1971; Ver- '
rill, 1882
17Y6.111-rayTOra
Verrill, 1882
U,IntOcünon
Verrill, 1882
rIyq)apb
Verr ► ll, 1-882
Ctq•u6pns
Verrill, 1882
IZO.ROCATbIQ Oxylib
KopoAcncxas pw6a
Verrill, 1882
Cc.11;1b aTaanruvecxaa
Verrill, 1882
Ce.ib:kb xpyraast
Blmorpajon, 1970
Mopcxae ltaincn
Vinogradov
Nesis
Key
1.
2.
3.
4.
5.
6.
7.
Voyage report, BrIRT (large refrigerator trawler) Atlant, 1965-1966;
Vinogradov and Stulova, 1971-1972 (annual reports);
Perlmutter, 1970; Vinogradov, 1970 (annual report); voyage report, BMRT
Atlant, 1965-1966;
Voyage report, SRTR (medium fishing trawler-refrigerator).Obdorsk, 1968;
Vinogradov, 1970 (annual report);
Ovchinnikov, 1970; Perlmutter, 1970;
Ovchinnikov, 1970; Torin, 1970.
and médium-sized (il to.18 cm) squids. The food relations, therefore, between
the squid and these items are very important and are modelled on the predatorvictim pattern.
The trophic interrelations of the longfin squid with the food items
belonging to the second trophic level of consumed items, are the most complex.
The second trophic level includes the largest number of food groups: from
the predatory copepods that are the main food of the juvenile squid to the
comparatively large planktophage fish that are the chief nutrient of the
large squids.
The different groups of invertebrates and fish that enter into
extremely complex trophic interrelationships among themselves on this level,
represent important food items for different squid size groups.
By way of an example of the trophic interrelationships on this level
we may analyze the pattern of food relations between the squids and the
herring (the round and Atlantic varieties). Squid juveniles, feeding on
zooplankton (copepodé and euphausiids) may emerge as a competitor for herring
food - an example of horizontal, competitive interrelationships. It is true
that competition must slacken if these items are separated in space and time
(stocks of herring rarely,coincide with those of squid).
small- and medium-size herring.
Large squids eat
Large herring in turn feed on stocks of
juvenile àquid during.the autumn [3].
In this case we see an example of a
close inverse relationship.
The relation between the squid and the silver hake (Merluccius
bilinearis) is manifested in even more complex trophic interrelationships.
We can-conclude from an analysis of data on the feeding of the silver hake in
autumn that its main food items have been the same four groups of, animals as
for the squid:
fish, shrimp, squids and euphausiids.
Hence hake and squid
In different size groups feeding on the one item, for example euphausiids,
may find themselves in competitive interrelationships.
The.
in its voyage reports for 1965-1966 and 1966-1967
noted large hake more than 25 cm long in the Norfolk region from November to
April.
- Stocks of this item were confined to depths of 120 to 160 meters and
bottom water temperatures of 10 to 11 0 C. [Its] main food items were longfin
/173
9
squid (58%) and fish ( 42%).
Immature specimens of silver hake less than 25
cm long fed mainly on macroplankton [3] and may be considered as competitors,
for the most part, of medium-sized squid.
The winter and spring accumulations of cephalopods largely coincide
in their regions and depths with those of hake.and are often confined to the
latter's spawning places. An analysis of squid feeding has shown that juvenile
hake play a significant part in the squid diet at this time.
Thus if we allow for a certain community of migration paths, habitats
and other biological features, we may assume that both species keep together
for a prolonged portion of the year and that their various size groups enter
into complex interrelationships simultaneously both as food competitors and
as enemies.
This increases the interdependence of these species and is one
of the factors regulating their abundance.
A disruption of the dynamic
equilibrium" in the abundance of one, due either to a weak year-class, overfishing or other causes may lead to an increase in the abundance of the other.
In this case, we still do not have accurate quantitative data at our disposal,
but we should note that in recent years, with the beginning of intensive
commercial hake fishing (from 1964 onwards), catches of this species in
relation to commercial fishing effort have been declining, while squid removals have been rising (statistics show that the squid catch has increased
'
10-fold).
The increase in squid abundance has inevitably led to more intensive
consumption of this item by predators on the third and fourth trophic levels.
From the example of the trophic interrelationships of the squid with
the herring and the hake, we see that our diagram cannot reflect the entire
camplexity of the relations between the squid and its food items and the
squid and its predators.
The food chains on the [various] trophic levels
must be looked at as a food "network" of interrelationships
g31.
10
The third level of consumed items includes cods, skates, ocean perch,
tautogs (blackfish), flounders, pejerreys., cynoscyons and-others.
If the
squid and the predatory fish of this trophic level can to a certain extent
be interrelated as enemies(using the example of the hake [Merluccius]), it
/174
is evident that trophic interrelationships of this kind with predators of a
higher order (sharks, swordfish, tunas, etc.) either do not occur at all or.
are insignificant.
The main interrelation between the predators of the fourth trophic
level and the squid follows the predator-victim pattern.
Thus unidirectional interrelationships of squid with other organisms
on the predator-victim pattern are most clearly manifested between the squid
a
and the food items of the first trophic level and between the squid and the
predators that belong to the fourth trophic level, i.e., at the ends of the
food chain.
predators.
In the second case, the-squid serves as the victim for its
The trophic interrelations of the squid on the second and third
trophic levels are the most complex and can be manifested in the "predatorvictim" (vertical interrelationships), "food competitor" (horizontal interrelationships) and "enemies" (inverse interrelationships) patterns.
-Ehwing to the age peculiarities of squid feeding, the different-sized
specimens of this item may be found correspondingly on three trophic levels.
Specimens 2 to 10 cm long, feeding mainly on plankton (copepods and euphausiids)
belong to the second troph'ic level; those 11 to 18 cm long (that feed on
planktophages) belong to the third trophic level, and large squids more than
18 cm long, which feed mainly on fish and squid of the same species, should be
assigned to the fourth trophic level. Since a squid population includes all
size groupings throughout the year, the types of trophic interrelationships
that we have described occur the year round.
11
Naturally, because of the seasonal variability of squid feeding and
•
the seasonal squid migrations, the structure of trophic interrelationships
changes as well. We still do not have [enough] information, however, to
plot feeding charts for individual seasons.
As regards the role of squid in the transmission of energy to higher
trophic levels, the simplest trophic chain is the following: phytoplankton
phytophages +squid +predators
on the fourth trophic level.
In this case,
there is no expenditure of energy on the third trophic level. We have no
doubt that energy enters the final chain in such a pattern, but in view of
the complexity of the trophic interrelationships and the different kinds of
. adaptations of the organisms, the scheme shown above of squid food interrelationships with organisms on four trophic levels is more common.
Thus the longfin squid represents one of the most important intermediate links in the transformation of organic matter between organisms on
the first two trophic levels and animals on the third and fourth trophic
levels.
The organisms belonging to the last links in the food chain are,
as a rule, large predators and valuable commercial items.
Conclusions
1.
--- •
On the basis of an analysis of the stomach contents of squids and of
animals that have fed on them, we have prepared a chart of the food interrelationships at different trophic levels.
2. Owing to the age variability of the nature bf their feeding, the
different squid size groups may be assigned to three trophic levels.
3. Within the trophic chains of shelf hydrobionts, squids represent an
important connecting link in the transmission of energy from the consumed
items of the first order to the animals of higher trophic levels.
/175
12
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. •
6