ISSN 0704-3716
Canadian Translation of Fisheries and Aquatic Sciences
No. 5018
1 Fii.:'H n & 0 c ea n s 1î Y Ë ! ,• Some data on the systematic position and biology oe a , Pacific member of the genus Reinhardtius Gill 5 i OCT 27 i_qp3
M.F. Vernidub, and K.I. Panin
Original title: Nekotoryye dannyye o sistematicheskom polozhenii i biologii tikhookeanskogo predstavitelya Reinhardtius Gill
In: Uch. zap. LGU (Scientific Journal of Leningrad State University) 15: 250-272, 1937
Original language: Russian
Available from: Canada Institute for Scientific and Technical Information National Research Council Ottawa, Ontario, Canada KlA 0S2
1983
35 typescript pages 4. ISSN 0704-3716
Canadian Translation of Fisheries and Aquatic Sciences
No. 5018
Sonie data on the systematic position and biology of a Pacific member of the genus Reinhardtius Gill
M.F. Vernidub, and K.I. Panin
Original title : Nekotoryye dannyye o sistematicheskom polozhenii i biologii tikhookeanskogo predstavitelya Reinhardtius Gill
In: Uch. Zap. LGU (Scientific Journal of Leningrad State University) 15: 250-272, 1937
Original language: Russian
Available from: Canada Institute for Scientific and Technical Information National Research Council Ottawa, Ontario, Canada KlA 0S2
1983
35 typescript pages • Secretary Secrétariat of State d'État
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Russian En g li s h Author - Auteur M.F. Vernidub and K.I. Panin
Title in English or French - Titre anglais ou français Some data on the systematic position and biology of a Pacific member of the genus Reinhardtius Gill.
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Nekotoryye dannyye o sistematicheskom polozhenii i biologii tikhookeanskogo predstavitelya Reinhardtius Gill.
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Uchonyye zapiski LGU (Scientific Journal of Leningrad State University), 1937, No. 15, p.250-272 ( 5-<0 ) Some data on the systematic position and biology of
a Pacific member of the germs Reinhardtius nil' by M.F. Vernidub and K.I. Panin
The so—called "Pacific black halibut", which is similar to
the Greenland halibut Reinhardtius hippoglossoides Walb., was
detected in a comparatively large number during an expedition of
the Pacific Institute of Fisheries (Vladivostok) in the Sea of
Okhotsk and the Bering Sea in 1931-1932.
The literature contains hardly any data on the black halibut
that inhabits the waters of these two seas. There is only one
E mention of the discovery of a Reinhardtius species in Sagami Bay
t VISE n (Japan, Pacific coast) where only one 375 mm specimen was caught E o RE leme SLATION (stuffed specimen, No. 456, Tokyo Museum). In 1897, it was de- o u se
scribed by Ischikawa and Matsuura as Hippoglossus granlandicus. n TRAN io o
e After studying this specimen, Jordan and Snyder described TION NON rm DITED io u. it as a new species, Reinhardtius matsuurae, in 1901. DUC In UNE
TRA According to these authors, the new species was established
on the basis of the fact that it could be distinguished from
*The numbers in the right—hand margin are the pages of the Russian text — translator
SEC 5-25 (Rev. 82/11) Canacrâ -2--
R. hippoglossoides Walb. by a greater abundance of scales and by a number of other characters which were not indicated. Therefore, the purpose of our investigation is to clarify the systematic position of the Pacific black halibut, and on the basis of the data available to us, to characterize it from the biological and
commercial point of view.
Our research was based on our own collections and observations, as well as on the collections of other expeditions of the Pacific
Institute of Fisheries.
We analyzed 36 specimens fixed with Formalin and alcohol (13 from the Sea of Okhotsk and 26 from the Bering Sea). In order to compare these Pacific black halibut with the Greenland halibut, we used the collections (8 specimens) of the Zoological Institute of the USSR Academy of Sciences.
The genus Reinhardtius belongs to the Hippoglossinae subfamily
(Pleuronectidae family). In many of its characters, it is inter- mediate between the Atheresthes and Hippoglossus genera of this
subfamily, which is clearly seen in table 1.
The following is a brief description of the genus.
Genus Reinhardtius Gill.
Pleuronectes Fabricius, Fauna Graenlandico, p. 163, 1870; Walbaum, Artedt piscium, p. 115, 1792. --Hippoglossus Reinhardt, Kg1.-Dansk, Vidensk. Selsk., p. 116, 1838; Günther, Cat. Fish. Brit. Mus., p. 404, 1862. --Reinhardtius Gill., Cat. Fish Coast N.A., p. 50, 1861; Gill., Proc. Ac. Nat. Sci. Phila., p. 218, 1864; Jordan a. Evermann, Bull. U.S. Nat. Mus., No. 47, p. 2610, 1898; Jordan a. Starks, Proc. U.S. Nat. Mus., v. XXXI, p. 195, 1907; Normann, Brit. Mus. Nat. Hist., V-1, 1934; Jensen, Mém. l'Acad. Roy. Sci, V-VI, No. 4, 1935. Platysomatichtys Bleeker, Bompt. Rendus Ac. Sci. Amsterdam, p. 426, 1862; Goode a. Bean, (! 51) Bull. Essex. Inst., 1879; Collett Norske Nord. Havs. Exped., p. 142, 1880; Jordan a. Goss, Rev. Flounders and Soles, p. 237, 1889; Smitt, Hist. Sci. Fish., 418, 1893. -3-
Table 1. Comparative characteristics of three related genera from the subfamily Hippoglossinae
Atheresthes Jord. et Reinhardtius Gill. Hippoglossus Cuvier. Gilb. Greenland halibut True halibut Arrowtooth flounder
Teeth large, arrow- Teeth in jaws large, not arrow-shaped, two rows shaped, two rows on on upper jaw, one row on lower jaw both jaws
Upper eye in notch of Upper eye in notch of Upper eye completely upper profile of head upper profile of head on pigmented side or on pigmented side
Gill rakers long, thin, Gill rakers massive, Gill rakers short, wide with a row of denticles short, thick, with at the base, pointed, along one edge accessory denticles on with denticles the end
Scales cycloid, quite Scales cycloid, spherical Scales cycloid, spherical large, fall off easily on body, elongated and on body, elongated and small on caudal peduncle small on caudal peduncle, very small accessory scales present Lateral line without Lateral line gently Lateral line highly arch in front sloping, without arch arched in front in front
Lower pharyngeal teeth Lower pharyngeal teeth Lower pharyngeal teeth small, in two rows very large, in a single in two rows TOW
Vertebrae (11)12+(35) Vertebrae 18+45=63 Vertebrae 16+35=51 36, 37(38)
Both eyes are found on the right side of the body. In com- parison with the flounders, its body is elongated and compressed from both sides. The greatest thickness of the body is equal to approximatley 1/3 of its greatest depth. The head is large, ap- proximately k of the length of the whole body. The mouth is large, the maxillary extends to the posterior margin of the eye, or beyond the eye. The teeth on the jaws are large and sharp; they are ar- ranged in two rows on the upper jaw, and there is a single row of very widely spaced teeth on the lower jaw. In addition, there are -4—
L fang—like teett... 2-3 pairs of large interiorly.d-iredtjC1 7-7—T'on the upper jaw in front. The teeth on the pigmented side and on the blind side are almost equally developed. The vomer and palatina have no teeth.
The gill rakers are short and stout, with a row of accessory den- ticles on the end. The scales are small and cycloid, spherical on the body, elongated on the caudal peduncle. The lateral line is gently sloping, with no arch in the anterior. The dorsal fin overlaps by 1/3 of the length of the upper eye along the orbital margin. There is no spine in front of or within the anal fin.
The caudal fin is slightly notched. The blind side of the body is pigmented. (252) Some data on the internal structure of Reinhardtius
The number of vertebrae is on the average greater than in other members of the subfamily Hippoglossinae, in which it varies from 46 to 51.
Due to the fact that the eye in Reinhardtius is found in the notch of the upper profile of the head, the bones of the skull and jaws are almost symmetrical; slight asymmetry is observed only in the anterior part of the skull. All of these characteristics make it similar to Atheresthes stomias Jord. a. Gilb, and the genus
Cleisthenes Jord. a. Starks (Protopsetta Schmidt), and indicate that this genus is an old one.
The upper eye of Reinhardtius lies in a totally closed bony orbit. The orbit is oval in young specimens and spherical in large ones. There are no infraorbitalia.
All of the cranial bones are unusually porous and contain cavities, which can be attributed to the deep—sea mode of life of Reinhardtius. We should also mention that not a single one of these bones is suitable for determining age.
The suboperculum has a very thin and brittle margin. There are four pairs of gill arches, and false gills are also present.
There are 7 branchiostegal rays; the urohyale is r—shaped, flat- tened from the sides, with highly developed cavities. The intes-
tine is short and there are 4 pyloric caeca.
Species of the genus Reinhardtius have long been known in
the northern waters of the Atlantic Ocean where they can be found everywhere north of Bergen to Bear Bank, including the western part of the Barents Sea, along the southern coast of Iceland and along the western coast of Greenland, including Baffin Bay. Allof
these areas -are inhabited by one and the same species, Rein- hardtius hippoglossoides Walbaum.
Systematic position of the Pacific black halibut 1 A detailed study of approximately 30 morphological characters
of Atlantic and Pacific representatives did not reveal any great
differences between them. Table 2 gives a comparison of the data
obtained for the main group of characters. The data for repre-
sentatives of the Okhotsk and Bering seas are given separately.
From this table, we can see that the Atlantic and Pacific repre-
sentatives differ significantly only in the following two charac-
ters:
1) the depth of the caudal peduncle, which is equal on the
average to 82.6% of its length in the Atlantic representatives,
1 Determination of morphological characters based on Duncker's system for flounders (1895).
-6-
Table 2. Comparison of morphological characters of the Atlantic and Pacific representatives of the genus Reinhardtius
Atlantic Ocean Sea of Okhotsk Bering Sea Morphological characters Av. 1 Maximum Av. 1 Maximum Av. Maximum Number of fish 8 13 23 Length, au 52.9 44.0-64.5 38.0 26.2-59.0 28.8 12.3-47.8
In 7,of body length Length of head 27.9 24.7-30.1 28.5 26.8-30.5 27.7 25.9-30.1 Greatest depth of body 33.8 30.5-35.5 32.0 30.3-33.4 32.1 29.6-34.4 Depth of caudal peduncle 9.2 8.6-9.7 8.3 7.6-9.2 8.1 6.5-8.9 Length of caudal peduncle 11.2 9.9-12.6 12.3 11.1-13.4 12.5 11.1-14.2 Pectoral fin of eyed side 11.8 10.8-13.3 11.3 10.0-12.7 10.8 8.3-13.0 Pectoral fin of blind side 10.8 9.9-11.9 10.4 9.2-11.1 9.85 8.2-11.1 Pelvic fin 6.45 5.5-7.6 7.1 6.5-8.0 7.1 5.3-8.0 Depth of caudal peduncle in 70 of its length 82.6 70.2-97.6 68.5 56.5-83.1 65.5 54.0-75.0
In 7. of head length
Length of snout 29.0 27.1-30.4 27.9 26.6-31.25 26.5 24.0-31.7 Postorbital region of head 60.2 58.8-61.9 59.3 57.5-61.3 59.8 52.3-63.9 Maxillary of eyed side 37.4 35.8-39.5 33.7 31.2-35.5 33.6 31.3-34.8 Maxillary of blind side 38.5 36.5-39.9 35.9 33.7-37.1 35.2 32.5-36.5 Lower jaw 52.9 51.3-55.6 50.5 48.4-52.3 49.8 47.8-54.8 Interorbital distance 14.65 13.5-15.5 14.1 12.4-15.7 11.5 7.1-13.9 Horizonal diameter of normal eye 13.9 12.9-15.1 14.2 12.5-16.1 15.3 12.3-20.9 Horizontal diameter of upper eye 15.4 14.0-16.8 15.9 13.4-19.0 16.6 13.8-20.3 Vertical diameter of normal eye 12.0 10.0-12.7 11.3 9.7-12.9 12.4 8.8-18.2 Vertical diameter of upper eye 13.2 12.4-15.0 11.6 10.0-13.0 12.4 10.9-16.7 - 7-
and to an average 68.3% and 65.6% in the Pacific representatives from the Sea of Okhotsk and the Bering Sea respectively.;
2) the length of the upper jaw on the eyed side, which is equal to not less than 35% of head length (average 37.4%) in the
Atlantic representatives, and not more than 35% of head length
(average 33.7%) in the Pacific representatives.
Insignificant differences are observed in the length of the upper jaw of the blind side and in the length of the lower jaw, which in the Pacific representatives are also slightly smaller than in the Atlantic ones.
The meristic characters are compared in table 3.
Table 3. Comparison of the meristic characters of the Atlantic and Pacific representatives of the genus Reinhardtius Number -of rays D
Number of 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 109 n rays
Atlantic 2 - 1 1 - 2 1 - - 1 (8) Ocean Sea of 1 - - 2 - - 2 - 2 2 1 1 1 - - - -(12) Okhotsk Bering Sea 1 - 1 1 4 2 2 1 1 - 3 2 1 1 - 1 - (21)
Average for Atlantic representatives - 98 rays Average for Okhotsk representatives - 95 rays Average for Bering representatives - 95 rays
Number of rays A
Number of rays 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 n
Atlantic Ocean 1 1 1 1 2 - - 1 1 8 Sea of Okhotsk - - - 2 - 1 1 2 1 1 1 1 2 - 1 13 Bering Sea 2 1 - 1 - 1 3 2 5 2 3 3 - - - 23
Average for Atlantic representatives - 74 rays Average for Ohkhotsk representatives - 72 rays Average for Bering representatives - 71 rays As we can see from the table, there is no significant dif- ference in heristic characters between the Atlantic and Pacific representatives.
Our data on both the Atlantic and Pacific representatives of (254) the genus Reinhardtius are compared with those of other authors
(Günther, 1862 and Smitt, 1893) in tables 4 and 5.
Table 4. Comparison of our data with Günther's data (1862) on Reinhardtius hippoglossoides Walb.
Atlantic Ocean Collécli -ons -of - Zoo- Günther's data Sea of Okhotsk Bering Sea logical-Museum of . USSR Academy of Sci.
av. av. av. D 100-102 98 95-109 95 88-100 95 89-103 A 75 74 69-78 72 67-78 71 74-75 Greatest depth of body 28.67 of body length 29.9 26.8-31.6 28.3 26.7-30.0 28.5 26.0-30.8 Length of head 25% of body length 24.7 21.9-26.4 25.3 23.7-26.8 24.6 23.2-26.3 Horizontal diameter of lower eye 50% of snout length 48.2 44.0-53.1 51.0 45.5-60.6 58.1 44.8-82.4 Horizontal diameter of lower eye 12.5% of head length 13.9 12.9-15.1 14.2 12.5-16.1 15.3 12.3-20.3 Length of P on eyed side 407 of head length 42.6 38.8-46.7 39.6 35.2-43.75 39.1 27.7-47.8
A comparison of our data with Günther's (1862) shows some
differences, but only very insignificant ones in the horizontal
diameter of the lower eye; in the Pacific representatives, it
is greater than in the Atlantic ones.
A comparison with Smitt's data (1893) shows an insignificant
difference in the length of the lower jaw (noted earlier), which
is smaller in the Pacific representatives. Table 5. Comparison of our data with Smitt's data (1893) for Reinhardtius hippoglossoides Walb.
Atlantic Ocean Collections of Smitt's data Sea of Okhotsk Bering Sea Zoological Museum of USSR Academy of Sciences
av. av. av. Greatest depth of body in specimens up to 18 cm, up to 25% of body length - 27.5 26.0-28.7 Greatest depth of body in specimens 40-75 cm in length, up to 26-28% of body length 29.9 26.8-31.6 28.3 26.7-30.0 28.7 26.6-30.8 Greatest thickness of body up to 31% of body depth 27.1 23.7-30.1 24.2 21.9-26.7 22.2 16.9-28.8 Greatest depth of body up to 297e of body length 27.2 25.2-28.8 26.0 24.5-27.8 25.3 21.9-26.8 Length of lower jaw greater than 50% of head length 52.9 51.3-55.6 50.5 48.4-52.3 49.8 47.8-54.8
Thus, our study of the morphological and meristic characters
of the Atlantic and Pacific representatives of the genus Reinhard-
tius did not reveal any great differences between them, i.e. dif-
ferences of species importance. Significant differemces were
observed only in two of the characters associated with migration
and feeding, namely, 1) the length of the lower jaw is smaller
in the Pacific representatives, and 2) the depth of the tail is
smaller in the Pacific representatives as compared with the
Atlantic ones. -1 0-
(255) On the basis of all this, we believe that the differences between the Atlantic and Pacific representatives of the genus
Reinhardtius lie within the limits of a species/ and are subspecific.
The Pacific form should be regarded as a subspecies of the
Atlantic species Reinhardtius hippoglossoides Walb.
We have already mentioned that only one specimen described by Jordan and Snyder (1901) as the new species Reinhardtius matsu- urae was known from the northern waters of the Pacific Ocean.
According to their description, it is very similar to our specimens
from the Okhotsk and Bering seas. Proceeding from this, we believe
that the Pacific form should be referred to as Reinhardtius hippo- glossoides matsuurae Jordan a. Snyder.
Characteristics of Reinhardtius hippoglossoides
matsuurae Jord. a Snyd.
Hippoglossus groenlandicus Ishikawa a. Matsuura, Prel-Cat. 1897, p. 25. Reinhardtius matsuurae Jordan a. Snyder, Journ. Coll. Sci. Imp. Univ., XV, 1901, p. 301; Jordan a. Starks, Proc. U.S. Nat. Mus., XXXI, 1907; Tanaka, Sh. Journ. Faculty Sci. Imp. Univ., Tokyo, V-III, 1931. Reinhardtius hippoglossoides Walb. Taranets A., Journal of the Far Eastern branch of the USSR Academy of Sciences, No. 2-3, 1933; Schmidt, P. Proc. 5th Pac. Sci. Con- gress, V-5, 1934; Andriyashev A. USSR Marine Research, No. 22, 1935; Andriyashev A. USSR Marine Research, No. 25, 1937.
D=88-103, A=64-78, P=14-15 11 - approximately 120, Br. I, 13-15 Number of vertebrae 18+45
Morphological characters In % of body length: Length of head 25.9-30.5, average 28.0; Length of caudal peduncle 11.1-14.2, average 12.4; Length of P on eyed side 8.3-13.0, average 10.0; Length of P on blind side 8.2-11.1, average 10.0; (256) Length of V 5.3-8.0, average 7.1; Greatest depth of body 29.6-34.4, average 32.0 Smallest depth of body 6.5-9.2, average 8.2; Depth of caudal peduncle in % of its length 54-83.1, average 67.0. In % of head length: Upper jaw of eyed side 31.2-35.5, average 33.6; Upper jaw of blind side 32.4-37.1, average 36.5; Lower jaw 47.8-54.8, average 50.0; Length of snout 24.0-31.7, average 27.0; Postorbital region of head 52.3-63.9, average 59.5; Interorbital distance 7.1-15.7, average 13.0; Horizontal diameter of lower eye 12.3-20.3, average 15.0; Horizontal diameter of upper eye 13.4-20.3, average 16.0; Vertical diameter of lower eye 8.8-18.2, average 12.0; Vertical diameter of upper eye 10.0-16.7, average 12.0.
The body is elongated and quite low. The dorsal fin overlaps by 1/3 of the upper eye along the edge of its left orbit (fig. 1 and 2). The anal fin begins under the 25-26th ray of D. There is no anal spine. The pectoral fins are of the same form and size on both sides of the body. The pelvic fins are symmetrical. The membrane connecting its rays does not extend to the end of them,
leaving the ends of the rays free. The caudal fin has a barely
perceptible notch.
Fig. 1. Reinhardtius hippoglossoides matsuurae
Fig. 2. The same from the left side -12-
(257) The entire body and head are covered with very small cycloid, tightly imbricated scales. The scales are spherical on the body, and elongated and larger on the caudal peduncle. The lateral line begins on the head behind the lower eye, slopes gently under the pectoral fins and then extends in a straight line along the median
line of the body onto the caudal fin. space The interorbital Vis flat and covered with scales. The mem- brane between the rays of the vertical fins is free of scales.
The upper jaw extends to or overlaps the vertical line of the posterior orbital margin of the lower eye. The upper eye is found in the notch of the upper profile of the head. There are teeth on the dentale and praemaxillare. The teeth on the lower jaw are in a single row; there are 11-15 large fang-like teeth on the eyed side and 8-11 weaker teeth on the blind side.
The number of teeth, their size and the spaces between them vary with age. The teeth on the upper jaw are in two rows. The outer row consists of large interiorly hooked teeth, about 25 on each side; the inner row consists of small closely spaced teeth set at a right angle to the teeth of the first row, about 50 on each side. In front, the praemaxillare bears 2-3 pairs of the
largest fang-like teeth which point into the mouth.
The teeth on the lower pharyngeal bones are arranged in a single row; they are slender, high and sharp, and number up to
12 on each side (fig. 3). The upper pharyngeal teeth are arranged in 3 rows (fig. 4). The gill rakers are short, widely spaced and massive, with a row of denticles on the end (Fig. 5). The gill rakers in young specimens are not as developed and are more widely -13-
spaced. The eyed side is light brown and grayish-brown. The *,eeg blind side is dark and bluish-gray.
Fig. 3. Lower pharyngeal teeth Fig. 4. Upper pharyngeal teeth of R. hippoglossoides matsuurae of R. hippoglossoides matsuurae Jord. a. Snyd. Jord. a. Snyd.
Fig. 5. First gill arch of R. hippoglossoides matsuurae Jord. a. Snyd.
Geographic distribution
One specimen was discovered in Sagami Bay (northern Japan) at the end of the 19th century, and one was found for the first time in the Okhotsk and Bering seas in 1931. According to 1931 and 1932 data, its distribution in these seas is as follows.
In theSea of Okhotsk, it is found along the entire 0 coast of Kamchatka from Cape Lopatka to Point Khariuzov (57 N) at a depth of 100-600 m. In the northern part of the sea, it is found within a 60-mile zone in the Koni Peninsula--Babushkin Bay (2(258) ■ area. In the central part, it is found at a depth of 592 m near
Tony Is. We have no data on the northwestern and western part of the sea. In the southwestern part of it, this form has been found off the eastern coast of Sakhalin Is. It is not encountered in the Shantar Sea.
In the Bering Sea, it has so far been discovered only in the northeastern part between Cape Navarin and Matveyev Is at a depth of 100 m and greater. It is not encountered south of Matveyev Is.
It is apparently not encountered in the northwestern part of the sea either. Here, we are obviously faced with the fact that this area of its habitat is completely isolated from the others, a fact which is impossible to explain because of insufficient research in this area.
Biological characteristics of R. hippoglossoides
matsuurae Jord. a. Snyd.
Conditions of existence
In the Okhotsk Sea, the black halibut is found at depths from
100 to 600 m, and possibly at greater depths. Almost all of the above—mentioned areas are characterized by a gradual lowering of the bottom relief, and only in the southernmost part of the Kam- chatka coast and on the eastern coast of Sakhalin Is. is there a drop in the continental terrace, beginning with a depth of 200m.
The near—bottom temperatures are above zero degrees in all of the indicated areas with the exception of the northern part
(Koni Peninsula--Babushkin Bay), and only in the northern part was the black halibut encountered at temperatures ranging from —0.5 °
° C. to +0.1
The areas with above—zero temperatures abound in thermophilic forms. They are especially abundant and increase in number in the bathyal region of the southern part of Kamchatka. This part
-probably St. Matthew Is. — transl. of the sea is most abundant in various Decapoda, Polychaeta, Echino-
dermata and various members of the Pleuronectidae and Gadidae.
All of the known habitats of the black halibut coincide mainly with the areas of above-zero temperatures, and are confined mainly
to stony-sandy bottoms near drops of the continental terrace, or
to silty-sandy bottoms at depths of 100-300m.
In the Bering Sea, the black halibut apparently inhabits only
the part adjacent to Anadyr Gulf. So far, it has been encountered
only at depths of 100-200m where the temperature varies from +1
to +2 ° C. Further research may show that it inhabits greater depths.
Below-zero temperatures have been established north of the
100m isobath, and the black halibut is apparently never encounter-
ed in this part. The region inhabited by the black halibut at
depths of 100-200m is confined to silty-sandy bottoms and abounds
in various Decapoda.
Because the black halibut is found at great depths with con-
stant above-zero temperatures, it should be assigned to the boreal
fauna, instead of the artic fauna.
Feeding
The literature contains no data on the feeding of the black
halibut. Only Jensen (1935) indicates that a Greenland halibut
(R. hippoglossoides) measuring 150-300 mm in length feeds on Gadus
saida and the young of Sebastes marinus and Pandalus borealis.
Our observations include only 25 specimens measuring more than (259) 50 cm in length. The predominant item_ in the diet of these
specimens is Theragra chalcogramma (Pall.); Lycodes are consumed
to a much smaller degree, and various crustaceans of the families -16-
Hippolytidae, Pandalidae and Crangonidae are preferred even less
According to the data of A.P. Andriyashev, exclusive- ly young Theragra chalcogramma were found in the stomachs of five specimens of different age. This type of feeding, which necessi- tates migrations irrespective of the type of bottom, is in step with the general appearance of the black halibut which has clearly defined predator features, namely 1) a strong muscular body with a well-developed caudal region; 2) an almost symmetrical mouth with well-developed teeth on the right and left strong and large jaw bones; 3) a branchial system with teeth on the branchial tubercles, and 4) a highly developed liver and short intestine.
We do not know whether there are intervals in their feeding during the year. The distribution of the sclerites in the scales is almost uniform, and if because of this we believe that the growth of the black halibut proceeds uniformly, then we should also expect its feeding to be more or less the same at all times. How- ever, we have seen otoliths with annuli, which may be an indication of the opposite, i.e. irregular feeding during different seasons of the year. Some of the otoliths had so-called "false annuli"
(true, weakly defined ones) beginning with the 4th year and on.
Perhaps, they are an indication of intervals in feeding during the spawning periods.
Reproduction and development
There are no specific data on the time and place of spawning of the Pacific black halibut in the Okhotsk and Bering seas. For the black halibut of the Sea of Okhotsk, the reports from expedi- tions of the State Hydrological Institute and the Pacific Institute of Fisheries (Generozova and Polutov) indicate that in August 1932,
females and males of every size were at the lst-2nd stage of sexual maturity, and only one female (80 cm long) was at the 5th stage
of maturity, i.e. had running eggs. For the black halibut of the
Bering Sea, we ourselves observed females and males at the 3rd-4th
and 4th stage of maturity at the beginning of September 1932, and
several male specimens were even at the 4-5th stage (the testes
contained a large amount of semen). For the same area, A.P. Andri-
yashev (1936) also indicates that adult females measuring 66-83 cm
in length had fairly large, light and transparent eggs at the 4th
stage of maturity in September. Thus, we have different data on
the stages of maturity for the two seas, and on the basis of these
data, we have no choice but to conclude that the spawning in the
Okhotsk and Bering seas takes place at different times, apparently
in July-August in the Sea of Okhotsk (which may be related to the
migration of the black halibut to depths exceeding 300 m at this
time) and in October-December in the Bering Sea.
Sexual maturity apparently sets in during the 9th-10th year
in females and slightly sooner in males. The first and second
stages of maturity are already observed in 35-50 cm specimens,
which corresponds to 5th--6th-year fish.
There are no data on the spawning grounds of the Pacific
black halibut, and so it was very interesting to read Jensen's
report (1925, 1935) that the Greenland halibut (R. hippoglossoides
Walb.), which is usually encountered at relatively small depths 260) in Baffin Bay (up to 300 m), was found spawning at depths ranging
from 384 to 1200 m. Its early larval stages were found at the same depths. Our data permit us to assume that the Pacific black
halibut probably spawns at great depths as well.
There is no data on the embryonic development of the Greenland halibut (R. hippoglossoides Walb.) either, but all of its subsequent development beginning with the early larva has been studied quite thoroughly by Schmidt (1904) and especially Jensen (1925, 1935).
Their data are of great interest to us, for the stages indicated by them can be applied in our own investigations.
According to Meek (1916), a R. hippoglossoides egg measuring
3-4 mm in diameter does not have a fat drop, and after spawning
the eggs remain in the near-bottom layer of water. Jensen (1925,
1935) also indicates that the ripe eggs of this halibut have a
diameter of 4-4.5 mm and spawning takes place at great depths.
The eggs apparently remain in the near-bottom layer up to the time
the larvae hatch out, and only later do they migrate to the upper
layers.
Jensen (1925, 1935) distinguishes the following larval stages:
1st instar - bathypelagic larvae. The larvae measure from 10
to 18 mm in length. The younger larvae have a yolk sac equal to
1/4-1/6 of the total length,and have no pigment at all; embryonic
pectoral fins are present. In the more mature larvae, the yolk is
almost resorbed, and the eyes and ventral part of the trunk is
well-pigmented (fig. 6).
Meeiemq.4.:\M!. ------
ci44;e7D7 .7. Fig. 6. Bathypelagic larva of R. hippoglossoides Walb. Length 22 mm (after Jensen, 1935) -19-
2nd instar - young pelagic larvae. These larvae measure from
16 to 24 mm in length. They are quite symmetrical, but the left
eye is somewhat higher than the right one. The body is elongated
and low; its greatest depth, including the vertical embryonic fin
folds, is equal to about 1/5 of the total length. P are large.
Rays appear in C. Mainly the posterior of the body and region of
the intestine are pigmented.
3rd instar - late pelagic larvae. These measure from 25 to
57 mm in length. At this stage, rays appear in D and A; D and A
separate. The embryonic pectoral fins are reduced. The depth of
the body and fins increases. V appears by the end of this stage,
and the left eye gradually moves higher. However, the larvae always
remains quite symmetrical and both sides are equally pigmented.
Towards the end of the stage, the pigment is abundantly distributed
in groups. Teeth develop on the praemaxillare and lower jaw begin-
ning with a body length of 33 mm (Fig. 7).
Fig. 7. Pelagic larva of R. hippoglossoides Walb. Length 32 mm (after Jensen, 1935) (261) Metamorphosis takes place when the larva has reached a
length of more than 57 mm, and ends when it is about 80 mm in
length. By this time, the left eye is in the upper profile of
the head on its right side. The larva now has scales (fig. 8).
From this point, the pigmentation on the right side of the body
increases. At the stage of metamorphosis, the larvae begin to -20-
settle on the bottom. At the pelagic stages, the larvae are car- ried by the current to different places, and so they settle on the bottom both in shallow and very deep places.
Fig. 8. Larva of R. hippoglossoides Walb. Length 65 mm (after Jensen, 1935)
Migrations
Migrations of the Pacific black halibut were observed on the western coast of Kamchatka in the summer of 1932. It was in June that this halibut was found everywhere at depths of 100-300 m, whereas in July and August it disappeared almost completely and only single individuals were caught in an otter trawl. These migrations to greater depths are apparently spawning migrations.
Other migrations by this halibut at some other time of the year are completely unknown to us.
It is also possible that the black halibut migrates to shallow-
er waters high in biomass productivity only in early summer, in which case the migrations would be related to feeding. Its early migration to greater depths is due to the nearing of the spawning
season.
The behavior of the Pacific black halibut in the Bering Sea
differs somewhat. Both in August and at the beginning of September, quite a large number of them (up to 100) were caught in an otter
trawl at depths of 100-200m (greater depths were not studied).
It is unlikely that its pre-spawning behavior would differ from
that of the Okhotsk black halibut.
Age and growth rate
The available material on age (in the form of scales and
otoliths) came from the catches of otter trawls. Seventy-nine
specimens came from the Sea of Okhotsk, and 75 from the Bering Sea.
However, a study of the scales showed that they were completely
unsuitable for determining age. Small elongated scales do not have
clearly defined annuli, and the latter do not become any clearer
after treatment with ammonium, or after staining of the scales with iron sulfate by Trempovich's method (1932). The scales
stained uniformly, which can serve as an indication of its uniform
growth. The odd form of an otolith, which is the result of its
peculiar growth mainly in length, and the splitting in its anterior
part make it brittle and very difficult to handle.
However, without preliminary processing, otoliths cannot be
used for determining age either. The most common method, polishing,
could not be used because of the brittleness of the otolith. Clari- 262) fication in glycerin and xylene did not improve the clarity of the
annuli by much; only clarification in oil of cloves with preliminary
dehydration in absolute alcohol made them much more distinct. We
therefore used this method in our investigation. The length of
time an otolith was kept in oil of cloves varied from 0.5 hr to
5 hr (increased with age). -22-
Table 6. Average lengths according to individual years for various age groups of the Pacific black halibut of the Sea of Okhotsk (cm)
.4.e...:..:,r.4,,i
tpreelieky4e4 '• - Leneh' e 0 U P . , • • 5., • • e- t.-!•7" •,' . - .!' -1/4'.--'. ii4;:e • '.71:rii' ":"1: -. - , xx. 1 ' XXI .3(XII . ix "; XII xtvi - ,...,-::-...--.-,,,,,..I. . .: • , . . :--• . .*:-.1:; 1 ,%, - - .. -..:•.: ::. :: . . ,- , - 'ef.' ..--.', .. •.:•••'::. . :T. ; • • . . : 11,6 16,8- -9,3- 7,5 11 3 9,3 16,8- 26,4 16,0 : 21,0. ' 26,4r-17,8 8.6 20,7' 20,5' - 18,6 ' 35,0-22,5 12,5 27$ r 27,0 26,3 35,0 22,5 _ 29,0 - •_28,1 [1:•24,4. 27,5 , 35,0 241,4-27,5 13,9 . 33,5. 8. 33,8 28,6 33,6 • 41,4 39,3 47,8 32,0 - 40,6 - 47,8-32,0 15,8 38,9: 40.0 38.8 . 46,2 •153,0-36,2 16,8 43,8 - 44,4 43,4 44,0 53,0 36Z e41.0: - 48,6 57.13 • 40,0 :51,0 -57,0-40,0 17,0 48,2- 45.5 43,5 55,8 61,8-43,5 18,3 ..52,1 . 51,2 . • 50,8 ,0: 53,0 61,8 18,2 56;7. , 1.1 r 57,0 65,0 46,8. 60,4 - 54,2" 54.0 15,55,8 . 50;2. :64,4 68,2-50,2 18,0 - 60,4 '57,4 59,0 68,2 17,5 . 63,9- 63,5 71,1- , 96 68,6. - . 611,6 57,2 •71,8 74,0-57,2 16,8 67,2" 64,4 66,4 74,Œ -61,0 15,2 69,8. , 69,4 76,2- 61p 74,5 76,2, 64,6 78,0 78,4-64,6 -13,8 72,7. ' , 72,3 78,4 76.2- 75,2, 80,8 : 69.0 80,4 80,8-69.0 11,8 " : 73,0 8Z5 82,8-73,Q " 9,8 79,1 78,0 82,8 - 7,5 81,2 80,4 84,8 •«76,3, . 84,3 84,4-76,3. 86,4 79,8 86,2 86,4-79,8 6,6 83,9- 83,0 , 88,6-82,6 6,0 86,2 85,5 8a6 .82,6 '88,0 - 85,0 89,6 - 90,3-85,0 .5,3 88,35 .90,3 .91.2-87,0 4,2 89,1; . . 87,0' ;91;2 89m. • 93,0: • 93,0-89,0 de 91,0 :94,8. "` 94,8: • • , 94,8
. "
r La 62 .•
Determination of age was carried out with the help of a magnifying glass.
Since the material was very limited, the yearly increment was determined by the method of inverse calculation using the Einar Lea board.
The limited material and the difficulty of age determination, especially in adult specimens, definitely increased the error in the determination of age and the growth rate significantly. Regard- less of this, the given material still cannot serve as initial data
for further investigations in this area.
*literal translation - transi.
Table 6. Average lengths according to individual years for various age groups of the Pacific black halibut of the Sea of Okhotsk (cm)