The Biology of Mesopelagic Fishes and Their Catches (1950–2018) by Commercial and Experimental Fisheries

Abstract

Following a brief review of their biology, this contribution is an attempt to provide a global overview of the catches of mesopelagic fishes (of which 2.68 million tonnes were officially reported to the FAO) throughout the world ocean from 1950 to 2018, to serve as a baseline to a future development of these fisheries. The overview is based on a thorough scanning of the literature dealing with commercial or experimental fisheries for mesopelagics and their catches, and/or the mesopelagic bycatch of other fisheries. All commercial (industrial and artisanal) fisheries for mesopelagic fishes were included, as well as experimental fisheries of which we were aware, while catches performed only to obtain scientific samples were omitted. The processes of generating bycatch and causing discards are discussed, with emphasis on Russian fisheries. From peer-reviewed and gray literature, we lifted information on mesopelagic fisheries and assembled it into one document, which we then summarized into two text tables with catch data, one by country/region, the other by species or species groups.

Full text

J. Mar. Sci. Eng. 2021, 9, 1057. https://doi.org/10.3390/jmse9101057 www.mdpi.com/journal/jmse
Article
The Biology of Mesopelagic Fishes and Their Catches (1950–
2018) by Commercial and Experimental Fisheries
Daniel Pauly 1, Chiara Piroddi 2, Lincoln Hood 3, Nicolas Bailly 1, Elaine Chu 1, Vicky Lam 1,
Evgeny A. Pakhomov 4,5, Leonid K. Pshenichnov 6, Vladimir I. Radchenko 7 and Maria Lourdes D. Palomares 1,*
1 Sea Around Us, Institute of Oceans and Fisheries, University of British Columbia, Vancouver, BC, V6T 1Z4
Canada; d.pauly@oceans.ubc.ca (D.P.); n.bailly@oceans.ubc.ca (N.B.); e.chu@oceans.ubc.ca (E.C.);
v.lam@oceans.ubc.ca (V.L.)
2 European Commission, Joint Research Centre, Ispra, 21027 Italy; chiara.piroddi@ec.europa.eu
3 Sea Around Us – Indian Ocean, Marine Futures Lab, School of Biological Sciences, University of Western
Australia, Crawley, WA, 6012, Australia; lincoln.hood@research.uwa.edu.au
4 Earth, Ocean and Atmospheric Sciences Department and the Institute for the Oceans and Fisheries, Univer-
sity of British Columbia, Vancouver, BC, V6T 1Z4, Canada; epakhomov@eoas.ubc.ca
5 Hakai Institute, Heriot Bay, BC, V0P 1H0, Canada
6 Institute of Fisheries and Ecology of the Sea (IFES) 8, Konsulskaya Str., Berdyansk, 71118, Ukraine;
lkpbikentnet@gmail.com
7 North Pacific Anadromous Fish Commission, Suite 502, 889 West Pender Street, Vancouver, BC V6C 3B2,
Canada; vlrad@npafc.org
* Correspondence: m.palomares@oceans.ubc.ca
Abstract: Following a brief review of their biology, this contribution is an attempt to provide a global
overview of the catches of mesopelagic fishes (of which 2.68 million tonnes were officially reported
to the FAO) throughout the world ocean from 1950 to 2018, to serve as a baseline to a future devel-
opment of these fisheries. The overview is based on a thorough scanning of the literature dealing
with commercial or experimental fisheries for mesopelagics and their catches, and/or the mesope-
lagic bycatch of other fisheries. All commercial (industrial and artisanal) fisheries for mesopelagic
fishes were included, as well as experimental fisheries of which we were aware, while catches per-
formed only to obtain scientific samples were omitted. The processes of generating bycatch and
causing discards are discussed, with emphasis on Russian fisheries. From peer-reviewed and gray
literature, we lifted information on mesopelagic fisheries and assembled it into one document (see
Online Supplementary Material), which we then summarized into two text tables with catch data,
one by country/region, the other by species or species groups.
Keywords: Myctophiformes; reconstructed fisheries catch; Sea Around Us; bycatch; discards; growth
Table S1. Species of fish in FishBase belonging to the Myctophiformes (n=254), Ne-
oscopelidae (n=6) and Myctophidae (n=248), and considered to contribute the bulk of mes-
opelagic fishes. Where available, the depth range (or a single depth of occurrence), maxi-
mum recorded length and trophic level are provided (see www.fishbase.org). Note: Lmax
is the maximum length in standard length (SL).
Citation: Pauly, D.; Piroddi, C.;
Hood, L.; Bailly, N.; Chu, E.; Lam,
V.; Pakhomov, E.A.; Pshenichnov,
L.K.; Radchenko, V.I.; Palomares,
M.L.D. The Biology of Mesopelagic
Fishes and Their Catches (1950–
2018) by Commercial and Experi-
mental Fisheries. J. Mar. Sci. Eng.
2021, 9, 1057.
https://doi.org/10.3390/jmse9101057
Academic Editor(s): Alexei M. Orlov
Received: 19 August 2021
Accepted: 12 September 2021
Published: 25 September 2021
Publisher’s Note: MDPI stays neu-
tral with regard to jurisdictional
claims in published maps and institu-
tional affiliations.
Copyright: © 2021 by the authors. Li-
censee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and con-
ditions of the Creative Commons At-
tribution (CC BY) license (http://crea-
tivecommons.org/licenses/by/4.0/).

J. Mar. Sci. Eng. 2021, 9, 1057 2 of 12
No.
Species
(Families, Subfamilies)
Depth range
(m)
Lmax
(SL, cm)
Trophic
level
Family Neoscopelidae
entry 1
data
data
1
Neoscopelus macrolepidotus
300–1180
25.0
4.2
2
Neoscopelus microchir
250–700
30.5
3.2
3
Neoscopelus porosus
454–642
18.3
3.6
4
Scopelengys clarkei
0–1000
--
3.2
5
Scopelengys tristis
400–1830
20.0
3.1
6
Solivomer arenidens
1241–2022
--
3.2
Family Myctophidae
Subfamily Diaphinae
7
Diaphus adenomus
180–600
18.0
3.2
8
Diaphus aliciae
489
6.0
3.1
9
Diaphus anderseni
100–560
3.2
3.1
10
Diaphus antonbruuni
500
5.5
3.1
11
Diaphus arabicus
0–468
--
3.1
12
Diaphus basileusi
120
16.4
3.2
13
Diaphus bertelseni
0–300
9.1
3.1
14
Diaphus brachycephalus
200–600
6.0
3.1
15
Diaphus burtoni
312
--
3.1
16
Diaphus chrysorhynchus
213–587
1.1
3.0
17
Diaphus coeruleus
457–549
13.7
3.9
18
Diaphus confusus
562
--
3.1
19
Diaphus dahlgreni
320
--
3.1
20
Diaphus danae
350
12.6
3.3
21
Diaphus dehaveni
247
--
3.1
22
Diaphus diadematus
350
4.2
3.1
23
Diaphus diademophilus
0–1808
4.9
3.1
24
Diaphus drachmanni
300
--
3.1
25
Diaphus dumerilii
0–805
8.7
3.0
26
Diaphus effulgens
0–6000
15.0
3.0
27
Diaphus ehrhorni
382
--
3.1
28
Diaphus faustinoi
540
--
3.1
29
Diaphus fragilis
15–1313
12.3
3.1
30
Diaphus fulgens
85–1000
4.5
3.1
31
Diaphus garmani
0–2091
6.0
3.1
32
Diaphus gigas
100–839
--
3.1
33
Diaphus handi
774
--
3.1
34
Diaphus holti
40–777
7.0
3.1
35
Diaphus hudsoni
0–840
8.4
3.3
36
Diaphus impostor
0–140
--
3.1
37
Diaphus jenseni
350–1389
5.0
3.1
38
Diaphus kapalae
0–290
--
3.1
39
Diaphus knappi
122–664
17.3
3.2
40
Diaphus kora
0–387
--
3.1
41
Diaphus kuroshio
100–1537
6.3
3.1
42
Diaphus lobatus
--
--
3.1
43
Diaphus lucidus
0–2999
11.8
3.0
44
Diaphus lucifrons
564
--
3.1
45
Diaphus luetkeni
40–750
6.0
3.8
46
Diaphus malayanus
1000–2000
4.5
3.1

J. Mar. Sci. Eng. 2021, 9, 1057 3 of 12
No.
Species
(Families, Subfamilies)
Depth range
(m)
Lmax
(SL, cm)
Trophic
level
47
Diaphus mascarensis
237–800
14.4
3.2
48
Diaphus meadi
250
5.4
3.0
49
Diaphus megalops
1–528
8.5
3.1
50
Diaphus metopoclampus
90–1085
7.5
3.3
51
Diaphus minax
476
--
3.1
52
Diaphus mollis
50–600
6.6
3.0
53
Diaphus nielseni
--
4.0
3.1
54
Diaphus ostenfeldi
350
12.0
3.2
55
Diaphus pacificus
--
--
3.1
56
Diaphus pallidus
310
--
3.1
57
Diaphus parini
320
--
3.1
58
Diaphus parri
350–1071
6.5
3.1
59
Diaphus perspicillatus
0–1500
7.1
3.1
60
Diaphus phillipsi
588–1330
7.7
3.1
61
Diaphus problematicus
40–820
10.5
3.0
62
Diaphus rafinesquii
40–2173
9.0
3.4
63
Diaphus regani
750
1.4
3.0
64
Diaphus richardsoni
350–1000
6.0
3.1
65
Diaphus rivatoni
0–152
9.0
3.1
66
Diaphus roei
558
--
3.1
67
Diaphus sagamiensis
549
--
3.1
68
Diaphus schmidti
100–1400
5.3
3.2
69
Diaphus signatus
1270
4.0
3.1
70
Diaphus similis
0–631
7.2
3.1
71
Diaphus splendidus
0–8000
9.0
3.0
72
Diaphus suborbitalis
387–1537
7.3
3.1
73
Diaphus subtilis
40–750
8.5
3.1
74
Diaphus taaningi
40–475
7.0
3.3
75
Diaphus termophilus
40–850
8.0
3.1
76
Diaphus theta
10–3400
9.3
3.2
77
Diaphus thiollierei
--
10.0
3.3
78
Diaphus trachops
100–686
6.3
3.1
79
Diaphus umbroculus
311
--
3.1
80
Diaphus vanhoeffeni
40–750
4.2
3.1
81
Diaphus watasei
100–2005
17.0
3.2
82
Diaphus whitleyi
311
--
3.1
83
Diaphus wisneri
50–375
--
3.1
84
Idiolychnus urolampus
124–582
11.0
3.2
85
Lobianchia dofleini
0–4000
5.0
3.0
86
Lobianchia gemellarii
25–800
6.0
3.0
Subfamily Gymnoscopelinae
87
Gymnoscopelus bolini
4200
28.0
3.3
88
Gymnoscopelus braueri
2700
13.2
3.2
89
Gymnoscopelus fraseri
50–250
8.8
3.2
90
Gymnoscopelus hintonoides
2200–2350
14.0
3.2
91
Gymnoscopelus microlampas
200–500
11.7
3.0
92
Gymnoscopelus nicholsi
300
15.7
3.4
93
Gymnoscopelus opisthopterus
550–900
16.2
3.3
94
Gymnoscopelus piabilis
--
14.6
3.2

J. Mar. Sci. Eng. 2021, 9, 1057 4 of 12
No.
Species
(Families, Subfamilies)
Depth range
(m)
Lmax
(SL, cm)
Trophic
level
95
Hintonia candens
--
13.0
3.2
96
Lampanyctodes hectoris
--
7.0
3.2
97
Lampichthys procerus
1–2000
8.2
3.1
98
Notoscopelus bolini
1–1300
10.2
3.1
99
Notoscopelus caudispinosus
1–360
14.0
3.2
100
Notoscopelus elongatus
45–1000
14.2
3.4
101
Notoscopelus japonicus
391–794
13.3
3.2
102
Notoscopelus kroyeri
0–1000
14.3
3.2
103
Notoscopelus resplendens
777–2121
9.5
3.0
104
Scopelopsis multipunctatus
3-2000
8.1
3.0
Subfamily Lampanyctinae
105
Bolinichthys distofax
100–690
9.0
3.1
106
Bolinichthys indicus
25–900
4.5
3.1
107
Bolinichthys longipes
50–1021
5.0
3.1
108
Bolinichthys nikolayi
25–1760
4.1
3.0
109
Bolinichthys photothorax
40–750
7.3
3.0
110
Bolinichthys pyrsobolus
60–778
9.2
3.1
111
Bolinichthys supralateralis
40–850
11.7
3.1
112
Ceratoscopelus maderensis
51–1480
8.1
3.3
113
Ceratoscopelus townsendi
100–500
15.1
3.5
114
Ceratoscopelus warmingii
391–2056
8.1
3.4
115
Lampadena anomala
330–2000
18.0
3.2
116
Lampadena atlantica
60–1000
20.0
3.2
117
Lampadena chavesi
40–800
8.0
3.1
118
Lampadena dea
1500–2390
8.9
3.1
119
Lampadena luminosa
50–1021
20.0
3.2
120
Lampadena notialis
1–800
13.9
3.2
121
Lampadena pontifex
1–750
11.0
3.1
122
Lampadena speculigera
1–1000
15.3
3.2
123
Lampadena urophaos
50–1000
20.0
3.2
124
Lampadena yaquinae
100–2056
13.0
3.2
125
Lampanyctus acanthurus
930–1537
13.0
3.3
126
Lampanyctus achirus
--
16.2
3.2
127
Lampanyctus alatus
40–1500
6.1
3.2
128
Lampanyctus ater
60–1100
14.0
3.2
129
Lampanyctus australis
--
13.1
3.3
130
Lampanyctus bristori
--
14.2
3.2
131
Lampanyctus crocodilus
1–1200
30.0
3.2
132
Lampanyctus crypticus
--
9.8
3.2
133
Lampanyctus cuprarius
40–1000
7.9
3.3
134
Lampanyctus fernae
1–750
9.1
3.2
135
Lampanyctus festivus
40–1052
13.8
3.3
136
Lampanyctus gibbsi
--
12.2
3.2
137
Lampanyctus hawaiiensis
300–850
8.1
3.1
138
Lampanyctus hubbsi
1–2500
3.0
3.1
139
Lampanyctus idostigma
100–500
9.6
3.2
140
Lampanyctus indicus
--
8.0
3.1
141
Lampanyctus intricarius
40–750
20.0
3.4
142
Lampanyctus isaacsi
0–2300
13.3
3.2

J. Mar. Sci. Eng. 2021, 9, 1057 5 of 12
No.
Species
(Families, Subfamilies)
Depth range
(m)
Lmax
(SL, cm)
Trophic
level
143
Lampanyctus iselinoides
64
--
3.2
144
Lampanyctus jordani
588–3400
14.0
3.3
145
Lampanyctus lepidolychnus
312–332
11.9
3.2
146
Lampanyctus lineatus
60–1150
23.7
3.0
147
Lampanyctus macdonaldi
60–1464
16.0
3.1
148
Lampanyctus macropterus
0–2091
6.8
3.2
149
Lampanyctus niger
100–1015
11.1
3.1
150
Lampanyctus nobilis
100–1000
12.4
3.1
151
Lampanyctus omostigma
3000
2.6
3.1
152
Lampanyctus parvicauda
100–500
--
3.2
153
Lampanyctus photonotus
40–1100
8.5
3.2
154
Lampanyctus phyllisae
--
15.1
3.2
155
Lampanyctus pusillus
40–850
4.3
3.4
156
Lampanyctus regalis
772–3400
17.2
3.2
157
Lampanyctus ritteri
20–1095
12.0
3.4
158
Lampanyctus simulator
0–500
9.3
3.2
159
Lampanyctus steinbecki
80–100
3.8
3.1
160
Lampanyctus tenuiformis
1537
15.3
3.3
161
Lampanyctus turneri
1757
7.0
3.2
162
Lampanyctus vadulus
0–370
9.9
3.2
163
Lampanyctus wisneri
600–650
8.8
3.1
164
Lepidophanes gaussi
0–850
5.0
3.1
165
Lepidophanes guentheri
40–750
7.8
3.0
166
Parvilux boschmai
--
--
3.2
167
Parvilux ingens
100–500
16.4
3.1
168
Stenobrachius leucopsarus
31–3400
10.7
3.2
169
Stenobrachius nannochir
441–3400
11.0
3.0
170
Taaningichthys bathyphilus
400–1550
8.0
3.1
171
Taaningichthys minimus
90–800
6.5
3.1
172
Taaningichthys paurolychnus
900–2000
9.5
3.2
173
Triphoturus mexicanus
25
5.7
3.3
174
Triphoturus nigrescens
100–1000
8.1
3.1
175
Triphoturus oculeum
770–3243
--
3.2
Subfamily Myctophinae
176
Benthosema fibulatum
1–2000
8.0
3.2
177
Benthosema glaciale
1–1407
10.3
3.1
178
Benthosema panamense
--
4.5
3.1
179
Benthosema pterotum
10–300
5.7
3.1
180
Benthosema suborbitale
50–2500
3.9
3.4
181
Centrobranchus andreae
650
6.5
3.4
182
Centrobranchus brevirostris
--
4.0
3.3
183
C. choerocephalus
1050
4.0
3.3
184
Centrobranchus nigroocellatus
1–700
5.0
3.4
185
Ctenoscopelus phengodes
--
9.3
3.4
186
Dasyscopelus asper
244–1948
6.5
3.7
187
Dasyscopelus obtusirostris
1–700
7.8
3.4
188
Dasyscopelus selenops
40–500
6.4
3.3
189
Dasyscopelus spinosus
1–700
9.0
3.5
190
Diogenichthys atlanticus
1–1050
2.9
3.1

J. Mar. Sci. Eng. 2021, 9, 1057 6 of 12
No.
Species
(Families, Subfamilies)
Depth range
(m)
Lmax
(SL, cm)
Trophic
level
191
Diogenichthys laternatus
1–2091
4.0
3.2
192
Diogenichthys panurgus
366
2.3
3.1
193
Electrona antarctica
1–1010
11.5
3.2
194
Electrona carlsbergi
1–1008
11.2
3.3
195
Electrona paucirastra
--
7.0
3.3
196
Electrona risso
90-1485
8.2
3.4
197
Electrona subaspera
--
12.7
3.3
198
Gonichthys barnesi
1–1000
5.0
3.2
199
Gonichthys cocco
1–1450
6.0
3.2
200
Gonichthys tenuiculus
--
4.1
3.2
201
Gonichthys venetus
--
--
3.2
202
Hygophum atratum
600–3132
4.9
3.2
203
Hygophum benoiti
51–700
5.5
3.0
204
Hygophum bruuni
--
--
3.2
205
Hygophum hanseni
57–728
6.7
3.2
206
Hygophum hygomii
1–1485
6.8
3.0
207
Hygophum macrochir
1–750
6.0
3.2
208
Hygophum proximum
1–1000
5.0
3.2
209
Hygophum reinhardtii
1–1050
6.0
3.2
210
Hygophum taaningi
250–1000
6.1
3.2
211
Krefftichthys anderssoni
2700
7.1
3.1
212
Loweina interrupta
60–800
3.9
3.2
213
Loweina rara
1–1050
4.5
3.2
214
Loweina terminata
1–825
3.0
3.1
215
Metelectrona ahlstromi
1–2000
--
3.3
216
Metelectrona herwigi
98
5.5
3.2
217
Metelectrona ventralis
0–426
10.7
3.3
218
Myctophum affine
0–600
7.9
3.0
219
Myctophum aurolaternatum
--
11.0
3.5
220
Myctophum brachygnathum
--
--
3.4
221
Myctophum fissunovi
--
7.0
3.4
222
Myctophum indicum
--
--
3.4
223
Myctophum lunatum
--
5.7
3.3
224
Myctophum lychnobium
1–1000
3.8
3.2
225
Myctophum nitidulum
412–1537
8.3
3.4
226
Myctophum orientale
--
--
3.4
227
Myctophum ovcharovi
40–90
7.2
3.4
228
Myctophum punctatum
1–1000
11.0
3.4
229
Protomyctophum andriashevi
50–332
6.0
3.4
230
Protomyctophum arcticum
90–1600
6.0
3.1
231
Protomyctophum beckeri
1–2100
3.5
3.2
232
Protomyctophum bolini
364–728
6.7
3.0
233
Protomyctophum chilense
1–400
--
3.3
234
Protomyctophum choriodon
--
9.5
4.2
235
Protomyctophum crockeri
100–500
3.7
3.2
236
Protomyctophum gemmatum
2000
8.6
3.4
237
Protomyctophum luciferum
2000
6.1
3.5
238
Protomyctophum mcginnisi
--
--
3.3
239
Protomyctophum normani
--
5.6
3.3

J. Mar. Sci. Eng. 2021, 9, 1057 7 of 12
No.
Species
(Families, Subfamilies)
Depth range
(m)
Lmax
(SL, cm)
Trophic
level
240
Protomyctophum parallelum
2500
5.0
3.3
241
P. subparallelum
350
3.6
3.2
242
Protomyctophum tenisoni
96
5.4
3.3
243
Protomyctophum thompsoni
785–1500
5.2
3.3
244
Symbolophorus barnardi
100–800
11.6
3.1
245
Symbolophorus boops
0–500
13.1
3.5
246
Symbolophorus californiensis
557–1497
11.0
3.1
247
Symbolophorus evermanni
100–500
8.0
3.4
248
Symbolophorus kreffti
1–150
11.2
3.2
249
Symbolophorus reversus
--
8.9
3.2
250
Symbolophorus rufinus
0–850
9.4
3.2
251
Symbolophorus veranyi
0–800
12.0
3.3
252
Tarletonbeania crenularis
0–710
10.4
3.1
253
Tarletonbeania taylori
0–1500
7.0
3.3
Subfamily Notolychninae
254
Notolychnus valdiviae
25–700
5.2
3.1
Quotes. The following consists of quotes with diverse information of mesopelagic fisher-
ies and their catches.
“During the late 1970s and early 1980s, the severe depletion of demersal fish stocks
(most notably Nothotenia rossii) was followed in the second half of the latter decade by
harvesting of benthopelagic species such as toothfish species with variable year class
strengths (C. gunnari) and mesopelagic species such as E. carlsbergi.[...] Economic consid-
erations effectively ended the E. carlsbergi fishery at the end of the 1991/92 season [1], while
other fishing grounds, such as the Ob and Lena Seamounts, were effectively closed from
the mid-1990s onwards” [2].
“Recently a fortuitous fishery for the lanternfish Lampanyctodes hectoris has developed
incidental to the anchovy/pilchard fishery off the western coast of South Africa [3]. An-
nual landings of lanternfishes (mostly L. hectoris) were 1,134 metric tons or 0.3 percent of
the pelagic fishery catch in this region in 1969 and increased to 42,560 metric tons or 10.45
percent of the catch in 1973” [4].
"There are reports of fishery for mesopelagics especially myctophids, the most well-
known is the purse seine fishery for Lampanyctodes hectoris off South Africa [5] and also in
erstwhile USSR where they fish off West Africa and off Southern Australia. Due to its high
lipid (wax esters) content most of the myctophids are unpalatable for consumption and is
used for the production of fish meal, fish oil and fish silage. But some species (Diaphus
coeruleus and Gymnoscopelus nicholski) have been fished for human consumption [6,7]. Dur-
ing the 70’s Gymnoscopelus bolini and G. nicholski, caught as bycatch in the Antarctic mar-
bled rock cod fishery has been smoked for human consumption. In India, however there
have been no reports of a myctophid fishery and its use for human consumption” [8].
“Commercial lanternfish fisheries include limited operations off South Africa, in the
sub-Antarctic and in the Gulf of Oman [9–12]. But majority of the myctophids are not used
for direct human consumption owing to their high lipid or wax ester content, therefore
they are used as predator fish feed, poultry feed, animal feed and crop fertilizers [8,13,14].
Exceptions to this are Diaphus coeruleus, Gymnoscopelus nicholski and G. bolini which were
considered edible in the Southwest Indian Ocean and Southern Atlantic in the late 1970s
[8,15–17]. There are no reports of human consumption of myctophids in India [8,17].
Lekshmy et al. [13] have carried out various methods for processing and utilization of
Benthosema pterotum. They have also carried out nutritional evaluation of fish meal, dry
fish and fish hydrolysate using casein protein as reference on rats for palability. However,
one cannot ignore the processing difficulties on a large scale. An industrial fishery for

J. Mar. Sci. Eng. 2021, 9, 1057 8 of 12
Lampanyctodes hectoris in South African waters closed in the mid-1980s due to processing
difficulties caused by the high oil content of the fish [17]. Interestingly, in eastern South
Atlantic, this particular species accounted for around 42,560 tones (10.45%) of pelagic
catch in 1973 [16]” [18].
“A single haul off Argentina yielded 30 tonnes (33 tons) of Diaphus dumerilii in one
hour. […]. Limited commercial exploitation occurs off South Africa, where annual purse
seine landings (mainly of Lampanyctodes hectoris) have fluctuated between 100 and 42,400
tonnes (110 to 46,725 tons). The lanternfishes are reduced to fish meal and fish oil. Because
of lanternfishes' high oil content, processing plants are forced to mix them with other spe-
cies to prevent clogging the machinery. Around South Georgia and Shag rocks, experi-
mental fishing on Electrona carlsbergi (mainly juveniles) averaged about 20,000 tonnes
(22,000 tons) per year between 1988 and 1990, but increased dramatically to 78, 488 tonnes
(86,494 tons) in 1991. The Commission for the Conservation of Antarctic Marine Living
Resources therefore introduced a 20,000 tonne (220,400 ton) TAC (total allowable catch)
for the species for the 1992 season” [19].
“During 1989–1990, 8 cruises were carried out using this vessel in the region, not only
for trial fishing but also for estimating the biomass of lantern fish (myctophids) resources”
[20].
“According to [21], fishermen in Suruga Bay who eat large quantities of Diaphus spp.
sort out and discard B. pterotum as inedible. That does not mean that this huge production
is useless; fish oil and protein have other uses than direct human consumption. Studies in
India [9,14] show that meal and hydrolysate from B. pterotum are excellent protein sup-
plements in fish and poultry feeds. These myctophids are readily fished; Norwegian re-
sults reached 100 tons hr-1 with a sonar-guided, 750 m2 (15 × 50 m) double warp trawl
(which is a seriously large piece of gear)” [22].
“Pearlside fishery of 2009 landed more than 46,000t; landing in 2010 was 18,000t and
decreased until 2013–2016 had 0 landings despite some trials” [23].
"The target species of the fishery are or have been marbled notothenia (Notothenia
rossii), mackerel icefish (Champsocephalus gunnari), grey notothenia (Lepidonotothen (= No-
tothenia) squamifrons), Günther's notothenia (Patagonotothen guntheri), sub-Antarctic lan-
ternfish (indiscriminately recorded as Electrona carlsbergi) and Patagonian toothfish (Dis-
sostichus eleginoides). […] Owing to their small size Gunther's notothenia and lanternfish
have been used for fish meal, while the other species have been fished primarily for direct
human consumption [24]. […] After the successive depletion of the demersal fish stocks,
harvesting of (benthopelagic) Patagonian toothfish and (pelagic) sub-Antarctic lanternfish
started in the second half of the 1980s […] Economical considerations prompted the ces-
sation of the fishery on Ianternfish after the 1991/92 season. […] The stock of sub-Antarctic
lanternfish has yet to be properly assessed following a tentative assessment in 1991 […],
although a substantial fishery with annual catches of several tens of thousand tonnes has
been conducted on the stock for a number of years” [25].
"After most of the demersal (bottom-dwelling) fish stocks were depleted, which hap-
pened before CCAMLR came into force, benthopelagic (living off the bottom) Patagonian
toothfish and mesopelagic (living in oceanic midwater) sub-Antarctic lanternfish began to
be harvested in the second half of the 1980s […]. By the end of the 1980s, fishing for most
species was either prohibited, as in the case of the marbled rockcod, or was limited by
total allowable catches (TACs). [...] Economic considerations prompted the cessation of
the fishery for lanternfish after the 1991/92 season. […] The Soviet Union began a trawl
fishery for lanternfish (reported indiscriminately as E. carlsbergi) in the Antarctic Polar
Front in the 1980s, with annual catches initially varying between 500 and 2,500 tonnes.
Catches increased from 1987/88 by 14,000 to 23,000–29,000 tonnes in the two subsequent
seasons, and peaked in 1990/91 (78,000 tonnes) and 1991/92 (51,000 tonnes) […]. The fish-
ery lapsed in the 1992/93 season, as it was no longer considered to be economically viable”
[26].

J. Mar. Sci. Eng. 2021, 9, 1057 9 of 12
“Iceland has in the last few years collected information on mesopelagic fish in the
Irminger Sea during their investigations on redfish and have also done some exploratory
fishing trials. In Faroese waters Russian trawlers fishing for blue whiting have occasion-
ally reported significant by-catches of mesopelagic fish, and the Faroese Fisheries Labor-
atory and the Marine Research Institute in Iceland have done some exploratory fishing,
but so far without any success” [27].
"Since 2002, the Federation of Vessel owners, in cooperation with the Marine Re-
search Institute in Reykjavík have conducted several experimental cruises. So far, none of
the trials have resulted in commercially exploitable catches. The experiments were per-
formed along the Reykjanes Ridge with commercial vessels, using a Gloria #1280 type
trawl. Modifications was made on the belly part and the cod end had 9 mm mesh size. In
summary there were low catch rate in all hauls, but also low acoustic recordings during
the surveys, according to the fishermen. Highest catch rate during these experiments was
3 t/h of Maurolicus muelleri” [27].
“In the Gulf of Oman, the only myctophid present is Benthosema pterotum and Iranian
fishers have started a commercial fishery for myctophids in their part of the Gulf of Oman
[28].
“In spite of its abundance in world oceans, currently only a few commercial mycto-
phid fisheries exist, which include limited operations off South Africa, in the sub- Antarc-
tic, and in the Gulf of Oman [5,19,29]. Global catch of myctophids during 1970–2010, var-
ied between a few tonnes to a maximum of 42,400 t reported during 1973 [30]. Though not
commercially exploited in India, these resources have been reported as bycatch of deep-
sea shrimp trawlers operating from southwest coast of India [31–33]. [It was reported that]
the annual catch of myctophids during 2010-11 was 2972 t and the catch was supported
mainly by five species viz., Diaphus watasei, D. garmani, Benthosema fibulatum, Myctophum
obtusirostre and Neoscopilus microchir. Boopendranath et al. [34] reported the annual catch
of myctophids, caught as bycatch in the deep-sea shrimp trawlers operating off southwest
coast of India, as 3676 t, with a catch rate of 19.87 kg h-1” [35].
“Myctophids are fairly abundant in Philippine waters, but are rarely caught by fish-
ermen except when they are attracted by light at night in the open seas” [36].
"Myctophids form bycatch in deep sea shrimp trawls with an annual average catch
of 2668 t during 2009–2011 in Kerala coast. Fishery occurred almost round the year with
peak during November - February. […] Along the south-west coast of India, lantern fish
(Order Myctophiformes) forms a major portion (20–35%) of the bycatch in the deep-sea
shrimp trawls [37]. These fishes, when landed are mostly used for fishmeal or manure
production” [33].
“Fishermen in Suruga Bay, Central Japan used Diaphus spp. as food [21]. Commercial
fishery for Diaphus coeruleus and Gymnoscopelus nicholski (edible species) in the south-west
Indian Ocean and southern Atlantic began in 1977 and catch by former USSR countries
reached 51,680 t in 1992, after which the fishery ceased due to decrease in catch. Despite
this, the Commission for Conservation of Antarctic Marine Living Resources (CCAMLR)
still permits Total Allowable Catch (TAC) of 200,000 t for this resource from the area under
its jurisdiction. Industrial purse seine fishery for Lampanyctodes hectoris was developed in
South African waters and closed in the mid-1980s due to processing difficulties caused by
the high oil content in the fish [17]. Lanternfishes are harvested commercially only off
South Africa and in the sub-Antarctic [19,38] […] Catch comprised of five species viz.,
Diaphus watasei (74.23%), Neoscopilus microchir (20.57%), Benthosema fibulatum (1.94%), Di-
aphus garmani (1.69%) and Myctophum obtusirostre (1.58%) […] D. watasei and N. microchir
were available round the year whereas, other species occurred only seasonally. D. watasei
was found to be dominant among the myctophids” [33].
“After a long period of high expectations, a commercial fishery for these mesopelagic
fishes was initiated in the Persian side of the Oman Sea” [39].

J. Mar. Sci. Eng. 2021, 9, 1057 10 of 12
“The federal government has prepared a draft Deep-Sea Fishing Policy for issuance
of 50 Licenses for Tuna long Liners, Squid Jigger, Mesopelagic fishing to foreign flagged
vessels and 6000 licenses to local fishing vessels” [40].
“Management measures: (1) TAC combined for lantern and lightfish: 50,000 t; (2)
Minimum mesh size of 28 mm; (3) Sardine bycatch limitation (anchovy-directed opera-
tions); (4) Closed season from 1 November to 14 January; (5) ‘Landings monitored and
estimated at factory landing sites” [41].
“During commercial fishing trials in 1995–1998, using a pelagic trawl with cod-end
mesh size of 10 mm, the average catch was between 24 and 28 t day-1 in Iranian waters
[…]. During trial commercial fishing in Oman waters in 1996, total monthly catches of
myctophids for the months of March, April and May were 446, 1563 and 1273 t, respec-
tively. Over 123 fishing days this gave an average catch of 20 t day)1. However, catches
declined during early summer and the trial was therefore discontinued” [42].
“[A] fishery for two species of myctophids which are considered edible viz., Diaphus
coeruleus and Gymnoscopelus nicholski existed in the Southwest Indian Ocean and Southern
Atlantic during 1977–1992 and catches up to 51,680 t has been reported in 1992. Shotton
[43] has reported regarding an industrial purse seine fishery for Lampanyctodes hectoris in
South African waters which was closed in the mid-1980s due to processing difficulties
caused by the high oil content of the fish. Qeshm Fish Process Company in Iran produces
fish meal and oil, mainly based on lantern fish and the plant has a nominal capacity of
3,600 tons of lanternfish per day, out of which approximately 700 tons of fish meal and 70
tons of fish oil are obtained (QFPCO 2011)” [44].
“Special attention should be paid here to numerous species from the group of Mycto-
phidae, pelagic Gobidae and other snail-sized fish (below 10 cm in length) forming dense
shoals identified as sound scattering layers. The exploitation of their stocks was begun by
the Republic of South Africa (Divisions 1.4 and 1.6) when 11- and 12.7-mm mesh purse
seines were introduced, although these fish inhabit the whole ICSEAF Area. At first their
catches were quite substantial, equaling, for instance, 42,000 tons (mostly L. hectoris) for
Division 1.6 in 1973. Between 1978 and 1983, the catches considerably, not exceeding 1,000
tons, with the exception of 1979 and 1981, when 10,000 tons were taken [5] (Newman,
1977)” [45].
“Lampanyctodes hectoris have accounted for 0.3–10.45% (1134–42,560 metric tons) of
the total fish landed by South African pelagic fishing boats operating in the cold water off
the west coast of south Africa during the years 1969–1973 [3]. Approximately 15 tons of
another species, Diaphus dumerilii, were taken in a single haul at a depth of 260–265 m off
Uruguay [46]” [38].
“Myctophids have been targeted by commercial fisheries in the Southern Ocean, no-
tably in the northern Scotia Sea area where ex-Soviet Union vessels targeted Electrona carls-
bergi at or just south of the Polar Front to the north of South Georgia [47]. Catches peaked
at around 30,000 tonnes in the 1988/89 season, with the fish converted to meal, but since
1990 there has not been a targeted fishery” [48].
“An annual PUCL for mesopelagic fish of 50,000 t was introduced in 2012, following
increased catches of lantern- and light fish by the experimental pelagic trawl fishery in
2011, when just over 7000 t of these species were landed. Since then, however, catches
have not exceeded 1000 t. It is anticipated that catches of mesopelagic fish may again in-
crease in 2014 with resumption of this experiment” [49].
“While under limited commercial exploitation in the southern Benguela, the meso-
pelagic catch has historically fluctuated between 100 and 42,400 tonnes and has accounted
for some 10% of the total annual catch made by South Africa’s small pelagic fishery in
some years […] However, the fishery intermittently closed during mid-80s due to pro-
cessing difficulties caused by the high wax ester content of the fish […]. In addition to the
commercial purse-seine fishery, DAFF granted two-year permits in 2010 for an experi-
mental mid-water trawl fishery targeting mesopelagic and pelagic stocks. Of the total

J. Mar. Sci. Eng. 2021, 9, 1057 11 of 12
catch reported for both years combined (9486.5 tonnes), 83% consisted of L. hectoris and
4% of M. walvisensis” [50].
“Some Icelandic companies are developing the maurolic fishery (Maurolicus muelleri)
in areas south of Iceland. While he is not always successful, at the end of January, there
were several successful days before main concentrations of maurolic migrated to the west.
According to the information of the First Officer and skipper of the "Faxi RE" trawler,
the fishery began in the area of the Grindavík Deeps, then moved south of the Eldey area.
All three HB Grandi trawlers were fishing. The catches were 70 to 80 tons for long trawls.
In the same area there were 12 other vessels of other companies. The "Faxi RE" used a
midwater trawl with a small-mesh insert. But it seems that for a more successful harvest,
a smaller mesh trawl and additional knowledge will be required. The fish is small enough.
The optimal time for catching it is daytime only.” [51].
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