158 
Fishery Bulletin 116(2) 
was represented by oogonia, primary growth oocytes, 
and cortical alveoli oocytes in all examined ovaries. 
Values of F abs ranged between 24,240 and 63,756 
oocytes per female with a mean of 42,187 oocytes per 
female (SD10,246). The F rel values ranged between 21 
and 67 oocytes/g of body weight, with a mean of 36 
oocytes/g of body weight (SD 9). 
There was a stronger correlation between fecun¬ 
dity and body weight (regression equation: y=25.187x 
+ 10,365, coefficient of determination [r 2 ]=0.73 for F abs ; 
y=55.454x+24,823, r 2 =0.76 for F rel ) than between fe¬ 
cundity and length (y=194.35x-15,374, r 2 = 0.62 for F abs ; 
y=469.3x-45,013, r 2 =0.65 for F re i). 
Discussion 
The results of our study prove that the smooth lump¬ 
sucker is an iteroparous, total spawning species ac¬ 
cording to the classifications of Murua and Saborido- 
Rey (2003), McBride et al. (2015), and Pavlov and 
Emel’yanova (2016). This reproductive strategy means 
that females of this species are multiple spawners and 
do not die after breeding, in contrast to previous opin¬ 
ions (Vinogradov, 1950; Fadeev, 2005). Some authors 
(Ilynsky and Radchenko, 1992; Orlov and Tokranov, 
2008) have questioned this assertion, suggesting that 
a proportion of the females survive and are involved 
in repeated spawning. Although the scientific literature 
documents that a large number of dead smooth lump¬ 
sucker are found after spawning, these dead fish are 
actually mostly males that were protecting egg clusters 
(Ul’chenko and Orlov, 2001). In the closely related spe¬ 
cies of the lumpfish, females survive after spawning, 
exit the spawning grounds, and return offshore when 
oviposition is complete (Cox and Anderson, 1922; Ken¬ 
nedy et al., 2015). However, for the lumpfish, the re¬ 
sults of Bagge 2 and Kasper et al. (2014) both indicate 
low levels of postspawning survival. 
The presence of several clearly grouped oocytes in 
ovaries (Fig. 3) indicated discontinuous oogenesis (de¬ 
terminate fecundity) and group-synchronous develop¬ 
ment of vitellogenic oocytes; oocytes were released one 
during the spawning season (total spawning). Similar 
reproductive characteristics have been observed in 
other teleost fishes of the families Clupeidae, Cyprini- 
dae, Salmonidae, Percidae, Scorpaenidae, and Cottidae 
(Munoz et al., 2002; Petersen et al., 2004; TenBrink 
and Aydin 3 ; McBride et al., 2015). In contrast, another 
member of the family Cyclopteridae, Lethotremus awae, 
is a multiple spawner during every mating season and 
is characterized by asynchronous development of vitel¬ 
logenic oocytes (Abe and Sato, 2009) — a reproductive 
2 Bagge, O. 1967. Some preliminary results from tagging 
of the lumpsucker (Cyclopterus lumpus) 1966. ICES CM 
1967/F:23, 7 p. [Available from website.] 
3 TenBrink, T., and K. Aydin. 2009. Life history traits of 
sculpins in the eastern Bering Sea and Aleutian Islands. Fi¬ 
nal Report 628, 215 p. [Available from the North Pac. Res. 
Board, 1007 W. 3 rd Ave., Ste. 100, Anchorage, AK 99501.] 
strategy that may be attributed to the more southern 
range of this species: eastern coast of central Japan 
and Yellow Sea near Chefoo (Pavlov and Emel’yanova, 
2016). 
The ovaries of the smooth lumpsucker are character¬ 
ized by the absence of sperm in all gonads, indicating 
gonochorism, oviparity, and external fertilization (Dev¬ 
lin and Nagahama, 2002). The size of mature oocytes 
and thick zona radiata are similar to those of some 
members of the families Salmonidae, Pleuronectidae, 
Hexagrammidae, Hemitripteridae, Cottidae, and Clu¬ 
peidae, which have adhesive demersal eggs that are 
deposited on a variety of substrates. A thick envelope 
protects these demersal eggs from mechanical dam¬ 
age during development in the coastal zone (Warfel 
and Merriman, 1944; Ivankov and Kurdyayeva, 1973; 
Stehr and Hawkes, 1979; Markevich, 2000; Kolpakov 
and Dolganova, 2006; TenBrink and Buckley, 2013). Oil 
droplets appear in the cytoplasm of chromatin nucleo¬ 
lar oocytes and are present until the embryonic stage 
(Kyushin, 1975). 
We found that smooth lumpsuckers from the Sea 
of Okhotsk and near the southern Kuril Islands had 
absolute fecundity values less than or equal to those 
reported elsewhere. Kyushin (1975) reported absolute 
fecundity values ranging from 45,500 to 80,000 eggs for 
smooth lumpsucker collected in February off the Shik- 
abe coast (Funka Bay, southern Hokkaido, Japan). Ab¬ 
solute fecundity values varying from 30,000 to 50,000 
eggs were reported off the southeastern coast of the 
Kamchatka Peninsula (Vinogradov, 1950). 
Our study has extended current knowledge on oo¬ 
genesis, reproductive strategy, fecundity and has 
identified specific features of the reproduction of the 
smooth lumpsucker. Snailfishes (Liparidae) have some 
features, such as large eggs, low fecundity, the laying 
of eggs in sheltered locations with the use of the fe¬ 
male ovipositor, and possibly the protection of clutches 
in species having a large sucking disk (Rass, 1950; 
Stein, 1980; Chernova, 2014), that are similar to fea¬ 
tures of the smooth lumpsucker, but members of Li¬ 
paridae have not been studied in detail. The data from 
our study may help to elucidate the reproductive biol¬ 
ogy of this closely related group of fishes. 
Acknowledgments 
We wish to recognize the contributions of the staff at 
the Pacific Research Fisheries Centre (TINRO-Centre) 
for assistance with collecting specimens. We thank the 
International Fisheries Section of the American Fisher¬ 
ies Society for their editorial assistance with the Eng¬ 
lish language. The article of A. Orlov in preparation of 
this contribution was partially supported by the Rus¬ 
sian Fund of Basic Research (grant no. 16-04-00456). 
