Despite the fact that the first information about representatives of ichthyofauna inhabiting Avacha Bay was obtained as far back as the 19th century (Tilesius, 1810; Pallas, 1814), only since 1930−1935 has the data of ichthyological surveys (Popov, 1933, 1935) and monitoring of seasonal changes in the ratio of fish species inhabiting the bay (Vinogradov, 1949) provided insight into the species composition of ichthyofauna in the bay. During the following period until the 1990s, purposeful faunistic studies on fish were not conducted in Avacha Bay, though the researchers of sectoral and academic institutions periodically carried out investigations there whose results supplemented the available information about the species composition of ichthyofauna (Matyushin, 1982, 1989; Vasilets et al., 1998). In the 1990s, the staff of the Kamchatka Institute of Ecology and Nature Management, Far East Branch, Russian Academy of Sciences (KIENM, presently Kamchatka Branch of Pacific Geographical Institute, Far East Branch, Russian Academy of Sciences (KB PGI FEB RAS)) captured fish with different fishing gear and investigated the intertidal zone. Generalization of the materials collected in 1990−2005 using published and inquiry data made it possible to generally characterize the modern composition of ichthyofauna in Avacha Bay and to compare it with that in the 1930s (Tokranov and Sheiko, 2015). However, the literature data on fishes inhabiting the intertidal zone are not numerous and scarce up to the present.

The aim of this study is to characterize ichthyofauna in the intertidal zone of Avacha Bay based on the generalization of all currently available information.

MATERIALS AND METHODS

The present report is based on the results of the analysis of the ichthyological collection of KB PGI FEB RAS, the published data, and the data of our surveys. In April–September 2014–2017, we monitored the species composition and abundance of fish in two areas of the intertidal zone in the northeastern part of Avacha Bay subjected to a strong anthropogenic impact (Fig. 1): the first area is located near the settlement of Seroglazka near the site of location of fishing vessels (it was studied regularly during all 4 years), the second area was located near Nikolskaya sopka in the center of the city of Petropavlovsk-Kamchatsky (surveys were conducted only in 2016–2017). Over the period of observations, more than 4700 specimens of different fishes were found and captured with hands under stones in intertidal puddles that made it possible to obtain a clear view of the modern species composition of ichthyofauna and a relative abundance of recorded species in pebble-boulder biotopes in the intertidal zone in the northeastern part of Avacha Bay.

figure 1

Fig. 1. Map of Avacha Bay and surveyed sites in the intertidal zone during different periods: (⚫) 1930−1935, (⚪) 1992−2000, (⚫) 2014−2017 ((1) near the settlement of Seroglazka, (2) near Nikolskaya sopka).

The belonging of some fish species to a particular ecological group (ichthyocene) was accepted according to Sheiko and Fedorov (2000). The degree of abundance of a particular species was determined based on the expert assessment of its occurrence in the intertidal zone: a dominant species is constantly recorded; a common species occurs periodically as solitary specimens, and a rare species is found only in some catches during the entire period of surveys. In order to have an idea for the stage of the life cycle at which each of the studied representatives of ichthyofaunal occurs in Avacha Bay, the specimens were subdivided into three groups: (1) juveniles (fingerlings and small immature fishes), (2) adults (large, mature fishes), (3) fishes of all ages. Earlier, such subdivision was used for the general analysis of the ichthyofauna composition in Avacha Bay (Tokranov and Sheiko, 2015).

RESULTS AND DISCUSSION

According to the available data, 27 species of fish from 12 families have been recorded for certain in the intertidal zone of Avacha Bay (Table 1). The highest species diversity is characteristic for representatives of the families Cottidae and Liparidae (six species each), which amount to more than 44% of the ichthyofauna in the intertidal zone. The other ten families include only one to two species each.

Table 1.  Species composition of ichthyofauna in the intertidal zone of Avacha Bay

Family, species

Ichthyocene

Abundance

Age category

(TL, mm)

Source of information

1. Salmonidae

    

1. Oncorhynchus gorbuscha

an ep

R

1 (30−40)

Collection of KB PGI;

Tokranov and Sheiko, 2015

2. O. kisutch

an ep

R

1 (30−40)

Ibid

2. Gasterosteidae

    

3. Gasterosteus aculeatus

an n

R

2 (72)

Matyushin, 1982

4. Pungitius pungitius

san

R

3 (nd)

Popov, 1933

3. Hexagrammidae

    

5. Hexagrammos octogrammus

sl

R

1 (50−70)

Collection of KB PGI; Vinogradov, 1949

6. Pleurogrammus monopterygius

el

R

1 (40)

Collection of KB PGI;

Tokranov and Sheiko, 2015

4. Cottidae

    

7. Enophrys diceraus

el

C

1 (20)

Vinogradov, 1949;

Tokranov and Sheiko, 2015

8. Megalocottus platycephalus

sl

R

1 (50)

Ibid

9. Microcottus sellaris

l

R

1 (90)

''

10. Myoxocephalus polyacanthocephalus

el

R

1 (10−70)

Collection of KB PGI;

Tokranov and Sheiko, 2015

11. M. stelleri

sl

C

1 (10−103)

Collection of KB PGI

Tokranov and Sheiko, 2015

Our data

12. Porocottus camtschaticus

sl

R

1 (20)

Tokranov and Sheiko, 2015

5. Agonidae

    

13. Hypsagonus quadricornis

el

R

1 (nd)

Vinogradov, 1949

6. Cyclopteridae

    

14. Aptocyclus ventricosus

n

R

2 (200−300)

Collection of KB PGI;

Vinogradov, 1950;

Tokranov and Sheiko, 2015

7. Liparidae

    

15. Liparis brashnikovi

sl

R

1 (<50)

Tokranov and Sheiko, 2015

16. L. callyodon

l

R

3 (<120)

Ibid

17. L. cf. kusnetzovi

l

R

2 (85)

Our data

18. L. cyclopus

el

R

2 (80−90)

Tokranov and Sheiko, 2015

19. L. miostomus

l

R

1 (<50)

Ibid

20. L. schantarensis

l

R

2 (80)

1 (30−40)

Vinogradov, 1949;

Tokranov and Sheiko, 2015

8. Bathymasteridae

    

21. Bathymaster signatus

el

R

1 (140)

Tokranov and Sheiko, 2015

9. Stichaeidae

    

22. Alectrias alectrolophus

l

N

3 (30−143)

Collection of KB PGI; Popov, 1933; Vinogradov, 1949; Matyushin, 1989; Tokranov and Sheiko, 2015; our data

23. Opistocentrus ocellatus

sl

R

1 (30−40)

Tokranov and Sheiko, 2015

10. Pholidae

    

24. Pholis fasciata

sl

R

2 (134)

Our data

25. Rhodymenichthys dolichogaster

l

C

2 (153−250)

Tokranov and Sheiko, 2015; our data

11. Ammodytidae

    

26. Ammodytes hexapterus

el

R

1 (58−123)

Matyushin, 1982

12. Pleuronectidae

    

27. Liopsetta glacialis

sl

R

2 (140)

Collection of KB PGI;

Tokranov and Sheiko, 2015

  1. Ichthyocenes: an ep—anadromous epipelagical; an n—anadromous neritic; san—semianadromous; n—neritic; l—littoral; sb—sublittoral; el—elittoral. Degree of abundance: (R) rare, (C) common, (N) numerous. Age category in samples: (1) only juveniles, (2) only adults, (3) specimens of all age groups. (TL) total length; (nd) no data.

Fishes recorded in the intertidal zone are components of seven ichthyocenes (Table 1), but the nucleus is formed by representatives of only three of them: littoral (seven species), sublittoral (eight species), and elittoral (seven species) ichthyocenes whose total proportion constitutes 81.5% of all species (Fig. 2). According to the expert assessment of the degree of abundance, most species are classified as rare (23 species or 85.2%): over the entire period of survey, only their solitary catches were recorded (Fig. 3). Three species, such as antlered sculpin Enophrys diceraus, frog sculpin Myoxocephalus stelleri, and stippled gunnel Rhodymenichthys dolichogaster, may be classified as common species (11.1%) that are captured periodically as isolated species. Only one species, stone cockscomb Alectrias alectrolophus (3.7%), is a numerous species constantly occurring in pebble-boulder biotopes in the intertidal zone. Juveniles constitute a prevailing part of fishes (17 species or 63%) recorded in the intertidal zone of Avacha Bay (Fig. 4, Table 1). Specimens of all age groups were found for three species (11%): in addition to stone cockscomb, they are ninespine stickleback Pungitius pungitius and spotted snailfish Liparis callyodon; only adult specimens were found for the other seven species (26%).

figure 2

Fig. 2. Number of species in different ichthyocenes in the intertidal zone of Avacha Bay; ichthyocenes: an ep—anadromous epipelagic; an n—anadromous neritic; san—semianadromous; n—neritic; l—littoral; sb—sublittoral; el—elittoral.

figure 3

Fig. 3. Ratio of species with a different degree of occurrence in the intertidal zone of Avacha Bay, % of the total number: () rare, (◼) common, (◻) numerous.

figure 4

Fig. 4. Ratio of species represented by specimens of different age groups in ichthyofauna in the intertidal zone of Avacha Bay, % of the total number: () juveniles, (◼) adults, (◻) specimens of all ages.

According to the data of monitoring of 2014–2017 conducted from April to September, only stone cockscomb was constantly the only representative of ichthyofauna in pebble-boulder biotopes in the intertidal zone in northeastern Avacha Bay during ebbs; its abundance was approximately 99.8% (Tokranov and Murasheva, 2016; Murasheva and Tokranov, 2017). In addition to stone cockscomb, solitary specimens of stippled gunnel and banded gunnel Pholis fasciata and juveniles of frog sculpin and Liparis cf. kusnetzovi were recorded there (Table 2).

Table 2.  Ratio of abundance of five fish species recorded during the ebb in the surveyed parts in the intertidal zone in northeastern Avacha bay in April–September 2014–2017

Parameter

Alectrias alectrolophus

Rhodymenichthys dolichogaster

Pholis fasciata

Myoxocephalus stelleri

Liparis cf. kusnetzovi

 

Near the settlement of Seroglazka, 2014−2017

Number of fishes, ind.

2647

2

1

1

1

Proportion, % of the abundance

99.811

0.075

0.038

0.038

0.038

Length (TL), mm

30−143

192−201

134

34

85

 

Near Nikolskaya sopka, 2016−2017

Number of fishes, ind.

2080

2

4

Proportion, % of the abundance

99.712

0.096

0.192

Length (TL), mm

33−134

153−193

56−103

 

Total in the both parts, 2014−2017

Number of fishes, ind.

4727

4

1

5

1

Proportion, % of the abundance

99.768

0.084

0.021

0.106

0.021

Length (TL), mm

30−143

153−201

134

34−103

85

The comparison of available data on the species composition of ichthyofauna in the intertidal zone of Avacha Bay in the 1930s with the present data makes it possible to conclude that the species diversity of fish in some parts of the intertidal zone most subjected to the anthropogenic impact reduced slightly by the beginning of the 1990s (Matyushin, 1989; Tranbenkova, 1999). Therefore, some representatives of ichthyofauna previously common in the intertidal zone (e.g., spawning specimens of smooth lumpsucker Aptocyclus ventricosus) occur solitary or are completely absent in recent years. The absence of some fish species at present in the intertidal zone that were registered in the littoral in the 1930s is probably caused by their rarity or inaccessibility of some areas occupied by port installations for investigations. On the other hand, findings of several previously unknown representatives of ichthyofauna in the 1990s–2000s (juveniles of Atka mackerel Pleurogrammus monopterygius, Kamchatka fringed sculpin Porocottus camtschaticus, spotted snailfish and smallmouth snailfish Liparis miostomus, Brashnikovi snailfish L.brashnikovi) clearly demonstrate—as it was reported early (Tokranov et al., 2000; Tokranov and Sheiko, 2002, 2015)—an insufficient knowledge of ichthyofauna both in Avacha Bay and in its intertidal zone.

Unlike the species composition, the abundance and frequency of different fish species in some parts of Avacha Bay have currently decreased compared to the 1930s. In our opinion, it is caused, first, by pollution of the intertidal zone by industrial and domestic wastes and disturbance of the natural state of many parts in the intertidal zone as a result of their anthropogenic transformation entailing destruction and complete disappearance of the belt of macrophytic algae (Klochkova and Berezovskaya, 2001), which was the habitat of quite a number of representatives of ichthyofauna, especially at early stages of ontogenesis.

CONCLUSIONS

At present, 27 fish species from 12 families have been recorded in the intertidal zone of Avacha Bay. Cottidae and Liparidae are characterized by the highest species diversity (more than 44% of recorded species); the other ten families are represented by one to two species. Fishes recorded in the intertidal zone are components of seven ichthyocenes (Table 1) though the nucleus is formed by representatives of only three of them: littoral (seven species), sublittoral (eight species) and elittoral (seven species) ichthyocenes whose total proportion constitutes 81.5% of all species. Among representatives of ichthyofauna found in the intertidal zone, 85.2% of species are rare: over the entire period of surveys, only solitary cases of their catches were recorded. Only stone cockscomb may be reasonably characterized as a dominant representative of ichthyofauna that constantly occurs in pebble-boulder biotopes of the intertidal zone. At present, its proportion reaches more than 99.8% of the abundance in some sites. Among fish specimens registered in the intertidal zone of Avacha Bay, 17 species are represented by juveniles, seven by adult specimens, and three species (ninespine stickleback, spotted snailfish, and stone cockscomb) by specimens of all age groups.