ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ
The hydrochemical characteristics of the Northwestern Part (NWBS) of the Black Sea offer a unique opportunity to study the development of meiofauna ecological features in hypoxic zones under field conditions. A comprehensive array of factual data allowed us to identify specific features of meiofauna...
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| author | Воробйова, Л.В. |
| author_facet | Воробйова, Л.В. |
| author_institution_txt_mv | [
{
"author": "Л.В. Воробйова",
"institution": "ДУ «Інститут морської біології НАН України»"
}
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| description | The hydrochemical characteristics of the Northwestern Part (NWBS) of the Black Sea offer a unique opportunity to study the development of meiofauna ecological features in hypoxic zones under field conditions. A comprehensive array of factual data allowed us to identify specific features of meiofauna structure development and its quantitative indicators, obtained through long-term, ongoing research. A total of 520 quantitative samples collected during regular annual scientific expeditions over more than 30 years were analyzed. Data analysis allowed us to determine the dynamics of quantitative indicators for both the general meiobenthos and its major large taxa (Foraminifera, Nematoda, Harpacticoida, Ostracoda, and Bivalvia), which are sensitive to changes in oxygen levels in the bottom layers of the northwestern shelf of the Black Sea. The quantitative indices of the aforementioned taxa were examined over a wide range of dissolved oxygen levels near the bottom in this marine region (from 0 to 12 mg·L-1 O2). The studies showed that only foraminifera maintain very high abundances even at very low dissolved oxygen levels. Crustaceans (harpacticoids and ostracods) and juvenile bivalves cannot survive under hypoxic conditions. The obtained data allowed us to determine two oxygen thresholds that determine both the biodiversity and abundance of the main representatives of the permanent and temporary components of the meiofauna. Long-term observations and analysis of the obtained data allow us to assess the environmental quality in the northwestern Black Sea using the meiobenthos. The nematode to harpacticoid abundance ratios presented in the paper reflect the structure, abundance, and biomass of the meiofauna under different oxygen conditions. He higher the values of this ratio, the worse the ecological state of the environment in the bottom layers of water. |
| doi_str_mv | 10.47143/1684-1557/2026.1.5 |
| first_indexed | 2026-06-08T01:00:18Z |
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49
МОРСЬКИЙ
ЕКОЛОГІЧНИЙ
ЖУРНАЛ
© Vorobyova L.V., 2026
UDC 591.524.11:543.272.1(262.5)(1-16)
DOI 10.47143/1684-1557/2026.1.5
INFLUENCE OF OXYGEN REGIME ON MEIOBENTHOS
OF UKRAINIAN SHELF OF THE BLACK SEA
Vorobyova L.V. – Dr., Prof., Leading Researcher
Institute of Marine Biology of the National Academy of Sciences of Ukraine
vorobyova.meio@gmail.com
The hydrochemical characteristics of the Northwestern Part (NWBS) of the Black Sea offer a unique opportunity to
study the development of meiofauna ecological features in hypoxic zones under field conditions. A comprehensive array
of factual data allowed us to identify specific features of meiofauna structure development and its quantitative indicators,
obtained through long-term, ongoing research. A total of 520 quantitative samples collected during regular annual scientific
expeditions over more than 30 years were analyzed. Data analysis allowed us to determine the dynamics of quantitative
indicators for both the general meiobenthos and its major large taxa (Foraminifera, Nematoda, Harpacticoida, Ostracoda,
and Bivalvia), which are sensitive to changes in oxygen levels in the bottom layers of the northwestern shelf of the Black
Sea. The quantitative indices of the aforementioned taxa were examined over a wide range of dissolved oxygen levels
near the bottom in this marine region (from 0 to 12 mg·L-1 O2). The studies showed that only foraminifera maintain
very high abundances even at very low dissolved oxygen levels. Crustaceans (harpacticoids and ostracods) and juvenile
bivalves cannot survive under hypoxic conditions. The obtained data allowed us to determine two oxygen thresholds that
determine both the biodiversity and abundance of the main representatives of the permanent and temporary components
of the meiofauna. Long-term observations and analysis of the obtained data allow us to assess the environmental quality
in the northwestern Black Sea using the meiobenthos. The nematode to harpacticoid abundance ratios presented in
the paper reflect the structure, abundance, and biomass of the meiofauna under different oxygen conditions. He higher
the values of this ratio, the worse the ecological state of the environment in the bottom layers of water.
Key words: Black Sea, Northwestern Shelf, meiofauna, oxygen regime.
Introduction
Extreme marine environments cover more than 50%
of the Earth's surface and provide numerous opportunities
to study biological responses and adaptations of organisms
to stressful living conditions. Extreme marine environ-
ments are sometimes associated with ephemeral and unsta-
ble ecosystems, but can host abundant, often endemic
and well-adapted meiofauna species (Zeppilli et al.
2018). Oxygen is the main driver among the abiotic
parameters determining habitat conditions and the pres-
ence of meiofauna (Coull 1985). Meiobenthic organisms
have relatively large surface areas and mostly high oxy-
gen demands: only a few specialized organisms will pre-
fer hypoxic conditions (Geer 2009).
Vertical distribution patterns showed that areas
with severe oxygen deficiency in the surface sediments
contained lower meiofauna than deeper areas. Because
of their small size, high abundances, short life cycles,
meiobenthic organisms are good test objects to deter-
mine the effects of perturbations in aquatic ecosystems.
There are species of meiobenthos, which not only can
tolerate hypoxia, but also prefer the conditions of acute
hypoxia (Sergeeva and Zaika 2013).
The water area of the NWBS, in terms of its hydro-
chemical characteristics, represents a unique opportu-
nity to study the formation of ecological characteristics
of the meiofauna at hypoxic zones in field conditions.
Analysis of data for 40 years of continuous research
in this part of the sea made it possible to determine
the dynamics of quantitative indicators of the whole
meiobenthos, as well as its main large taxa under differ-
ent oxygen conditions (0–13 mg·L-1 O2) in the bottom
layers of water.
The NWBS is an area of particularly high risk
of anthropogenic damage to ecosystems. There are many
reasons for this, but the fundamental one is apparently
the unprecedented size of the specific area of the catch-
ment basin. The total area of the basins of rivers, entering
the northwestern part of the Black Sea is 1,456,000 km2,
while the water area is equal to 278,000 km2. Conse-
quently, each square kilometer of shelf water area is subject
to a load from 52 square kilometers of highly urbanized
50 ISSN 1684-1557 Морський екологічний журнал, № 1. 2026
Vorobyova L.V.
territory of the basin. Initially this ratio of areas was a pos-
itive factor mobilizing great volumes of entry of nutrients
from the hinterland of the basin. Thus, the high produc-
tivity of the shelf ecosystem was guaranteed. In condi-
tions of highly elevated technogenic chemical stress on
the watershed, it has become one of the main prerequisites
for anthropogenic degradation (Bogatova et al. 1990).
Since the mid-1980s, the main problem of the NWBS
has been associated with a constant deficiency of dis-
solved oxygen in the bottom layers of water. This was
first described in publications in 1977 (Zaitsev 1977 a, b).
Many researchers have noted that hypoxia and anoxia on
the NW shelf are associated with significantly increased
anthropogenic eutrophication (Garkavaya, Bulanaia
and Bogatova 1982). Large-scale bottom hypoxia over
several decades covered 1/3 or more of the entire NWBS
water area in the summer and partly in the autumn. This
phenomenon was especially acute in the 1980s and 1990s.
The need to oxidize large additional masses
of autochtonous organic matter in conditions
of restrained vertical water exchange usual for the warm
period of the year has determined the active formation
of wide zones of hypoxia/anoxia near bottom waters
on the shelf. This has been observed previously, but it
has been episodic-temporally and local-spatiality. Since
1972, when hypoxia was first recorded at a large scale
the oxygen deficiency in the near bottom layers has
been observed almost annually, occupying the area from
depths of 4 to 20 m between the Dnieper-Bug Estuar-
ies. Towards autumn this process spreads to a depth
of 40 m in the open sea. Recently, a tendency has been
noted for a decrease in the amount of oxygen in the near
bottom layer in summer and spreading on the hypoxia
zone. On average, in the summer period of 1988-1989,
the amount of dissolved oxygen in the near bottom layer
made up1/9 mg·l-1, degree of saturation 22%.
The localization of the benthos biomass is gener-
ally determined by the distribution of the plankton bio-
mass in the upper layers of the pelagic zone. As it is
known (Zenkevich et al. 1971), the main source of food
resources for bottom animals is the population of the sur-
face layers of the ocean. The abundance of organisms
of various meiobenthos taxa in a given biotope is deter-
mined by the presence of a sufficient amount of food
available to them and, accordingly, by a more active col-
onization of various types of substrates.
The development and functioning of meiobenthic
animals are closely dependent on biological processes
occurring in the surface layers of the pelagic zone (Coull
1999). Therefore, shifts in the structure of pelagic com-
munities subject to eutrophication have a mirror image
in the benthal. The concentrations of various com-
pounds in the upper sediment layer, the oxygen regime,
and the rate of chemical exchange between sediments
and bottom water change (Zaika 1992).
The aim of this work is to demonstrate the main
patterns of formation of the numbers of the main
taxa of meiofauna under different oxygen conditions
based on long-term research in the northwestern part
of the Black Sea.
Materials and Methods
The results obtained are an analysis of long-term
observations (1983–2018) of the formation of quanti-
tative meiobenthos parameters on the Ukrainian shelf
of the Black Sea under varying oxygen conditions.
Methods for collecting and processing samples are
described in numerous publications by the author
(Vorobyova 2000).
A total of 520 quantitative samples collected in
the northwestern part of Black Sea were included in
present analysis. Dissolved oxygen values in the bottom
water layers during the study period ranged from 0–1 to
11 mg·L-1 O2.
Results and discussion
Numerous studies have shown that the oxygen
regime is of key importance in the formation of biologi-
cal diversity and quantitative indicators of meiobenthos
representatives (Coull 1985; Zeppilli et al. 2018 et al.).
As noted above, analyses of data for 40 years
of continuous research in NWBS made it possible
to determine the dynamics of quantitative indicators
of both the general meiobenthos and its main large taxa
under different oxygen conditions (0–13 mg·L-1 O2) in
the bottom layers of water.
In the NWBS, in the zones of formation of periodic
large-scale hypoxia zones, meiobenthos is character-
ized by very low biodiversity, but the highest possible
abundance due to the mass development of small-sized
forms with short-cycle development. Scientific publica-
tions indicate that the lower level of tolerance of ben-
thic forms in the coastal zone is observed at an oxygen
concentration of less than 2 mg·L-1 O2, and this value
was taken as the boundary between normal conditions
and hypoxia (Zaika, Konovalov and Sergeeva 2011;
Rosenberg 1980). Sometimes the dissolved oxygen con-
tent of 2.8–3 mg·L-1 O2 is also considered as the bound-
ary (Modig and Olafsson 1998).
Our studies in the NWBS have established that
when the hydrochemical regime deteriorates, the dom-
inance of individual species or groups of meiobenthos
increases (Vorobyova 2000). Under current conditions,
the dominant nematode-harpacticoid complex of organ-
isms has been replaced by a Foraminifera-nematode
complex on the entire northwestern shelf. At the same
time, Foraminifera often accounted for 79.7–80%
of the total population density in the NWBS (1982–
1995). With dissolved oxygen content in the bottom
layers of up to 6–9 mg·L-1 O2, the quantitative indicators
of meiobenthos are formed by almost all groups charac-
teristic of a given community (Fig. 1, 2).
51Морський екологічний журнал, № 1. 2026 ISSN 1684-1557
Influence of oxygen regime on meiobenthos of Ukrainian shelf of the black sea
The high biomass of meiobenthos under favorable
oxygen conditions is an indicator that it is mainly
formed by crustaceans and representatives of the tem-
porary component (pseudomeiobenthos).
With low oxygen content in the meiobenthos,
small-sized individuals with a short development cycle
develop. With oxygen levels from 3.5 to 5 mg·L-1 O2,
representatives of five to seven groups are usually pres-
ent in the meiobenthos. In the meiobenthos of water
areas where the benthic ecosystem experiences signif-
icant pressure from various types of pollution, quan-
titative indicators are 87–100% formed by one or two
groups. This is especially typical for estuary areas, bays,
estuaries, and gulfs.
There are species of meiobenthos, which not only
can withstand hypoxia, but prefer the conditions of acute
hypoxia. Our long-term research in the NWBS allowed
us to obtain extensive factual material on the ecolog-
ical characteristics of meiobenthos at various oxygen
regime indicators in the bottom layers of water. From
1982 to 1998, systematic annual collection of material
(in spring, summer and autumn) was carried out to study
the meiobenthos of the Ukrainian shelf of the NWBS in
complex marine expeditions on research scientific ves-
sels (standard grid – 50 stations). From 1994 to 2017,
the reaction of the meiobenthos to the dissolved oxy-
gen content was studied using materials collected in
the OMR (Odesa Marine Region).
Foraminifera. Studies in the NWBS (Voroby-
ova 2000) showed that with repeated extensive zones
of hypoxia and anoxia, the species diversity of Foraminif-
era sharply decreases, but their population density can
reach up to 1-2 million. ind.·m-2. Scientists have known
for a long time that many bottom Foraminifera can live
in anoxic conditions (Risgaard-Petersen et al. 2006).
Research in the North Sea has shown (Moodley
and Hess 1992) that Ammonia beccarii is irregularly dis-
tributed in the subtidal sediment of the southern North
Sea, with substantial numbers occurring as deep as
35 cm below the water-sediment interface. Deep infau-
Fig. 1. Average total abundance of meiobenthos at different oxygen levels at the bottom
Fig. 2. Average total biomass of meiobenthos at different oxygen levels at the bottom
52 ISSN 1684-1557 Морський екологічний журнал, № 1. 2026
nae specimens are insensitive to oxygen concentrations,
and all specimens isolated from different depth intervals
continued their normal activities (feeding and growth)
when exposed to dysaerobic oxygen content. Specimens
of E. excavatum, Q. seminulum, and E. scabra, when
subjected to the same conditions, behave similarly to
A. beccarii. These benthic foraminifera have very low
oxygen requirements. The chambers of A. beccarii that
are formed in situ at different depth intervals in the sed-
iment have a wide range in the porosity (i.e., % of area
occupied by pores) which is adequate for gas exchange
under both high and low oxygen conditions. Foraminif-
era live at the oxic-anoxic boundary throughout the sed-
iment and therefore must occasionally be subjected to
completely anoxic conditions. A. beccarii, E. excava-
tum, and Q. seminulum actively survived at least 24 h
without oxygen, indicating that they are capable of fac-
ultative anaerobic metabolism (Moodley et al. 1997).
Many benthic Foraminifera associated with
oppressed oxygen environments retain chloroplasts.
They provide a metabolic advantage that allows
foraminifers to inhabit these habitats. Thus, it has been
suggested that many benthic foraminifers are facultative
anaerobes. It has now been established that foraminifers
can dwell in soil to a depth of 12–15 cm and periodically
transfer oxygen-free conditions (Basov and Chusid
1983; Moodley and Hess 1992). When studying
meiobenthos in the northwestern part of the Black Sea,
we recorded live Foraminifera in the thickness of the soil
up to 10 cm in water areas at depths of 25–90 m (Voro-
byova and Kulakova 2009). The analysis of long-term
data allowed us to show the dynamics of the formation
of average abundance of Foraminifera at various levels
of dissolved oxygen in the bottom layers of water (Fig. 3).
An analysis of the quantitative indicators of the main
taxa of meiobenthos, which form its total abundance
and biomass, showed the regularity of this process with
different parameters of dissolved oxygen in the bottom
layers of water.
To form the ecological characteristics of Foraminif-
era (abundance, biomass, share in the total number
of meiobenthos, etc.), there are two indicator values
of dissolved oxygen in the bottom layers of the sea,
which determine both the species diversity and the total
number of protozoa. The first, which is characteristic
of hypoxia (1–4 mg·L-1 O2), in which foraminifera are
present in the meiobenthos, capable of living at low
O2 levels. The second – 8 mg·L-1 O2 and above, at which
oxyphilic species develop.
As can be seen, a significant concentration
of Fоraminifera is confined to both very low and high
levels of dissolved oxygen. Based on the results obtained,
it can be assumed that the first peak is formed by species
that continue to develop during hypoxia. The second
peak should relate to oxyphilic species. A similar picture
is given for the Sea of Okhotsk (Saidova 1960), where
the author describes two groups of Foraminifera, one
of which forms the maximum abundance at a dissolved
oxygen content of 4‑5 mg·L-1 O2 or more, and the sec-
ond, forming the southern maximum, lives with less dis-
solved oxygen (about 2–4 mg·L-1 O2).
To determine the significance of foraminifers in
the formation of quantitative indicators of small bot-
tom invertebrate animals (meiobenthos) of the OMR
ecosystem, we calculated the average indices of por-
tion of Foraminifera in the total number of meiobenthos
under different oxygen conditions (Fig. 4).
As can be seen, from the presented figure,
foraminifers are constantly an essential component
of the meiobenthic community of eutrophic water
areas of the Black Sea. To some extent they can reflect
the quality of the marine environment in the benthal
Fig. 3. Average indices of the density of foraminiferal settlements (ind.·m-2)
under different oxygen conditions (Vorobyova 2021)
Vorobyova L.V.
53Морський екологічний журнал, № 1. 2026 ISSN 1684-1557
and the formation of quantitative indicators of the rest
of the meiobenthic community, which is valuable as
a food for young fish and some species of macrozoo-
benthos.
Nematoda. In order to clarify the distribu-
tion boundaries of life on the shelf and beyond, in
1984–1986 we collected samples at depths from 75 to
600 m, covering the lower shelf area and the upper part
of the adjacent continental slope of the western Black Sea
(Zaitsev et al. 1987). Representatives of various hyd-
robionts typical of the Black Sea meiobenthos were
found in the samples. However, while most of them
were found at depths of 100 m, nematodes of the orders
Desmoscolecida and Chromadorida were discovered by
I.I. Kulakova at the maximum depth studied – 600 m
(Data from Kulakova I.I. (Zaitsev et al. 1987). It is evi-
dent from the data presented (Table 1) that traces of oxy-
gen are still present at a depth of 203 m, and the oxy-
gen-free zone begins deeper. At present, it can be stated
that the lower boundary of the meiobenthos distribution
is 600–645 m.
We have proposed several hypotheses. One of them
is based on the following conclusions. At all deep-sea
stations, increased amounts of carotenoid pigments were
found in the surface sediment layer up to 1 cm thick data.
The main source of these pigments is phytoplankton
cells that have settled to the bottom. However, at depths
of less than 100 m, carotenoids are destroyed faster than
at 200–600 m; they accumulate, are preserved, and in
some cases form concentrations an order of magnitude
higher than in shallow, oxygen-rich shelf area.
Another reason for the introduction of free-liv-
ing marine nematodes into the anaerobic zone is their
assimilation of new food sources, such as, for exam-
ple, sulfur bacteria and newly synthesized organic sub-
stances (Galtsova 1976, 1991). Living in a new envi-
Fig. 4. Proportion (%) of Foraminifera in the total abundance (ind.·m-2)
of meiobenthos under different oxygen conditions (Vorobyova 2021)
Table 1
Vertical distribution of some deep-sea bottom animals in the Black Sea and the conditions
of their existence (Zaitsev et al. 1987)
D
ep
th
, m
t, ºС S, ‰ pH O2,
mg L⁻¹
H2S,
ml·l-1
Pigments in the surface
layer of sediments, mg·g-2
Nematodа
Number of
species
Number
ind.·m-2
76 7.68 17.86 8.15 4.90 – 0.6 10 12 500
100 7.90 18.02 8.08 4.13 – 0.5–0.9 12 2 400
145 8.55 19.08 7.80 0.27 0.02 0.2–0.3 10 1 900
203 8.73 20.57 7.80 0.10 0.20 7.0 2 1 500
250 9.24 20.89 7.72 0 0.30 2.1 2 500
307 8.83 21.01 7.80 0 0.66 5.9 3 1 500
400 8.90 21.22 7.82 0 2.50 2.1–3.5 2 400
500 8.91 21.30 7.75 0 3.20 2.3 3 500
600 8.90 21.30 7.75 0 3.60 4.1–5.1 2 400
Influence of oxygen regime on meiobenthos of Ukrainian shelf of the black sea
54 ISSN 1684-1557 Морський екологічний журнал, № 1. 2026
ronment led to adaptations. As V.V. Galtsova points
out, they developed a new type of “oxygen-free respira-
tion”, and at the same time, thiobiont forms are capable
of switching to the usual aerobic type of respiration in
an oxygen environment.
In relation to oxygen, four groups of nematodes
have been identified (Keegan, Boaden and Ceidigh
1977): 1st – oxybionts, 2nd – inhabitants of the transi-
tion layer with a reduced content of free oxygen, 3rd –
anoxybionts and 4th – facultative anoxybionts. Repre-
sentatives of the Chromadorida and Desmoscolicida
orders, which we found at a depth of 250–600 m, belong
to the 3rd group.
For more than a century, the world of science has
considered the anoxic zone of the Black Sea to be azoic,
or lifeless. But data (Sergeeva 1988; Sergeeva et al.
2011) indicate that the deep-water bottom sediments
of the Black Sea, which possesses permanent hydrogen
sulfide pollution, are the natural habitats for some spe-
cies of eukaryotic fauna (Protozoa and Metazoa).
Therefore, the completion of the work on the nem-
atodes, the most abundant group metazoan meioben-
thos will reveal interesting details of their distribution
in the gradient of hypoxia observed with increasing
depth of the Black Sea. Now it was detected at depths
of 120–240 m more than 40 forms of nematodes
belonging to one order Desmoscolecida. The maximum
size of the group of nematodes honored at a depth of
120–130 m. Permanent hypoxic zone of the Black Sea
is an interesting research polygon. Later, in 1987, rep-
resentatives of meiobenthos (9 species) were discovered
at a depth of 300–645 m (Mikhailova and Sergeeva
1987; Sergeeva 1988).
There is information (Sergeeva and Kolesnikova
1996) about the findings of nematodes, harpacticoids
and other representatives of the meiobenthos at a depth
of more than 600 m, up to 2000 m. The authors can-
not state whether these organisms were alive at the time
of their discovery or whether they had not yet under-
gone the process of decomposition.
Our studies have shown that for nematodes, as well
as Foraminifera, there are two maximum oxygen levels
(4 and 8 mg·L-1 O2), which determine the formation
of the density of nematode taxocene settlements. The max-
imum average number of nematodes was noted at 1 mg ·
mg·L-1 O2, 207,056.3 ± 51,388.2 ind.·m-2, the minimum –
at 4 mg·L-1 O2 – 47,854.5 ± 7,255.7 ind.·m-2 (Fig. 5).
Their share in the total number of organisms
at oxygen concentration below 4 mg·L-1 O2 varied from
31 to 68% and averaged 47.3%. The number of nema-
todes increases due to the inhabitants of the transition
layer with a reduced content of free oxygen (the sec-
ond group). The maximum indicator is 157,108.1 ±
24,745.0 ind.·m-2.
The minimum density of nematodes was character-
istic for 8 mg·L-1 O2 (101,572.1 ± 14,527.3 ind.·m-2).
The proportion of worms in the total number of meioben-
thos varied from 54 to 64.5%. At high oxygen content,
the density of nematodes increases due to the develop-
ment of oxybiont species. Their proportion in the total
number of meiobenthos remains high – on average
53.5%.
Harpacticoida. All meiofauna appear to have some
sensitivity to extended periods of hypoxia, but a wide
range in tolerance (from hours to days, weeks or months
of exposure) have been observed. Within a given habitat
certain species of Foraminifera and Nematoda are typi-
cally most tolerant to hypoxia/anoxia while crustacean
meiofauna often are the least tolerant. More specifically,
the present study harpacticoid copepods as they repre-
sent, after nematodes, the second-most-abundant meio-
fauna group (Giere 2009) and are known to respond rap-
idly to hypoxic/anoxic conditions (Modig and ́Olafsson
1998). Moreover, harpacticoid copepods form an essen-
tial link between primary producers and higher trophic
levels (Giere 2009).
Fig. 5. Average indices of the density of Nematoda settlements (ind.·m-2) under different oxygen conditions
Vorobyova L.V.
55Морський екологічний журнал, № 1. 2026 ISSN 1684-1557
With a low content of dissolved oxygen in the bot-
tom layers of water (1–3 mg·L-1 O2), the average den-
sity of crustacean settlements was 11,689 ind.·m-2.
It increased sharply at 4 mg·L-1 O2 to 31,637.9 ind.·m-2.
The average value of the copepod abundance at oxygen
of 4–7 mg·L-1 O2 was 28,005.1 ind.·m-2. The highest aver-
age density of crustacean settlements (Fig. 6) was asso-
ciated with oxygen values of 8–9 mg·L-1 O2 (76,399.6 ±
10,961.0 ind.·m-2 and 93,890.7 ± 36,112 ind.·m-2, respec-
tively). On average, for the O2 range of 8–12 mg·L-1 O2,
the abundance of harpacticoids reached an average
of 50,934.0 ind.·m-2.
When considering the significance of harpacti-
coids in the formation of the total number of meioben-
thos, it was shown that their share in it differs signifi-
cantly and is highest at 4 and 8 mg·L-1 O2 of dissolved
oxygen (Fig. 7). This confirms our conclusion that
for harpacticoids, as well as foraminifera and nema-
todes, these two indicators are borderline, determining
the formation of the main ecological characteristics
of these taxa. Most harpacticoid species are very sensi-
tive to the conditions of the surrounding marine envi-
ronment. In areas experiencing high anthropogenic
load, their species diversity and quantitative indica-
tors are extremely poor. Thus, in the Sukhoi Estuary,
which is in a critical ecological state, Harpacticoida
representatives are very rare throughout the entire
water area of the estuary. Their numbers do not exceed
2,000 ind.·m-2. In the Grigorievsky Estuary, the eco-
system of which experiences significant anthropogenic
loads, the density of harpacticoid settlements differed
sharply and averaged 1.600–24.470 ind.·m-2. Anal-
ysis of long-term data on the formation of the num-
ber of representatives of the Harpacticoid taxocene
showed the following.
Ostracoda. The populations of the Ostracoda
taxocene are mostly inhabitants of the upper sublit-
toral and prefer rocky substrates with an abundant
amount of macrophytes on them. They develop well in
high-quality environments (Fig. 8).
Fig. 6. Average indices of the density of Harpacticoida settlements (ind.·m-2) under different oxygen conditions
Fig. 7. Share (%) of Harpacticoidа different dissolved oxygen content in the bottom layers of water
Influence of oxygen regime on meiobenthos of Ukrainian shelf of the black sea
56 ISSN 1684-1557 Морський екологічний журнал, № 1. 2026
The overall abundance and biomass of meiobenthos
under varying oxygen conditions in the sea's bottom lay-
ers depend not only on the population density of the per-
manent (eumeiobenthos). This is particularly true for
the larvae and juveniles of mollusks and meiobenthic
polychaetes (temporary meiobenthos).
Under favorable conditions for metamorpho-
sis, the abundance and biomass of juvenile mollusks
and polychaetes can play a significant role in shaping
the overall density and biomass indicators for the entire
meiobenthic community. In critical situations for
the marine benthos (high anthropogenic load, oxygen
deficiency in the bottom water layers, low salinity, etc.),
their share in the overall indicators decreases sharply,
even during favorable seasonal periods for the mass
settling of benthic invertebrate larvae from the pelagic
zone to the benthal.
The settling of mollusk larvae from the pelagic zone
to the benthal zone and the successful completion of their
metamorphosis occurs at depths of up to 50 meters.
This is typically characteristic of most of the Black
Sea shelf. In areas of increased eutrophication, charac-
terized by the periodic formation of extensive hypoxic
zones, the potential for successful larval development
from the velikoncha stage to a size category that allows
them to transition into macrozoobenthos is sharply
reduced. Even under favorable temperature conditions
and substrate quality, larvae that settle to the bottom
die massively due to oxygen deficiency. An analysis
of their population density distribution in various areas
of the Black Sea (mid-1990s) revealed that the highest
abundance of larval and juvenile mollusks, as well as
polychaetes, is on the Caucasian shelf, somewhat lower
on the Crimean shelf, and lowest on the northwestern
shelf. Their shares in the total meiobenthos abundance
are distributed accordingly: on the Caucasian shelf, they
averaged 29.7 %, on the Crimean shelf 7.5%, and on
the NWBS 2.1% (Vorobyova 2009).
A large database has made it possible to demonstrate
the formation of quantitative indicators of meiobenthos
in the NWBS (Table 2).
Thus, regular annual sampling on the NWBS for
40 years allowed us to establish the main patterns of for-
mation of the number and biomass of not only the total
meiobenthos, but also its main large taxa, which play
the main role in the formation of the biodiversity
Fig. 8. Average indices of the density of Ostracoda settlements (ind.·m-2) under different oxygen conditions
Table 2
Indices (ind. m-2) of the main taxa of the temporary component of meiofauna under different oxygen conditions
O2, mg·L-1 Number, ind.·m-2
Oligochaeta
Number, ind.·m-2
Polychaeta
Number, ind.·m-2
Bivalvia
Number, ind.·m-2
Temporary meiofauna, total
0-1 262.5 ± 171.5 1362,5 ± 555.5 300 ± 248.1 2706 ± 874.6
2 1589,2 ± 1064.4 2050.1 ± 1274.9 828,6 ± 533.2 4467.8 ± 1984.6
3 982,8 ± 537.7 1518.5 ± 443.6 960.2 ± 533.3 4524.0 ± 1097.6
4 2237,5 ± 1463.5 3387,5 ± 1002.3 529,0 ± 248.4 9300.1 ± 5278.3
5 727.0 ± 582.5 202.0 ± 703.4 835.4 ± 3145 4675.3 ± 1379.1
6 2361,4 ± 633.4 2326.1 ± 658.5 940.9 ± 359.5 5798.9 ± 1223.8
7 3435.6 ± 984.7 4646.3 ± 244.0 1975.5 ± 683.9 10407.2 ± 2558.0
8 2794.1 ± 694.6 3828.4 ± 957.2 3296.4 ± 819.2 9529.3 ± 1768.4
9 4322,55 ± 1031.1 6397.0 ± 2591.6 9721.0 ± 4781.6 21164.6 ± 6179.7
10 1366.1 ± 301.7 8668.3 ± 2196.1 10317.1 ± 4180.2 5798.4 ± 2087.4
11 1734.5 ± 452.2 6003.2 ± 1601.0 5708.5 ± 2196.3 13670.6 ± 3446.2
Vorobyova L.V.
57Морський екологічний журнал, № 1. 2026 ISSN 1684-1557
of the meiobenthos and its quantitative indicators. In
addition, a large amount of factual material allows us to
assess the quality of the marine environment in the ben-
thal. For this, we used the indices of the ratio of nema-
todes and harpacticoids at different oxygen regimes in
the bottom layers of water. As we can see (Fig. 9) on
the presented graph, they are highest at a dissolved oxy-
gen content of 1–3 mg·L-1 O2.
Thus, the data presented demonstrate that we
are examining the formation of meiofauna abun-
dance at the following dissolved oxygen levels: 1)
0–3 to 4 mg·L-1 O2; 2) 5–7 mg·L-1 O2; 3) 7 or more
to 12 mg·L-1 O2. In the first case, the meiobenthos
contains a few species, primarily from two large taxa
(foraminifera and nematodes, with minimal numbers
of polychaetes and oligochaetes). Thus, our long-
term research has shown that under hypoxia and even
anoxia, the structure of the meiobenthic community
of the NWBS changes dramatically. At oxygen levels
up to 3 mg·L-1 O2, foraminifera and nematodes usually
dominate, i.e., the total abundance of meiobenthos is
formed by a foraminifera-nematode or nematode-fo-
raminifera complex of organisms (Vorobyоva 2009).
In the second case, meiobenthic biodiversity increases
sharply with the development of euryoxybiont forms.
However, the total biomass increases only slightly due
to the high density of nematodes and Foraminifera. Har-
pacticoida account for 10–12% of the total meiobenthic
abundance. In the third case, the meiobenthos is typi-
cally represented by stenooxybiont species, and this is
where the highest total biomass of the meiobenthic com-
munity of organisms occurs.
Conclusions
In the vast waters of the northwestern shelf,
which has been negatively impacted by various types
of anthropogenic influences for many decades, large
areas of hypoxia in the bottom water layers are con-
stantly forming.
Analysis of long-term observations has revealed
the main patterns of formation of the structure and quanti-
tative indicators of meiofauna across a wide range of oxy-
gen levels in the bottom water layers. It has been established
that the maximum indicators of the total meiobenthos
are formed at oxygen levels of 7 mg·L-1 O2 and above.
Of the major large taxa, only foraminifera are very abun-
dant, even under severe hypoxia. Moreover, their share
of the total meiobenthos could be over 50%.
The abundance of crustaceans increases only in
good environmental conditions, with oxygen levels
above 6 mg·L-1 O2.
The ratio of total nematode abundance to total har-
pacticoid abundance indicates that small-sized forms
with short life cycles develop in the meiobenthos under
hypoxic conditions. At high oxygen levels, the total
biomass of the meiobenthos is formed by crustaceans
and juvenile mollusks.
Fig. 9. Indices of the ratio of nematodes and harpacticoids at different oxygen indices of the ratio of nematodes
and harpacticoids at different oxygen
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2018. Vol. 48. PP. 35–71. https://doi.org/10.1007/
s12526-017-0815-z
ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО
ШЕЛЬФУ ЧОРНОГО МОРЯ
Воробйова Л.В., д.б.н., проф.
ДУ «Інститут морської біології НАН України», vorobyova.meio@gmail.com
Гідрохімічні характеристики північно-західного шельфу Чорного моря надають унікальну можливість для
вивчення розвитку екологічних особливостей мейофауни в гіпоксичних зонах у польових умовах. Комплексний
набір фактичних даних дав змогу виявити специфічні особливості розвитку структури мейофауни та її кількісні
показники, отримані під час тривалих непереривних досліджень. Було проаналізовано загалом 520 кількісних
проб, зібраних під час регулярних щорічних наукових експедицій протягом понад 30 років. Аналіз даних дав
можливість визначити динаміку кількісних показників як для мейобентоса в цілому, так і для його основних
великих таксонів (форамініфери, нематоди, гарпактикоїди, остракоди та бівальвій), чутливих до змін рівня
кисню в придонних шарах північно-західного шельфу Чорного моря. Кількісні показники вказаних таксонів
були вивчені в широкому діапазоні рівнів розчиненого кисню в цьому морському регіоні (від 0 до 12 мг л⁻¹ O2).
Дослідження показали, що тільки форамініфери зберігають дуже високу чисельність навіть за дуже низького
рівня розчиненого кисню. Ракоподібні (гарпактикоїди та остракоди) та молоді двостулкові молюски не
можуть вижити в умовах гіпоксії. За отриманими даними визначено два кисневі пороги, що визначають як
біорізноманіття, так і чисельність основних представників постійних та тимчасових компонентів мейофауни.
Довгострокові спостереження й аналіз отриманих даних дають змогу оцінити якість довкілля в північно-західній
частині Чорного моря з використанням мейобентосу. Представлені у статті співвідношення чисельності нематод
і гарпактикоїд відображають структуру, чисельність та біомасу мейофауни в різних кисневих умовах. Чим вище
значення цього співвідношення, тим гірший екологічний стан середовища в придонних шарах води.
Ключові слова: Чорне море, північно-західний шельф, мейофауна, кисневий режим.
Дата першого надходження статті до видання: 25.03.2026
Дата прийняття статті до друку після рецензування: 27.04.2026
Дата публікації (оприлюднення) статті: 29.05.2026
Стаття поширюється на умовах
ліцензії відкритого доступу (CC BY 4.0)
Vorobyova L.V.
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| id | oai:ojs2.mej.od.ua:article-730 |
| institution | Marine Ecological Journal |
| keywords_txt_mv | keywords |
| language | English |
| last_indexed | 2026-06-09T01:00:18Z |
| publishDate | 2026 |
| publisher | Marine Ecological Journal |
| record_format | ojs |
| resource_txt_mv | mejodua/ed/8539756c735d72c4a209a9e5dc16a3ed.pdf |
| spelling | oai:ojs2.mej.od.ua:article-7302026-06-08T05:24:58Z INFLUENCE OF OXYGEN REGIME ON MEIOBENTHOS OF UKRAINIAN SHELF OF THE BLACK SEA ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ Воробйова, Л.В. Чорне море, північно-західний шельф, мейофауна, кисневий режим Black Sea, Northwestern Shelf, meiofauna, oxygen regime The hydrochemical characteristics of the Northwestern Part (NWBS) of the Black Sea offer a unique opportunity to study the development of meiofauna ecological features in hypoxic zones under field conditions. A comprehensive array of factual data allowed us to identify specific features of meiofauna structure development and its quantitative indicators, obtained through long-term, ongoing research. A total of 520 quantitative samples collected during regular annual scientific expeditions over more than 30 years were analyzed. Data analysis allowed us to determine the dynamics of quantitative indicators for both the general meiobenthos and its major large taxa (Foraminifera, Nematoda, Harpacticoida, Ostracoda, and Bivalvia), which are sensitive to changes in oxygen levels in the bottom layers of the northwestern shelf of the Black Sea. The quantitative indices of the aforementioned taxa were examined over a wide range of dissolved oxygen levels near the bottom in this marine region (from 0 to 12 mg·L-1 O2). The studies showed that only foraminifera maintain very high abundances even at very low dissolved oxygen levels. Crustaceans (harpacticoids and ostracods) and juvenile bivalves cannot survive under hypoxic conditions. The obtained data allowed us to determine two oxygen thresholds that determine both the biodiversity and abundance of the main representatives of the permanent and temporary components of the meiofauna. Long-term observations and analysis of the obtained data allow us to assess the environmental quality in the northwestern Black Sea using the meiobenthos. The nematode to harpacticoid abundance ratios presented in the paper reflect the structure, abundance, and biomass of the meiofauna under different oxygen conditions. He higher the values of this ratio, the worse the ecological state of the environment in the bottom layers of water. Гідрохімічні характеристики північно-західного шельфу Чорного моря надають унікальну можливість для вивчення розвитку екологічних особливостей мейофауни в гіпоксичних зонах у польових умовах. Комплексний набір фактичних даних дав змогу виявити специфічні особливості розвитку структури мейофауни та її кількісні показники, отримані під час тривалих непереривних досліджень. Було проаналізовано загалом 520 кількісних проб, зібраних під час регулярних щорічних наукових експедицій протягом понад 30 років. Аналіз даних дав можливість визначити динаміку кількісних показників як для мейобентоса в цілому, так і для його основних великих таксонів (форамініфери, нематоди, гарпактикоїди, остракоди та бівальвій), чутливих до змін рівня кисню в придонних шарах північно-західного шельфу Чорного моря. Кількісні показники вказаних таксонів були вивчені в широкому діапазоні рівнів розчиненого кисню в цьому морському регіоні (від 0 до 12 мг л⁻¹ O2). Дослідження показали, що тільки форамініфери зберігають дуже високу чисельність навіть за дуже низького рівня розчиненого кисню. Ракоподібні (гарпактикоїди та остракоди) та молоді двостулкові молюски не можуть вижити в умовах гіпоксії. За отриманими даними визначено два кисневі пороги, що визначають як біорізноманіття, так і чисельність основних представників постійних та тимчасових компонентів мейофауни. Довгострокові спостереження й аналіз отриманих даних дають змогу оцінити якість довкілля в північно-західній частині Чорного моря з використанням мейобентосу. Представлені у статті співвідношення чисельності нематод і гарпактикоїд відображають структуру, чисельність та біомасу мейофауни в різних кисневих умовах. Чим вище значення цього співвідношення, тим гірший екологічний стан середовища в придонних шарах води. Marine Ecological Journal Морський екологічний журнал 2026-05-29 Article Article Рецензована Стаття application/pdf https://mej.od.ua/index.php/mej/article/view/730 10.47143/1684-1557/2026.1.5 Marine Ecological Journal ; No. 1 (2026): Marine ecological journal; 49-60 Морський екологічний журнал; № 1 (2026): Морський екологічний журнал; 49-60 10.47143/1684-1557/2026.1 en https://mej.od.ua/index.php/mej/article/view/730/727 https://creativecommons.org/licenses/by/4.0 |
| spellingShingle | Чорне море північно-західний шельф мейофауна кисневий режим Воробйова, Л.В. ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ |
| title | ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ |
| title_alt | INFLUENCE OF OXYGEN REGIME ON MEIOBENTHOS OF UKRAINIAN SHELF OF THE BLACK SEA |
| title_full | ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ |
| title_fullStr | ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ |
| title_full_unstemmed | ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ |
| title_short | ВПЛИВ КИСНЕВОГО РЕЖИМУ НА МЕЙОБЕНТОС УКРАЇНСЬКОГО ШЕЛЬФУ ЧОРНОГО МОРЯ |
| title_sort | вплив кисневого режиму на мейобентос українського шельфу чорного моря |
| topic | Чорне море північно-західний шельф мейофауна кисневий режим |
| topic_facet | Чорне море північно-західний шельф мейофауна кисневий режим Black Sea Northwestern Shelf meiofauna oxygen regime |
| url | https://mej.od.ua/index.php/mej/article/view/730 |
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