Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary
The Black Sea hypoxic layer, situated near the boundary of the hydrogen sulfide zone, is inhabited by the polychaetes Vigtorniella zaikai, Kisseleva, 1992 and Protodrilus sp., which can serve as indicators of the interface between oxic and anoxic water. The maximum number of polychaetes in the north...
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Інститут зоології ім. І.І. Шмальгаузена НАН України
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irk-123456789-1094152016-11-29T03:02:18Z Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary Zaika, V.E. Sergeeva, N.G. Экология The Black Sea hypoxic layer, situated near the boundary of the hydrogen sulfide zone, is inhabited by the polychaetes Vigtorniella zaikai, Kisseleva, 1992 and Protodrilus sp., which can serve as indicators of the interface between oxic and anoxic water. The maximum number of polychaetes in the northern part of the Black Sea occur at a depth of 150 m, while near Bosporus, where the flow of the Marmara Sea water enters the Black Sea, the peak depth varies from 150 to 250 m. В Чёрном море, в гипоксическом слое, расположенном у границы сероводородной зоны, обитают полихеты Vigtorniella zaikai, Kisseleva, 1992 и Protodrilus sp., которые могут служить индикаторами интерфейса оксических/аноксических вод. Пик поселений полихет в северной половине моря постоянно регистрируется на глубинах около 150 м, тогда как у Босфора, где в Чёрное море впадают воды Мраморного моря, глубина пика неустойчива и меняется от 150 до 250 м. 2012 Article Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary / V.E. Zaika, N.G. Sergeeva // Вестник зоологии. — 2012. — Т. 46, № 4. — С. 327–335. — Бібліогр.: 13 назв. — англ. 0084-5604 DOI 10.2478/v10058-012-0026-9 http://dspace.nbuv.gov.ua/handle/123456789/109415 595.142: 546.221. 1 (262.5) en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Экология Экология |
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Экология Экология Zaika, V.E. Sergeeva, N.G. Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary Вестник зоологии |
| description |
The Black Sea hypoxic layer, situated near the boundary of the hydrogen sulfide zone, is inhabited by the polychaetes Vigtorniella zaikai, Kisseleva, 1992 and Protodrilus sp., which can serve as indicators of the interface between oxic and anoxic water. The maximum number of polychaetes in the northern part of the Black Sea occur at a depth of 150 m, while near Bosporus, where the flow of the Marmara Sea water enters the Black Sea, the peak depth varies from 150 to 250 m. |
| format |
Article |
| author |
Zaika, V.E. Sergeeva, N.G. |
| author_facet |
Zaika, V.E. Sergeeva, N.G. |
| author_sort |
Zaika, V.E. |
| title |
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary |
| title_short |
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary |
| title_full |
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary |
| title_fullStr |
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary |
| title_full_unstemmed |
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary |
| title_sort |
deep-water benthic polychaetes (vigtorniella zaikai and protodrilus sp.) in the black sea as indicators of the hydrogen sulfide zone boundary |
| publisher |
Інститут зоології ім. І.І. Шмальгаузена НАН України |
| publishDate |
2012 |
| topic_facet |
Экология |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/109415 |
| citation_txt |
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators of the Hydrogen Sulfide Zone Boundary / V.E. Zaika, N.G. Sergeeva // Вестник зоологии. — 2012. — Т. 46, № 4. — С. 327–335. — Бібліогр.: 13 назв. — англ. |
| series |
Вестник зоологии |
| work_keys_str_mv |
AT zaikave deepwaterbenthicpolychaetesvigtorniellazaikaiandprotodrilusspintheblackseaasindicatorsofthehydrogensulfidezoneboundary AT sergeevang deepwaterbenthicpolychaetesvigtorniellazaikaiandprotodrilusspintheblackseaasindicatorsofthehydrogensulfidezoneboundary |
| first_indexed |
2025-07-07T23:01:39Z |
| last_indexed |
2025-07-07T23:01:39Z |
| _version_ |
1837031018152853504 |
| fulltext |
UDC 595.142: 546.221. 1 (262.5)
DEEP-WATER BENTHIC POLYCHAETES (VIGTORNIELLA ZAIKAI
AND PROTODRILUS SP.) IN THE BLACK SEA AS INDICATORS
OF THE HYDROGEN SULFIDE ZONE BOUNDARY
V. E. Zaika, N. G. Sergeeva
A. O. Kovalevsky Institute of Biology of the Southern Seas of the NAS of Ukraine
2, Nakhimov av., Sevastopol, 99011 Ukraine
E-mail: vzaika36@gmail.com, nserg05@mail.ru
Received 12 November 2011
Accepted 28 May 2012
Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea as Indicators
of the Hydrogen Sulfide Zone Boundary. Zaika V. E., Sergeeva N. G. – The Black Sea hypoxic layer,
situated near the boundary of the hydrogen sulfide zone, is inhabited by the polychaetes Vigtorniella zaikai,
Kisseleva, 1992 and Protodrilus sp., which can serve as indicators of the interface between oxic and anox-
ic water. The maximum number of polychaetes in the northern part of the Black Sea occur at a depth
of 150 m, while near Bosporus, where the flow of the Marmara Sea water enters the Black Sea, the peak
depth varies from 150 to 250 m.
Key wo rd s: Black Sea, hypoxia, Polychaeta, biological indicators of hydrogen sulfide, abundance, dis-
tribution.
Ãëóáîêîâîäíûå äîííûå ïîëèõåòû (Vigtorniella zaikai è Protodrilus sp.) â ׸ðíîì ìîðå êàê èíäèêàòî-
ðû ñåðîâîäîðîäíîé çîíû. Çàèêà Â. Å., Ñåðãååâà Í. Ã. –  ׸ðíîì ìîðå, â ãèïîêñè÷åñêîì ñëîå,
ðàñïîëîæåííîì ó ãðàíèöû ñåðîâîäîðîäíîé çîíû, îáèòàþò ïîëèõåòû Vigtorniella zaikai, Kisseleva,
1992 è Protodrilus sp., êîòîðûå ìîãóò ñëóæèòü èíäèêàòîðàìè èíòåðôåéñà îêñè÷åñêèõ/àíîêñè÷åñêèõ
âîä. Ïèê ïîñåëåíèé ïîëèõåò â ñåâåðíîé ïîëîâèíå ìîðÿ ïîñòîÿííî ðåãèñòðèðóåòñÿ íà ãëóáèíàõ
îêîëî 150 ì, òîãäà êàê ó Áîñôîðà, ãäå â ׸ðíîå ìîðå âïàäàþò âîäû Ìðàìîðíîãî ìîðÿ, ãëóáèíà
ïèêà íåóñòîé÷èâà è ìåíÿåòñÿ îò 150 äî 250 ì.
Êëþ÷åâûå ñëîâ à: ׸ðíîå ìîðå, ãèïîêñèÿ, Polychaeta, áèîëîãè÷åñêèå èíäèêàòîðû ñåðîâîäî-
ðîäà, ÷èñëåííîñòü, ðàñïðåäåëåíèå.
Introduction
In the Black Sea the deep waters are permanently anoxic and contaminated with hydrogen sulfide. The
hypoxic layer, where the oxygen deficit increases with depth, is situated above the hydrogen sulfide zone. Changes
in the composition of benthic communities can be observed where the hypoxic waters impinge on the seafloor
and create a gradient of increasing hypoxia. Approaching the boundary of the hydrogen sulfide zone the species
composition of macrofauna becomes poorer; large forms occur only occasionally and finally vanish. Gradually,
metazoan meiofauna begin to dominate, and together with protozoans, create a distinctive benthic communi-
ty. For example, in the northern part of the Black Sea at 150 m depth Nematoda dominate the meiobenthos
community along with the deep-water polychaetes Vigtorniella zaikai Kisseleva, 1992 and Protodrilus sp., the
largest of which reaching a length of up to 1—1.5 mm (Kisseleva, 1998). The representatives of Hydrozoa and
protozoans (Gromida, Foraminifera and Ciliophora) also constitute part of the meiobenthic community. Because
the meiobenthic community is characterized by particular species of polychaetes living under conditions of per-
manent oxygen deficit, we term it the V. zaikai community (Zaika et al., 2009).
The consistent presence of V. zaikai in every sample of the benthic community bordering the sulfidic zone,
and its abundance peak (2—8 thousand specimens/m-2) at depths of about 150 m (Sergeeva, Zaika, 2000; Zaika,
1999; Zaika et al., 2008, 2009), indicate a good match between this species and the environmental conditions.
The second polychaete species, Protodrilus sp., possibly has somewhat different environmental requirements,
but nevertheless generally is distributed across the same transitional zone. The question therefore arises: can
the polychaetes V. zaikai and Protodrilus sp. be used as biological indicators of the region adjacent to the hydro-
Vestnik zoologii, 46(4): e-19—e-27, 2012
DOI 10.2478/v10058-012-0026-9
Ýêîëîãèÿ
Unauthenticated
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gen sulfide zone? To test this idea, samples were obtained during 2009—2010 across a wide range of depths in
three areas of the Black Sea, including the near-Bosporus and north-western regions. Here we present the results
of our analyses of these samples.
The present paper was written within the framework of the EC 7th Framework Project “In situ moni-
toring of oxygen depletion in hypoxic ecosystems of coastal and open seas, and land-locked water bodies” (HYPOX,
{226213) and national theme of the NAS of Ukraine.
Material and methods
The material was collected in the northern part of the Black Sea during R/V “Maria S. Merian” cruise
15/1 (April—May 2010). Meiobenthos sampling along the Ukrainian north-western shelf was conducted in two
areas: 1 (north-western part of the sea, NWP) and II (Crimean area) (fig. 1).
In the region of the Bosporus Strait (Black Sea), samples were obtained on two occasions: in November
2009 during a cruise of the R/V “Arar” of the Istanbul Technical University and in April 2010 during cruise
15/1 of the R/V “Maria S. Merian” (fig. 2).
Along the Ukrainian shelf sediments samples were taken using a TV multi-corer (TV MUC) in Area I
and with a TV MUC and a push-corer (PshC) deployed from the remote operated vehicle “Jago” in Area II.
In the Turkish area sediment samples for meiobenthos were taken with a gravity corer and a TV MUC. During
sampling, the characteristics of the seafloor (color, presence of detritus and epibionts, etc), as well as sedimen-
tological features in the core samples (color, presence and thickness of oxidized and reduced layers, smell of
hydrogen sulfide, presence of macrobenthos, dead shells, etc), were documented.
The sediment cores were sliced into 1-cm-thick layers down to a depth of 5—10 cm depth in order to
study the vertical distribution of the fauna. Each layer was washed through two sieves, the upper one with a
mesh size of 1 mm, the lower one with a mesh size of 63 μm. All the organisms found in the samples were
picked out and recorded. Specimens of the polychaetes Protodrilus sp. and V. zaikai were counted and sepa-
rated for further study. Adult and non-mature specimens were distinguished in the population of the former
species; in the case of the latter species, the number of segments in every specimen was counted.
e-20 V. E. Zaika, N. G. Sergeeva
Fig 1. Meiobenthos sampling stations along NW shelf of Ukraine (R/V “«Maria S. Merian»”, April—May 2010):
I – NW area, II – Crimean shelf.
Ðèñ. 1. Còàíöèè îòáîðà ìåéîáåíòîñà íà ÑÇ øåëüôå Óêðàèíû (ÍÈÑ «Maria S. Merian», àïðåëü—ìàé,
2010): I – ÑÇ àêâàòîðèÿ, II – êðûìñêèé øåëüô.
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Results
Along NW shelf of Ukraine, V. zaikai was present in sediment samples from 14 sta-
tions while Protodrilus sp. occurred at 11 stations within the 120—175 m depth range
(table 1). The density distribution of these species was not homogeneous along the
depth gradient. The maximum abundance of V. zaikai was 4.07 and 4.55 thousand spec-
e-21Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea...
Fig. 2. Meiobenthos stations in the region of the Bosporus: � – stations 1—10 (R/V “Arar”, November, 2009),
î – stations 198—302 (R/V “Maria S. Merian”, April, 2010).
Ðèñ. 2. Ìåéîáåíòîñíûå ñòàíöèè â ðàéîíå Áîñôîðà: � – ñòàíöèè 1—10 (ÍÈÑ «Arar», íîÿáðü, 2009),
î – ñòàíöèè 198—302 (ÍÈÑ «Maria S. Merian», àïðåëü, 2010).
Ta b l e 1. Abundance of V. zaikai and Protodrilus sp. (thousands specimens/m2) along the depth profile on the
Crimean margin (RV “Maria S. Merian”, April—May 2010)
Ò à á ëèö à 1. Ðàñïðåäåëåíèå ÷èñëåííîñòè (òûñ. ýêç./ì2) ïîëèõåò V. zaikai è Protodrilus sp. ïî ãëóáèíå íà
Êðûìñêîì ïîëèãîíå (ÍÈÑ «Maria S. Merian», àïðåëü—ìàé 2010 ã.)
¹ station
Depth, m
459
120
412
124
460
130
489
138
398
144
424
148
512
150
378
155
379
155
405
157
374
162
372
163
425
163
426
175
V. zaikai 0.07 1.56 4.07 1.06 4.55 2.47 0.72 1.14 0.28 3.24 1.80 3.59 1.77 1.27
Protodrilus sp. 0.14 0.96 0.36 1.20 0 5.86 0.12 0.42 1.27 0.24 0 0 0.07 0.14
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imens/m2 at 130 and 144 m depth, respectively, compared with 3, 24 and 3, 59 thou-
sand specimens/m2 at 157 and 163 m depth, respectively, and 1. 27 thousand specimens/m2
at 175 m depth. Densities of V. zaikai were 2. 98 and 1.98 thousand specimens/m2 in
two sub-samples from 148 m depth (st. 424). This variation could be explained by the
differences of sediment structure, especially the surface layers.
At the 148 m depth the abundance of Protodrilus sp. (5.65 —6.07 thousand speci-
mens/m2; mean 5.86 thousand specimens/m2) was significantly higher than that of
V. zaikai. At two stations located at 163 m depth, Protodrilus sp. was either absent
(Station 372) or present in low abundance (0.07 thousand specimens/m2; Station 425),
rising slightly to 0.14 thousand specimens/m2 at the deepest (175 m) site. Hence, peaks
of abundance of these polychaetes species were located at particular water depths.
Specimens of V. zaikai were collected along the NW shelf of Ukraine on three pre-
vious occasions. In February—March 2007 the abundance of this species was 2.6 thou-
sand specimens/m2 (Zaika et al., 2009), which was similar to its abundance in 2010.
Probably, these values were local seasonal abundance minima for V. zaikai. Such low den-
sities of this species were not observed in other regions. However, according to the lit-
erature data, the maximum abundance during the summer increased to 7.9—8.0 thousand
specimens/m2 in July 1994 and May—June 2007 (Sergeeva, 2004; Zaika et al., 2009).
In the Crimean water area in April—May 2010 V. zaikai and Protodrilus sp. were
observed for the first time within this depths range. The abundances of both species are
shown in table 1 and fig. 3.
These data revealed an unusual distribution of the two boundary polychaete species
with water depth across the Crimea shelf. Each of the species exhibited its own abun-
dance peak. The abundance of V. zaikai increased from a minimum at 120 m depth up
to maximum values at 130 and 144 m depth (4.06 and 4.55 thousand specimens/m2, respec-
tively). This species was also abundant at somewhat deeper sites (148—163 m). Generally,
therefore, it can be concluded that the maximum abundance of V. zaikai across the
Ukrainian shelf in April—May was between 130 and 163 m water depth, although there
were sharp decreases at some stations within this depth range, perhaps due to small-scale
differences in the nature of the substrate. Protodrilus sp. was also most common within
the same depth range, although this species was generally much less abundant that V. zaikai.
The density peak was observed at 148 m depth, where Protodrilus was more abundant than
V. zaikai.
e-22 V. E. Zaika, N. G. Sergeeva
Fig. 3. Distribution of boundary polychaete species. along depth transects of the Ukrainian shelf in the Areas
I and II (April–May 2010): 1 – V. zaikai; 2 – Protodrilus sp.
Ðèñ. 3. Ðàñïðåäåëåíèå ïîãðàíè÷íûõ âèäîâ ïîëèõåò V. zaikai, è Protodrilus sp. ïî ãëóáèíàì óêðàèíñêî-
ãî øåëüôà íà ïîëèãîíàõ I è II (àïðåëü—ìàé 2010 ã.): 1 – V. zaikai; 2 – Protodrilus sp.
0
1000
2000
3000
4000
5000
6000
110 120 130 140 150 160 170 180
Depth, m
A
bu
nd
an
ce
,i
nd
./m
2
V. zaikai Protodrilus sp
1
2
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A comparison between peaks and troughs of abundance suggests that the two species
have somewhat different environmental requirements. It could also be indicative of the
local or temporary nature of the boundary zone structure. However, the distribution of
Protodrilus sp. and V. zaikai within the range of depths where oxygen-deficient bottom
water impinges on the seafloor emphasizes the role of these species as indicators of hypox-
ia and the proximity of the hydrogen sulfide zone boundary.
Analysis of all V. zaikai data collected from northern part of the Black Sea shows
that the abundance of this species was 2.5—8 thousand specimens/m2 in the north-west-
ern area, 4.5 thousand specimens/m2 across the Crimean shelf, and 11.4—17.1 thousand
specimens/m2 across the Kerch shelf (Zaika et al., 2010). Thus, the highest abundance
of V. zaikai was recorded in May—June 2007 in the region of Kerch (Zaika et al., 2010).
We compared the age structure of V. zaikai populations from the two areas (NW
part and Kerch shelf) sampled in May—June 2007. The Kerch area yielded an unusual-
ly high proportion (28 %) of juvenile worms with only three body segments as well as
specimens with a maximum number of 10 body segments. In the NW area, the polychaetes
had up to 14 body segments and the proportion of specimens with three body segments
was only 4 % (fig. 4).
The distribution of V. zaikai was studied on two occasions in the southern part of
the Black Sea near the Bosporus. During the cruise of R/V “Arar” (November 2009),
this species was found for the first time at 250 m depth, although it was absent at shal-
lower sites (75—122, 160 and 190 m). A live gastropod mollusk was also found at 250
m, which suggests that the boundary between hypoxic and anoxic waters and the lower
depth limit of the macrobenthos correspond to this depth (Sergeeva et al., 2011). The
population density of V. zaikai at 250 m depth was 7280 specimens/m2. The polychaete
penetrated into the sediment to a greater depth than in the northern half of the Black
Sea; 25 % of specimens inhabited the 1—2 cm layer and a few were even found in the
5—7 cm layer.
During the cruise of the R/V “Maria S. Merian” in May 2010 both polychaete species
were found at a wide range of depths (V. zaikai – at 134—250 m, Protodrilus sp. –
116—159 m depths) (table 2, fig. 5) in the near-Bosporus region.
There are three notable features of the results from the 2010 samples: I – the wide
range of water depths where V. zaikai occurred; II – confirmation of the existence of
this species at 250 m depth; III – the “core habitat”, with the maximum of abundance
of each species, was located at 150–159 m depth for V. zaikai and 134 m depth for
e-23Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea...
Fig. 4. Distribution of V. zaikai population according to the number of body segment during May – June 2007
in two areas: I – NW area; II – Kerch shelf.
Ðèñ. 4. Ðàñïðåäåëåíèå ïîïóëÿöèè V. zaikai ïî êîëè÷åñòâó ñåãìåíòîâ â ìàå – èþíå 2007 ã. íà äâóõ ïîëè-
ãîíàõ: I – ÑÇ×; II – Êåð÷åíñêèé ïîëèãîí.
0
5
10
15
20
25
30
35
40
0 2 4 6 8 10 12 14 16
Number of segments
P
er
ce
n
t
o
f
in
d
iv
id
u
al
s
1
2
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Protodrilus sp. The term “core of habitat” was first used with reference to the boundary
community by Zaika et al. (2008) and is similar to the term “community core” of Kisseleva
(1981).
In 2010 V. zaikai was found in the 134—250 m depth range while in 2009 it was only
recorded at 250 m depth. This depth variation could reflect unstable conditions in the
near—Bosporus region. The habitat core of V. zaikai was located at a similar depth near
the northern shores in 2010 and in other years (Zaika, 2009).
In 2009 the density of V. zaikai at 250 m depth was 7.3 thousand specimens/m2, which
was similar to the maximum value (7.2—7.8 thousand specimens/m2) recorded in 2010
in this region. It was suggested that the population core moved down during the year. In
the NW area the maximum densities (mean values) were also 7.9—8.0 thousand speci-
mens/m2 in 1994 and 2007.
The 2009 population consisted mostly of individuals with 3—4 body segments; the
largest had only 5—6 segments. In 2010, individuals with 3—5 body segments dominat-
ed in the community, while those with 7—8 body segments were rare. These observations
suggest that the population was mainly represented by newly settled juveniles, which peri-
odically migrated into the water column. Probably, even juveniles that penetrated into
the upper centimeters of the soft sediments of the Bosporus region could easily migrate
in case of emergency. It seems likely that the shifting direction of oxygenated bottom cur-
rents from the Marmara Sea can change the structure of hypoxic benthic biotopes. In
this case it is possible that V. zaikai can migrate within the depth range of 134—250 m,
forming a habitation core where the oxygen regime is optimal. However, this hypothe-
sis can be proved only by further interdisciplinary studies.
e-24 V. E. Zaika, N. G. Sergeeva
Tab l e 2. Abundance (thousand specimens/m2) of the two polychaete species inhabiting the hypoxic/anoxic bound-
ary zone in the near-Bosporus region (RV “Maria S. Merian”, May 2010)
Ò à á ëèö à 2. ×èñëåííîñòü ïîãðàíè÷íûõ ïîëèõåò (òûñ. ýêç./ì2) â ïðèáîñôîðñêîì ðàéîíå (ÍÈÑ «Maria S.
Merian», ìàé 2010 ã.)
Fig. 5. Abundance of the boundary zone species with depth in the near-Bosporus region: 1 – V. zaikai; 2 –
Protodrilus sp.
Ðèñ. 5. Ðàñïðåäåëåíèå ÷èñëåííîñòè ïîãðàíè÷íûõ âèäîâ ïî ãëóáèíàì â ïðèáîñôîðñêîì ðàéîíå: 1 –
V. zaikai; 2 – Protodrilus sp.
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Depth,m
A
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¹ station / Depth, m 285/116 275/134 264/150 243/153 235/159 228/174 263/250
V. zaikai 0 0.14 7.20 5.22 7.78 0.14 0.14
Protodrilus sp. 0.33 2.61 0 0 0.14 0 0
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Discussion
All data concerning the bathymetric distributions of Vigtorniella zaikai and Protodrilus
sp. are summarized in table 3.
The data obtained in different research cruises were not averaged. In table 3 we only
include the depths that define the upper and lower limits of occurrence of the two poly-
chaete species. For example, with one exception, the upper occurrence boundary of both
species was located in the depth range 116—134 m (usually at 120—130 m). The only excep-
tion was an occurrence of Protodrilus sp. at 78 m depth, but this could not be consid-
ered typical. Unlike the upper boundary, the lower boundary of both species was vari-
able. In general, it was located at a depth of 151—163 m or somewhat deeper at 170—190
m; in the Bosporus region, however, V. zaikai penetrated down to 250 m depth.
Although the maximum densities of the two species coincide rarely, their core
habitats have the same overall depth limits (table 3). In addition, in the near—Bosporus
region one survey has shown that their core habitats are located at approximately the same
depths as on the north—western shelf, while another survey revealed a shift in the lower
boundary of V. zaikai to 250 m depth, and Protodrilus sp. entirely absent. Of these two
polychaete species, V. zaikai is usually more numerous than Protodrilus sp. and can be
found closer to water containing hydrogen sulfide.
A large number of specimens of an unknown polychaete species, and a Protodrilus
species identified as P. flavocapitatus, were reported earlier at three stations located at 150
m depth along the Romanian coast (Bacescu, 1963). We speculate that under oxygen-
depleted conditions on the Romanian shelf, the species referred to were V. zaikai and
Protodrilus sp. The ecological and biological characteristics of P. flavocapitatus suggest that
it was not the representative of the genus Protodrilus found on the Romanian shelf (Kisseleva
1998).
We can suggest some preliminary conclusions regarding the influence of physical and
chemical parameters. The upper occurrence boundary of the bordering polychaetes
species is situated somewhere near the inflection point observed in oxygen profile in the
lower part of the oxycline (Stunzhas, 2000). The benthic region under discussion lies
beneath the coastal convergence zone, where a deepening of the profiles of all physical
parameters takes place during severe winters (Titov, 2000). Thus variations in climatic
conditions can shift biological boundaries. The special conditions in the near-Bosporus
region, associated with the influence of water flowing in from the Marmara Sea, has already
been mentioned.
We cannot explain the size-age structure of V. zaikai populations. As already dis-
cussed, juveniles dominated at the study sites in the northern and southern parts of the
Black Sea. Based on the data in fig. 4, it appears that the samples from the Kerch area
were taken at a period of mass settlement of the polychaete larvae from the plankton.
However, the prevalence of younger stages is also characteristic of populations in the near-
e-25Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea...
Ta b l e 3. Bathymetric occurrence of boundary polychaete species in NW and near-Bosporus regions (according
to Sergeyeva, 2004; Zaika et al., 1999; Zaika, Sergeeva, 2008; Zaika et al., 2009)
Ò à á ëèö à 3. Ãðàíèöû âñòðå÷àåìîñòè è ãëóáèíû ÿäåð ïîñåëåíèé äëÿ ïîãðàíè÷íûõ âèäîâ ïîëèõåò â ÑÇ è
ïðèáîñôîðñêîì ðàéîíàõ (Sergeyeva 2004; Zaika et al., 1999; Zaika, Sergeeva, 2008; Zaika et al., 2009)
Boundary / Depth, m
North-Western region Near-Bosporus region
V. zaikai Protodrilus sp. V. zaikai Protodrilus sp.
Upper occurrence boundary 117—120 78, 124—130 134 116
Lower occurrence boundary from
to
151—163
175—190
151—157
170—175
250 159
Depth of habitat core from
to
139—148 138—148 150—160
250
134
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Bosporus region during November 2009 as well as in May 2010. A high percentage of
mature forms has never been observed in populations of V. zaikai.
There are two possible explanations for this size-age structure. 1) Maturation with
further active reproduction followed by the death of mature specimens occurs quickly over
a very short time period. 2) The habitats of mature specimens and settled juveniles are
spatially separated. Additional data are needed in order to test these hypotheses. We must
emphasise that the habitats of V. zaikai were not studied during the periods December–
January and August–October. Three autumn months are quite long enough for repro-
duction to occur and the mature specimens to die. However, it implies that the mature
stage has a very short mean life span, which does not seem consistent with the long life
of V. zaikai plankton larvae, which live at the same temperature of 8°C.
At first sight, the second hypothesis seems to be unlikely. Zaika et al. (2009) sug-
gested that the decaying remains of dolphins could serve as a habitat for mature V. zaikai.
This idea should be tested by analyzing the fauna associated with the cadavers of dol-
phins decaying on the bottom, especially at the hypoxic depths. The remain of dolphins
were not examined in the Black Sea at any depth. Time will tell which of these hypoth-
esis is correct.
Despite these uncertainties about the life histories of V. zaikai and Protodrilus sp.,
it is now possible to answer the main question posed in this paper; the two polychaete
species considered here can indeed be used in the Black Sea as biological indicators of
the boundary of the hydrogen sulfide zone. These species occur only in close proximity
to this boundary. Some hydrophysical data suggest that the boundary may fluctuate over
time. However, these data are fragmentary, and biological indicators often provide the
only method for determining the actual location of the hydrogen sulfide boundary.
Isolated occurrences of the polychaete species can be difficult to interpret, and there-
fore it is better to base conclusions on mass occurrences of V. zaikai, which was record-
ed closer to sulfidic waters than Protodrilus sp. While the lower occurrence boundary of
these bordering polychaete species clearly varies with time, fluctuations in the depth of
their habitat core provides a more reliable basis for interpreting the long-term dynamics
of water-column stratification, mainly in relation to oxygen availability.
e-26 V. E. Zaika, N. G. Sergeeva
Fig. 6. Position of benthic boundary belt occupied by the meiobenthic community dominated by V. zaikai.
Ðèñ. 6. Ïîëîæåíèå ïîãðàíè÷íîãî ïîÿñà áåíòàëè, çàíÿòîãî ñîîáùåñòâîì ìåéîáåíòîñà ñ äîìèíèðîâà-
íèåì V. zaikai.
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Thus, the data obtained on the distribution of the polychaetes species in our study
areas allows us to map the approximate location of the boundary belt occupied by a
meiobenthic community characterized by a high abundance of V. zaikai (fig. 6). At the
scale of this map, the hydrogen sulfide zone begins immediately below the belt occupied
by the meiobenthic boundary community. The location of this belt should be studied near
the western and eastern coasts of the Black Sea, especially in the region of the Georgian –
Turkish border.
A part of this work was funded by the EU 7th FP project HYPOX (“In situ monitoring of oxygen dep-
letion in hypoxic ecosystems of coastal and open seas, and land-locked water bodies”) EC Grant 226213. We
are grateful to Prof. Namik Çagatay and Prof. Antje Boetius for the opportunity to take part in cruises of the
RV “Arar” and “Maria S. Merian”. We particularly thank A. J. Gooday for checking English and the anony-
mous reviewers for very helpful comments in the manuscript. The authors are grateful to colleagues of Ecology
Benthos Department of IBSS, Dr. S. A. Mazlumyan and Dr. Igor Bondarev, for taking part in benthic sam-
pling during the scientific cruises and also to the principal technician L. F. Lukyanova for primary treatment
of the bottom sediments and V. G. Kopy for the help in preparing the figures.
Bacescu M. Contribution a la biocoenologie de la mer Noire l’etage periazoique et le facies dreissenifere leurs
caracteristiques // Rapp. Proc.-verb. Reun. C. I. E. S. M. – 963. – 17, 2. – P. 107—122.
Kisseleva M. I. Benthos of soft bottom sediments of the Black Sea. – Kyiv : Naukova Dumka, 1981. – 168 p. –
Russian : Êèñåëåâà Ì. È. Áåíòîñ ðûõëûõ ãðóíòîâ ׸ðíîãî ìîðÿ.
Kisseleva M. I. New genus and species of the family Chrysopetalidae Polychaeta from the Black Sea // Zool.
journ. – 1992. – 71, fasc. 11. – P. 128—132. – Russian : Êèñåëåâà Ì. È. Íîâûé ðîä è âèä ñåìåé-
ñòâà Chrysopetalidae Polychaeta â ׸ðíîì ìîðå.
Kisseleva M. I. Peculiarity of vertical distribution of polychaetes family Protodrilidae and Nerillidae in the Black
Sea // Zool. journ. – 1998. – 77, fasc. 35. – P. 533—539. – Russian : Êèñåëåâà Ì. È. Oñîáåííîñòè
âåðòèêàëüíîãî ðàñïðåäåëåíèÿ ïîëèõåò ñåìåéñòâ Protodrilidae è Nerillidae â ׸ðíîì ìîðå.
Sergeyeva N. G. Structure and Distribution of Meiobenthos in the Region of Methane Gas Seeps from the Black
sea bottom // Hydrobiological J. – 2004. – 40, N 6. – P. 45—56.
Sergeeva N. G., Zaika V. E. Ecology of Polychaeta from bordering pelagic and benthic communities of the Black
Sea // Rep. Acad. Sci. Ukraine. – 2000. – N 1. – P. 197—201. – Russian : Ñåðãååâà Í. Ã., Çàèêà Â. Å.
Ýêîëîãèÿ ïîëèõåò èç ïîãðàíè÷íûõ ñîîáùåñòâ ïåëàãèàëè è áåíòàëè ׸ðíîãî ìîðÿ.
Sergeeva N. G., Zaika V. E., Bondarev I. P. The lowest zoobenthos border in the Black Sea Near-Bosporus
region // Marine Ecological J. – 2011. – 10, N 1. – P. 65—72. – Russian : Ñåðãååâà Í. Ã.,
Çàèêà Â. Å., Áîíäàðåâ È. Ï. Íèæíÿÿ ãðàíèöà çîîáåíòîñà â Ïðèáîñôîðñêîì ðàéîíå ׸ðíîãî ìîðÿ.
Stunzhas P. A. About composition of the interaction zone of aerobic and anaerobic waters of the Black sea accord-
ing to measurements with the help of membraneless sensor of oxygen // Oceanology. – 2000. – 40, N 4. –
P. 539—545. – Russian : Ñòóíæàñ Ï. À. Î ñòðîåíèè çîíû âçàèìîäåéñòâèÿ àýðîáíûõ è àíàýðîá-
íûõ âîä ×åðíîãî ìîðÿ ïî èçìåðåíèÿì áåçìåìáðàííûì äàò÷èêîì êèñëîðîäà.
Titov V. Â. Formation of winter hydrological structure of the Black sea depending on severe winter //
Oceanology. – 2000. – 40, N 6. – P. 826—832. – Russian : Òèòîâ Â. Á. Ôîðìèðîâàíèå çèìíåé ãèä-
ðîëîãè÷åñêîé ñòðóêòóðû ׸ðíîãî ìîðÿ â çàâèñèìîñòè îò ñóðîâîñòè çèìû.
Zaika V. E. Specific pelagic and benthic communities of the Black Sea in the hydrogen sulfide zone // Biology
of the Sea. Vladivostok. – 1999. – 25, N 6. – P. 480—482. – Russian : Çàèêà Â. Å. Ñïåöèôè÷åñêèå
ñîîáùåñòâà ïåëàãèàëè è áåíòàëè ׸ðíîãî ìîðÿ ó ãðàíèöû ñåðîâîäîðîäíîé çîíû.
Zaika V. E., Sergeeva N. G. The boundary change of benthic settlement of polychaetes Protodrilus sp. and
Vigtorniella zaikai in the Black Sea // Marine Ecological J. – 2008. – 7, N 2. – P. 49—53. – Russian :
Çàèêà Â. Å., Ñåðãååâà Í. Ã. Èçìåíåíèå ãðàíèö äîííûõ ïîñåëåíèé ó ïîëèõåò Protodrilus sp. è
Vigtorniella zaikai â ׸ðíîì ìîðå.
Zaika V. E., Sergeeva N. G., Gulin M. B. The population structure of polychaete Vigtorniella zaikai (Kisseleva,
1992) in the Black Sea and the characteristics of a community in which it is the dominating species //
Marine Ecological J. – 2009. – 8, N 4. – P. 59—66. – Russian : Çàèêà Â. Å., Ñåðãååâà Í. Ã., Ãóëèí Ì. Á.
Ñòðóêòóðà ïîïóëÿöèè ïîëèõåòû Vigtorniella zaikai (Kisseleva, 1992) â ׸ðíîì ìîðå è õàðàêòåðè-
ñòèêà ñîîáùåñòâà, â êîòîðîì îíà äîìèíèðóåò.
Zaika V. E., Sergeeva N. G., Kisseleva M. I. Two polychaetes species bordering deep anoxic waters in the Black
Sea // Tavrichesky med.-biol. vestnik. – 1999. – N 1—2. – P. 56—60.
e-27Deep-Water Benthic Polychaetes (Vigtorniella zaikai and Protodrilus sp.) in the Black Sea...
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