Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna
The marbled goby, Pomatoschistus marmoratus (Risso, 1810), the Mediterranean immigrant in the Black Sea fauna, was examined for the metazoan parasites. In total, 12 parasite species were found to occurre. The component community consists of 7 trematode species, two nematodes, two acanthocephalans, a...
Saved in:
| Published in: | Вестник зоологии |
|---|---|
| Date: | 2010 |
| Main Author: | |
| Format: | Article |
| Language: | English |
| Published: |
Інститут зоології ім. І.І. Шмальгаузена НАН України
2010
|
| Subjects: | |
| Online Access: | https://nasplib.isofts.kiev.ua/handle/123456789/65734 |
| Tags: |
Add Tag
No Tags, Be the first to tag this record!
|
| Journal Title: | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| Cite this: | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna / Yu. Kvach // Вестник зоологии. — 2010. — Т. 44, № 6. — С. 509–518. — Бібліогр.: 28 назв. — англ. |
Institution
Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1859904702892736512 |
|---|---|
| author | Kvach, Yu. |
| author_facet | Kvach, Yu. |
| citation_txt | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna / Yu. Kvach // Вестник зоологии. — 2010. — Т. 44, № 6. — С. 509–518. — Бібліогр.: 28 назв. — англ. |
| collection | DSpace DC |
| container_title | Вестник зоологии |
| description | The marbled goby, Pomatoschistus marmoratus (Risso, 1810), the Mediterranean immigrant in the Black Sea fauna, was examined for the metazoan parasites. In total, 12 parasite species were found to occurre. The component community consists of 7 trematode species, two nematodes, two acanthocephalans, and one cestode species. Two parasite species (Aphalloides coelomicola and Dichelyne minutus) were represented by both adults and larvae, four parasite species were presented by adults (Asymphylodora pontica, Paratimonia gobii, Acanthocephaloides propinquus, and Telosentis exiguus), and six species were presented by larvae (Bothriocephalus gregarius, Cryptocotyle concavum, C. lingua, Pygidiopsis genata, Timoniella imbutiforme, and Contracaecum microcephalum). T. imbutiforme has the great tendency to join the infracommunity. The core of parasite fauna of the marbled goby consists of two specialists, A. coelomicola and P. gobii, which immigrated to the Black Sea together with their hosts. The co-immigration is a result of co-evolution of the parasites’ and host’s life cycles.
Приведены описания 4 видов гименолепидоидных цестод от луговой тиркушки юга Украины: Glareolepis poralae (Meggitt, 1927) Spassky, 1967; Wardium (s. l.) tauricum Kornyushin et Greben, sp. n.; Wardium (s. l.) sp.; Echinocotyloides longirostris (Rudolphi, 1819) Kornyushin, 1983. Детально описана морфология G. poralae. Установлено, что в гермафродитных члениках у G. poralae семенники отсутствуют, а структуры, описанные в литературе как семенники в гермафродитных члениках — это лопасти яичника. Описан новый вид цестод Wardium (s. l.) tauricum sp. n. с аплопараксоидными крючьями 13–15 мкм длиной и конусовидным вооруженным циррусом 20–22 мкм длиной. Еще один вид цестод Wardium (s. l.) sp. из-за отсутствия сколекса определен только до рода. Он отличается от всех известных видов этого рода размерами и вооружением цирруса.
|
| first_indexed | 2025-12-07T15:58:55Z |
| format | Article |
| fulltext |
UDC 597.585.1:616.99(262.55)
HELMINTHS OF THE MARBLED GOBY (POMATOSCHISTUS
MARMORATUS) A MEDITERRANEAN IMMIGRANT
IN THE BLACK SEA FAUNA
Yu. Kvach
Odessa Branch of the Institute of Biology of the Southern Seas, NAS of Ukraine,
Pushkinska str., 37, Odessa, 65125 Ukraine
E-mail: kvach@localka.net
Received 7 June 2010
Accepted 20 October 2010
Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black
Sea Fauna. Kvach Yu. — The marbled goby, Pomatoschistus marmoratus (Risso, 1810), the
Mediterranean immigrant in the Black Sea fauna, was examined for the metazoan parasites. In total,
12 parasite species were found to occurre. The component community consists of 7 trematode species,
two nematodes, two acanthocephalans, and one cestode species. Two parasite species (Aphalloides
coelomicola and Dichelyne minutus) were represented by both adults and larvae, four parasite species
were presented by adults (Asymphylodora pontica, Paratimonia gobii, Acanthocephaloides propinquus, and
Telosentis exiguus), and six species were presented by larvae (Bothriocephalus gregarius, Cryptocotyle
concavum, C. lingua, Pygidiopsis genata, Timoniella imbutiforme, and Contracaecum microcephalum).
T. imbutiforme has the great tendency to join the infracommunity. The core of parasite fauna of the
marbled goby consists of two specialists, A. coelomicola and P. gobii, which immigrated to the Black Sea
together with their hosts. The co-immigration is a result of co-evolution of the parasites’ and host’s life
cycles.
Ke y wo r d s: marbled goby, Black Sea, parasites, Mediterranean immigrants.
Ãåëüìèíòû áû÷êà-ëûñóíà (Pomatoschistus marmoratus) — ñðåäèçåìíîìîðñêîãî èììèãðàíòà â ôàóíå
×åðíîãî ìîðÿ. Êâà÷ Þ. — Áû÷îê-ëûñóí, Pomatoschistus marmoratus (Risso, 1810), —
ñðåäèçåìíîìîðñêèé èììèãðàíò â ÷åðíîìîðñêîé ôàóíå, îáñëåäîâàí íà íàëè÷èå ìíîãîêëåòî÷íûõ
ïàðàçèòîâ. Ìíîãîêîìïîíåíòíîå ñîîáùåñòâî ëûñóíà ñîñòàâëÿþò 12 âèäîâ ïàðàçèòîâ: 7 âèäîâ
òðåìàòîä, äâà — íåìàòîä, äâà — ñêðåáíåé è îäèí âèä öåñòîä. Äâà âèäà ïàðàçèòîâ (Aphalloides
coelomicola è Dichelyne minutus) ïðåäñòàâëåíû êàê âçðîñëûìè ñòàäèÿìè, òàê è ëè÷èíêàìè, ÷åòûðå
(Asymphylodora pontica, Paratimonia gobii, Acanthocephaloides propinquus è Telosentis exiguus) —
âçðîñëûìè îñîáÿìè, 6 (Bothriocephalus gregarius, Cryptocotyle concavum, C. lingua, Pygidiopsis genata,
Timoniella imbutiforme è Contracaecum microcephalum) — ëè÷èíêàìè. Âåðîÿòíîñòü îáíàðóæåíèÿ â
ñîñòàâå èíôðàñîîáùåñòâ íàèáîëåå âûñîêà äëÿ T. imbutiforme. ßäðî ïàðàçèòîôàóíû âêëþ÷àåò â
ñåáÿ äâà ñïåöèôè÷íûõ ê ëûñóíó âèäà, A. coelomicola è P. gobii, êîòîðûå èììèãðèðîâàëè â ×åðíîå
ìîðå âìåñòå ñî ñâîèìè õîçÿåâàìè. Ñîâìåñòíàÿ èõ èììèãðàöèÿ ÿâëÿåòñÿ ñëåäñòâèåì êîýâîëþöèè
æèçíåííûõ öèêëîâ ðàññìàòðèâàåìûõ ïàðàçèòîâ è õîçÿèíà.
Êëþ÷åâûå ñ ëîâ à: áû÷îê-ëûñóí, ×åðíîå ìîðå, ïàðàçèòû, ñðåäèçåìíîìîðñêèå èììèãðàíòû.
Introduction
The Black Sea fauna consists of the group of Mediterranean immigrants which entered this water body
probably within the past 12 000 years (Miller, 1965). One of the common representatives of this faunistic
group is the marbled goby Pomatoschistus marmoratus (Risso, 1810), which is widely distributed in whole
Mediterranean basin (Miller, 1986).
Because of their ecological plasticity, the gobiids are a convenient model for studying the processes of
colonization by parasites (Zander, Kesting, 1998). In the Black Sea the gobiid parasite fauna is more
heterogeneous than the gobiid fish fauna itself (Naidenova, 1974). The specific parasite fauna, which is not
similar to the ones of other gobiid species in the Black Sea, is typical for the marbled goby (Kvach, 2005 a).
It consists of the specialists such as Aphalloides coelomicola Dollfus, Chabaud et Golvan, 1957 and
Paratimonia gobii Pre´vôt et Bartoli, 1967.
Vestnik zoologii, 44(6): 509–518, 2010 Ýêîëîãèÿ
The aim of the presented study is to analyse the parasite community of the marbled goby and the role
of this fish in the parasite life cycles in the Northwestern Black Sea.
Material and Methods
The gobies were sampled by deep net (100 cm x 50 cm; cell /◎ 5 mm) in 2002–2006 at 8 localities of
the Northwestern Black Sea (NWBS): 1. Tyligul Estuary, 46°53' N 31°03' E; 2. Tyligul Estuary, 46°41' N
31°09' E; 3. Tyligul Estuary; 4. Hryhorivsky Estuary, 46°37' N 31°00' E; 5. Khadzhibey Estuary, 46°33' N
30°41' E; 6. Gulf of Odessa, 46°28' N 31°45' E; 7. Sukhyi Estuary, 46°21' N 30°38' E; 8. Budaki Lagoon,
46°01' N 30°24' E (fig. 1). Two water bodies, the Tyligul Estuary and the Budaki Lagoon are isolated from
the sea by natural sandbars. The sampling places in the Gulf of Odessa and the Sukhyi Estuary were located
in lagoons, artificially isolated from the sea by mooring lines. Only in the Hryhorivsky Estuary the gobies
were sampled in the southern part, connected with the open sea.
In total, 108 gobies were examined for metazoan parasites (table 1). The fish were transported alive to
the laboratory, put in aquarium, and then executed in two days. The standard length (SL, cm) of fish
individual was measured first.
All parasitological nomenclature follows the levels of Bush et al. (1997) and Zander (1998). The
prevalence (P, %), intensity (presented as intensity range, IR), mean intensity (MI), and abundance (A)
were calculated according to Bush et al. (1997). The importance of the host-parasite relationship, expressed
as the status as core or satellite species, is described by abundance: > 2 — core species, 0.6–2 — secondary
species; 0.2–0.6 — satellite species; < 0.2 — rare species (Zander et al., 2000; Zander, 2003). The tendency
to join the infracommunity was evaluated according to the Infracommunity Index, ICI (Zander, 2004):
,
where Mij was the ratio (numbers) of multiple infected hosts j with parasite i and other parasites, Nj was the
ratio (numbers) of infected hosts j, and Ij was the mean of parasite species in host j (= mean
510 Yu. Kvach
Fig. 1. Schematic map of the investigated area (black circles are sampling localities).
Ðèñ. 1. Ñõåìàòè÷åñêàÿ êàðòà ðàéîíà èññëåäîâàíèé (÷åðíûìè êðóæêàìè îáîçíà÷åíû ìåñòà ñáîðà ìàòå-
ðèàëà).
infracommunity). The great tendency to join the community was determined by the ICI level more then 0.30
(Zander, 2004).
The Species Evenness (J) was calculated according to Zander et al. (1993):
,
where Hs is Shannon-Wiener Index, S is number of species.
Evenness values > 0.6 mean a greater part of the homogeneity, of > 0.7 a high homogeneity.
Abbreviations of names of parasites life stages were used as follows: pl — plerocercoid, met —
metacercaria, L3 — 3rd stage larva.
Results
The maximal number of fishes was caught in the Budaki Lagoon, Khadzibey and
Tyligul estuaries (table 1). In the Gulf of Odessa, Hryhorivsky and Sukhyi estuaries the
marbled goby was rather rare (5, 7 and 8 fishes respectively).
No metazoan parasites occurred in the marbled goby in the Khadzhibey Estuary.
All fishes caught in the Budaki Lagoon, Sukhyi and Tyligul estuaries were infected with
parasites (table 1). Four from five examined fishes were infected in the Gulf of Odessa,
6 from 7 in the Hryhorivsky Estuary. The maximal number of parasites and abundance
were noted in the Budaki Lagoon.
In total, 12 metazoan parasite species occurred in the marbled goby in the NWBS
(table 2). The component community consisted of 7 digenean species, two nematodes,
two acanthocephalans, and one cestode species. Two parasite species (Aphalloides
coelomicola and Dichelyne minutus (Rudolphi, 1819)) were presented by both adults and
larvae (table 2). Four parasite species were presented by adults, and six by larvae.
Most of parasite species, namely Bothriocephalus gregarius Renaud et al., 1983 pl,
Asymphylodora pontica (Tschernyschenko, 1949), Paratimonia gobii, D. minutus,
Acanthocephaloides propinquus (Dujardin, 1845), and Telosentis exiguus von Linstow,
1901, were found in the gut of fish. Four parasite species, A. coelomicola met,
Cryptocotyle concavum (Creplin, 1825) met, C. lingua (Creplin, 1825) met, T. imbuti-
forme met, occurred in muscles of gobies. Two species from this list (C. concavum met
and C. lingua met) were also found in fins and skin, but one (A. coelomicola met) in
ovaries. The parasites located in tissues (metacercariae of A. coelomicola, Cryptocotyle
spp., Timoniella imbutiforme (Molin, 1859)) were most numerous in the marbled goby
(table 2). Three parasite species were found encysted in intestinal walls: Pygidiopsis
genata Looss, 1907 met, D. minutus L3, Contracaecum microcephalum (Rudolphi, 1819)
L3. Only adults of A. coelomicola occurred in the body cavity of gobies.
Two species, T. imbutiforme met and A. propinquus, occurred in all the water bod-
ies studied (table 2). Such parasites as B. gregarious pl, C. microcephalum L3, T. exigu-
us were found only in one water body (table 2).
511Parasites of the marbled goby (Pomatoschistus marmoratus)…
Ta b l e 1. The number of examined hosts and its infestation parameters
Ò à á ëèö à 1. Êîëè÷åñòâî èññëåäîâàííûõ õîçÿåâ è ïàðàìåòðû èõ çàðàæåííîñòè
‘Total’ column includes the data from the Khadzhibey Estuary, where no metazoan parasites occurred
in the marbled goby.
Host number 33 8 5 18 7 37 108
SL, cm 3.8 ± 1.0 3.9 ± 2.4 4.2 ± 0.8 2.6 ± 0.2 3.7 ± 0.8 2.3 ± 0.6 3.2 ± 1.2
Prevalence (number / %) 33 / 100 8 / 100 4 / 80 0 / 0 6 / 85.7 37 / 100 88 / 81.5
Number of uninfected hosts 0 0 1 18 1 0 20
Total number of parasites
occurred 4586 2638 81 0 57 923 8285
Mean intensity 139.0 329.8 20.3 0 9.5 24.9 94.1
Abundance 139.0 329.8 16.2 0 8.1 24.9 76.7
Localities
Budaki
Lagoon
Sukhyi
Estuary
Gulf of
Odessa
Khadzhi-
bey Estuary
Hryhorivsk
y Estuary
Tyligul
Estuary
Total
512 Yu. Kvach
Ta b l e 2. Component community of parasites of the marbled goby in the NWBS
Òàáëèö à 2. Ìíîãîêîìïîíåíòíîå ñîîáùåñòâî ïàðàçèòîâ áû÷êà-ëûñóíà â ñåâåðî-çàïàäíîé ÷àñòè ×åðíîãî
ìîðÿ
CESTODA
Bothriocephalus
gregarius pl
P 3 from 7 2.8
IR 2–6 2–6
MI 4.0±2.0 4.0±2.0
A 1.7 0.1
TREMATODA
Aphalloides
coelomicola
P 45.5 3 from 8 10.8 20.4
IR 1–10 16–36 1–11 1–36
MI 3.5±3.2 26.0±10.0 4.3±4.6 6.7±9.0
A 1.6 9.8 0.5 1.4
A. coelomicola met P 27.3 10.8 12.0
IR 1–10 1–20 1–20
MI 4.0±3.0 7.0±8.8 4.9±5.2
A 1.1 0.8 0.6
Asymphylodora
pontica
P 2 from 8 13.5 6.2
IR 1 1–15 1–15
MI 1.0±0.0 6.8±6.3 5.1±5.9
A 0.3 0.9 0.3
Cryptocotyle con-
cavum met
P 69.7 7 from 8 2 from 7 54.1 48.1
IR 2–70 9–1000 1 1–41 1–1000
MI 21.0±16.8 319.7±465.2 1.0±0.0 6.4±9.0 54.8±191.8
A 14.6 279.8 0.3 3.4 26.4
C. lingua met P 57.6 32.4 28.7
IR 1–9 1–13 1–13
MI 4.7 ± 2.2 3.6 ± 3.8 4.3 ± 2.9
A 2.8 1.2 1.2
Paratimonia gobii P 6.1 5 from 8 1 from 5 32.4 18.5
IR 1–3 1–50 44 1–80 1–80
MI 2.0 ± 1.4 19.8 ± 20.4 44.0 17.3 ± 22.3 17.7 ± 20.9
A 0.1 12.4 8.8 5.6 3.3
Pygidiopsis genata
met
P 57.6 3 from 8 20.4
IR 3–24 31–46 3–46
MI 10.6±5.8 40.0±7.9 14.6 ± 11.9
A 6.1 15.0 3.0
Timoniella imbuti-
forme met
P 100.0 3 from 8 2 from 5 6 from 7 94.6 73.1
IR 57–210 30–36 3–19 2–6 2–61 2–210
MI 112.3 ± 37.8 33.0 ± 3.0 11.0 ± 11.3 4.5 ± 1.4 12.9 ± 14.7 54.5 ± 60.1
A 112.4 12.4 4.4 3.9 12.2 39.9
NEMATODA
Dichelyne minutus P 1 from 8 3 from 5 4 from 7 2.7 8.3
IR 1 3–5 1–6 2 1–6
MI 1.0 4.0 ± 1.0 3.0 ± 2.2 2.0 3.0 ± 1.7
A 0.1 2.4 1.7 0.05 0.3
D. minutus L3 P 1 from 7 0.9
IR 1 1
MI 1.0 1.0
A 0.1 0.01
Contracaecum micro-
cephalum L3
P 3.0 0.9
IR 1 1
MI 1.0 1.0
A 0.03 0.01
ACANTHOCEPHALA
Acanthocephaloides
propinquus
P 18.2 1 from 8 2 from 5 1 from 7 8.1 12,0
IR 1–2 1 1–2 1 1–12 1–12
MI 1.3 ± 0.5 1.0 1.5 ± 0.7 1.0 4.7 ± 6.4 2.1 ± 3.0
A 0.2 0.1 0.6 0.1 0.4 0,3
Localities
Budaki
Lagoon
Sukhyi
Estuary
Gulf of
Odessa
Hryhorivsky
Estuary
Tyligul
Estuary
Total
The parasite component communities of the Tyligul and Hryhorivsky estuaries are
presented by all types by its host-parasite relationships: core, secondary, satellite, and
rare (fig. 2). In the Sukhyi Estuary and the Budaki Lagoon the secondary species were
absent, but in the Gulf of Odessa no satellite and rare species occurred. The core
species were found in all water bodies (except the Khadzhibey Estuary where no meta-
zoan parasite was found). The number of the core species in the component commu-
nity varied from one (in the Hryhorivsky Estuary) to five (in the Sukhyi Estuary and
the Budaki Lagoon) (fig. 2). In the Budaki Lagoon a part of core was taken by dige-
neans C. concavum, C. lingua, P. genata, and T. imbutiforme (table 2). In addition, sum-
513Parasites of the marbled goby (Pomatoschistus marmoratus)…
‘Total’ column includes the data from the Khadzhibey Estuary, where no metazoan parasites occurred
in the marbled goby.
Table 2 (continued).
Îêîí÷àíèå òàáëèöû 2.
ACANTHOCEPHALA
Acanthocephaloides
propinquus
P 18.2 1 from 8 2 from 5 1 from 7 8.1 12,0
IR 1–2 1 1–2 1 1–12 1–12
MI 1.3 ± 0.5 1.0 1.5 ± 0.7 1.0 4.7 ± 6.4 2.1 ± 3.0
A 0.2 0.1 0.6 0.1 0.4 0,3
Telosentis exiguus P 1 from 7 0,9
IR 2 2
MI 2.0 2,0
A 0.3 0,02
Species richness 8 8 4 6 8 12
Shannon Index 0.72 0.64 1.09 1.33 1.45 1.20
Species Evenness 0.35 0.31 0.79 0.74 0.70 0.48
Localities
Budaki
Lagoon
Sukhyi
Estuary
Gulf of
Odessa
Hryhorivsky
Estuary
Tyligul
Estuary
Total
Fig. 2. The number of core, secondary, satellite, and rare species in the parasite component community of
the marbled goby in the various water areas of the NWBS.
Ðèñ. 2. Êîëè÷åñòâî îñíîâíûõ, âòîðîñòåïåííûõ, ñàòåëëèòíûõ è ðåäêèõ âèäîâ â ìíîãîêîìïîíåíòíîì
ñîîáùåñòâå ïàðàçèòîâ áû÷êà-ëûñóíà â ðàçëè÷íûõ âîäîåìàõ ñåâåðî-çàïàäíîé ÷àñòè ×åðíîãî ìîðÿ.
0,00 5,00 10,00 15,00 20,00 25,00 30,00 35,00 40,00 45,00
Contracaecum microcephalum
Telosentis exiguus
Bothriocephalus gregarius
Acanthocephaloides propinquus
Dichelyne minutus
Asymphylodora pontica
Cryptocotyle lingua
Aphalloides coelomicola
Pygidiopsis genata
Paratimonia gobii
Cryptocotyle concavum
Timoniella imbutiforme
R
ar
e
S
at
el
lit
e
S
ec
on
da
ry
C
or
e
marising the adults and larvae of A. coelomicola, this species took a part of core, too
(A = 3.46). C. microcephalum and A. propinquus were rare in the Budaki Lagoon. Four
parasite species (C. concavum, P. gobii, P. genata, and T. imbutiforme) were core in the
Sukhyi Estuary, but A. pontica and A. propinquus were satellite (table 2). In the Gulf of
Odessa the digeneans P. gobii, T. imbutiforme, and the nematode D. minutus were core,
but the acanthocephalan A. propinquus was secondary species in the parasite communi-
ty. Only T. imbutiforme was core in the Hryhorivsky Estuary, B. gregarius and D. min-
utus (for adults and larva the summarised A = 1.9) were secondary, C. concavum and
T. exiguus were satellite, and A. propinquus was rare. In the Tyligul Estuary C. con-
cavum, P. gobii, and T. imbutiforme were core species, A. coelomicola (for adults and
larvae the summarised A = 1.2), A. pontica, and C. lingua were secondary, A. propin-
quus was satellite, and D. minutus was rare (table 2). In total, in the NWBS the core
species were C. concavum, P. gobii, P. genata, and T. imbutiforme, secondary species
were A. coelomicola and C. lingua, satellite species were A. pontica, D. minutus, and
A. propinquus, other species were rare (fig. 3).
The parasite fauna was presented by five types of infracommunities with the species
numbers from one to six (fig. 4). In the Budaki Lagoon the parasite fauna was present-
ed by all types of infracommunities and most of examined fishes were infected with four
and more parasite species. In the Tyligul Estuary most of examined fishes were infect-
ed with one to three parasite species. The fishes infected with only one parasite species
were relatively rare, from two individuals in the Hryhorivsky and Sukhyi estuaries and
the Gulf of Odessa, to six individuals in the Tyligul Estuary. Only one species, T. imbu-
tiforme, had great tendency to join the infracommunity in the Tyligul Estuary (table 3).
This parasite also had high ICI in other water bodies, except the Sukhyi Estuary, and
maximal tendency to join the community for the total NWBS (ICI = 0.29).
The greater part of the homogeneity of the parasite component community was
mentioned in the Gulf of Odessa, Hryhorivsky, and Tyligul estuaries (table 2). In the
514 Yu. Kvach
Fig. 3. The abundance of different parasite species in the Pomatoschistus marmoratus parasite fauna by
categories: core, secondary, satellite, and rare.
Ðèñ. 3. Îáèëèå ðàçëè÷íûõ âèäîâ ïàðàçèòîâ â ïàðàçèòîôàóíå Pomatoschistus marmoratus ïî êàòåãîðèÿì:
îñíîâíûå, âòîðîñòåïåííûå, ñîïóòñòâóþùèå, ðåäêèå.
Budaki Lagoon and Sukhyi Estuary it was low. In total, the low homogeneity was typ-
ical for the parasite component community of the marbled goby in the NWBS.
Discussion
Twenty-seven metazoan parasite species have been mentioned in the Black Sea
basin according to data of different authors (Radulescu, 1953; Chernyshenko, 1960,
1962, 1966; Chernyshenko, Sventsytskaya, 1970; Naidenova, 1974; Machkevsky et al.,
1990). We found 17 parasite species, 15 of which had already been mentioned in pub-
lished data, but two species, Àsymphylodora pontica and Ñryptocotyle lingua met, were
mentioned from the marbled goby for the first time.
The absence of metazoan parasites in the Khadzhibey Estuary was caused by its
low salinity (~ 3‰). All of parasites typical for the marbled goby have marine or brack-
ish water origin. The absence of digeneans, which are core of the marbled goby para-
site fauna, was caused by lack of mollusks in the fauna of the Khadzhibey Estuary
(Kvach, 2004).
The relative abundance of the various parasite species was the highest in the Gulf
of Odessa, where the species richness was the lowest (table 2). All parasite species in
this water body, except Paratimonia gobii, had similar tendency to join the communi-
ty, ICI = 0.25 (table 3).
The digenean Asymphylodora pontica is a specific parasite of gobiids in the Black
Sea, infecting mostly neogobiins, such as the round goby Apollonia melanostoma, and
the monkey goby A. fluviatilis (Gaevskaya et al., 1975; Kvach, 2005 a). However, the
case of infestation of gobiin fish (a grass goby Zosterisessor ophiocephalus from the
Hryhorivsky Estuary) was mentioned by Yu. Kvach (2005 b). In both cases (occurrence
in the marbled and grass gobies) we can consider it as an occasional infestation.
515Parasites of the marbled goby (Pomatoschistus marmoratus)…
Fig. 4. The part of different infracommunities in the parasite fauna of the marbled goby in different water
bodies of the NWBS.
Ðèñ. 4. Äîëÿ ðàçëè÷íûõ èíôðàñîîáùåñòâ â ïàðàçèòîôàóíå áû÷êà-ëûñóíà â ðàçëè÷íûõ âîäîåìàõ
ñåâåðî-çàïàäíîé ÷àñòè ×åðíîãî ìîðÿ.
The marbled goby plays important role in the realization of the life cycle of the
cestode Botriocephalus gregarius and the digenean Timoniella imbutiforme. Both species
are generalists on the larval stage. In the Mediterranean Sea B. gregarius uses the mar-
bled goby as main paratenic host, but the definitive host is the turbot Psetta maxima
(Robert et al., 1988). In the NWBS this species occurred in the Hryhorivsky Estuary
only. This is a water body connected with the sea by 15-m depth navigation canal,
which is a suitable habitat for turbots. The T. imbutiforme met is a numerous parasite
species occurring in all the water bodies examined (table 2). This is only parasite species
which has great tendency to join the infracommunity, but only in the Tyligul Estuary,
where the highest diversity of parasites was noted according to the Shannon Index
(table 2, 3). The gobiids serve as second intermediate hosts of this parasite, together
with other small-sized fishes, such as Atherinidae (Maillard, 1973 a).
The marbled goby is an annual fish species, which dies after spawning (Miller,
1986; Smirnov, 1986). The small sizes of this fish makes it the suitable pray item for
many species of predatory fish and fish-eating birds. This fact influents the parasite
fauna of this goby. So, the core species in the parasite component community are most-
ly metacercariae, such as Cryptocotyle spp., Pygidiopsis genata, T. imbutiforme (table 2).
Also, in the core of the parasite fauna are two parasites specific for genera Pomato-
schistus and Knipowitschia: Aphalloides coelomicola and P. gobii (table 2). The marbled
goby plays a role of both definitive and second intermediate host for A. coelomicola
(Maillard, 1973 b). The adults are located in the body cavity, and the metacercariae
are in the ovaries and muscles of the same fish. This parasite is well adapted to the life
cycle of the gobies and the necessary condition for the transmission of the digenean is
the host death (Pampoulie et al., 2000). The infestation of Pomatoschistus with P. gobii
is a case of adaptation of the parasite life cycle to the specific diet of the host. The
metacercariae of P. gobii are located in inhaling siphons of mussels Abra ovata and
Cerastoderma glaucum (Maillard, 1975). The small-sized gobies, such as the marbled
goby, can use the cut-off siphons as the pray item.
So, among the core species of the parasite fauna of the marbled goby in the stud-
ied NWBS lagoons and estuaries are not only generalists, as Cryptocotyle spp. or T. im-
butiforme, but also specialists, such as A. coelomicola and P. gobii. Both specialists have
Mediterranean origin, as well as their hosts. This is a marked difference between the
marbled goby and other Black-Sea gobiids, the core of which parasite fauna consists of
generalists only (Kvach, 2005 a; Kvach, Og
�
uz, 2009). The infestation of the marbled
goby with two above mentioned specialist trematode species is a result of co-evolution,
516 Yu. Kvach
Ta b l e 3. Infracommunity indices
Ò à á ëèö à 3. Èíäåêñû èíôðàñîîáùåñòâà
No t e. Bold: the great tendency to join the infracommunity.
Mean infracommunity 3.7 ± 1.4 3.1 ± 1.6 2.0 ± 1.2 2.8 ± 1.8 2.6 ± 1.1 3.1 ± 1.5
Bothriocephalus gregarius pl 0.18 0.01
Aphalloides coelomicola 0.16 0.12 0.07 0.11
Asymphylodora pontica 0.08 0.05 0.03
Cryptocotyle concavum met 0.19 0.28 0.12 0.21 0.19
C. lingua met 0.15 0.13 0.11
Paratimonia gobii 0.02 0.20 0.13 0.13 0.07
Pygidiopsis genata met 0.15 0.12 0.08
Timoniella imbutiforme met 0.27 0.12 0.25 0.29 0.37 0.29
Dichelyne minutus 0.04 0.25 0.24 0.01 0.004
Contracaecum microcephalum L3 0.01 0.03
Acanthocephaloides propinquus 0.04 0.04 0.25 0.06 0.02 0.05
Telosentis exiguus 0.06 0.004
Localities
Budaki
Lagoon
Sukhyi
Estuary
Gulf of
Odessa
Hryhorivsky
Estuary
Tyligul
Estuary
Total
supporting the ‘Red Queen Hypothesis’ of L. Van Valen (1973). During the co-evolu-
tion the adaptation of the life cycles of parasites to the specific life cycle of host (the
marbled goby is annual small-sized fish) was completed. It was a cause of successful
co-immigration of the marbled goby and its specific parasites to the Black Sea.
Bush A. O., Lafferty K. D., Lotz J. M., Shostak A. W. Parasitology meets ecology on its own terms: Margolis
et al. revisited // J. Parasitol. — 1997. — 83. — P. 575–583.
Chernyshenko A. S. Parasite fauna of the Hryhorivsky Estuary // Proceedings of 1st Ichthyological Conference
on the Study of the North-wstern part of the Black Sea / V. S. Chepurnov. — Chisinau, 1960. —
P. 205–211. — Russian : ×åðíûøåíêî À. Ñ. Ïàðàçèòîôàóíà ðûá Ãðèãîðüåâñêîãî ëèìàíà // Òð. I
èõòèîë. êîíô. ïî èçó÷åíèþ ìîðñêèõ ëèìàíîâ ñåâåðî-çàïàäíîé ÷àñòè ×åðíîãî ìîðÿ /
Â. Ñ. ×åïóðíîâ. — Êèøèíåâ, 1960. — Ñ. 205–211.
Chernyshenko A. S. Parasites of fish of the Khadzhibey Estuary // Proceedings of the University of Odessa.
Seria of Biol. Sci. — 1962. — 152, N 12. — P. 39–43. — Ukrainian : ×åðíèøåíêî À. Ñ. Ïàðàçèòè
ðèá Õàäæèáåéñüêîãî ëèìàíó // Ïðàö³ Îäåñüêîãî óí-òó. Ñåð. Á³îë. — 1962. — 152, N 12. —
Ñ. 39–43.
Chernyshenko A. S. Parasite fauna of fish of the estuaries of the north-western Black sea region (ecological
faunistic review) // Helminth fauna of animals of the southern seas / V. A. Vodianitskiy et al. — Kyiv :
Naukova dumka, 1966. — P. 105–113. — Russian : ×åðíûøåíêî À. Ñ. Ïàðàçèòîôàóíà ðûá ëèìàíîâ
Ñåâåðíîãî Ïðè÷åðíîìîðüÿ (ýêîëîãî-ôàóíèñòè÷åñêèé î÷åðê) // Ãåëüìèíòîôàóíà æèâîòíûõ
þæíûõ ìîðåé / Â. À. Âîäÿíèöêèé è äð.
Chernyshenko A. S., Sventsytskaya G. A. Comparative ecological characteristic of the helminth fauna of
gobiids (Fam. Gobiidae) of the estuaries of the North-Western Black Sea region and the Gulf of
Odessa // Questions of the Marine Parasitology: Abstracts of the 1st Symposium on Parasites and
Diseases of Marine Animals (Sevastopol, 1970) / V. A. Vodianitskiy. — Kyiv : Naukova dumka,
1970. — P. 135–137. — Russian : ×åðíûøåíêî À. Ñ., Ñâåíöèöêàÿ Ã. À. Ñðàâíèòåëüíî-ýêîëîãè÷åñêàÿ
õàðàêòåðèñòèêà ãåëüìèíòîôàóíû áû÷êîâ (ñåì. Gobiidae) ëèìàíîâ Ñåâåðî-Çàïàäíîãî
Ïðè÷åðíîìîðüÿ è Îäåññêîãî çàëèâà // Âîïðîñû ìîðñêîé ïàðàçèòîëîãèè: Ìàòåðèàëû I Ñèìï. ïî
ïàðàçèòàì è áîëåçíÿì ìîðñêèõ æèâîòíûõ (Ñåâàñòîïîëü, 1970) / Â. À. Âîäÿíèöêèé.
Gaevskaya A. V., Gusev A. V., Delamure S. L. et al. The keys to the Parasites of Vertebrates of the Black Sea
and the Sea of Azov / V. N. Greze. — Kyiv : Naukova dumka, 1975. — 551 p. — Russian : Ãàåâ-
ñêàÿ À. Â., Ãóñåâ À. Â., Äåëÿìóðå Ñ. Ë. è äð. Îïðåäåëèòåëü ïàðàçèòîâ ïîçâîíî÷íûõ ×åðíîãî è
Àçîâñêîãî ìîðåé / Â. Í. Ãðåçå.
Kvach Yu. The changes of the fauna of helminths of gobiids (Gobiidae) of the Khadzhibey Estuary (1996–
2001) // Vestnik zoologii. — 2004. — Suppl. 18. — P. 66–68. — Ukrainian : Êâà÷ Þ. Çì³íà ôàóíè
ãåëüì³íò³â áè÷ê³â (Gobiidae) Õàäæèáåéñüêîãî ëèìàíó (1996–2001 ðð.) // Âåñòíèê çîîëîãèè: Îòä.
âûï. ¹ 18.
Kvach Yu. A comparative analysis of helminth faunas and infection parameters of ten species of gobid fishes
(Actinopterygii: Gobiidae) from the north-western Black Sea // Acta Ichthyol. Piscat. — 2005 a. —
35. — P. 103–110.
Kvach Yu. Helminth fauna of the grass goby Zosterisessor ophiocephalus of the water bodies of the North-
western Black Sea // Ecological safety of the coastal and shelf zones and complex using of shelf sources
/ V. A. Ivanov et al. — 2005 b. — Is. 12. — P. 603–609. — Russian : Êâà÷ Þ. Ãåëüìèíòîôàóíà áû÷êà
òðàâÿíèêà Zosterisessor ophiocephalus âîäîåìîâ Ñåâåðî-Çàïàäíîãî Ïðè÷åðíîìîðüÿ //
Ýêîëîãè÷åñêàÿ áåçîïàñíîñòü ïðèáðåæíîé è øåëüôîâîé çîí è êîìïëåêñíîå èñïîëüçîâàíèå
ðåñóðñîâ øåëüôà / Â. À. Èâàíîâ è äð.
Kvach Yu., Og ˘uz M. C. Communities of metazoan parasites of two fishes of Proterorhinus genus
(Actinopterygii: Gobiidae) // Helminthologia. — 2009. — 46, N 3. — P. 168–176.
Machkevsky V. K., Mordvinova V. K., Parukhin A. M. Helminth fauna of gobies of the Gulf of Yegorlyk and
Lake Donuzlav, the places of mussels farms location // Ekologiya Morya. — 1990. — Is. 36. —
P. 69–75. — Russian : Ìà÷êåâñêèé Â. Ê., Ìîðäâèíîâà Ò. Í., Ïàðóõèí À. Ì. Ãåëüìèíòîôàóíà
áû÷êîâ Åãîðëûöêîãî çàëèâà è îç. Äîíóçëàâ — ìåñòàõ ðàçìåùåíèÿ ìèäèéíûõ õîçÿéñòâ //
Ýêîëîãèÿ ìîðÿ.
Maillard C. Etude du cycle e´volutif du Tre´matode: Acanthostomum imbutiforme (Molin, 1859) Gohar, 1934,
parasite de Morone labrax (Linne´, 1758) // Annales Parasitologie humaine et compare´e. — 1973 a. —
48, N 1. — P. 33–46.
Maillard C. Mise en e´vidence du cycle e´volutif abre´ge d’Aphalloides coelomicola Dollfus, Chabaud et
Golvan, 1957 (Tre´matode). Notion d’«hôte historique» // Compte Rendu de l’Acade´mie des sciences
de Paris, Se´rie D. — 1973 b. — 277. — P. 317–320.
Maillard C. Cycle e´volutif de Paratimonia gobii Pre´vot et Bartoli 1967 (Trematoda-Monorchiidae) // Acta
Tropica. — 1975. — 32, N 4. — P. 327–323.
Miller P. J. Relictogobius kryzhanovskii and the penetration of Mediterranean gobies into the Black Sea //
Nature. — 1965. — 208. — P. 474–475.
517Parasites of the marbled goby (Pomatoschistus marmoratus)…
Miller P. J. Gobiidae // Fishes of the North-eastern Atlantic and the Mediterranean / P. J. P. Whitehead,
M.-L. Bauchot, J.-C. Hureau et al. — Paris : UNESCO, 1986. — Vol. 3. — P. 1019–1086.
Naidenova N. N. Parasite fauna of fish of the gobiid family of the Black Sea and the Sea of Azov. — Kyiv :
Naukova dumka, 1974. — 182 p. — Russian : Íàéäåíîâà Í. Í. Ïàðàçèòîôàóíà ðûá ñåìåéñòâà
áû÷êîâûõ ×åðíîãî è Àçîâñêîãî ìîðåé.
Pampoulie Ch., Lambert A., Rosecchi E. et al. Host death: A necessary condition for the transmission of
Aphalloides coelomicola Dollfus, Chabaud, and Golvan, 1957 (Digenea, Cryptogonimidae)? //
J. Parasitol. — 2000. — 86, N 2. — P. 416–417.
Ra˘dulescu I. Contribut�iuni la cunoas �terea parasitilor guvidului Gobius microps Kro �yer din Marea Neagra //
Bul. Inst. Cerc. Pisc. — 1953. — 12, N 4. — P. 49–54.
Robert F., Renaud F., Mathieu E., Gabrion C. Importance of the paratenic host in the biology of
Bothriocephalus gregarius (Cestoda, Pseudophyllidea), a parasite of the turbot // Int. J. Parasitol. —
1988. — 18, N 5. — P. 611–621.
Smirnov A. I. Perch-likes (gobiids), scorpionfishes, flatfishes, clingfishes, anglerfishes // Fauna of Ukraine. —
Kyiv : Naukova dumka, 1986. — Vol. 8, fasc 5. — 320 p. — Russian : Ñìèðíîâ À. È. Îêóíåîáðàçíûå
(áû÷êîâûå), ñêîðïåíîîáðàçíûå, êàìáàëîîáðàçíûå, ïðèñîñêîîáðàçíûå, óäèëüùèêîîáðàçíûå //
Ôàóíà Óêðàèíû. — Êèåâ : Íàóê. äóìêà, 1986. — Ò. 8, âûï. 5. — 320 ñ.
Van Valen L. A new evolutionary law // Evol. Theory. — 1973. — 1. — P. 1–30.
Zander C. D. Ecology of host parasite relationships in the Baltic Sea // Naturwissenschaften. — 1998. —
85. — P. 426–436.
Zander C. D. Four-year monitoring of parasite communities in gobiid Fishes of the south-western
Baltic. I. Guild and component community // Parasitol. Res. — 2003. — 90. — P. 502–511.
Zander C. D. Four-year monitoring of parasite communities in gobiid fishes of the south-western Baltic. II.
Infracommunity // Parasitol. Res. — 2004. — 93. — P. 17–29.
Zander C. D., Strohbach U., Groenewold S. The importance of gobies (Gobiidae, Teleostei) as hosts and
transmitters of parasites in the SW Baltic // Helgola �nder Meeresuntersuchengen. — 1993. — 47. —
P. 81–111.
Zander C. D., Kesting V. Colonization and seasonality of goby (Gobiidae, Teleostei) parasites from the
southwestern Baltic Sea // Parasitol. Res. — 1998. — 84. — P. 459–466.
Zander C. D., Reimer L. W., Barz K. et al. Parasite communities of the Salzhaff (Northwest Mecklenburg,
Baltic Sea). II. Guild communities, with special regard to snails, benthic crustaceans, and small-sized
fish // Parasitol. Res. — 2000. — 86. — P. 359–372.
518 Yu. Kvach
<<
/ASCII85EncodePages false
/AllowTransparency false
/AutoPositionEPSFiles true
/AutoRotatePages /None
/Binding /Left
/CalGrayProfile (Dot Gain 20%)
/CalRGBProfile (sRGB IEC61966-2.1)
/CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2)
/sRGBProfile (sRGB IEC61966-2.1)
/CannotEmbedFontPolicy /Error
/CompatibilityLevel 1.4
/CompressObjects /Tags
/CompressPages true
/ConvertImagesToIndexed true
/PassThroughJPEGImages true
/CreateJDFFile false
/CreateJobTicket false
/DefaultRenderingIntent /Default
/DetectBlends true
/DetectCurves 0.0000
/ColorConversionStrategy /CMYK
/DoThumbnails false
/EmbedAllFonts true
/EmbedOpenType false
/ParseICCProfilesInComments true
/EmbedJobOptions true
/DSCReportingLevel 0
/EmitDSCWarnings false
/EndPage -1
/ImageMemory 1048576
/LockDistillerParams false
/MaxSubsetPct 100
/Optimize true
/OPM 1
/ParseDSCComments true
/ParseDSCCommentsForDocInfo true
/PreserveCopyPage true
/PreserveDICMYKValues true
/PreserveEPSInfo true
/PreserveFlatness true
/PreserveHalftoneInfo false
/PreserveOPIComments false
/PreserveOverprintSettings true
/StartPage 1
/SubsetFonts true
/TransferFunctionInfo /Apply
/UCRandBGInfo /Preserve
/UsePrologue false
/ColorSettingsFile ()
/AlwaysEmbed [ true
]
/NeverEmbed [ true
]
/AntiAliasColorImages false
/CropColorImages true
/ColorImageMinResolution 300
/ColorImageMinResolutionPolicy /OK
/DownsampleColorImages true
/ColorImageDownsampleType /Bicubic
/ColorImageResolution 300
/ColorImageDepth -1
/ColorImageMinDownsampleDepth 1
/ColorImageDownsampleThreshold 1.50000
/EncodeColorImages true
/ColorImageFilter /DCTEncode
/AutoFilterColorImages true
/ColorImageAutoFilterStrategy /JPEG
/ColorACSImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/ColorImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/JPEG2000ColorACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000ColorImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasGrayImages false
/CropGrayImages true
/GrayImageMinResolution 300
/GrayImageMinResolutionPolicy /OK
/DownsampleGrayImages true
/GrayImageDownsampleType /Bicubic
/GrayImageResolution 300
/GrayImageDepth -1
/GrayImageMinDownsampleDepth 2
/GrayImageDownsampleThreshold 1.50000
/EncodeGrayImages true
/GrayImageFilter /DCTEncode
/AutoFilterGrayImages true
/GrayImageAutoFilterStrategy /JPEG
/GrayACSImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/GrayImageDict <<
/QFactor 0.15
/HSamples [1 1 1 1] /VSamples [1 1 1 1]
>>
/JPEG2000GrayACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000GrayImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasMonoImages false
/CropMonoImages true
/MonoImageMinResolution 1200
/MonoImageMinResolutionPolicy /OK
/DownsampleMonoImages true
/MonoImageDownsampleType /Bicubic
/MonoImageResolution 1200
/MonoImageDepth -1
/MonoImageDownsampleThreshold 1.50000
/EncodeMonoImages true
/MonoImageFilter /CCITTFaxEncode
/MonoImageDict <<
/K -1
>>
/AllowPSXObjects false
/CheckCompliance [
/None
]
/PDFX1aCheck false
/PDFX3Check false
/PDFXCompliantPDFOnly false
/PDFXNoTrimBoxError true
/PDFXTrimBoxToMediaBoxOffset [
0.00000
0.00000
0.00000
0.00000
]
/PDFXSetBleedBoxToMediaBox true
/PDFXBleedBoxToTrimBoxOffset [
0.00000
0.00000
0.00000
0.00000
]
/PDFXOutputIntentProfile ()
/PDFXOutputConditionIdentifier ()
/PDFXOutputCondition ()
/PDFXRegistryName ()
/PDFXTrapped /False
/Description <<
/CHS <FEFF4f7f75288fd94e9b8bbe5b9a521b5efa7684002000410064006f006200650020005000440046002065876863900275284e8e9ad88d2891cf76845370524d53705237300260a853ef4ee54f7f75280020004100630072006f0062006100740020548c002000410064006f00620065002000520065006100640065007200200035002e003000204ee553ca66f49ad87248672c676562535f00521b5efa768400200050004400460020658768633002>
/CHT <FEFF4f7f752890194e9b8a2d7f6e5efa7acb7684002000410064006f006200650020005000440046002065874ef69069752865bc9ad854c18cea76845370524d5370523786557406300260a853ef4ee54f7f75280020004100630072006f0062006100740020548c002000410064006f00620065002000520065006100640065007200200035002e003000204ee553ca66f49ad87248672c4f86958b555f5df25efa7acb76840020005000440046002065874ef63002>
/DAN <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>
/DEU <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>
/ESP <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>
/FRA <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>
/ITA <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>
/JPN <FEFF9ad854c18cea306a30d730ea30d730ec30b951fa529b7528002000410064006f0062006500200050004400460020658766f8306e4f5c6210306b4f7f75283057307e305930023053306e8a2d5b9a30674f5c62103055308c305f0020005000440046002030d530a130a430eb306f3001004100630072006f0062006100740020304a30883073002000410064006f00620065002000520065006100640065007200200035002e003000204ee5964d3067958b304f30533068304c3067304d307e305930023053306e8a2d5b9a306b306f30d530a930f330c8306e57cb30818fbc307f304c5fc59808306730593002>
/KOR <FEFFc7740020c124c815c7440020c0acc6a9d558c5ec0020ace0d488c9c80020c2dcd5d80020c778c1c4c5d00020ac00c7a50020c801d569d55c002000410064006f0062006500200050004400460020bb38c11cb97c0020c791c131d569b2c8b2e4002e0020c774b807ac8c0020c791c131b41c00200050004400460020bb38c11cb2940020004100630072006f0062006100740020bc0f002000410064006f00620065002000520065006100640065007200200035002e00300020c774c0c1c5d0c11c0020c5f40020c2180020c788c2b5b2c8b2e4002e>
/NLD (Gebruik deze instellingen om Adobe PDF-documenten te maken die zijn geoptimaliseerd voor prepress-afdrukken van hoge kwaliteit. De gemaakte PDF-documenten kunnen worden geopend met Acrobat en Adobe Reader 5.0 en hoger.)
/NOR <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>
/PTB <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>
/SUO <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>
/SVE <FEFF0041006e007600e4006e00640020006400650020006800e4007200200069006e0073007400e4006c006c006e0069006e006700610072006e00610020006f006d002000640075002000760069006c006c00200073006b006100700061002000410064006f006200650020005000440046002d0064006f006b0075006d0065006e007400200073006f006d002000e400720020006c00e4006d0070006c0069006700610020006600f60072002000700072006500700072006500730073002d007500740073006b00720069006600740020006d006500640020006800f600670020006b00760061006c0069007400650074002e002000200053006b006100700061006400650020005000440046002d0064006f006b0075006d0065006e00740020006b0061006e002000f600700070006e00610073002000690020004100630072006f0062006100740020006f00630068002000410064006f00620065002000520065006100640065007200200035002e00300020006f00630068002000730065006e006100720065002e>
/ENU (Use these settings to create Adobe PDF documents best suited for high-quality prepress printing. Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.)
>>
/Namespace [
(Adobe)
(Common)
(1.0)
]
/OtherNamespaces [
<<
/AsReaderSpreads false
/CropImagesToFrames true
/ErrorControl /WarnAndContinue
/FlattenerIgnoreSpreadOverrides false
/IncludeGuidesGrids false
/IncludeNonPrinting false
/IncludeSlug false
/Namespace [
(Adobe)
(InDesign)
(4.0)
]
/OmitPlacedBitmaps false
/OmitPlacedEPS false
/OmitPlacedPDF false
/SimulateOverprint /Legacy
>>
<<
/AddBleedMarks false
/AddColorBars false
/AddCropMarks false
/AddPageInfo false
/AddRegMarks false
/ConvertColors /ConvertToCMYK
/DestinationProfileName ()
/DestinationProfileSelector /DocumentCMYK
/Downsample16BitImages true
/FlattenerPreset <<
/PresetSelector /MediumResolution
>>
/FormElements false
/GenerateStructure false
/IncludeBookmarks false
/IncludeHyperlinks false
/IncludeInteractive false
/IncludeLayers false
/IncludeProfiles false
/MultimediaHandling /UseObjectSettings
/Namespace [
(Adobe)
(CreativeSuite)
(2.0)
]
/PDFXOutputIntentProfileSelector /DocumentCMYK
/PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged
/UntaggedRGBHandling /UseDocumentProfile
/UseDocumentBleed false
>>
]
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [612.000 792.000]
>> setpagedevice
|
| id | nasplib_isofts_kiev_ua-123456789-65734 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0084-5604 |
| language | English |
| last_indexed | 2025-12-07T15:58:55Z |
| publishDate | 2010 |
| publisher | Інститут зоології ім. І.І. Шмальгаузена НАН України |
| record_format | dspace |
| spelling | Kvach, Yu. 2014-07-01T17:01:32Z 2014-07-01T17:01:32Z 2010 Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna / Yu. Kvach // Вестник зоологии. — 2010. — Т. 44, № 6. — С. 509–518. — Бібліогр.: 28 назв. — англ. 0084-5604 https://nasplib.isofts.kiev.ua/handle/123456789/65734 597.585.1:616.99(262.55) The marbled goby, Pomatoschistus marmoratus (Risso, 1810), the Mediterranean immigrant in the Black Sea fauna, was examined for the metazoan parasites. In total, 12 parasite species were found to occurre. The component community consists of 7 trematode species, two nematodes, two acanthocephalans, and one cestode species. Two parasite species (Aphalloides coelomicola and Dichelyne minutus) were represented by both adults and larvae, four parasite species were presented by adults (Asymphylodora pontica, Paratimonia gobii, Acanthocephaloides propinquus, and Telosentis exiguus), and six species were presented by larvae (Bothriocephalus gregarius, Cryptocotyle concavum, C. lingua, Pygidiopsis genata, Timoniella imbutiforme, and Contracaecum microcephalum). T. imbutiforme has the great tendency to join the infracommunity. The core of parasite fauna of the marbled goby consists of two specialists, A. coelomicola and P. gobii, which immigrated to the Black Sea together with their hosts. The co-immigration is a result of co-evolution of the parasites’ and host’s life cycles. Приведены описания 4 видов гименолепидоидных цестод от луговой тиркушки юга Украины: Glareolepis poralae (Meggitt, 1927) Spassky, 1967; Wardium (s. l.) tauricum Kornyushin et Greben, sp. n.; Wardium (s. l.) sp.; Echinocotyloides longirostris (Rudolphi, 1819) Kornyushin, 1983. Детально описана морфология G. poralae. Установлено, что в гермафродитных члениках у G. poralae семенники отсутствуют, а структуры, описанные в литературе как семенники в гермафродитных члениках — это лопасти яичника. Описан новый вид цестод Wardium (s. l.) tauricum sp. n. с аплопараксоидными крючьями 13–15 мкм длиной и конусовидным вооруженным циррусом 20–22 мкм длиной. Еще один вид цестод Wardium (s. l.) sp. из-за отсутствия сколекса определен только до рода. Он отличается от всех известных видов этого рода размерами и вооружением цирруса. en Інститут зоології ім. І.І. Шмальгаузена НАН України Вестник зоологии Экология Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna Гельминты бычка-лысуна (Pomatoschistus marmoratus) — средиземноморского иммигранта в фауне Черного моря Article published earlier |
| spellingShingle | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna Kvach, Yu. Экология |
| title | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna |
| title_alt | Гельминты бычка-лысуна (Pomatoschistus marmoratus) — средиземноморского иммигранта в фауне Черного моря |
| title_full | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna |
| title_fullStr | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna |
| title_full_unstemmed | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna |
| title_short | Helminths of the Marbled Goby (Pomatoschistus marmoratus), a Mediterranean Immigrant in the Black Sea Fauna |
| title_sort | helminths of the marbled goby (pomatoschistus marmoratus), a mediterranean immigrant in the black sea fauna |
| topic | Экология |
| topic_facet | Экология |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/65734 |
| work_keys_str_mv | AT kvachyu helminthsofthemarbledgobypomatoschistusmarmoratusamediterraneanimmigrantintheblackseafauna AT kvachyu gelʹmintybyčkalysunapomatoschistusmarmoratussredizemnomorskogoimmigrantavfaunečernogomorâ |