Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov
Two new species were identified among cestodes collected from Dasyatis pastinaca (Linnaeus, 1758) in the Black Sea and the Sea of Azov. Their morphology corresponds in the most characters to the descriptions of Phyllobothrium lactuca van Beneden, 1850 and P. gracilis Wedl, 1855 by L. Borcea (Borcea,...
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| Zitieren: | Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov / V.V. Kornyushin, T.A. Polyakova // Вестник зоологии. — 2012. — Т. 46, № 4. — С. 291–308. — Бібліогр.: 36 назв. — англ. |
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irk-123456789-1094122016-11-29T03:02:13Z Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov Kornyushin, V.V. Polyakova, T.A. Фауна и систематика Two new species were identified among cestodes collected from Dasyatis pastinaca (Linnaeus, 1758) in the Black Sea and the Sea of Azov. Their morphology corresponds in the most characters to the descriptions of Phyllobothrium lactuca van Beneden, 1850 and P. gracilis Wedl, 1855 by L. Borcea (Borcea, 1934) and T. P. Pogorel’tseva (Pogorel’tseva, 1960) from the same host and seas. В сборах цестод от Dasyatis pastinaca (Linnaeus, 1758) из Черного и Азовского морей идентифицировано два вида цестод, которые по большинству морфологических признаков соответствуют описаниям Phyllobothrium lactuca van Beneden, 1850 и P. gracilis Wedl, 1855 в публикациях Л. Борче (Borcea, 1934) и Т. П. Погорельцевой (1960) по материалам, собранным от этого же хозяина и в этом районе. 2012 Article Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov / V.V. Kornyushin, T.A. Polyakova // Вестник зоологии. — 2012. — Т. 46, № 4. — С. 291–308. — Бібліогр.: 36 назв. — англ. 0084-5604 DOI 10.2478/v10058-012-0025-x http://dspace.nbuv.gov.ua/handle/123456789/109412 595.121:597.317.1(262.5) en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Фауна и систематика Фауна и систематика |
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Фауна и систематика Фауна и систематика Kornyushin, V.V. Polyakova, T.A. Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov Вестник зоологии |
| description |
Two new species were identified among cestodes collected from Dasyatis pastinaca (Linnaeus, 1758) in the Black Sea and the Sea of Azov. Their morphology corresponds in the most characters to the descriptions of Phyllobothrium lactuca van Beneden, 1850 and P. gracilis Wedl, 1855 by L. Borcea (Borcea, 1934) and T. P. Pogorel’tseva (Pogorel’tseva, 1960) from the same host and seas. |
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| author |
Kornyushin, V.V. Polyakova, T.A. |
| author_facet |
Kornyushin, V.V. Polyakova, T.A. |
| author_sort |
Kornyushin, V.V. |
| title |
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov |
| title_short |
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov |
| title_full |
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov |
| title_fullStr |
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov |
| title_full_unstemmed |
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov |
| title_sort |
cairaeanthus gen. n. (cestoda, rhinebothriidea), with the description of two new species from dasyatis pastinaca in the black sea and the sea of azov |
| publisher |
Інститут зоології ім. І.І. Шмальгаузена НАН України |
| publishDate |
2012 |
| topic_facet |
Фауна и систематика |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/109412 |
| citation_txt |
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis pastinaca in the Black Sea and the Sea of Azov / V.V. Kornyushin, T.A. Polyakova // Вестник зоологии. — 2012. — Т. 46, № 4. — С. 291–308. — Бібліогр.: 36 назв. — англ. |
| series |
Вестник зоологии |
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| first_indexed |
2025-07-07T23:01:24Z |
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2025-07-07T23:01:24Z |
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1837031002431553536 |
| fulltext |
ÓÄÊ 595.121:597.317.1(262.5)
CAIRAEANTHUS GEN. N. (CESTODA, RHINEBOTHRIIDEA),
WITH THE DESCRIPTION OF TWO NEW SPECIES
FROM DASYATIS PASTINACA IN THE BLACK SEA
AND THE SEA OF AZOV
V. V. Kornyushin1, T. A. Polyakova2
1 Institute of zoology NAS of Ukraine, B. Chmielnicky str., 15, Kyiv, 01601 Ukraine
E-mail: vadikorn@izan.kiev.ua
2 Institute of Biology of the Southern Seas, NAS of Ukraine, Nakhimov av., 2, Sevastopol, 99011, Ukraine
dE-mail: polyakova-acant@yandex.ru
Received 25 May 2012
Accepted 28 May 2012
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species from Dasyatis
pastinaca in the Black Sea and the Sea of Azov. Kornyushin V. V., Polyakova T. A. – Two new species
were identified among cestodes collected from Dasyatis pastinaca (Linnaeus, 1758) in the Black Sea and
the Sea of Azov. Their morphology corresponds in the most characters to the descriptions of Phyllobothrium
lactuca van Beneden, 1850 and P. gracilis Wedl, 1855 by L. Borcea (Borcea, 1934) and T. P. Pogorel’tseva
(Pogorel’tseva, 1960) from the same host and seas. However, some morphological characters of these
cestodes (namely, the marginal loculi on the pedicellate bothridia, cephalic peduncle, absence of the api-
cal sucker and post-poral testes, genital pore opening marginally, close to posterior end of proglottid)
correspond to diagnosis of Rhinebothriidea. Since the species from the Black Sea and the Sea of Azov
did not fit diagnoses of any known genera currently belonging to Rhinebothriidea, the new genus,
Cairaeanthus Kornyushin et Polyakova, gen. n., with two species, C. ruhnkei Kornyushin et Polyakova,
sp. n. and C. healyae Kornyushin et Polyakova, sp. n., is established. Cairaeanthus gen. n. differs from
Anthocephalum Linton, 1890 in absence of apical sucker, posteriorly bifid bothridia and vitelline fields
interrupted by the ovary, not extending in the posterior end of the proglottid. The new genus can be dis-
tinguished from Rhinebothrium Linton, 1890, Echeneibothium van Beneden, 1850, Rhabdotobothrium Euzet,
1953, Rhinebothroides Mayes, Brooks et Thorson, 1981, Scalithrium Ball, Neifar et Euzet, 2003 by the
absence of facial loculi on the bothridia and by numerous vitelline follicles, stopping at the anterior mar-
gin of the ovary; from Rhodobothrium Linton, 1889 – by absence of post-poral testes, and from
Spongiobothrium Linton, 1889 – by numerous vitelline follicles, stopping at the level of ovary. C. ruhnkei
sp. n. differs from C. healyae sp. n. in the lengths of the strobila, the cephalic peduncle and the both-
ridia pedicel, in size of marginal loculi and the ovary, in the number of the proglottids and the testes in
the proglottid, and in the structure of genital atrium. Two new species are differentiated also from the
morphologically related species Phyllobothrium pastinacae Mokhtar-Mocamouri, Zamali, 1981. Examined
type specimens of P. pastinacae were attributed to Rhinebothiidea based on a set of morphological char-
acters. However, they appear to differ from all known Rhinebothriidea, including the species from the
genera Anthocephalum Linton, 1890 and Cairaeanthus gen. n. Taxonomic position of P.pastinacae needs
further identification.
Key wo rd s: Cairaeanthus gen. n., Rhinebothriidea, Cairaeanthus ruhnkei sp. n., Cairaeanthus healyae sp. n.,
Phyllobothrium pastinacae, Dasyatis pastinaca, the Mediterranean Sea, the Black Sea, the Sea of Azov.
Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), îïèñàíèå äâóõ íîâûõ âèäîâ îò Dasyatis pastinaca èç
×åðíîãî è Àçîâñêîãî ìîðåé. Êîðíþøèí Â. Â., Ïîëÿêîâà Ò. À. –  ñáîðàõ öåñòîä îò Dasyatis pasti-
naca (Linnaeus, 1758) èç ×åðíîãî è Àçîâñêîãî ìîðåé èäåíòèôèöèðîâàíî äâà âèäà öåñòîä, êîòî-
ðûå ïî áîëüøèíñòâó ìîðôîëîãè÷åñêèõ ïðèçíàêîâ ñîîòâåòñòâóþò îïèñàíèÿì Phyllobothrium lactu-
ca van Beneden, 1850 è P. gracilis Wedl, 1855 â ïóáëèêàöèÿõ Ë. Áîð÷å (Borcea, 1934) è
Ò. Ï. Ïîãîðåëüöåâîé (1960) ïî ìàòåðèàëàì, ñîáðàííûì îò ýòîãî æå õîçÿèíà è â ýòîì ðàéîíå.
Îäíàêî îïðåäåëåííûå ìîðôîëîãè÷åñêèå ïðèçíàêè ýòèõ öåñòîä (íàëè÷èå êðàåâûõ ëîêóë íà ñòå-
áåëü÷àòûõ áîòðèäèÿõ è ñòåáëÿ ñêîëåêñà, îòñóòñòâèå àïèêàëüíûõ ïðèñîñîê íà ñêîëåêñå è ïîñòâà-
ãèíàëüíîé ãðóïïû ñåìåííèêîâ, íàðÿäó ñ ðàñïîëîæåíèåì ïîëîâîãî îòâåðñòèÿ ìàðãèíàëüíî âáëè-
Vestnik zoologii, 46(4): e-1—e-18, 2012
DOI 10.2478/v10058-012-0025-x
Ôàóíà è ñèñòåìàòèêà
Unauthenticated
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çè çàäíåãî êðàÿ ïðîãëîòòèäû) ñîîòâåòñòâóþò äèàãíîçó îòðÿäà Rhinebothriidea. Ïîñêîëüêó öåñòî-
äû îò ÷åðíîìîðñêèõ ñêàòîâ íå ñîîòâåòñòâóåò äèàãíîçó íè îäíîãî èç èçâåñòíûõ íà ñåãîäíÿ ðîäîâ
îòðÿäà Rhinebothriidea, äëÿ íèõ îáîñíîâàí íîâûé ðîä Cairaeanthus Kornyushin et Polyakova,
gen. n. ñ äâóìÿ íîâûìè âèäàìè C. ruhnkei Kornyushin et Polyakova, sp. n. è C. healyae Kornyushin
et Polyakova, sp. n. Íîâûé ðîä îòëè÷àåòñÿ îò ðîäà Anthocephalum Linton, 1890 îòñóòñòâèåì àïè-
êàëüíûõ ïðèñîñîê íà ñêîëåêñå, ðàçäâîåííûìè â çàäíåé ÷àñòè áîòðèäèÿìè è òåì, ÷òî ïîëÿ æåë-
òî÷íûõ ôîëëèêóëîâ ïðåðûâàþòñÿ íà óðîâíå ÿè÷íèêà, íå äîñòèãàÿ çàäíåãî êðàÿ ïðîãëîòòèäû. Îò
Echeneibothium van Beneden, 1850, Rhinebothrium Linton, 1890, Rhabdotobothrium Euzet, 1953,
Rhinebothroides Mayes, Brooks et Thorson, 1953, Scalithrium Ball, Weifar, Euzet, 2003 íîâûé ðîä îòëè-
÷àåòñÿ îòñóòñòâèåì ëèöåâûõ ëîêóë íà áîòðèäèÿõ è òåì, ÷òî æåëòî÷íûå ïîëÿ äîñòèãàþò òîëüêî ïåðåä-
íåãî êðàÿ ÿè÷íèêà; îò Rhodobothrium Linton, 1889 – îòñóòñòâèåì ïîñòâàãèíàëüíîé ãðóïïû ñåìåí-
íèêîâ, îò Spongiobothrium Linton, 1889 – ìíîãî÷èñëåííûìè æåëòî÷íûìè ôîëëèêóëàìè, ïîëÿ êîòî-
ðûõ ïðåðûâàþòñÿ íà óðîâíå ÿè÷íèêà. Âèäû C. ruhnkei sp. n. è C. healyae sp. n. îòëè÷àþòñÿ äðóã
îò äðóãà äëèíîé ñòðîáèëû, ñòåáëÿ ñêîëåêñà è ñòåáåëüêà áîòðèäèé, êîëè÷íåñòâîì ëîêóë íà áîò-
ðèäèÿõ, êîëè÷åñòâîì ïðîãëîòòèä, ñåìåííèêîâ â ïðîãëîòòèäå, ôîðìîé ÿè÷íèêà, ñòðîåíèåì ïîëî-
âîãî àòðèóìà. Íîâûå âèäû äèôôåðíåöèðîâàíû òàêæå îò Phyllobothrium pastinacae Mokhtar-
Mocamouri et Zamali, 1981 îò D. pastinaca èç Ñðåäèçåìíîãî ìîðÿ íà îñíîâàíèè ïåðåèçó÷åíèÿ òèïî-
âîãî ìàòåðèàëà. Ïî íàëè÷èþ îïðåäåëåííûõ ìîðôîëîãè÷åñêèõ ïðèçíàêîâ ýòè öåñòîäû òàêæå
îòíåñåíû ê îòðÿäó Rhinebothriidea. Îäíàêî îíè îòëè÷àþòñÿ îò ïðåäñòàâèòåëåé âñåõ èçâåñòíûõ íà
ñåãîäíÿ ðîäîâ Rhinebothriidea, â òîì ÷èñëå îò Anthocephalum Linton, 1890 è Cairaeanthus
gen. n. Òàêñîíîìè÷åñêèé ñòàòóñ ýòèõ öåñòîä òðåáóåò îòäåëüíîãî óòî÷íåíèÿ.
Êëþ÷åâûå ñ ëîâ à: Cairaåanthus gen. n., Rhinebothriidea, Cairaåanthus ruhnkei sp. n., Cairaåanthus
healyae sp. n., Phyllobothrium pastinacae, Dasyatis pastinaca, Ñðåäèçåìíîå ìîðå, ×åðíîå ìîðå,
Àçîâñêîå ìîðå.
Introduction
Cestodes of the genus Phyllobothrium van Beneden, 1849 from the dasyatid stingray Dasyatis pastinaca (L.,
1758) in the Black Sea were first reported by L. Borcea (Borcea, 1934). In his article on the cestode fauna
from the Black Sea elasmobranch fishes two species were recognized as Phyllobothrium lactuca van Beneden,
1850 and P. gracilis Wedl, 1855, as new for the Black Sea. Afterwards, many researchers (e. g. Chulkova, 1939;
Osmanov, 1940; Chernyshenko, 1949, 1955; Pogorel’tseva, 1952, 1960, 1964, 1970; Reshetnikova, 1955;
Kurashvili, 1960; Gayevskaya et al., 1975; Mange, 1993; Miroshnichenko, 2004) reported these parasites from
stingrays D. pastinaca and Raja clavata L., 1758 in the Black Sea and the Sea of Azov. After L. Borcea (Borcea,
1934) the descriptions of these species and their schematic figures were presented only in one article
(Pogorel’tseva, 1960). So far as original collections of phyllobothriid of T. P. Pogorel’tseva were lost, identi-
fications of cestodes from the Black Sea stingrays in subsequent publications were based only on the above-
mentioned descriptions.
Among the cestodes collected from D. pastinaca in the Black Sea we identified two species correspond-
ing to the descriptions of P. lactuca van Beneden, 1850 sensu L. Borcea (1934) and T. P. Pogorel’tseva (1960)
and P. gracilis Wedl, 1855 sensu L. Borcea (1934) and T. P. Pogorel’tseva (1960) by most morphological char-
acters. However, comparison of morphology of these cestodes with the type description of P. lactuca from the
shark Mustelus mustelus (L., 1758) in the North Sea (Ruhnke, 1996a) revealed significant differences and uncon-
formity of the cestodes from D. pastinaca in the Black Sea to the present diagnosis of Phyllobothrium.
Moreover, species from the Black Sea stingrays are neither members of the family Phyllobothriidae Braun, 1900
nor members of the order Tetraphyllidea Carus, 1863. According to such morphological characters as presence
of marginal loculi in pedicellate bothridia, cephalic peduncle, absence of apical suckers and post-poral testes
these species was attributed to order Rhinebothriidea Healy, Caira, Jensen, Webster, Littlewood, 2009 (Healy
et al., 2009). P. gracilis have been transferred to Anthocephalum Linton, 1890 (Ruhnke, 1994 b) and then to
the order Rhinebothriidea (Healy et al., 2009), but species parasitizing D. pastinaca from the Black Sea and
the Sea of Azov significantly differ from the description of this species as well as from the diagnosis of
Anthocephalum in general. Subsequent analysis shown that these species do not completely coincide with the
diagnosis of any known genera presently belonging to Rhinebothriidea.
Therefore, cestodes parasitizing D. pastinaca in the Black Sea and the Sea of Azov, previously identified
as P. lactuca and P. gracilis (Borcea, 1934; Pogorel’tseva, 1960), are determined herein as two new species from
the new genus belonging to the order Rhinebothriidea. In this paper, the descriptions of Cairaeanthus
Kornyushin et Polyakova, gen. n. and two new species are presented.
Material and methods
Cestodes were collected from dasyatid stingray D. pastinaca captured in 1977 and during 2000—2011 in
the coastal water of the Black Sea and the Sea of Azov. The spiral intestines from 112 fish, two of which were
from the Sea of Azov, were examined. 437 cestodes belonging to the new genus were found in 64 stingray spec-
imens and there were 229 mature worms among them. Worms were removed from the spiral intestine, washed
e-2 V. V. Kornyushin, T. A. Polyakova
Unauthenticated
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in fresh water to relax their musculature and everted cirrus, and fixed in 70° ethanol (Bychowskaya-Pavlovskaya,
1969). Cestodes were stained with acetocarmine, dehydrated in an ethanol series (70-100°), cleared in clove
oil and mounted in Canada balsam (Roskin, Levinson, 1957). For comparison type specimens of Phyllobothrium
pastinacae Mokhtar-Mocamouri, Zamali, 1981 from D. pastinaca from the Mediterranean Sea (holotype
MNHN 90HB148 c VII and paratypes MNHN90HB149 c VII, MNHN90HB150 c VII) deposited in the
Museum Nationale d’Histoire Naturelle parasitological collection (Paris, France) were investigated.
Measurements in the text are given as the range, followed in parentheses by the mean, the standard error,
the number of worms examined (n) and the total number of observations if more than one observation per
worm was taken (n). Measurements are reported in micrometres, unless otherwise stated. Measuring and light
micrographs were made using the light microscope Zeiss Axio Imager M 1. All figures were produced in scal-
able vector graphics using the program Inkscape 0.48.2.—1 (Scalable Vector Graphics, 2011). Data analysis was
carried out using independent t-tests and Principal Component Analysis (StatSoft Inc., 2001). The statistical
analyses and their graphical representation were produced using the Statistica 6 for Windows software pack-
age.
Cairaåanthus Kornyushin et Polyakova, gen. n.
Rh inebo th r i i d e a. Mid-size, apolytic worms. Scolex tetrabothridiate. Each both-
ridia posteriorly bifid. Myzorhynchus absent. Bothridia pedicellate, each with numerous
marginal loculi. Apical sucker absent. Cephalic peduncle present. Strobila craspedote, mul-
tiproglottided. Testes numerous, median, arranged in several layers; post-poral testes absent.
Vas deferens convoluted; seminal vesicle absent. Cirrus armed with spinitriches. Genital
pores opening marginally close to posterior end of proglottids, irregularly alternating. Vagina
opening into genital atrium anterior to cirrus-sac. Vaginal sphincter absent. Ovary near
posterior end of proglottid, median, H-shaped, tetralobed in cross-section; lobes asym-
metrical; lobulated to follicular. Vitellarium follicular, numerous, lateral; consisting of 2 wide
lateral bands, extending from lateral margin to median line of proglottid, surrounding excre-
tory ducts. Vitelline fields not interrupted at level of genital atrium, becoming thin, stop-
ping at anterior margin of ovary in mature proglottids, not extending posteriorly to ovary.
Uterus ventral, reaching anterior margin of proglottid, with inner pocket-like partition-
ing. Parasites of Dasyatidae (Pisces).
T yp e s p e c i e s: Cairaeanthus ruhnkei Kornyushin et Polyakova, sp. n.
O t h e r s p e c i e s: Cairaeanthus healyàå Kornyushin et Polyakova, sp. n.
E t ymo l o g y: The genus is named for Dr. Janine N. Caira, for her invaluable con-
tributions to the investigation of tetraphyllidean and rhinebothriidean cestodes; and for
the Greek anthus – “flower”.
D i f f e r e n t i a l d i a g no s i s. Cairaeanthus gen. n. corresponds to the diagnosis of
Rhinebothiidea (Healy et al., 2009). Among eight valid genera of this order, Anthocephalum
most closely resembles Cairaeanthus in the presence of the numerous marginal loculi on
the slightly pedicellate bothridia and folded bothridia, in absence of the post-poral
testes, and in the marginal position of the genital pore near the posterior end of the proglot-
tid. The new genus differs from Anthocephalum in the absence of apical sucker, in pos-
teriorly bifid bothridia, and in vitelline fields interrupted by the ovary not extending in
the posterior end of the proglottid. Cairaeanthus can be distinguished from other rhinoboth-
riidean genera as follows: from Echeneibothium van Beneden, 1850; Rhinebothrium
Linton, 1890; Rhabdotobothrium Euzet, 1953; Rhinebothroides Mayes, Brooks, Thorson,
1981; Scalithrium Ball, Neifar, Euzet, 2003 – by absence of facial loculi on bothridia and
by numerous vitelline follicles, stopping at the anterior margin of the ovary; from
Rhodobothrium Linton, 1889 – by absence of post-poral testes, and from Spongiobothrium
Linton, 1889 – by numerous vitelline follicles, stopping at the level of ovary (Linton,
1889; Euzet, 1994; Ruhnke, 1994; Ball et al., 2003).
Among the other cestode taxa, parasitizing elasmobranches, representatives of the
family Phyllobothriidae from the order Tetraphyllidea are similar to Cairaeanthus. The
new genus resembles Phyllobothrium (Ruhnke, 1996 à) in the presence of posteriorly bifid
bothridia, but differs from it in the presence of the marginal loculi in the pedicellate both-
ridia and the absence of the neck, the apical sucker, the glandular apical organ, the vagi-
nal sphincter and the post-poral testes. Moreover, cestodes of the new genus are apoly-
e-3Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
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tic vs anapolytic worms in Phyllobothrium spp. By absence of apical sucker Cairaeanthus
gen. n. differs also from the following phyllobothriidean genera: Anthobothrium van
Beneden, 1850; Orygmatobothrium Diesing, 1863; Monorygma Diesing, 1863; Dinobothrium
van Beneden, 1889; Crossobothrium Linton, 1889; Calyptrobothrium Monticelli, 1893;
Marsupiobothrium Yamaguti, 1952; Gastrolecithus Yamaguti, 1952; Chimaerocestos Williams,
Bray, 1984; Clistobothrium Dailey, Vogelbein, 1990; Paraorygmatobothrium Ruhnke,
1994; Anindobothrium Marques, Brook, Lasso, 2001; Orectolobicestus Ruhnke, Caira,
Carpenter, 2006 (Yamaguti, 1959; Euzet, 1994; Ruhnke, 1994, 1996b; Marques et al.,
2001; Ruhnke et al., 2006; Ruhnke, Caira, 2009). The new genus can be distinguished
from the genera Thysanocephalum Linton, 1889; Myzocephalus Shipley, Hornell, 1906;
Myzophyllobothrium Shipley, Hornell, 1906; Pithophorus Southwell, 1925; Scyphophyllidium
Woodland, 1927; Caulobothrium Baer, 1948; Glyphobothrium Williams, Campbell, 1977;
Duplicibothium Williams, Campbell, 1978 by the absence of post-poral testes. From
Trilocularia Olsson, 1867; Tritaphros Loennberg, 1889; Carpobothrium Shipley, Hornell,
1906; Pseudanthobothrium Baer, 1956; Pentaloculum Alexander, 1963; Phormobothrium
Alexander, 1963; Zyxibothrium Hayden, Campbell, 1981 new genus can be differentiat-
ed by the presence of marginal loculi, and from Clydonobothrium Euzet, 1959 – by the
absence of the myzorhynchus (Euzet, 1994; Yamaguti, 1959; Randhawa et al., 2008).
Cairaeanthus ruhnkei Kornyushin et Polyakova, sp. n. (fig. 1—8)
Typ e ho s t: Dasyatis pastinaca (L., 1758) (Dasyatidae).
T y p e l o c a l i t y: Karantinnaya Bay, Sevastopol, the Black Sea (44°36.41'N,
33°29.54'E).
Add i t i ona l l oca l i t i e s: the Black Sea (off Sevastopol, Streletskaya and Martynova
Bay; off the cape Sarych and Phoros; Karkinitskiy Bay; the Black Sea Biosphere Reserve,
Yagorlytskiy Bay; water area of Karadag Nature Reserve; Kerch Channel) and the Sea
of Azov (Obitochny Bay).
S i t e o f i n f e c t i o n: spiral intestine.
Type spec imens: holotype Ñ. 186.002.32 (06.11.2003; 1349/157; spiral valve N 17;
1 specimen; Karantinnaya Bay, Sevastopol, the Black Sea; IBSS, DEP; coll. T. A. Po -
lyakova) and paratypes C. 177.002.19 (11.09.2003; 1154/146; spiral valve N 8; 1 speci-
men; Streletskaya Bay, Sevastopol, the Black Sea; IBSS, DEP; coll. T. A. Polyakova),
Ñ. 178.002.20 (11.09.2003; 1154/147; spiral valve N 18; 1 spe cimen; Streleckaya bay,
Sevastopol, the Black Sea; IBSS, DEP; coll. T. A. Polyakova), Ñ. 179.002.21 (11.09.2003;
1154/148; spiral valve N 15; 1 specimen; Streleckaya Bay, Sevastopol, the Black Sea; IBSS,
DEP; coll. T. A. Polyakova), C. 199.002.47 (29.09.2005; 2297/165; spiral valve N 11; 1
specimen; Karantinnaya Bay, Sevastopol, the Black Sea; IBSS; Department Ecological
Parasitology; coll. T. A. Polyakova), Ñ. 206.002.67 (29.09.2005; 2297/166; spiral valve
N 14; 1 specimen; Karantinnaya Bay, Sevastopol, the Black Sea; IBSS, DEP;
coll. T. A. Polyakova) ex D. pastinaca from the Black Sea deposited in the collection of
the Department of Ecological Parasitology, Institute of Biology of the Southern Seas
NASU, Sevastopol. Additional paratypes CP 35.1 and CP 35.2 (04.07.1977; mk 12; Black
Sea, Biosphere Reserve, Yagorlytskiy Bay; Institute of Zoology; coll. V. V. Kornyushin)
ex D. pastinaca from the Black Sea are in the collection of the Department of Parasitology,
I. I. Schmalhausen Institute of Zoology, NASU, Kyiv.
E t ymo l o g y: This species is named for Dr Timothy R. Ruhnke, for his contribu-
tion to the study of phyllobothriidean cestodes.
De s c r i p t i on. Based on holotype, 5 paratypes and 24 specimens from D. pastinaca
in the Black Sea and 6 specimens from the same host in the Sea of Azov.
Worms mid-size, craspedote, apolytic, 56—255 mm (123 ± 8.3; n = 36) long,
300—2200 (957 ± 57; n = 36; n = 72) wide. Proglottids 495—1803 (981 ± 48; n = 36)
in number. Scolex large, 800—3130 (1472 ± 76; n = 33) long, 1100—2800 (1832 ± 71;
n = 33) wide, tetrabothridiate (fig. 1, 5). Myzorhynchus absent. Apical sucker absent.
Bothridia 110—1500 (836 ± 50; n = 33; n = 34) long, 200—1500 (622 ± 40; n = 33; n =
e-4 V. V. Kornyushin, T. A. Polyakova
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51) wide; posteriorly bifid; with numerous marginal loculi, each 50—100 (59 ± 3; n =
31) long, 30—100 (55 ± 3; n = 31) wide; with short and wide pedicel, 100—500 (279 ±
19; n = 32; n = 37) long, 100—500 (242 ± 18; n = 32; n = 34) wide (fig. 6). Cepalic
peduncle indistinct, short, 200—680 (460 ± 22; n = 36) long, 280—900 (546 ± 23; n =
36) wide. Lateral excretory ducts 10—70 (35 ± 2; n = 36) in diameter.
Immature proglottids rectangular, 20—200 (117 ± 7; n = 36; n = 48) long, 280—1550
(688 ± 35; n = 36; n = 49) wide (fig. 2). Mature proglottids trapezoid, almost square
e-5Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
Fig. 1—4. Cairaeanthus ruhnkei ex Dasyatis pastinaca from the Black Sea: 1 – scolex, paratype C. 178.002.20;
2 – immature proglottid, holotype C. 186.002.23; 3 – cirrus, holotype C. 186.002.23; 4 – mature proglot-
tid, holotype C. 186.002.23. Scale bars: 1 – 300 μm; 2 – 100 μm; 3, 4 – 200 μm.
Ðèñ. 1—4. Cairaeanthus ruhnkei îò Dasyatis pastinaca èç ×åðíîãî ìîðÿ: 1 – ñêîëåêñ, ïàðàòèï C178.002.20,
2 – íåïîëîâîçðåëàÿ ïðîãëîòòèäà, ãîëîòèï C. 186.002.23; 3 – öèððóñ, ãîëîòèï C. 186.002.23; 4 – ïîëî-
âîçðåëàÿ ïðîãëîòòèäà, ãîëîòèï C. 186.002.23. Ìàñøòàáíàÿ ëèíåéêà: 1 – 300 ìêì; 2 – 100 ìêì; 3, 4 –
200 ìêì.
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130—630 (315 ± 18; n = 36; n = 41) long, 550—1760 (1166 ± 45; n = 36; n = 39) wide.
Terminal proglottids 330—2200 (1027 ± 86; n = 35; n = 40) long, 630—2200 (1207 ±
46; n = 36; n = 39) wide (figs 4, 8). Mature proglottid with 51—98 (83 ± 2; n = 36; n =
43) testes. Testes round, 30—130 (57 ± 3; n = 36; n = 41) in diameter, median,
arranged in two layers; post-poral testes absent (fig. 2, 4, 8). Cirrus-sac pyriform or spher-
ical, 50—300 (204 ± 12; n = 36; n = 38) long, 40—270 (158 ± 10; n = 36) wide (fig. 3,
7). Cirrus armed with spinitriches. Total length of everted cirrus 800—850 (833 ± 17; n =
3), of inverted 220—750 (482 ± 23; n = 34; n = 44) (fig. 4). Vas deferens 10—100 (61 ± 5;
e-6 V. V. Kornyushin, T. A. Polyakova
Fig. 5—8. Photomicrographs of Cairaeanthus ruhnkei ex Dasyatis pastinaca from the Black Sea: 5 – scolex, paratype
C178.002.20; 6 – marginal locules, paratype C178.002.20; 7 – cirrus, holotype C186.002.23; 8 mature
proglottid, holotype C186.002.23. Scale bars: 5 – 300 μm; 6 – 50 μm; 7, 8 – 200 μm.
Ðèñ. 5—8. Ìèêðîôîòîãðàôèÿ Cairaeanthus ruhnkei îò Dasyatis pastinaca èç ×åðíîãî ìîðÿ: 5 – ñêîëåêñ,
ïàðàòèï C. 178.002.20, 6 – êðàåâûå ëîêóëû, ïàðàòèï C. 178.002.20; 7 – öèððóñ, ãîëîòèï C. 186.002.23;
8 – ïîëîâîçðåëàÿ ïðîãëîòòèäà, ãîëîòèï C. 186.002.23. Ìàñøòàáíûå ëèíåéêè: 5 – 300 ìêì; 6 – 50 ìêì;
7, 8 – 200 ìêì.
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n = 36; n = 38) in diameter, coiled, extending anteriorly to cirrus-sac, displaced to poral
side in mature proglottids; seminal vesicle absent. Genital pores lateral, 15—57 % (32 ±
2; n = 36; n = 44) of proglottid length from posterior end, irregularly alternating. Vagina
extending from Mehlis’ gland anteriorly, initially median, strongly wriggle, then lateral
to cirrus-sac in mature proglottids, opening anterior to cirrus-sac in genital atrium; vagi-
nal sphincter absent. Genital atrium simple, tubular, 50—300 (110 ± 9; n = 36) long,
30—200 (91 ± 7; n = 36) wide (fig. 3, 7). Ovary near posterior end of proglottids, medi-
an, H-shaped, tetralobed in cross-section; lobes asymmetrical. In mature proglottids, ovary
lobes surrounding excretory ducts, extending almost to lateral margin of proglottid, sep-
arating vitellarium from posterior end of proglottid. Aporal ovary lobe 100—650 (268 ±
27; n = 36) long, 110—700 (329 ± 21; n = 36) wide, poral lobe 100—600 (250 ± 25; n
= 36) long, 100—600 (312 ± 20; n = 36) wide (fig. 4, 8). Ovicapt at posterior margin
of isthmus, large, spherical or oviform, 50—200 (84 ± 5.4; n = 36; n = 39) in diameter.
Mehlis’ gland 20—200 (61 ± 8; n = 32) wide, invisible in some proglottids. Uterus ven-
tral to vagina, extending from isthmus to median line of proglottid, not reaching its ante-
rior end (fig. 4, 8). Vitellarium follicular; follicles numerous, 10—90 (43 ± 4; n = 36) in
diameter in mature proglottids (fig. 2, 4, 8); in 2 lateral fields, 100—720 (246 ± 19; n =
36; n = 39) wide, each consisting of one dorsal and one ventral row of follicles, between
which excretory ducts passes; extending from anterior end of proglottid to anterior mar-
gin of ovary, not interrupted at level of genital atrium, becoming thin, not extending pos-
teriorly to ovary. Eggs oval, 25—40 (36 ± 3; n = 4; n = 6) long, 20 wide (n = 4).
Rema r k s. Absence of apical sucker is considered as a main differential character
of the new genus. For the proof of this fact, the living specimens of Cairaeanthus were
studied. So far as the marginal loculi of C. ruhnkei are small and the apical suckers may
be only slightly larger than the marginal loculi, that makes their recognition difficult, espe-
cially on fixed material (e. g. in many species of Anthocephalum), movements of both-
ridia in the living specimens were examined. Additional structures on the bothridia were
not found in both alive and fixed specimens.
Di f f e r en t i a l d i agnos i s. Cairaeanthus ruhnkei sp. n. differs from all known species
of the closely related genus Anthocephalum, namely A. gracile (Wedl, 1855), A. centru-
rum Ruhnke, 1994, A. alicae Ruhnke, 1994, A. cairae Ruhnke, 1994, A. duszynskii
Ruhnke, 1994, A. kingae Ruhnke, Seaman, 2009, A. michaeli Ruhnke, Seaman, 2009,
A. lukei Ruhnke, Seaman, 2009 and A. currani Ruhnke, Seaman, 2009 in a number of
characters. In addition to differences mentioned above in the generic diagnosis, C. ruhnkei
can be distinguished from them by larger total length (56—255 mm vs 40—50 mm,
11—20 mm, 4—9 mm, 8—14 mm, 18—31 mm, 8.6—13 mm, 5.7—16.3 mm, 7.9—17.2 mm,
6.6—14.4 mm, respectively) and by more proglottids per worm (495—1803 vs 500—600,
35—65, 9—15, 80—110, 120—160, 33—50, 23—41, 28—56, 35—70, respectively). Moreover,
C. ruhnkei differs from A. gracile in smaller number of testes (51—98 vs 100—130) and
from A. cairae, A. kingae, A. michaeli, A. lukei and A. currani, on the contrary, in larg-
er number of testes (51—98 vs 28—52, 30—37, 30—49, 32—48, 37—50 respectively).
Cairaeanthus healyae Kornyushin et Polyakova, sp. n. (fig. 9—17)
Typ e ho s t: Dasyatis pastinaca (L., 1758) (ñåì. Dasyatidae).
T yp e l o c a l i t y: Martynova Bay, Sevastopol, Black Sea (44°37.08'N, 33°30.28'E).
Add i t i o n a l l o c a l i t i e s: the Black Sea (off Sevastopoloff; the cape Sarych and
Phoros; Karkinitskiy Bay, of the Black Sea Biosphere Reserve, Yagorlytskiy Bay).
S i t e o f i n f e c t i o n: spiral intestine.
Type spec imens: holotype Ñ. 137.001.163 (10.07.2002; 982/139; spiral valve N 8;
1 specimen; Martynova Bay, Sevastopol, the Black Sea; IBSS, DEP; coll. T. A. Polyakova)
and paratypes Ñ. 26.001.27 (21.07.2001; 87/30à; spiral valve N 11; 1 specimen; Martynova
Bay, Sevastopol, the Black Sea; IBSS, DEP; coll. T. A. Polyakova), Ñ. 48.001.54
(28.12.2001; 186/45; spiral valve N 10; 1 specimen; cape Phoros, the Black Sea; IBSS,
DEP; coll. T. A. Polyakova), Ñ. 49.001.55 (28.12.2001; 186/46; spiral valve N 10; 1 spec-
e-7Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
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imen; cape Phoros, the Black Sea; IBSS, DEP; coll. T. A. Polyakova), Ñ. 59.001.65
(28.12.2001; 186/57; spiral valve N 10; 1 specimen; cape Phoros, the Black Sea; IBSS,
DEP; coll. T. A. Polyakova) ex D. pastinaca from the Black Sea deposited in the col-
lection of the Department of Ecological Parasitology, Institute of Biology of the Southern
Seas NASU, Sevastopol. Additional paratypes CP 36.1 (04.07.1977; mk 12; Black Sea,
Biosphere Reserve, Yagorlytskiy Bay; Institute of Zoology; coll. V. V. Kor nyushin) ex
e-8 V. V. Kornyushin, T. A. Polyakova
Fig. 9—12. Cairaeanthus healyae ex Dasyatis pastinaca from the Black Sea: 9 – scolex, paratype Ñ. 26.001.27;
10 immature proglottid, holotype Ñ. 137.001.163; 11 – inverted cirrus, holotype Ñ. 137.001.163; 12 – mature
proglottid, holotype Ñ. 137.001.163 (vitelline follicles in poral side proglottides have been omitted). Scale bars:
9 – 300 μm; 10, 12 – 200 μm; 11 50 μm.
Ðèñ. 9—12. Cairaeanthus healyae îò Dasyatis pastinaca èç ×åðíîãî ìîðÿ: 9 – ñêîëåêñ, ïàðàòèï Ñ. 26.001.27; 10 –
íåïîëîâîçðåëàÿ ïðîãëîòòèäà, ãîëîòèï Ñ. 137.001.163; 11 – èíâàãèíèðîâàííûé öèððóñ, ãîëîòèï Ñ. 137.001.163;
12 – ïîëîâîçðåëàÿ ïðîãëîòòèäà, ãîëîòèï Ñ. 137.001.163 (æåëòî÷íûå ôîëëèêóëû íà ïîðàëüíîé ñòîðîíå ïðî-
ãëîòòèäû áûëè îïóùåíû). Ìàñøòàáíàÿ ëèíåéêà: 9 – 300 ìêì; 10, 12 – 200 ìêì; 11 – 50 ìêì.
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D. pastinaca from the Black Sea are in the collection of the Department of Parasitology,
I. I. Schmalhausen Institute of Zoology, NASU, Kyiv.
E t ymo l o g y: This species is named for Dr Claire J. Healy, for her contribution to
the study of tetraphyllidean and rhinobothriidean tapeworms.
De s c r i p t i on. Based on holotype, 4 paratypes and 26 specimens from D. pastinaca
from the Black Sea.
Worms mid-size, craspedote, apolytic, 108—315 mm (188 ± 8.8; n = 31) long,
200—1700 (687 ± 49; n = 31; n = 62) wide. Proglottids 1003—2395 (1450 ± 74; n = 31)
in number. Scolex large, 720—3000 (1650 ± 75; n = 31; n = 34) long, 1250—3000 (2133 ±
67; n = 31; n = 35) wide, tetrabothridiate (figs 9, 13). Myzorhynchus absent. Bothridia
600—1620 (1002 ± 42; n = 30; n = 33) long, 260—1440 (631 ± 42; n = 30; n = 54) wide;
posteriorly bifid; with numerous marginal loculi, each 10—30 (25 ± 1; n = 31; n = 51)
long, 10—30 (24 ± 1; n = 31; n = 51) wide; with pedicel 180—500 (318 ± 14; n = 31;
n = 38) long, 130—540 (244 ± 14; n = 31; n = 41) wide (fig. 14). Apical sucker absent.
Cephalic peduncle very long, 2.2—11.7 mm (8.1 ± 0.3; n = 31) in length, 200—750 (465 ±
20; n = 31; n = 43) wide. Lateral excretory ducts 10—90 (31 ± 2.2; n = 31; n = 43) in
diameter.
Immature proglottids square, 30—380 (130 ± 8; n = 31; n = 71) long, 230—950 (516 ±
19; n = 31; n = 70) wide, initially wider than long (fig. 10), becoming longer than wide
with maturity. Mature proglottids trapezoid, 230—1170 (445 ± 28; n = 31; n = 38) long,
400—1500 (918 ± 39; n = 31; n = 41) wide. Terminal proglottids, 520—2100 (1024 ±
63; n = 31; n = 50) long, 320—1700 (859 ± 44; n = 31; n = 51) wide, greatly elongate,
sometimes oval, tapered to posterior end (fig. 17). Testes numerous, 103—157 (127 ± 2;
n = 31; n = 51) in number (fig. 10, 12, 17), median, arranged in two layers, round, 20—80
(46 ± 2.3; n = 31; n = 44) in diameter; post-poral testes absent. Cirrus-sac big, oval,
50—280 (148 ± 8.5; n = 31; n = 46) long, 50—450 (169 ± 13; n = 31; n = 40) wide (fig.
11, 15, 16). Cirrus armed with spinitriches. Total length of everted cirrus 850—1350 (1052 ±
92; n = 5), of inverted 260—1030 (427 ± 20; n = 30; n = 53). Vas deferens 10—180 (63 ±
5; n = 31; n = 43) in diameter, coiled, extending anteriorly to cirrus-sac; median, slight-
ly displaced to poral side in mature proglottids; seminal vesicle absent. Genital pores lat-
eral, 20—50 % (30 ± 1.2; n = 31; n = 47) of proglottid length from posterior end, irreg-
ularly alternating. Vagina extending from Mehlis’ gland to genital atrium, initially medi-
an, straightened, then lateral to cirrus-sac in mature proglottids, opening anterior to
cirrus-sac in genital atrium; wall of distal end of vagina dense, glandular, with inside folds
almost closing vaginal duct; vaginal sphincter absent. Genital atrium 40—210 (98 ± 9;
n = 31; n = 33) long, 30—180 (71 ± 5; n = 31; n = 35) wide; its proximal part near to
opening of cirrus-sac covered by numerous spinitriches (fig. 11, 15, 16). Ovary near pos-
terior end of proglottids, median, H-shaped, tetralobed in cross-section; lobes asymmet-
rical. In mature proglottids, ovary lobes surrounding excretory ducts, extending almost
to lateral margin of proglottid, separating vitellarium from posterior end of proglottid. Aporal
ovary lobe 100—640 (197 ± 20; n = 31; n = 38) long, 110—500 (262 ± 13; n = 31; n =
38) wide, poral ovary lobe 100—550 (166 ± 16; n = 31; n = 38) long, 110—440 (248 ±
13; n = 31; n = 38) wide. Ovicapt 50—200 (94 ± 5; n = 31; n = 34) in diameter. Mehlis’
gland cariniform, fabiform or spherical, 10—60 (28 ± 2; n = 29) wide. Uterus ventral to
vagina, extending from isthmus to median line of proglottid, reaching its anterior end (figs
12, 17). Vitellarium follicular; follicles numerous, 10—90 (43 ± 3; n = 31; n = 38) in
diameter in mature proglottids; in 2 lateral fields, 90—400 (173 ± 11; n = 31; n = 37)
wide, each consisting of one dorsal and one ventral row of follicles, between which excre-
tory ducts passes; extending from anterior end of proglottid to anterior margin of ovary,
not interrupted at level of genital atrium, becoming thin, not extending posteriorly to ovary.
Eggs oval, 40 long, 20 wide (n = 1).
Remark s. In the living specimen of C. healyae bothridia are very mobile and resem-
ble the movement of the octopus feelers. The specimens of C. healyae are very delicate
in comparison with the scolex and the strobila of C. ruhnkei. Spinitriches were found in
e-9Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
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the genital atrium of C. healyae near opening of cirrus-sac, but their shape was not good
examined in the present work. However, size of these spinitriches exceeds the size of the
spinitriches of the cirrus. In immature proglottids the spinitriches of the genital atrium
are located in its central part on equal distance from the cirrus-sac and the vagina
(fig. 15, Â) looking as «muff».
e-10 V. V. Kornyushin, T. A. Polyakova
Fig. 13—17. Photomicrographs of Cairaeanthus healyae ex Dasyatis pastinaca from the Black Sea: 13 – scolex,
paratype Ñ. 26.001.27; 14 – marginal locules, paratype Ñ. 26.001.27; 15 – inverted cirrus, holotype
Ñ. 137.001.163; 16 – everted cirrus, holotype Ñ. 137.001.163; 17 – mature proglottid, holotype Ñ. 137.001.163.
Scale bars: 13 – 300 μm; 14, 15 – 50 μm; 16, 17 – 200 μm; A – vagina; B – spinitriches of genital atri-
um; C – cirrus; D – cirrus-sac.
Ðèñ. 13—17. Ìèêðîôîòîãðàôèÿ Cairaeanthus healyae îò Dasyatis pastinaca èç ×åðíîãî ìîðÿ: 13 – ñêî-
ëåêñ, ïàðàòèï Ñ. 26.001.27; 14 – êðàåâûå ëîêóëû, ïàðàòèï Ñ. 26.001.27; 15 – èíâàãèíèðîâàííûé öèð-
ðóñ, ãîëîòèï Ñ. 137.001.163; 16 – ýâàãèíèðîâàííûé öèððóñ, ãîëîòèï Ñ. 137.001.163; 17 – ïîëîâîçðå-
ëàÿ ïðîãëîòòèäà, ãîëîòèï Ñ. 137.001.163. Ìàñøòàáíàÿ ëèíåéêà: 13 – 300 ìêì; 14, 15 – 50 ìêì; 16, 17 –
200 ìêì; A – âàãèíà; B – ñïèíèòðèõèè ïîëîâîãî àòðèóìà; C – öèððóñ; D – áóðñà öèððóñà.
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Di f f e r e n t i a l d i a g no s i s: Cairaeanthus healyae sp. n. significantly differs from
C. ruhnkei sp. n. in longer strobila (108—315 mm vs 56—255 mm), smaller sizes of marginal
loculi (10—30�10—30 vs 50—100�30—100), larger length of cephalic peduncle (2.2—11.7
mm vs 200—680), smaller diameter of testes (20—80 vs 30—130) and smaller width of
vitelline fields (90—400 vs 100—720), number of testes (103—157 vs 51—98) and proglot-
tids (1003—2395 vs 495—1803). Additionally, the wall of the genital atrium is armed by
the spinitriches in C. healyae, but is not in C. ruhnkei.
In addition to characters mentioned in generic diagnosis, C. healyae can be distin-
guished from all known Anthocephalum spp. by larger total length (108—315 mm vs
40—50 mm in A. gracile, 11—20 mm in A. centrurum, 4—9 mm in A. alicae, 8—14 mm in
A. cairae, 18—31 mm in A. duszynskii, 8.6—13 mm in A. kingae, 5.7—16.3 mm in A. michaeli,
7.9—17.2 mm in A. lukei, 6.6—14.4 mm in A. currani), more proglottids per worm
(1003—2395 vs 500—600, 35—65, 9—15, 80—110, 120—160, 33—50, 23—41, 28—56,
35—70, respectively) and by larger number of the testes (103—157 vs 100—130, 35—75,
30—67, 28—52, 35—71, 30—37, 30—49, 32—48, 37—50, respectively).
Morphometric analysis of Cairaåanthus ruhnkei sp. n. and C. healyae sp. n.
The choice of diagnostic characters for species determination is one of the princi-
pal questions in generic taxonomy. In order to deicide which characters and which dif-
ferences are adequate for the species differentiation, it is necessary to have information
on the range of intraspecific variability. For this purpose 38 characters (36 measured and
2 discrete) from 67 specimen of Cairaeanthus spp. were analysed using t-tests for inde-
pendent variables (table 1) and Principal Component Analysis (ÐÑÀ).
e-11Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
Fig. 18. ÐÑÀ plot of the contributions made by 24 characters to the first two factors calculated from 66 spec-
imens of two species of Cairaeanthus spð. ex. Dasyatis pastinaca from the Black Sea and the Sea of Azov. Separated
are 14 characters making the greatest contribution to separation of this species: 1 – number of proglottids; 2 –
length of cephalic peduncle; 3 – number of testes; 4 – length of mature proglottids; 5 – length of pedicels
bothridia; 6 – length of strobila; 7 – width of scolex; 8 – length of bothridia; 9 – length of cirrus-sac; 10 –
length of ovary; 11 – width of mature proglottids; 12 – width of strobila; 13 – length of marginal loculi; 14 –
width of marginal loculi.
Ðèñ. 18. ÐÑÀ ãðàôèê ôàêòîðíûõ íàãðóçîê 26 ïðèçíàêîâ, èñïîëüçîâàííûõ äëÿ äèôôåðåíöèàöèè 66 ýêçåì-
ïëÿðîâ Cairaeanthus spð. îò Dasyatis pastinaca èç ׸ðíîãî è Àçîâñêîãî ìîðåé. Âûäåëåíû 14 ïðèçíàêîâ,
äåëàþùèõ íàèáîëüøèé âêëàä â ðàçäåëåíèå âèäîâ: 1 – êîëè÷åñòâî ïðîãëîòòèä; 2 – äëèíà ãîëîâíîãî
ñòåáåëüêà; 3 – êîëè÷åñòâî ñåìåííèêîâ; 4 – äëèíà ïîëîâîçðåëûõ ïðîãëîòòèä; 5 – äëèíà ñòåáåëüêà áîò-
ðèäèÿ; 6 – äëèíà ñòðîáèëû; 7 – øèðèíà ñêîëåêñà; 8 – äëèíà áîòðèäèé; 9 – äëèíà áóðñû öèððóñà;
10 – äëèíà ÿè÷íèêà; 11 – øèðèíà ïîëîâîçðåëûõ ïðîãëîòòèä; 12 – øèðèíà ñòðîáèëû; 13 – äëèíà êðàå-
âûõ ëîêóë; 14 – øèðèíà êðàåâûõ ëîêóë.
-1,0 -0,5 0,0 0,5 1,0
Factor 1 : 35,5 %
-1,0
-0,5
0,0
0,5
1,0
F
ac
to
r
2
:
15
,5
%
1
7
13
9
11
12
10
8
14
56
4
3
2
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C. ruhnkei significantly differs from C. healyi in 26 of the 38 investigated charac-
ters. These different characters describe size of strobila, marginal loculi, cephalic pedun-
cle, immature and mature proglottids and ovary, as well as width of scolex, terminal proglot-
tids, genital atrium, vitelline fields, Mehlis’ gland, length of bothridia and pedicel, cir-
rus-sac, diameter of excretory ducts and testes, number of testes and proglottids (table 1).
These 26 characters were also analysed by PCA, they were reduced to two principal com-
ponents (i. e. Factors) which explained more than 50 % of the overall variance, and
e-12 V. V. Kornyushin, T. A. Polyakova
Ta b l e 1. Morphological characters of Cairaåanthus ruhnkei sp. n. and C. healyae sp. n. ex Dasyatis pastinaca
from the Black Sea and the Sea of Azov
Ò à á ëèö à. 1 Ìîðôîëîãè÷åñêèå ïðèçíàêè Cairaåanthus ruhnkei sp. n. è C. healyae sp. n. îò Dasyatis pasti-
naca èç ×åðíîãî è Àçîâñêîãî ìîðåé
* X – mean, μm; ± SE – standard error, μm; T-test – independent t-test.
** T-test: significant differences at the p ≤ 0,05
Strobila Length, mm 123.0 ± 8.3 182.2 ± 9.6 4.7**
Width 960 ± 29 677 ± 31 —6.7
Scolex Length 1447 ± 72 1642 ± 83 1.8
Width 1789 ± 70 2096 ± 75 3.0
Bothridia Length 839 ± 47 986 ± 42 2,3
Width 639 ± 34 622 ± 24 —0.4
Pedicels bothridia Length 273 ± 17 357 ± 19 3.3
Width 251 ± 20 247 ± 15 0.1
Marginal loculi Length 57 ± 2 25 ± 1.1 —11.6
Width 52 ± 2 24 ± 1.1 —12.2
Cephalic peduncle Length 460 ± 22 8117 ± 354 23.7
Width 558 ± 26 459 ± 21 —2.9
Excretory ducts Diameter 35 ± 2 28 ± 2 —2.3
Immature proglottids Length 121 ± 8 165 ± 9,4 3.5
Width 703 ± 35 557 ± 23 —3.6
Mature proglottids Length 331 ± 19 447 ± 26 3.7
Width 1198 ± 44 867 ± 45 —5.2
Terminal proglottids Length 1071 ± 95 1105 ± 81 0.3
Width 1194 ± 50 796 ± 52 —5.5
Position of genital pores, % 32 ± 2 31 ± 1.2 —0.2
Number of testes 86 ± 2 126 ± 2.1 13.03
Testes Diameter 62 ± 4 47 ± 3 —2.9
Vas deferens Diameter 61 ± 5 60 ± 5 —0.13
Cirrus-sac Length 204 ± 13 159 ± 10 —2.8
Width 156 ± 11 176 ± 17 1.03
Cirrus Length 526 ± 27 549 ± 48 —1.3
Genital atrium Length 109 ± 9 93 ± 9 —1.2
Width 91 ± 7 68 ± 5 —2.6
Ovary Length 255 ± 28 138 ± 16 —2.8
Width 343 ± 22 237 ± 14 —4.3
Ovicapt Diameter 86 ± 6 92 ± 5 0.8
Mehlis’ gland Width 61 ± 8 28 ± 2 —3.7
Vitellaruim filds Width 241 ± 17 164 ± 10 —3.7
Vitelline follicles Diameter 43 ± 4 42 ± 3 0.1
Eggs Length 34 40 34.2
Width 20 20 20.0
Number of proglottids 981 ± 48 1447 ± 69 5.7
Characters
Black Sea and Sea of Azov Black Sea
T-testX ± SE*
Cairaåanthus ruhnkei sp. n.
(n = 36)
Cairaåanthus healyae sp. n.
(n = 30)
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e-13Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
Fig. 19. ÐÑÀ plot of the scores of the first two factors calculated from 14 characters for 67 specimens belong-
ing to two species of Cairaeanthus spð. ex. Dasyatis pastinaca from the Black Sea and the Sea of Azov (
direction of increasing characters separating the specimens).
Ðèñ. 19. ÐÑÀ ãðàôèê ðàñïîëîæåíèÿ 67 ýêçåìïëÿðîâ Cairaeanthus spð. îò Dasyatis pastinaca èç ×åðíîãî
è Àçîâñêîãî ìîðåé â ñèñòåìå êîîðäèíàò ïåðâûõ äâóõ ôàêòîðîâ, âû÷èñëåííûõ íà îñíîâå 14 ìîðôîëî-
ãè÷åñêèõ ïðèçíàêîâ ( íàïðàâëåíèå óâåëè÷åíèÿ çíà÷åíèÿ ïðèçíàêîâ).
Cairaeanthus healyae Cairaeanthus ruhnkei
-5 -4 -3 -2 -1 0 1 2 3 4 5
Factor 1: 44 %
-3
-2
-1
0
1
2
3
F
ac
to
r
2:
1
6
%
Number of proglottids and
testes, length of cephalic
peduncle
Length and width of marginal
loculi, width of strobile
and mature proglottids
14 characters, which made the greatest contributions for this Factors were chosen
(fig. 18). These were the total number of proglottids and testes, the length of cephalic
peduncle, mature proglottids, bothridium pedicel, strobila, bothridia, cirrus sac, ovary and
marginal loculi, width of scolex, mature proglottids, strobila and marginal loculi.
As a result of PCA done on metrical data for 14 selected characters, 67 measured
specimens were clearly divided on the PCA plots based on their scores in the first plane
of the PCA (explaining 60 % of the overall variance) into two groups corresponding to
each species (fig. 19). The number of proglottids and testes and the length of cephalic
peduncle contributed most to separation of C. ruhnkei sp. n. from C. healyae sp. n. by
Factor 1, explaining 44 % of the overall variance (fig. 19). The size of marginal loculi,
the width of strobila and mature proglottids made the greatest contributions for Factor
2, however, explained only 16 % of the overall variance. Thus, using the methods of the
multivariate statistics permits successful differentiation of morphologically related con-
generic cestodes.
Discussion
New species, C. healyae and C. ruhnkei, resemble Phyllobothrium pastinacae Mokhtar-
Mocamouri, Zamali, 1981 (Mokhtar-Mocamouri, Zamali, 1981) in the following char-
acters: bothridia posteriorly bifid; myzorhynchus absent; bothridia pedicellate, cephalic
peduncle present; post-poral testes absent; genital pore opening marginally, close to pos-
terior end of proglottid; vitelline follicles numerous, extending from anterior end of proglot-
tid to ovary, interrupted at level of ovary, not extending posteriorly to it. Studied mate-
rial on Phyllobothrium pastinacae (MNHN 90HB148 c VII; MNHN 90HB149 c VII;
MNHN 90HB150 c VII) included two total specimens and one detached terminal
proglottid. Since the authors of P. pastinacae did not specify the holotype specimen, we
supposed that the specimen from the slide MNHN 90HB148 c VII, which was repre-
sented on fig. 1 in Mokhtar-Mocamouri, Zamali (1981: p. 377) as the holotype. Cestodes
from the both slides differed from Cairaeanthus ruhnkei sp. n. and C. healyae sp. n. by
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the presence of the apical sucker and by the facial loculi in the specimen from MNHN
90HB148 c VII.
F. Mokhtar-Mocamouri and Z. Zamali assigned P. pastinacae to the genus
Phyllobothrium based on the shape of the scolex, presence of 4 pedicel bothridia, which
are posteriorly bifid and have the marginal loculi, of long cephalic peduncle and absence
of the post-poral testes in the proglottids (Mokhtar-Mocamouri, Zamali, 1981: p. 379).
However, mentioned morphological characters, except for posteriorly bifid bothridia, do
not correspond to the present diagnosis of the genus Phyllobothrium (Ruhnke, 1996 a).
e-14 V. V. Kornyushin, T. A. Polyakova
Fig. 20—23. Photomicrographs of Phyllobothrium pastinacae (MNHN 90HB148 c VII): 20 – scolex; 21 – facial
locules; 22 – inverted cirrus; 23 – mature proglottid. Scale bars: 20 – 500 μm; 21 – 100 μm; 22, 23 – 200 μm.
Ðèñ. 20—23. Ìèêðîôîòîãðàôèÿ Phyllobothrium pastinacae (MNHN 90HB148 c VII): 20 – ñêîëåêñ; 21 –
ëèöåâûå ëîêóëû; 22 – èíâàãèíèðîâàííûé öèððóñ; 23 – ïîëîâîçðåëàÿ ïðîãëîòòèäà (æåëòî÷íûå ôîë-
ëèêóëû íà ïîðàëüíîé ñòîðîíå ïðîãëîòòèäû áûëè îïóùåíû). Ìàñøòàáíàÿ ëèíåéêà: 20 – 500 ìêì; 21 –
100 ìêì; 22, 23 – 200 ìêì.
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New morphological characters were found during the reinvestigation of the type mate-
rial of P. pastinacae. On the bothridia of the holotype specimen numerous facial loculi
are present, and the paratype specimen has marginal loculi; in all type specimens of P. pasti-
nacae the vaginal sphincter is absent and the vitelline fields stop at the anterior margin
of the ovary, not reaching the posterior end of the proglottid. These facts confirm that
P. pastinacae is not a representative of Phyllobothrium.
Moreover, the presence of facial loculi on bothridia in one of the two examined type
specimens of P. pastinacae distinguishes it from the following phyllobothriideans:
e-15Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
Fig. 24—28. Photomicrographs of Phyllobothrium pastinacae (24, 25, 26, 27 – MNHN 90HB149 c VII, 28 –
MNHN 90HB150 c VII): 24 – scolex; 25 – marginal loculi; 26 – apical sucker; 27 – mature proglottid;
28 – gravid proglottid. Scale bars: 24 – 400 μm; 25, 26, 27 – 200 μm; 28 – 100 μm.
Ðèñ. 24—28. Ìèêðîôîòîãðàôèÿ Phyllobothrium pastinacae (24, 25, 26, 27 – MNHN 90HB149 c VII, 28 –
MNHN 90HB150 c VII): 24 – ñêîëåêñ; 25 – êðàåâûå ëîêóëû; 26 – àïèêàëüíûå ïðèñîñêè; 27 – ïîëî-
âîçðåëàÿ ïðîãëîòòèäà; 28 – çðåëàÿ ïðîãëîòòèäà. Ìàñøòàáíàÿ ëèíåéêà: 24 – 400 ìêì; 25, 26, 27 –
200 ìêì; 28 – 100 ìêì.
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Anthobothrium van Beneden, 1850; Monorygma Diesing, 1863; Orygmatobothrium Diesing,
1863; Dinobothrium van Beneden, 1889; Thysanocephalum Linton, 1889; Crossobothrium
Linton, 1889; Calyptrobothrium Monticelli, 1893; Carpobothrium Shipley, Hornell, 1906;
Myzocephalus Shipley, Hornell, 1906; Myzophyllobothrium Shipley, Honell, 1906;
Pithophorus Southwell, 1925; Scyphophyllidium Woodland, 1927; Gastrolecithus Yamaguti,
1952; Marsupiobothrium Yamaguti, 1952; Pseudanthobothrium Baer, 1956; Clydonobothrium
Euzet, 1959; Clistobothrium Dailey, Vogelbein, 1990; Paraorygmatobothrium Ruhnke, 1994;
Orectolobicestus Ruhnke, Caira, Carpenter, 2006 (Yamaguti, 1959; Euzet, 1994; Ruhnke,
1994, 1996 à, b; Ruhnkeetal., 2006; Marquesetal., 2001; Randhawaetal., 2008). The pres-
ence of the apical sucker on the scolex of the both type specimens of P. pastinacae dis-
tinguishes them from Trilocularia Olsson, 1867; Tritaphros Lonnberg, 1889; Pentaloculum
Alexander, 1963; Phormobothrium Alexander, 1963; Zyxibothrium Hayden, Campbell, 1981
and the absence of post-poral testes – from Caulobothrium Baer, 1948; Glyphobothrium
Williams, Campbell, 1977; Duplicibothium Williams, Campbell, 1978; Chimaerocestos
Williams, Bray, 1984 and Anindobothrium Marques, Brook, Lasso, 2001 (Euzet, 1994).
Finally, type specimens of P. pastinacae clearly differ in the scolex shape, absence
of armaments of the scolex and post-poral testes, and also in the position of the genital
pore near the posterior end of the proglottid from the following taxa of the order
Tetraphyllidea: Onchobothriidae Braun, 1900; Disculicipitidae Joyeux, Baer, 1936;
Prosobothriidae Baer, Euzet, 1955; Litobothriidae Dailey, 1969; Dioecotaeniidae Schmidt,
1969; Cathetocephalidae Dailey, Overstreet, 1973; Chimaerocestidae Williams, Bray, 1984
(Euzet, 1994).
Thus, the cestodes in the type material of P. pastinacae belong neither to the genus
Phyllobothrium nor to the order Tetraphyllidea.
The two whole type specimens of P. pastinacae should be assigned to the order
Rhinebothiidea (Healy et al., 2009) based on the presence of the apical sucker on the
pedicel bothridia, of the facial loculi in one specimen and the marginal loculi in the other,
by the presence of cephalic peduncle and also by the absence of post-poral testes and by
the distal position of the genital pore near the posterior end of proglottid.
Moreover, the two specimens from the slides examined differ from each other by
some substantial qualitative characters (e. g., presence of the marginal, or the facial loculi)
(fig. 20—28) and a number of metrical characters, and apparently belong to two sepa-
rate species of Rhinebothriidea.
Reinvestigation of the type material of Phyllobothrium pastinacae, which has some
similarity with Cairaeanthus spp., showed that these specimens do not belong to
Phyllobothrium and the order Tetraphyllidea, but are the members of the order
Rhinebothriidea. However, the quality of available slides was found to be insufficient for
the finally decision on the taxonomic belonging of these cestodes. Additional material from
the same sea and host is required for more detailed description of these species and iden-
tification of their generic status. It might be even supposed that these species and
Cairaeanthus spp. may be members of different genera from one family, but for confir-
mation of this assumption further study is necessary.
During the recent years, the interest of helminthologists to the lowest cestodes par-
asitizing elasmobranches, namely tetraphyllideans, lecanicephalideans, trypanorhyncha etc.,
has taken a remarkable start. At present, a tendency to the restriction of orders, families
and genera with the separation of new or the restoration of “old” taxa presents in the
works (Ruhnke, 1994 b, 1996 à—b; Ruhnke, Seaman, 2009; Ruhnke et al., 2006;
Ruhnke, Carpenter, 2008). The results of the molecular phylogenetic analysis of rhinoboth-
riines confirmed previous hypotheses on their independence from Tetraphyllidea and proved
monophyly of the order Rhinebothriidea (Healy et al., 2009). Eight valid genera, previ-
ously the members of Tetraphyllidea, were included in the restored order Rhinebothriidea
in the new combination. In connection with this revision the taxonomic status of most
cestode species from elasmobranches in the Black Sea and the Sea of Azov needs to be
defined more precisely.
e-16 V. V. Kornyushin, T. A. Polyakova
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Before the present study, cestodes from stingrays D. pastinaca and R. clavata from
the Black Sea and the Sea of Azov have been identified as Phyllobothrium lactuca and
P. gracilis (Borcea, 1934; Pogorelceva, 1960; Gaevskaya et al., 1975). We showed that
those were representatives of the order Rhinebothriidea and belonged to a new genus
Cairaeanthus. The order Rhinebothriidea thus comprises nine genera now.It should be
noted that in the previous articles (Borcea, 1934; Pogorelceva, 1960; Gaevskaya et al., 1975)
Phyllobothrium lactuca and P. gracilis in the Black Sea and the Sea of Azov were reported
from both stingray species occurring in these seas. However, we found both species of
Cairaeanthus only in D. pastinaca, although cestodes of R. clavata were examined too. The
latter fish species was parasitized by four species of cestodes: Grillotia erinaceus (van
Beneden, 1858), Acanthobothrium dujardini (van Beneden, 1850), Echeneibothrium variabile
(van Beneden, 1850), and Echinobothrium typus (van Beneden, 1849).
Thus, D. pastinaca in the Black Sea off Crimea is parasitized by C. ruhnkei and
C. healyae, and in the Sea of Azov – only by C. ruhnkei. These species have been found
only in these seas and only in this host so far. Moreover, in the Black Sea the genus
Phyllobothrium s. str., is currently represented by one species, Phyllobothrium squali
(Yamaguti, 1952) parasitizing Squalus acanthias L., 1758 (Vasileva et al., 2002).
The authors are grateful to Dr Janine N. Caira (Department of Ecology and Evolutionary Biology, University
of Connecticut, USA) and Dr Timothy R. Ruhnke (Department of Biology, West Virginia State University,
Institute, USA) for reading the manuscript and making numerous useful suggestions. The authors thank Dr
E. V. Dmitrieva (Institute of Biology of the Southern Seas, Sevastopol, Ukraine) for critical comments on the
final version of this manuscript and help in statistical data processing. Authors bring their gratitude to Dr Jimmy
Cassone, curator of the parasitological collection of the Museum Nationale d’Histoire Naturelle (Paris,
France) for loaning the type material of Phyllobothrium pastinacae and to Y. I. Kuzmin (Institute of Zoology,
Kiev, Ukraine) for delivery of this material; Dr V. A. Kharchenko (Institute of Zoology, Kiev, Ukraine) for
possibility to work on the microscope Zeiss Axio Imager M 1. The authors also thank N. A. Skidan
(Melitopol’skii Institute of Ecology and Social Technology, Melitopol, Ukraine) for tapeworm collection from
stingrays in the Sea of Azov and anonymous referees for their help and suggestions.
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type Scalithrium minimum (Van Beneden, 1850) n. comb. parasite de Dasyatis pastinaca (Elasmobranchii,
Dasyatidae) // Parasite. – 2003. – 10. – P. 31—37.
Borcea L. Note preliminaire sur les cestodes des Elasmobranches ou Sélaciens de la Mer Noire // Annls sci-
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Îäåññêîãî çàëèâà.
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Gaevskaya A. V., Gusev A. V., Delyamure S. L. et al. Key to the parasites of vertebrates of the Black Sea and
the Sea of Azov / Parasitic invertebrates of fishes, fish-eating birds and marine animals. – Kyiv : Nauk.
Dumka. – 1975. – P. 296—364. – Russian : Ãàåâñêàÿ À. Â., Ãóñåâ À. Â., Äåëÿìóðå Ñ. Ë. è äð.
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e-17Cairaeanthus gen. n. (Cestoda, Rhinebothriidea), with the Description of Two New Species...
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