9
Instructions for use Title 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫吸管虫亜綱について Author(s) 山口, 篤 Citation 日本プランクトン学会報, 53(1), 29-36 Issue Date 2006-02-25 Doc URL http://hdl.handle.net/2115/52374 Rights © 2006 日本プランクトン学会 Type article File Information yamaguchi_2006.pdf Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP

北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

  • Upload
    others

  • View
    0

  • Download
    0

Embed Size (px)

Citation preview

Page 1: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

Instructions for use

Title 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫吸管虫亜綱について

Author(s) 山口, 篤

Citation 日本プランクトン学会報, 53(1), 29-36

Issue Date 2006-02-25

Doc URL http://hdl.handle.net/2115/52374

Rights © 2006 日本プランクトン学会

Type article

File Information yamaguchi_2006.pdf

Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP

Page 2: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

infection. J. Eukaryot. Microbiol. 51: 145�155.Saldarriaga, J. F., F. J. R. Taylor, P. J. Keeling & T. Cavalier-

Smith 2001. Dinoflagellate nuclear SSU rRNA phylogenysuggests multiple plastid losses and replacements. J. Mol.Evol. 53: 204�213.

Saldarriaga, J. F., F. J. R. M. Taylor & T. Cavalier-Smith2004. Molecular data and the evolutionary history ofdinoflagellates. Eur. J. Protistol. 40: 85�111.

Saunders, G. W., D. R. A. Hill, J. P. Sexton & R. A. Andersen1997. Small-subunit ribosomal RNA sequences from se-lected dinoflagellates: testing classical evolutionary hy-potheses with molecularsystematic methods. Pl. Syst.Evol. 11 (Suppl): 237�259.

Sewell, R. B. S. 1951. The epibionts and parasites of theplanktonic copepoda of the Arabian Sea. Johan MurrayExpedition. Sci. Rep. Br. Mus. Nat. Hist. 9: 255�394.

Shields, J. D. 1994. The parasitic dinoflagellates of marinecrustaceans. Ann. Rev. Fish Dis. 4: 241�271.

Skovgaard, A. 2005. Infection with the dinoflagellate para-site Blastodinium spp. in two Mediterranean copepods.Aquat. Microb. Ecol. 38: 93�101.

Skovgaard, A., R. Massana, V. Balague, & E. Saiz 2005.Phylogenetic position of the copepod-infesting parasiteSyndinium turbo (Dinoflagellata, Syndinea). Protist 156:

413�423Stentiford, G. D. & J. D. Shields 2005. A review of the

parasitic dinoflagellates Hematodinium-like infection inmarine crustaceans. Dis. Aquat. Org. 66: 47�70.

Takano,Y. & T. Horiguchi 2004. Surface ultrastructure andmolecular phylogenetics of fourunarmored heterotrophicdinoflagellates, including the type species of the genusGymnodinium. Phycol. Res. 52: 107�116.

Taylor, F. J. R. 1968. Parasitism of the toxin-producingdinoflagellate Gonyaulax catenella by the endoparasiticdinoflagellate Amoebophrya ceratii. J. Fis. Res. Bd. Canada25: 2241�2245.

Taylor, F. J. R. (ed.). 1987. The Biology of Dinoflagellates.Blackwell Sci. Publ., Oxford, 785 pp.

Yamaguchi, A. & T. Horiguchi 2005. Molecular phyloge-netic study of the heterotrophic dinoflagellate genus Pro-toperidinium (Dinophyceae) inferred from small subunitrRNA gene sequences Phycol. Res. 53: 30�42.

Zhang, Z., B. R. Green & T. Cavalier-Smith 1999. Single genecircles in dinoflagellate chloroplast genomes. Nature 400:155�159.

2005� 12� 2���� 2006� 1� 13���

���������� ��������������������� !"

�� ������������ �041�8611 ����� 3�1�1

Suctorian ciliate epibionts on calanoid copepods in the subarctic Pacific

AIHJH=> Y6B6<J8=>

Graduate School of Fisheries Sciences, Hokkaido University, 3�1�1 Minato-cho, Hakodate, Hokkaido 041�8611,

Japan

E-mail: [email protected]

Abstract From zooplankton samples collected from subarctic Pacific and adjacent seas (JapanSea and Okhotsk Sea), 10 calanoid copepod species including Gaetanus armiger, G. simplex, G.variabilis, Heterorhabdus tanneri, Heterostylites major, Metridia pacifica, Paraeuchaeta birostrata, P.elongata, P. rubra and Scaphocalanus magnus were found to host epibiont suctorian ciliates.Suctorian ciliates were observed on 5 developmental stages (C4M, C5F/M, C6F/M) among thecopepodid stages. Seasonal observations on the prevalence of suctorians on M. pacifica in theOyashio region western subarctic Pacific showed high incidence in summer to autumn whenambient temperatures were high. Within the subarctic Pacific, the prevalence of suctorians on M.pacifica was found to be higher at the high latitude stations. Suctorian ciliates on M. pacifica wereclassified into two types; type A characterized by short stalk and spherical cell (diameter: 50�100mm), and type B, by enlarged stalk and petaloid-form tentacles (cell diameter: 300�500 mm). With

� �

��� !�"#$%&'()'�*+,-./01. 29

Page 3: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

regard to the substrate of suctorians on M. pacifica, they were observed most frequently on theurosome (89�), followed by the metasome (60�), cephalosome (19�) and swimming legs (5�),while they were never observed on the antennules, feeding appendages or caudal rami.

Key words: parasite, host, Suctoria, calanoid copepods, Metridia pacifica

� � � ������������������ � (Pe-

ritrichia)� � � (Chonotrichia)������ �(Suctoria)����������� (Fernan-

dez-Leborans & Tato-Porto 2000 a, b, Fernandez-

Leborans 2001)� � !"������#��� ����$%&' 2 7(�)#� 11) 49$��*�� (Fernandez-Leborans & Tato-Porto 2000a)��� ��+,�-�./�01234 56781914 � !"�:� �;<4 � !"�:�=>?� Acartia, Boeckella, Centropages, Diaptomus���Pleuromamma )���@����A:B�(Fernandez-Leborans & Tato-Porto 2000a)� CD� � !"������#�����EF� �� 2 3

(�)#� 4) 8$��*���A:� -�+,�#G6781914 � !"�:B� (Fernandez-

Leborans 2001)�H���I���3;J1KJL MN4��O#��;<4 � !"�����������-�./���� ��)P�Q� � !"���������� ��RSC:���$%&' 2 10(�)#�21) 39$��*�� (Fernandez-Leborans &

Tato-Porto 2000b)� �;<4 � !"����?*� 2 4(�)#� 5) 11$��@���Q�Ephelota)� Calanus finmarchicus, Candacia armata,

Pleuromamma borealis��� P. gracilis���?*�Paracineta)� Gaetanus antarcticus, G. curvicornis,

Pleuromamma abdominalis, P. xiphias, Metridia lu-

cens���M. longa ���?*� Pelagacineta)�Calanus helgolandicus, Euchaeta spp., Metridia spp.����Q� Pseudocorynophrya)� Rhincalanus spp.

����Q� Brachyosoma )� Scottocalanus daug-

lishi���?*-�T��@��� UFernandez-

Leborans & Tato-Porto 2000b%VWX������� ��Y�������/� .Z�[\

(tentacle)%]^�� [\�_`:a��b�/c`�d/��Q� e�[\%f������ghP���%�i��jkl%P� (Kitching 1952,

Lilly 1953)� ����������mS�+,�i^

�n�opq��irs*��Q� �����A�t�B���uk (commensalism):B�irs*�� UNicol 1984%VWX� v^iv� �������BCQ�v./��w� +,:B�� !"���xy���z��� (Henerby & Ridgeway 1979)� CD�������{��I�� !"����|}�Y~���%vD*#eiv�@��� (Turner et al.

1979)�����;<4 � !"������������#���� !��� "!�� #$H U!��!HX ���!;�!H��I�v�:BQ� �%���;<4 � !"�����i������� &��:� �%�'� �(�� )&H���M���1H�=D��*�H����RPD�;<4 � !"���������� U��� �X ���@#�� CD� ����;<4 � !"�Metridia paci-

fica���PD�������� �%� �(��+���I���,�� ���J�H?* �%�41�NC:�#-./� CD-����{����%y^D�

� ����� ���� ����

19960 9�?* 19970 10��?I �%�'� �(�i-��1H U)&H���M���1HX� (Fig. 1)�  ¡ 2000 mC:% ¢ 0.1 mm�2£¤¥¦K%3 5��§�¨.©ªP4D«5 U5�$��7N�J¬�X %3� ���E��;<4 � !"���������� U��� �X��R#��?%­®PD U©ª)¯���©ª��°Y� Kobari & Ikeda 2001%VWX� e�*«5$��RPD�;<4 � !"�������67%9±¦1P� $���8²9³´i�-���:(prevalence: �)%µ¶D�

Metridia pacifica�������������;<4 � !"�� Metridia pacifica��PwPw·�����:��*�D� -���:���,��B��?%;*?�#�D¶� 20020 5�

¸ ¸

)&3;J1KJ¹º� < 53» < 1¼ (2006)30

Page 4: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

�� 2004� 12�������� �������� Site H (41�30�N, 145�47�E, Fig. 1)����� 0.1

mm� NORPAC������ 500 m !"#$%&�'()* +5,-./0123456 �7��8 )*-�9:&(M. pacifica;<=�>��?�9:@=A�BA��CCD�EFG�HIJ�KLMN�OP&(Q RSTUVW 1X2Y�Z[�Q

Metridia pacifica���������2004� 6��� 8�������� \]^�

(165�E�_`( 41�NX47�N)8 a�� \]^�(165�W�_`( 41�NX50�N)b cde23fg(53X30�N, 177X00�W)��hT���� 0.33 mmi(V 0.10 mm� NORPAC������ 50 m !gjkiZ"#$%&�'()* +5,-./0123456�7��8 )*-�9:&(Metridia pacifica;<=�>��?�9:@=A�BA��CCD�EFG�HIJ�lmMN�OP&(Q

Metridia pacifica������� ����nolmpq�OP-ZrDs�HIJ�t�u(de23fg�)*- !8 EFG�HI�t�u�Metridia pacifica;<= 56@=�>��8 ?�HI��OP&(Q v=��� 7Yw +x [cephalo-

some]8 y [metasome]8 z [urosome]8 {|}~[antennule]8 {|}~�����H�� [feeding ap-

pendage except antennule]8 ��� [swimming legs]

b cz� [caudal rami]6 �p��8���EFGHI����OP8 BA&(Q EFG����V������D�C������������(D�� 2��

�����u(�Z (Fig. 2)8 BAV�����C��`(Q

Fig. 1. Location of sampling stations in the subarctic Pacific and adjacent seas. Stars indicate stations where thevertical stratified samplings were done down to �2000 m for details of sampling data, see Table 1 of Kobari & Ikeda(2001). At Site H located in the NW Pacific, seasonal changes of epizoic suctorian ciliates on Metridia pacifica (Fig. 2)were observed during 2002�2004 (Fig. 3). Open circles indicate stations where the spatial changes in prevalence ofepizoic suctorian ciliates on M. pacifica were studied during June�August 2004 (Table 2).

Fig. 2. Two forms of epizoic suctorian ciliates onMetridia pacifica. Type A: small forms being attachedon the urosome (A). Type B: Large petaloid-formsbeing attached on the urosome (B). Small forms atenlarged magnification (C). Large petaloid-forms atenlarged magnification (D). Scale bars: 500 mm (A, B);100 mm (C, D).

� �

��3��3�b��� ¡¢ �?�£¤.C¥¦. 31

Page 5: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

� ��������� ���������������� ������������� ����������� !� 6" 10# 5

$%&'����(�)*+)�,-./0123456 (Table 1)7 *+)/018956#:;�<�=>?@AB� CD1�E5F 3 mmGH�IMetri-

dia pacificaJKLMN�#1OPQRS#?@AB�I T6$%&'�<�UVW-1XFYLI C4Z C5

��[\]$%&'FYLI C3]G^_-�U/01895`ab6B�1:;�<�@c95A7 T6IdeJfb6�������������MN�����*+)gh13i9567������������jk��lMA������� !�UMicrocalanus pygmaeus?I m�?Metri-

dia pacifica, Paracalanus parvus, Pseudocalanus minu-

tus ?@nI o��p�?�lMAU Neocalanus

cristatus, N. plumchrus, N. flemingeri Z Eucalanus

bungii?@A1 (Yamaguchi et al. 2002)I *+)-./018956UB59jk���o��p��l#[qI M. pacificar?@b6 (Table 1)7 BM.

pacifica �-./0MA*+)�U 2s�t Paraci-

neta"u� 1de4567 M`vqwx?y�=z41 50{100 mm?|�} (stalk)J~�F (type A:

Fig. 2A, C)�I ���fL����`D�}J~qI y

�U=>? (300{500 mm)�����1�������AF (type B: Fig. 2B, D)?@A7 M. pacifica�UB 2s�t*+)G��I Paracineta pleuro-

mammae���95A*+)1 41�N, 165�E����gh<61I �/0_-U 1_-r?@b67 �������������A������� !������M. pacificaU_-j? 2�I o��p�? 5����MA�l#?@n (Yamaguchi et al. 2002)I �-./0S*+)��J9a�MAB�U�����95A?I G��y�deJfb67

Metridia pacifica������������������������ Metridia pacifica-.�8956*+)U^� 2s�t[q type A

(Fig. 2A, C)r1gh<67 �����<6 0.1 mm

����n ¡<�I M. pacifica C1]G¢£ deJfb61I *+)-./0UVW-�r89567�������M. pacificaVW-U¤ 3{4¥��¦{§¨ 8{10¥� ��5©5ghª��J~b��6 (Fig. 3)7 *+)/0�U 0{70«¬J­®<I�¯/0�U 9.4«?@b67 °±J²<�deJfb6 2003±� 2004±����rA�/0�U�5©5M. pacificaVW-ghª��³]�´LI XF´�/0�18956U�E5F¦{§¨?@b67

Table 1. List of calanoid copepod species/stages on which epizoic suctorian ciliates were observed, its prevalenceand geographical locations. �: no data.

Species Stage Prevalence («) Location

Gaetanus armiger C6F 0�50 NW PacificGaetanus simplex C5F 0�7 Okhotsk Sea

C5M 0�25 NW Pacific, Okhotsk SeaC6F 0�50 NE, NW Pacific, Okhotsk Sea

Gaetanus variabilis C5F 0�6 NW PacificC6F 0�10 NW Pacific

Heterorhabdus tanneri C6F 0�50 NW Pacific, Okhotsk SeaHeterostylites major C6F � NW Pacific

C6M � NE PacificMetridia pacifica C6F 0�68 NE, NW Pacific, Okhotsk SeaParaeuchaeta birostrata C5M 0�50 NW Pacific, Okhotsk Sea

C6F 0�100 NC, NE, NW Pacific, Okhotsk SeaParaeuchaeta elongata C4M 0�100 Okhotsk Sea

C5F 0�25 NW Pacific, Okhotsk SeaC5M 0�6 Okhotsk Sea, Japan SeaC6F 0�100 NC, NE, NW Pacific, Okhotsk Sea, Japan SeaC6M 0�11 Okhotsk Sea

Paraeuchaeta rubra C5F 0�25 NW PacificC5M 0�33 NW PacificC6F 0�100 NE, NW Pacific, Okhotsk Sea

Scaphocalanus magnus C6F 0�100 NW Pacific

µ µ

��t�¶�·¶¸¹� º 53» º 1¼ (2006)32

Page 6: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

Metridia pacifica���������Metridia pacifica�������������� ���������������� ��������������� !�"#$ %&�' ()

%&��*"�+,-M. pacifica./0�123���4����5 0668789:;��3� ������� 45630�N< -5=;123>�&� (Table 2)'������5���������8��������8?�@A��+,BC@DEF G3� H?@D����#$IJ��5H?@A��KL����� !-&�' ���� !-5� MN��O�PQ�-123� type A�R��ST" type B�R�(Fig. 2B, D)?123�' U� type B�R�5���������@A�+, (47VN, 165VE; 49VN, 165VW)��D"?#$IJ�' WX"������5YZ�[;� M. pacifica./0�12\0]?^D@A�!���D"@DEF�&� (Table 2)'

Metridia pacifica����� ���H?�������@�&����� !�_��`D" Metridia pacifica./0a�0�������K #$3�' b0��K 7cd��e"#$3�8Uf� H?@D����ghJ��5��-=\0�897��D"������ij8>&"D� (Fig. 4)'k�@D����ghJ��5l� (607)-<m���(197)��no (57)�p-&�' q�rs�tou�o�5������5ghJ>�&�' ��R��9�5 type A-v�hJ"D��� type B����K?��� l�����no-w� R��xyz��&"���K5{>h>�&�' q��|\0��D"type A8 type B�R������3"D\0?�]123�'

� �����������������0���}~������������}5����-9:;{>U8��hJ"D (Fernandez-

Leborans 2004)' ����5�DYZ��� pH����123� �}��D�� �����5Y� �!������]u�������"�#� �9:;�� �e� | $ � � � � } - (Fernandez-Leborans

2004)' ����������84��%! �&'!��� ¡�¢£!� ��D"¤¥¦§(¤y¨©� 6t10ª� 5)«¬­��D"0���}~��������12�®*IJ� (Table 1)' U�O¯ Paraeucha-

eta birostrata, P. elongata��� P. rubra�`D"54J°J{>����0���3"�w� ±²�{}�U8�G³IJ"D (Yamaguchi & Ikeda

2001)' ´!}+µ�80���}~������?±

Fig. 3. Seasonal changes in abundance of Metridiapacifica adult females (open symbols), and prevalenceof epizoic suctorian ciliates (closed symbols) at Site Hin the western subarctic Pacific (cf. Fig. 1) during May2002 to December 2004.

Table 2. Spatial changes in integrated meantemperature (0�50 m), density of Metridia pacificaadult females, and prevalence of epizoic suctorianciliates on them.

Location/PositionTemperature

(¶)Density

(inds. m�3)Prevalence

(7)

Bering Sea (177VW)53V30�N 8.1 63.0 67.6

NW Pacific (165VE)47V00�N 6.4 4.5 30.844V00�N 7.3 4.0 17.442V30�N 7.4 36.4 3.241V00�N 14.2 7.7 3.1

NE Pacific (165VW)50V00�N 7.9 33.1 48.149V00�N 8.2 108.4 23.348V00�N 8.0 4.0 4.047V00�N 9.2 5.9 4.845V30�N 10.4 9.2 0.044V00�N 10.8 19.7 0.042V30�N 11.0 1.7 0.041V00�N 13.8 0.5 0.0

· ·

z¥�£¸���e¹�º»��4�^,}8¼"} 33

Page 7: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

��������� ���� � ������������ ��� ������ ��������� ��������� !"#$�%�" (O≈lafsdot-

tir & Svavarsson 2002)&�� �'(�)#*+��,-�%�����.������/0123�0456���#7%� 8� 3 mm9:�;< � �=>?�<��@� AB�C�D#7/EF�G�H,�@���"I#�JK��" (Table 1)& >����,-�%L� !M�,LN;O<,P�����.������Q� I�P.�����R"ST�N$RU"/V !"#WX$�%�" (O≈lafsdottir & Svavarsson 2002)& �=>?�<���Y/I#,Z�%P� �[ ���.��\]^_�`aP� bc#de�!"I#�!��%-� (Coppellotti & Matarazzo 2000)� �=>?�<P� bcfN? ��� �'(�) Pg> 800 m�h

,�"R"� ,�@#$7%�"I##ei�!"�L7�O�& �����.���j��/0456�\]� �?,L`aR"Metridia pacifica� Paraeucha-

eta elongataPklO%mn&o�C�\< !"I#L��#7%JK��"& '<��g>Pp�q(#�#$7%�/)r (D25stD75s) �R# 40t240

m (M. pacifica)# 60t500 m (P. elongata) !" (Ya-

maguchi et al. 2002)&*uv�w,��R"0123�0456� Limno-

calanus macrurus B���,��R"^_� Toko-

phrya quadripartita P+�G�H,P���Q� L.

macrurusx�,yR"+�-z1{|}{<,P���O�I#�!��%�" (Evans et al. 1979)& M/� ��~��4{.,-�%0123�0456�Metridia lucens#M. longa��,��R"^_�'� Paracineta sp.��P�#���B�,���"/V� Sherman & Schaner (1965)P/0�, Paraci-

neta sp.�/17%7M\� L7@PST#7%B�C�� �" PO��#��7%�"& ��,-�%L��/0��P�����.�������2CR"��O�,OY%�"#����%�"(O≈lafsdottir & Svavarsson 2002)& ��� �@0456�,-�%^_�������"�G�H�EF�G�H,��7%�/P (Table 1)� EF�G�H,-�%/0�� ��3���34��� ���I#�Mauchline 1998C��� 5� !"#WX �"&

Metridia pacifica���������Metridia ������^_�,Z�%P� Sher-

man & Schaner (1965) ���~��4{.,-�%M. lucens#M. longa�^_� Paracineta sp.����,P����!�� ���6,��%��I#C7 7%�"& ��~8�� �¡9) M. pacifica

^_���:P¢£;�AB�,"R"�F (Pad-

mavati et al. 2004),�@� �?�¤g¥,O"¦t�� �Kobari & Ikeda 2001 Fig. 2C��� ,¤�Y/(Fig. 3)& Evans et al. (1979)Lv�w Limnocalanus

macrurus,Z�%^_���:P¢£;�¡,"R"�F,§L¤�I#C7 7%�"& M/� Ohtsuka

et al. (2004),x�¨©ª<�,-�%«��.����¬'�� 0456����P 3­t6­,P���Q� =®O�������"& �¬'�¯°±{} (phoront)��}²�� (tomite)�>³P6�g¥´µ¶ P¦g¥´µ¶,?·%fV%����CR"I#�*� !"#����%�" (Oh-

Fig. 4. Incidence of the two types ciliates (bottom)on various parts of the body of Metridia pacifica adultfemales (top). Ce: cephalosome, Me: metasome, Ur:urosome, A1: antennules, FA: feeding appendagesexcept antennules, SL: swimming legs and CR: caudalrami.

¸ ¸

%@z1{|}{¹º7 A 53» A 1¼ (2006)34

Page 8: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

tsuka et al. 2004)� ������ ������������ �������������

Metridia pacifica���������������� Metridia pacifica������������ �� !�"� #$��%&�#'� �$�%&��(�)*��+�,�,��!��-���.(Table 2)� ����/01��2,�M. lucens�M.

longa���������� Paracineta sp.����! �������%&��'� Sherman & Schaner

(1965)���������! 43�"#'���4���-�� ����/01����-�� �!.,Nova Scotia Water�5�"#$%!������,-�&6�,�� ����7819:��2;�<=>?' Meganyctiphanes norvegica ���������@���ABC �D,����� ���EF!�"�GH�"�()��#, (8I82J)��*+�� K��()���, @2JLMC ��!+NO��,� (Nicol

1984)� ���,�+�� Nicol (1984)�GH�"�()���P�Q�!�'� ��������!-.����-/ ���RS( �T� GH�"�()�<=>?'��P�-.UV!0-�..W� �����XYZO�[ -�.���\+�,�� ��1D��� �]^���_2�"#$%!���`���,��� (Sherman & Schaner 1965)� ����-.UV�a0!�������R#W��� (Nicol 1984)

�34-�bc�M. pacifica� �"# @#$�%&�#,���� Table 2C R5� ���!d6�����2� Sherman & Schaner (1965)!Metridia lucens

�M. longa�����������+�+N+.Para-

cineta sp.�e78f! 23I66 mm�[[9P������ bcg\O�.��� type A @e78f 50I100 mm, Fig. 2A, CC �h:����'i�,�� +-+ Sherman & Schaner (1965)� type B�; �����P�j<=e7 (Fig. 2B, D)�>k�D,��&6�,�,� ���AB�; �?�lmno� Curds

(1985a, b, c, 1986, 1987)���p@����2,�Ae�q]��,�� �k���r����2,�g\O�.���AB����?B(st�CD"������ buEvFwR��?B(stRGW�,'H!���

Metridia pacifica������� �bcMetridia pacificax������yIJz�+��KJ� {J2�|LJ!��M��� }�Np

O~� ���;�[K����yI����,�,��!"-�. (Fig. 4)� Ephelota;����� 0'PQR��SRZ��,�������!�+,��Jz��� �����"� T�Ue7VW�* �XY��RZ[ ��!���������,� (O≈lafsdottir &

Svavarsson 2002)� �.� ���!%[\k��]�'���,�O��,���� 0'PQR��S (stalk)

RZD����]�'!�R����^_�`����,.W���������,� (O≈lafsdottir & Sva-

varsson 2002)� bcM. pacifica�2,�����!�-�.KJ� {J2�|LJ��0>�'2��ab�zc+�2"� d������D"�e;[ ,Jz���� p4� �������-�.NpO~� ���;�[K��d�Xf����g��hO��Jz @ �]^^_Re;[ ,JzC ���.W� ������yI�+�i+�,�,���r��

� � � �Kitching (1952)[ Lilly (1953)����� ����Oj��������*RX���kY������!��-�O��,�� ���!�0>�'������ ����l)�"#&���� ���m�Z�2�|XY-��cn�*!������ op����0>�'�e;�l��(�)*�,��]��!� ����q�0>�' Chiridius armatus���������!��+�,�����"f��!Z� ���+NO��,� (Olsen et al. 2000)� ����������"��r !Z+�� d�� �¡¢£¤omRs"��t �H!��.Wf��RZ�+.��¥uO��,� (Olsen et al. 2000)� �0>�'�e;�Cl��+����"��v�e7!�� �����"d�6w!/^�� ¦8§V!x[-�q�¨� ©�1ª«1XYy�XYO�[ '����!�����(Weissman et al. 1993)�zW�?U¬­R������ ®z[������{l¯z�°W�±²oª�?U¬­���!� ��©�1ª«1�®zzW�¬ �|��� 5���v}[?v³�¬ ���`���2"� 5�z:´����!�~O�������-�. (Ho & Perkins 1985,

Theodoride◊s 1989)� +-+µ�� �K�©�1ª«1¶��2,�®zzW���·��!�¸���,(*���,��!+NO�DD�" (Gomez-Gutierrez et

al. 2003, Ohtsuka et al. 2004)� ��©�1ª«1�·���+�������.XY�¹º����� 5r5r®z�,��R�~ 6��»���,��,�

¼ ¼

©�1ª«1�2;�®zX¯z�5������ 35

Page 9: 北太平洋亜寒帯域のカラヌス目カイアシ類に体表付着する繊毛虫 … · regard to the substrate of suctorians on M. pacifica,they were observed most frequently

�� ������������ ���� ��� ��������� ������� !"#$%&'()*+,'�-./�0�� 1&� 23����45�678/,9:;<=>?� ��@ABCD/ EFG����HIJ9KLMNO0�,9P��

� �Q!RP��0&S� TUVW�0&>X&� 4YZ[\� ]^\� _3\� ]`\� ]a\� bc\ defgh,diKLfgh�Jjkl9&Y1$� mQ�no'pq�!rS1Y&]2stutuvwxyuKLv z{ |}~,��#�}~��=>kl9&Y1$� 1&� mQ!�j��=5!��1Y&��tutuv���yuKLy t� �}~�kl9&Y1$�

� � � �

Coppellotti, O. & P. Matarazzo 2000. Ciliate colonization ofartificial substrates in the Lagoon of Venice. J. Mar. Biol.Ass. U.K. 80: 419�427.

Curds, C. R. 1985a. A revision of the Suctoria (Ciliophora,Kinetofragminophora) 1. Acineta and its morphologicalrelatives. Bull. British Mus. Nat. Hist. (Zool.) 48: 75�129.

Curds, C. R. 1985b. A revision of the Suctoria (Ciliophora,Kinetofragminophora) 2. An addendum to Acineta. Bull.British Mus. Nat. Hist. (Zool.) 49: 163�165.

Curds, C. R. 1985c. A revision of the Suctoria (Ciliophora,Kinetofragminophora) 3. Tokophrya and its morphologi-cal relatives. Bull. British Mus. Nat. Hist. (Zool.) 49: 167�193.

Curds, C. R. 1986. A revision of the Suctoria (Ciliophora,Kinetofragminophora) 4. Podophrya and its morphologi-cal relatives. Bull. British Mus. Nat. Hist. (Zool.) 50: 59�91.

Curds, C. R. 1987. A revision of the Suctoria (Ciliophora,Kinetofragminophora) 5. The Paracineta and Corynophryaproblem. Bull. British Mus. Nat. Hist. (Zool.) 52: 71�106.

Evans, M. S., L. M. Sicko-Goad & M. Omair 1979. Seasonaloccurrence of Tokophrya quadripartita (Suctoria) as epi-bionts on adult Limnocalanus macrurus (Copepoda: Cala-noida) in southeastern Lake Michigan. Trans. Am. Mi-crosc. Soc. 98: 102�109.

Fernandez-Leborans, G. 2001. A review of the species ofprotozoan epibionts on crustaceans. III. Chonotrich cili-ates. Crustaceana 74: 581�607.

Fernandez-Lebotans, G. 2004. Protozoan epibionts on Mysusrelicta Loven, 1862 (Crustacea, Mysidacea) from LakeLusiai (Lithuania). Acta Zoologica 85: 101�112.

Fernandez-Leborans, G. & M. L. Tato-Porto 2000a. A reviewof the species of protozoan epibionts on crustaceans. I.Peritrich ciliates. Crustaceana 73: 643�683.

Fernandez-Leborans, G. & M. L. Tato-Porto 2000b. A reviewof the species of protozoan epibionts on crustaceans. II.Suctorian ciliates. Crustaceana 73: 1205�1237.

Gomez-Gutierrez, J., W. T. Peterson, A. De Robertis & R. D.

Brodeur 2003. Mass mortality of krill caused by parasi-toid ciliates. Science 301: 339.

Henerby, M. S. & B. T. Ridgeway 1979. Epizoic ciliatedprotozoa of planktonic copepods and cladocerans andtheir possible use as indicators of organic water pollution.Trans. Am. Microsc. Soc. 98: 495�508.

Ho, J.-S. & Perkins 1985. Symbionts of marine Copepoda: Anoverview. Bull. Mar. Sci. 37: 586�598.

Kitching, J. A. 1952. Observations on the mechanism offeeding in the suctorian Podophrya. J. Exp. Biol. 29: 255�268.

Kobari, T. & T. Ikeda 2001. Life cycle of Neocalanus flemin-geri (Crustacea: Copepoda) in the Oyashio region, westernsubarctic Pacific, with notes on its regional variations.Mar. Ecol. Prog. Ser. 209: 243�255.

Lilly, D.M. 1953. The nutrition of carnivorous Protozoa.Ann. N.Y. Acad. Sci. 56: 910�926.

Mauchline, J. 1998. The biology of calanoid copepods. Adv.Mar. Biol. 33: 1�710.

Nicol, S. 1984. Ephelota sp., a suctorian found on the eupha-usiid Meganyctiphanes norvegica. Can. J. Zool. 62: 744�746.

Ohtsuka, S., M. Hora, T. Suzuki, M. Aikawa, G. Omura & K.Yamada 2004. Morphology and host-specificity of theapostome ciliate Vampyrophrya pelagica infecting pelagiccopepods in the Seto Inland Sea, Japan. Mar. Ecol. Prog.Ser. 282: 129�142.

O≈lafsdottir, S.H. & J. Svavarsson 2002. Ciliate (Protozoa)epibionts of deep-water asellote isopods (Crustacea): pat-tern and diversity. J. Crustacean Biol. 22: 607�618.

Olsen, E.M., T. Jorstad & S. Kaartvedt 2000. The feedingstrategies of two large marine copepods. J. Plankton Res.22: 1513�1528.

Padmavati, G., T. Ikeda & A. Yamaguchi 2004. Life cycle,population structure and vertical distribution of Metridiaspp. (Copepoda: Calanoida) in the Oyashio region (NWPacific Ocean). Mar. Ecol. Prog. Ser. 270: 181�198.

Sherman, K. & E. G. Schaner 1965. Paracineta sp., an epizoicsuctorian found on Gulf of Maine Copepoda. J. Protozool.12: 618�625.

Theodoride◊s, J. 1989. Parasitology of marine zooplankton.Adv. Mar. Biol. 25: 117�177.

Turner, J. T., M. T. Postek & S. B. Collard 1979. Infestationof the estuarine copepod Acartia tonsa with the ciliateEpistylis. Trans. Am. Microsc. Soc. 98: 136�138.

Weissman, P., D. J. Lonsdale & J. Yen 1993. The e#ect ofperitrich ciliates on the production of Acartia hudsonicain Long Island Sound. Limnol. Oceanogr. 38: 613�622.

Yamaguchi, A. & T. Ikeda, 2001. Abundance and populationstructure of three mesopelagic Paraeuchaeta species (Co-pepoda: Calanoida) in the Oyashio region, western subarc-tic Pacific Ocean with notes on their carcasses and epizoicciliates. Plankton Biol. Ecol. 48: 104�113.

Yamaguchi, A., Y. Watanabe, H. Ishida, T. Harimoto, K.Furusawa, S. Suzuki, J. Ishizaka, T. Ikeda & M.M. Taka-hashi, 2002. Community and trophic structures of pelagiccopepods down to greater depths in the western subarcticPacific (WEST-COSMIC). Deep-Sea Res. I 49: 1007�1025.

2005� 11� 24��C� 2006� 1� 18���

� �

�m���u�� � 53� � 1� (2006)36