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Oodinium Inlandicum Sp. Nov. (Blastodiniales, Dinophyta), a New Ectoparasitic Dinoflagellate Infecting a Chaetognath, Sagitta Crassa

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Oodinium Inlandicum Sp. Nov. (Blastodiniales, Dinophyta), a New Ectoparasitic Dinoflagellate Infecting a Chaetognath, Sagitta Crassa Plankton Biol. Ecol. 48 (2): 85-95, 2001 plankton biology & ecology t> The Plnnkton Society of Japan 2001 Oodinium inlandicum sp. nov. (Blastodiniales, Dinophyta), a new ectoparasitic dinoflagellate infecting a chaetognath, Sagitta crassa Takeo Horiguchi1 & Susumu Ohtsuka2 'Division of Biological Sciences, Graduate School of Science, Hokkaido University, Sapporo 060-08 JO, Japan ^Fisheries Laboratory, Hiroshima University, 5-8-1 Minato-machi, Takehara, Hiroshima 725-0024, Japan Received 13 February 2001; accepted 26 April 2001 Abstract: A new ectoparasitic dinoflagellate, Oodinium inlandicum Horiguchi et Ohtsuka sp. nov. in festing a neritic chaetognath, Sagitta crassa Tokioka is described from the Seto Inland Sea, western Japan. This is the first record of the genus Oodinium in the western North Pacific. The new species is readily distinguished from other congeners by: (1) host specificity; (2) the shape and location of the nucleus; and (3) the morphology of the peduncle. The present study has revealed that as develop ment proceeds, the nucleus and cytoplasm change markedly. The cytoplasm of the mature trophont consists of a central endoplasm, rich in cytosol and a highly membranous exoplasm. The nucleo- plasm is uniform and lacks electron dense structures, such as chromosomes. In contrast, the young trophont has an elongated dinokaryotic nucleus, which almost completely occupies the anterior half of the cytoplasm. The peduncle penetrates host tissue, but mechanical damage caused by the para site seems not to be extensive. The comparative ultrastructure and ecology of the new parasite are also discussed. Key words: dinoflagellate, Oodinium inlandicum sp. nov., ectoparasite, chaetognath, Sagitta crassa During our survey of parasites of zooplankton in the Seto Introduction Inland Sea, western Japan, the most common neritic A large number of dinoflagellates are known to parasitize chaetognath encountered was Sagitta crassa Tokioka, and it marine vertebrates and invertebrates (e.g., Chatton 1920; was heavily infested by an Oodinium-\ikc dinoflagellate. Cachon & Cachon 1987; Shields 1994; Ohtsuka et al. Light and electron microscopical observations have re 2000). Those parasitic dinoflagellates only displaying a typ vealed that this is an undescribed species of the genus Ood ical dinokaryotic nucleus during a part of their life cycle are inium. The taxonomy of parasitic dinoflagellates in Japan classified into the order Blastodiniales, class Blastodini- ese waters is poorly understood (Ohtsuka et al. 2000), and phyceae (Fensome et al. 1993). Currently, six families are thus biodiversity studies centering on parasitic dinoflagel recognized in the order Blastodiniales. One of these, the lates are badly needed. family Oodiniaceae, has members which are characterized by being ectoparasitic and possessing suboval to fusiform Materials and Methods cells with well-developed absorption apparatuses in the trophont stage. This results in very atypical dinoflagellate Sampling morphology, unlike the trophonts of the Protoodiniaceae, Host zooplankton were collected on three separate occa which are clearly recognisable as belonging to this division. sions (September and October 1999 and October 2000) by The type genus of the family, Oodinium was established by nocturnal surface net tows using a conical net (diameter 30 Chatton (1912) and consists of ectoparasites of metazoans cm; mesh size 0.1 mm) in the center of the Seto Inland Sea, such as annelids, appendicularians, and chaetognaths (Chat western Japan, (34°19'N, 132°55'E). Chaetognaths with at ton 1912; Cachon 1964; McLean & Nielsen 1989). tached dinoflagellates collected on 9 September 1999 were used for the current descriptions and measurements. Corresponding author: Takeo Horiguchi; e-mail, [email protected] 86 T. Horiguchi & S. Ohtsuka Light microscopy cytoplasm yellow-brown, granular in appearance; peduncle bulbous to rod-shaped, 6-500//m in length, almost trans Chaetognaths infested by Oodinium cells were fixed in parent, with many longitudinal striations; nucleus in mature 10% neutralized formalin/sea-water and observed using a cell non-dinokaryotic, hemispherical in shape, situated in Nikon Eclipse E600 microscope. For observation of the the upper extremity of the cell; nucleus in immature cell di- thecal plates, fixed dinoflagellate trophonts were stained with 1% Fluorescent Brightener 28 (Sigma) and observed with a Nikon Optiphoto-2 epifluorescent microscope, equipped with a Nikon EFD2 epiflourescent device and with an UV-2A filter set). The nucleus was visualized by staining fixed cells with DAPI (4'-6-Diamidino-2-phenylin- dole) (0.5 g ml"1) and observing them with the same epiflu orescent microscope. Measurement of the body length of Sagitta crassa, from the anterior tip of the head to the pos terior end of the caudal fin, was made on all undamaged in dividuals in the sample collected on 9 September 1999. The degree of infection relative to host size and the sites of at tachment of Oodinium to the body of S. crassa were noted. The frequency of each attachment site in each size class of the host was calculated as a percentage of the total number of sites. Transmission electron microscopy (TEM) 2-3 3-4 4-5 5-6 6-7 7< Parasitized specimens of Sagitta crassa were fixed in the Body length (mm) field in 2.5% glutaraldehyde made up with seawater, and Fig. 1. Relationships between prevalence of infection of para then transported at about 5°C to the laboratory. The fixed site Oodinium inlandicum sp. nov. and body length of host Sagitta specimens were rinsed with seawater and postfixed in 2% crassa. Number of chaetognaths examined is shown above each Osmium tetroxide for 1 h. These were dehydrated through column. an ethanol series and finally embedded in Spurr's resin (Spurr 1969). The chaetognaths were individually mounted in a small amount of resin between two sheets of overhead projector (OHP) transparency film prior to polymerization so as to obtain thin flat embedded samples. After polymer ization, Oodinium cells were located using a light micro scope and excised (size 1X2mm), together with a small piece of the host from the thin resin plate using a razor blade. These excisions were then stacked on to a plastic block for sectioning. Observations were made with a JEOL 100S transmission electron microscope. Results Description Oodinium inlandicum Horiguchi et Ohtsuka sp. nov. 2-3 3-4 4-S S-6 6- 7 7< Ectoparasita chaetognathae, Sagittae crassae. Cellula Body length (mm) vegetativa (trophont) ellipsoidea vel bacilliformis, 30-150 Head /im longa; cytoplasma luteoli-brunneola granulataque; pe- dunculus bulbiformis vel bacilliformis, 6-500/im longus et cu Trunk fere hyalinus, cum striis longitudinalibus numerosique; nu nm Tall cleus cellulae maturae non-dinokaryonticus hemisphaeri- m Anterior fin cusque, in parte extremitas supera cellulae situs; nucleus in Posterior fin cellula immatura, dinokaryonticus ellipsoideaque; thecam Tall fin habens; sine chloroplasti et stigmate. Fig. 2. Distribution of Oodinium inlandicum sp. nov. on body of Ectoparasite on chaetognath, Sagitta crassa. Vegetative host Sagitta crassa. Number of individuals examined is same as in cell (trophont) oval or rod-shaped, 30-150//m in length; Fig. I. A New Ectoparasittc Dinoflsgellate 87 Fig, 3, Chaetognatha, Sagiiia crassa heavily infected by Ood- iiiium inlattdicum sp. nov. A. An terior part of Sagitta crassa. Ar rowheads indicate Oodiniitm trophonts, B. Posterior part, in fected by larger clinofkigellaics. Scalebar=I00/im. nokaryotic, oval; thecal plates present; no chloroplasts and Trends of infection eyespot present. Infestations of Oodinium inkuulicum were apparent in Holotypc: An embedded specimen in cpoxy resin collections of Sagitki crassa made during September and (mounted on a slide glass) has been deposited in the herbar October 1999 and again in October 2000. The occurrence ium of the Graduate School of Science. Hokkaido Univer of this parasitic dinoflagellate seems to have been restricted sity as SAP 089327. Isotype: SAP 089328. to warm water seasons, i.e. water temperature between 18-26°C, although Sagitta crassa persisted year-round in Type locality the Seto Inland Sea. It is interesting to note that a coexist Takehara (34°19'N, 132°55'E), Hiroshima Prefecture, ing species of chaetognath, Sagitta enftaia Grassi, at all western Japan (Seto Inland Sea). times was found to be unaffected by the new dinoflagellate. Larger specimens (more than 6 mm long) of 5. crassa Etymology tended to be infected more heavily by the dinoflageilate than small ones (Fig. 1). Ninety eight percent ofchaetog- The epithet refers to the type locality, viz. the Seto Inland naths larger than 6 mm were infected by the dinoflagellate, Sea. The new dinoflagellatc has hitherto been known only while less than 15% of specimens smaller than 2-3 mm in this inland sea region. long were infected (Fig. 1). Attachment sites of the parasite on the body of S, crassa arc shown in Fig. 2 according to host size. Trophonts of O. iniandicum preferentially at- 88 T. Horiguchi & S. Ohtsuka Fig. 4. Oodinium in- sp. nov. A. ts on body trunk. Note long peduncle (P). Nucleus (N) and nuclco- lus (arrowhead) can be seen. B. A large and two young trophonts on cau dal fin. C. Trophonts as sociated with ciliary fence receptor (arrowhead). D. Two young trophonts on caudal fin. Scale bar= 100/mi (ached to the body (head, trunk and tail) of S. crassa, but (Fig. 5). The significance of a long peduncle is, however, as were also less frequently observed on the fins (Figs 3, 4). yet unknown. Mature trophonts are tinged yellow-brown The head, trunk and tail of ihc host were almost equally uti and are uniformly granular, except for the upper extremity lized as attachment sites except in small (less than 2 mm) where a nucleus is located (Figs 3, 4). This nuclear region chaelognaths. The anterior, posterior and tail fins were less is hemispherical and appears to be more or less transparent. frequently (2-15%) colonised by the parasite. The In younger trophonts. the color of the cell is paler and the trophonts were often found to associate with the ciliary distinction between the nuclear region and the remainder of fence receptors on the body of 5.
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