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Morphology and infraciliature of two new marine ciliates

European Journal of Protistology xxx(2011)xxx–xxx

Morphology and infraciliature of two new marine ciliates, Paracyrtophoron tropicum nov.gen.,nov.spec.and Aegyria rostellum nov. spec.(Ciliophora,Cyrtophorida),isolated from tropical waters in southern China

Xiangrui Chen a,b,Xiaozhong Hu b,?,Jun Gong c,Khaled A.S.Al-Rasheid d,Saleh A.Al-Farraj d, Weibo Song b

a College of Life Science and Biotechnology,Ningbo University,Ningbo315211,China

b Laboratory of Protozoology,Institute of Evolution and Marine Biodiversity,Ocean University of China,Qingdao266003,China

c Yantai Institute of Coastal Zone Research,Chinese Academy of Sciences,Yantai264003,China

d Zoology Department,King Saud University,Riyadh11451,Saudi Arabia

Received11October2010;received in revised form20July2011;accepted21July2011

Abstract

The morphology and infraciliature of two new marine cyrtophorid ciliates,Paracyrtophoron tropicum nov.gen.,nov.spec. and Aegyria rostellum nov.spec.,isolated from tropical waters in southern China,were investigated using live observation and protargol impregnation methods.Paracyrtophoron nov.gen.differs from the closely related Cyrtophoron by lack of fragment kinety at anterior ends of right somatic kineties and thigmotactic cilia in posterior portion of ventral surface,while from the well-de?ned Chlamydodon by lack of the cross-striped band around the periphery of the somatic?eld.Paracyrtophoron tropicum nov.spec.,the type of the new genus,can be recognized by the combination of the following characters:cell size about 150–175×70–90?m in vivo;elliptical to kidney-shaped in outline,dorsoventrally?attened about2.5:1;conspicuous cortical granules;one canal-like depression extending from postoral area to subcaudal region of cell;ca.90somatic kineties;12–16 nematodesmal rods;one or two terminal fragments on dorsal side.Aegyria rostellum is characterized by the following features: size about90–150×40–70?m in vivo,triangular or ear-shaped body with broad anterior end,having a rostriform structure and pigment spots,56–63somatic kineties,one preoral kinety,three or four circumoral kineties,and32–42nematodesmal rods.Based on previous and current studies,the de?nition for the genus Aegyria is updated:body dorsoventrally?attened;oral ciliature consisting of one preoral and several circumoral kineties;podite located in posterior ventral region and surrounded by somatic kineties;no obvious gap between right and left somatic kineties;postoral and left somatic kineties progressively shortened posteriorly from right to left.Additionally,two new combinations were proposed.

Keywords:Aegyria;Marine ciliates;Morphology;New species;Paracyrtophoron

?Corresponding author.

E-mail address:xiaozhonghu@https://www.wendangku.net/doc/651076355.html,(X.Hu).

doi:10.1016/j.ejop.2011.07.005

2X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx

Introduction

Cyrtophorid ciliates are commonly found in aquatic bio?lms or periphytons on a variety of immersed sur-faces such as stones,macrophytes,and arti?cial substrates (Baldock et al.1983;Borror1972;Cairns and Yongue1968; Deroux1970;Gismervik2004;Gong et al.2005;Kahl 1931).As an extensive pioneer work on marine cyrtophorids, Deroux(1975,1976a,b,c)described ca.50species from French coast off the Atlantic using arti?cial substrate and silver staining methods,contributing greatly to the taxon-omy and biogeography of this group of ciliates.Subsequently, more species were added(Dragesco and Dragesco-Kernéis 1986;Hu and Suzuki2005;Petz et al.1995;Song and Packroff1997;Song and Wilbert2002)and some molecu-lar investigations were performed regarding the phylogenetic relationships among this highly diverse group(Gong et al. 2008;Li and Song2006).During the last ten years’survey on ciliate fauna in the Bohai Sea and Yellow Sea,northern China,over40cyrtophorid species were identi?ed(Gong and Song2003,2004,2006;Gong et al.2002,2003,2005; Shao et al.2008;Pan et al.2011).Among these,most belong to species-rich genera,for example,Dysteria Huxley,1857, Chlamydodon Ehrenberg,1835,and Hartmannula Poche, 1913.While some other genera are relatively inadequately studied and thus need extensive sampling and investigation. Recently we isolated two cyrtophorid species during inves-tigation on ciliate diversity in tropical mangrove wetland waters in Shenzhen,southern China.Detailed observations have demonstrated that both are hitherto unknown species, one belonging to the little known genus Aegyria Claparède and Lachmann,1859while the other cannot be assigned to any known cyrtophorid genus.Based on current observations a new diagnosis for the genus Aegyria is provided. Material and Methods

Paracyrtophoron tropicum nov.spec.and Aegyria rostel-lum nov.spec.were collected from Shenzhen Bay mangrove wetland in Shenzhen(114?04 E,22?37 N),southern China in April2008.Sediments and roily seawater were taken,with locale temperature about25?C and salinity about16‰.The upper layer of sediment was collected together with some water from the site,and raw cultures have been maintained for10days at room temperature,with the addition of several rice grains during that period to enrich bacteria as food for the cultures.Clone culture was attempted but failed.Living cells were observed at100–1250×magni?cation using differen-tial interference contrast microscopy.The infraciliature was revealed by protargol impregnation(Li et al.2010;Song and Wilbert1995).Drawings of stained specimens were made with the help of a camera lucida.Measurements were per-formed under100–1250×magni?cation.Systematics and terminology are mainly according to Corliss(1979)and Lynn (2008).

Results and Discussion

Family Chlamydodontidae Stein,1859 Paracyrtophoron nov.gen.

Diagnosis:Rightmost kineties extending dorso-apically and bending to the left,forming a conspicuous suture with left kineties;three circumoral kineties obliquely arranged; nematodesmal rods sturdy and prominent;cross-striped band, transpodial segments and anterior terminal fragment absent; single heteromerous macronucleus;marine habitat. Etymology:The name Paracyrtophoron is a composite of the Greek pre?x para+(beside,at,along,during)and the name of the cyrtophorid genus Cyrtophoron and refers to the similar ventral ciliature of Cyrtophoron and Para-cyrtophoron.For Cyrtophoron no derivation is given in the original description.Like Cyrtophoron,neuter gender. Type species:Paracyrtophoron tropicum nov.spec. Comparison with closely related genera:Considering the general living morphology and infraciliature,the new genus belongs to the family Chlamydodontidae,which includes three genera,i.e.Cyrtophoron Deroux,1975,Chlamydodon Ehrenberg,1835and Lynchellodon Jankowski,1980(Lynn 2008).Paracyrtophoron differs from Cyrtophoron mainly by the lack of a fragment near anterior ends of right kineties and transpodial fragments in the posterior portion of the ven-tral surface(Aliev1991;Deroux1975).Chlamydodon can be distinguished clearly from the new genus by having a critical generic character,i.e.cross-striped band(CSB)around the periphery of the somatic?eld(Deroux1975,1994;Dragesco and Dragesco-Kernéis1986;Ehrenberg1835;Fauré-Fremiet 1950;Gong et al.2005;Lynn2008;Kahl1931;Kurth and Bardele2001;Song et al.2009).Lynchellodon was estab-lished for Chlamydodon lynchelliformis Borror,1972,which has a CSB but lacks an anterior suture,thus can not be con-fused with the new genus.

Paracyrtophoron tropicum nov.spec.(Figs.1A–D,2,3; Table1)

Diagnosis:Body size about150–175×70–90?m in vivo; cell elliptical to kidney-shaped in outline,dorsoventrally?at-tened about2.5:1;cortical granules large,scattered on dorsal side and lined along the kineties;one canal-like depression extending from postoral area to subcaudal region of cell;ca. 48right,four postoral,and37left kineties;12–16nematodes-mal rods;one or two terminal fragments on dorsal side. Etymology:The species-group name tropicum(Latin adjective;tropical)refers to the fact that this species was discovered in a tropical biotope(South China Sea).

Type locality:Mangrove wetland waters in Shenzhen (114?04 E,22?37 N),southern China;salinity approx.16‰, water temperature about25?C,pH ca.7.6.

X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx3 Fig.1.(A–J)Morphology and infraciliature of Paracyrtophoron tropicum nov.gen.,nov.spec.(A–D,I,J),Cyrtophoron poljanskyi(E, from Aliev1991)and Cyrtophoron apsheronica(F,from Aliev1991),Chlamydodon obliquus(G,from Gong et al.2005),and Cyrtophoron isagogicum(H,from Deroux1975).(A)Ventral view of a typical individual.(B)Ventral view showing the detailed infraciliature of anterior part,arrows indicate the postoral kineties,arrowheads mark the circumoral kineties.(C)Dorsal view showing the cortical granules of anterior dorsal part,arrow indicates the densely packed cortical granules along the terminal fragment.(D)Lateral view.(E–H)Ventral view of infraciliature of similar species.(I,J)Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen,arrow indicates the dorsal depression.LK,left kineties;PoK,postoral kineties;RK,right kineties;Su,suture;TF,terminal fragments.Scale bars=100?m(A, G,I,J);30?m(E,F,H).

Type deposition:The slide containing the protargol-impregnated holotype specimen marked with ink circle (Figs.1I,J,3A,B)is deposited in the Laboratory of Pro-tozoology,Institute of Evolution and Marine Biodiversity, OUC,China,with registration number CXR-20080408-01. One paratype slide is deposited in the collection of the Natural History Museum,London,UK,with the registration number 2010:2:7:1.

Description:Body oval and kidney-shaped in outline, mostly about150–175×70–90?m in vivo,ratio of length to width about2:1.Dorsoventrally?attened,ratio of width to thickness about2.5:1;ventral side?attened,dorsal side unevenly humped,anterior end higher than remaining body;

a conspicuous canal-like depression on dorsal side,4–5?m in width,extending from postoral area to subcaudal region of cell(Figs.1D,2A,B).In ventral view,cell outline reni-form;anterior end evenly rounded and slightly projecting to left(Fig.2A).Pellicle thick and?exible.Cortical gran-ules primrose yellow,about0.6–0.8?m across,irregularly arranged on dorsal side and lined along ventral ciliary rows. On dorsal side,mass of cortical granules,which are smaller and more yellow than others,packed in a row adjacent to terminal fragments(Figs.1C,2D,E,G).Endoplasm with mass of grayish-green,brownish-yellow and purple-red granules3–5?m in diameter(Fig.2H).Cytostome oval, conspicuous in vivo,positioned at about25%of body length (Figs.1A,2A,C).No contractile vacuole observed.Macronu-cleus juxtaposed heteromerous,rounded to oval,approx. 35?m in diameter after protargol impregnation,located in body center;micronucleus not observed owing to coverage and strong staining of macronucleus.Cilia ca.6–8?m long in vivo.Movement by gliding on substratum or swimming. 78–97densely spaced somatic kineties.Four postoral kineties commence anteriorly below cytostome,terminate at rear cell end.42–55right kineties extend anteriorly beyond the level of cytostome and bend to left;31–39left kineties extend subapically,forming a prominent suture with right kineties(Figs.1B,3A,C).Usually the outermost15–18right kineties extend onto dorsal surface;the innermost one or two are interrupted by oral kineties.Terminal fragment is contin-uous or divided into two parts,adjacent to the outermost right kinety on the dorsal side(Figs.1J,3E).

Three circumoral kineties parallel to each other,closely arranged and obliquely oriented(Figs.1B,3F).Cyrtos

4X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx

Fig.2.(A–H)Microphotographs of Paracyrtophoron tropicum nov.gen.,nov.spec.from life.(A–C)Dorsal(A),lateral(B)and ventral(C) side of typical individuals,arrows in(A)mark the dorsal depression,arrow in(C)indicates the cytostome.(D)Dorsal view,showing the yellow-belt,combined by mass of small cortical granules.(E)Dorsal view of anterior body portion,showing the cortical granules lined along ciliary rows.(F)Cytostome in ventral view,arrows indicate the nematodesmal rods.(G)Showing the cortical granules on ventral side lined along ciliary rows.(H)Dorsal view of a pressed individual,showing the colourful endoplasm.Ma,macronucleus.Scale bars=100?m.(For interpretation of the references to color in this?gure legend,the reader is referred to the web version of the article.)

relatively short,extends posteriorly and rightward,com-posed of12–16nematodesmal rods(Figs.1A,2C,F); one circle of kinetosome-like dots always observed around cyrtos in protargol impregnated specimens(Fig.3G,arrow-heads).

Comparison with related species:In terms of the dorsoven-trally?attened body and oral apparatus,Paracyrtophoron tropicum nov.spec.is very similar to members of the genus Cyrtophoron Deroux,1975(Aliev1991;Deroux1975). However,compared with the latters,our organism can be dis-tinguished by the lack of transpodial segments in the posterior portion of the ventral surface and a fragment near anterior ends of right kineties,and possession of large body size (150–175?m vs.ca.55–100?m)and more somatic kineties (78–97vs.23–32)(Fig.1E,F,H).

With reference to body shape and size and the arrangement of somatic and oral kineties,this new species also resembles Chlamydodon obliquus Kahl,1931.The former,however, differs from the latter in the presence(vs.absence)of cortical granules,the number of somatic kineties(78–97vs.66–72), and the lack of cross-striped band and contractile vacuole (Fig.1G;Gong et al.2005;Kahl1931).Family Hartmannulidae Poche,1913

Aegyria Claparède and Lachmann,1859

Improved diagnosis:Body dorsoventrally?attened.Oral ciliature consisting of one preoral and several circumoral kineties,which are positioned anterior left and right of the cytostome respectively.Podite located in posterior ventral region and surrounded by somatic kineties.No obvious gap between right and left somatic kineties.Postoral and left somatic kineties progressively shortened posteriorly from right to left.

Type species:(by subsequent designation in present paper): Aegyria oliva Claparède and Lachmann,1859 Remarks:Claparède and Lachmann(1859)did not give a clear de?nition.Based on previous and present studies,we supply a new diagnosis(as above).For details on complicated nomenclature,see below.

Considering the general living morphology and infracilia-ture,Aegyria is very similar to Hartmannula Poche,1913and Brooklynella Lom and Nigrelli,1970(Deroux1975;Gong et al.2005).However,it differs from the latters in having preoral kinety and circumoral kineties widely separated(vs.

X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx5 Fig.3.(A–G)Microphotographs of Paracyrtophoron tropicum nov.gen.,nov.spec.after protargol impregnation.(A,B)Ventral and dorsal view of infraciliature of holotype specimen;arrowheads indicate the suture,arrow marks the circumoral kineties.(C)Detail of infraciliature of ventral anterior part,arrow indicates the suture,arrows mark basal body-like granules,double-arrowheads denote the cytostome.(D)Macronucleus, arrow marks the ortomere area.(E)Anterior portion of dorsal infraciliature,arrow marks the terminal fragment.(F)Infraciliature of oral?eld, arrows indicate the circumoral kineties.(G)Focusing on the surface of cytostome,one circle of kinetosome-like dots always observed around cytos in protargol-impregnated specimens(arrowheads).CK,circumoral kineties;Ma,macronucleus.Scale bars=100?m.

obliquely arranged)and podite surrounded(vs.near,but not surrounded)by posterior somatic kineties.

Aegyria rostellum nov.spec.(Figs.4A–F,5,Table1) Diagnosis:Marine Aegyria about90–150×40–70?m in vivo;triangular or ear-shaped body with broad ante-rior end,having a rostriform structure and pigment spots; 56–65somatic kineties including34–37left and22–27 right kineties;one preoral and three or four circumoral kineties,four or?ve transpodial segments;cyrtos promi-nent,consisting of ca.32–42nematodesmal rods;single oval macronucleus.

Etymology:The Latin noun rostellum(diminutive of ros-trum)alludes to the prominent rostriform structure in the left side of the anterior end.

Type locality:Mangrove area near Shenzhen(114?04 E, 22?37 N),southern China;salinity approx.16‰,water tem-perature approx.25?C,approx.pH7.6.

Type deposition:The slide containing the protargol-impregnated holotype specimen(Figs.4D,E,5C,E)is deposited in the Laboratory of Protozoology,Institute of Evolution and Marine Biodiversity,OUC,China,with reg-istration number CXR-20080408-04.One paratype slide is deposited in the collection of the Natural History Museum, London,UK,with the registration number2010:2:7:2. Description:Cell in vivo mostly about 100–130×50–60?m.Body shape variable,but generally triangular or ear-shaped,right margin convex,left slightly curved near the anterior end;a conspicuous rostriform projection on anterior left,packed with many dark-green pigments underneath forming spots(Figs.4A,5A,B). Dorsoventrally rather thick,about 1.7–2.0:1?attened, ventral side?at,dorsal side strongly vaulted on anterior and middle parts.Cytoplasm colourless or grayish,containing lots of greasily shining globules(5–6?m in diameter),which rends cells highly opaque or even dark gray(Figs.4A,5B). Contractile vacuole not observed.Macronucleus ellipsoidal,

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Table1.Morphometric characteristics of Paracyrtophoron tropicum nov.gen.,nov.spec.(?rst line)and Aegyria rostellum nov.spec.(second line).

Characters a Min Max Mean SD SE CV n Body length(?m)128171148.312.6 2.88.520

81132108.515.5 4.314.313 Body width(?m)6510377.28.1 1.810.520

345544.0 6.4 1.814.513 No.of somatic kineties789788.4 4.9 1.1 5.520

566359.5 2.00.6 3.413 No.of right kineties425547.8 2.90.6 6.120

222723.9 1.00.3 4.213 No.of left kineties313936.5 2.30.5 6.320

343735.30.90.2 2.513 No.of postoral kineties44 4.000020

–––––––No.of circumoral kineties33 3.000020

34 3.70.50.113.513 No.of transpodial segments–––––––

45 4.90.30.1 6.112 Length of nematodesmal rods(?m)–––––––

304036.8 3.10.98.413 Length of opening of the cyrtos(?m)–––––––

182521.7 2.30.610.613 No.of nematodesmal rods121614.1 1.10.27.820

324236.5 3.6 1.09.913 Macronuclear length(?m)304536.4 4.5 1.012.420

255037.9 6.8 1.917.913 Macronuclear width(?m)224030.4 5.1 1.116.820

224031.5 6.0 1.719.013 Abbreviations:CV,coef?cient of variation in%;Max,maximum;Mean,arithmetic mean;Min,minimum;n,number of specimens investigated;No.,number; SD,standard deviation;SE,standard error of arithmetic mean.

a Data based on protargol-impregnated specimens.

juxtaposed heteromerous,about40×30?m after protargol impregnation;micronuclei invisible in prepared speci-mens possibly owing to strong staining and coverage of macronucleus.Podite inconspicuous,8–10?m long in vivo, subcaudally positioned(Fig.4A).Locomotion typically very slow.When disturbed,cells stick?rmly to the substrate. Ciliation basically on ventral side with densely spaced cilia.Cilia about8?m in vivo.Infraciliature as shown in Figs.4D,E,5C,E–G.About56–63somatic kineties,includ-ing22–27right and34–37left kineties.Right kineties extend anteriorly beyond level of cytostome,approximately17right-most rows extend onto dorsal surface;the innermost three to ?ve rows end at circumoral kineties;about10right kineties extending posteriorly to vicinity of podite,four or?ve of which interrupted by podite(Figs.4D,5C,G).Left kineties commence at level of oral area,ca.10outermost rows form a suture with right kineties;posterior end of left kineties progressively shortened from right to left.About?ve to six kineties at posterior end“interrupted”by podite and hence forming a bald area,leaving?ve transpodial segments (Fig.4D).Terminal fragment composed of more than30 basal bodies,positioned in left portion of frontal end of body (Figs.4E,5F).

Oral kineties composed of three or four circumoral kineties and one preoral kinety.Circumoral kineties almost in paral-lel with right kineties,positioned right ahead of cytostome; inmost circumoral kinety slightly shorter than the rest;pre-oral kinety arranged in direction of suture and left ahead of cytostome(Figs.4D,5E).Buccal basket genus-typical and considerably wide,cyrtos conspicuous and directed leftwards,composed of32–42nematodesmal rods,about 30–40?m in length(Figs.4B,5H).

Nomenclature and comparison with congeners:Claparède and Lachmann(1859)established Aegyria with four included species,but he did not?x its type.Because of this, Diesing(1866)established Glenotrochilia with Aegyria oliva Claparède and Lachmann,1859as type species by mono-typy.But for the same species,Deroux(1975)established Aegyriana,a fact obviously overlooked by Song and Wilbert (2002)and Song et al.(2009).Thus,both Aegyriana Deroux, 1975and Aegyriana Deroux in Song and Wilbert,2002 are junior objective synonyms of Glenotrochilia.Jankowski (2007)accepted Glenotrochilia but erected Anemadesma for Aegyriana minuta Deroux,1975.We do not agree with him as these two genera lack critical features for distinction. The original report and subsequent redescriptions showed

X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx7 Fig.4.(A–G)Morphology and infraciliature of Aegyria rostellum nov.spec.(A–F)and two similar species(G–J).(A)Ventral view of a typical individual,arrow indicates rostriform projection,packed with dark-green granules.(B)Cyrtos,to show the nematodesmal rods.(C) Left lateral view.(D,E)Infraciliature of ventral and dorsal side and nuclear apparatus of holotype specimen,arrow in(D)indicates the transpodial segments,the arrows in(E)mark a detail of the heteromerous macronucleus.(F)Dorsal view of a living cell.(G,H)Aegyria oliva (from Song et al.2009,after permission).(I,J)Aegyria paroliva(from Song and Wilbert2002,after permission).CK,circumoral kineties; FvK,frontoventral kineties;Ma,macronucleus;P,podite;Pr,preoral kinety;S,suture;TF,terminal fragment.Scale bars=100?m.

that Aegyria oliva has different body shape and infracilia-ture from many well-de?ned Dysteria species(Claparède and Lachmann1859;Deroux and Dragesco1968;Deroux1975, 1994);therefore Aegyria is no longer a synonym of Dysteria Huxley,1857despite a few species of Aegyria reported by Claparède and Lachmann(1859)have been transferred into the latter(Aescht2001;Bütschli1889;Corliss1979;Gong et al.2002,2003;Gourret and Roeser1886;Kahl1931;Song et al.2009).Thus,Aegyria becomes valid and Glenotricha is a junior objective synonym of Aegyria.Moreover,accord-ing to Art.12of the ICZN(1999),Aegyria Claparède and Lachmann,1859is still available.Since no one has?xed a type species for Aegyria so far we?x here Aegyria oliva as the type.According to Aescht(2001),Fromentel(1875,p.171) has?xed Aegyria oliva as type of Aegyria.However,this statement by Aescht(2001)is obviously incorrect.Based on the above de?nition,only three species can be assigned to this genus:Aegyria oliva Claparède and Lachmann,1859(type species),A.minuta(Deroux1975)https://www.wendangku.net/doc/651076355.html,b.(basionym Aegyriana minuta Deroux1975)and A.paroliva(Song and Wilbert2002)https://www.wendangku.net/doc/651076355.html,b.(basionym Aegyriana paroliva Song and Wilbert2002).Aegyria minuta is small(ca.55?m long)and has fewer somatic kineties(ca.18vs.56–63in A. rostellum),and hence can be clearly separated from this new species(Deroux1975).

Aegyria rostellum resembles the type species A.oliva in terms of its body length(90–150?m vs.40–120?m)and marine habitat.However,the former can be distinguished from the latter in the following features:body shape(tri-angular or ear-shaped vs.oval or elliptical);the rostriform projection on anterior left side(present vs.absent);the num-ber of somatic kineties(56–63vs.35–44),nematodesmal

8X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx

Fig.5.(A–H)Microphotographs of Aegyria rostellum nov.spec.from life(A,B)and after protargol impregnation(C–H).(A)Ventral view of a typical individual,arrows indicate the belt-shaped depression.(B)Dorsal view to show the prominent rostriform projection packed with mass of dark-green granules.(C,D)Ventral(C)and dorsal(D)view of infraciliature of holotype specimen,arrow in(C)marks the bald area which is due to the podite.(E)Detailed ventral view of anterior part,arrow indicates the circumoral kineties,arrowhead marks the preoral kinety.(F)Detailed dorsal view of anterior part,arrow marks the terminal fragment.(G)Detailed ventral view of posterior part,arrowheads mark the transpodial segments.(H)Buccal basket,arrows indicate the cyrtos.Ma,macronucleus.Scale bars=100?m.(For interpretation of the references to color in this?gure legend,the reader is referred to the web version of the article.)

rods(32–42vs.20–30),and contractile vacuoles(absent vs. 4–6)(Fig.4G,H;Deroux and Dragesco1968;Song et al. 2009).

Aegyria paroliva is similar to A.rostellum in terms of basic pattern of infraciliature and habitat.However,it can be clearly separated from the new species by body shape (elliptical or D-shaped vs.triangular or ear-shaped),body size(60–80×30–50?m vs.90–150×40–70?m);number of somatic kineties(33–40vs.56–63)(Fig.4I,J;Song and Wilbert2002).

Acknowledgements

This work was supported by the Natural Science Foun-dation of China(Project number:41006086,40976075),the Center of Biodiversity Research,King Saud Univer-sity,Saudi Arabia,Ningbo Natural Science Foundation (No.2011A610014),Scienti?c Research Fund of Zhejiang Provincial Education Department(No.Y201017789),the Discipline Project of Ningbo University(No.xkc11006),the Key Laboratory of Mariculture of Ministry of Education, Ocean University of China(No.2009002)and K.C.Wong Magna Fund in Ningbo University.We greatly appreciate the comments of the two anonymous reviewers and Dr.Helmut Berger who helped to improve our manuscript signi?cantly. References

Aescht,E.,2001.Catalogue of the generic names of ciliates(Pro-tozoa Ciliophora).Denisia1,1–350.

X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx9

Aliev, A.R.,1991.New infusorian species from the genus Cyrtophoron(Ciliata Hypostomata)from hyperhaline and min-eralized water bodies of Azerbaijan.Zool.Zh.70,126–130(in Russian).

Baldock,B.,Baker,J.,Sleigh,M.A.,1983.Abundance and produc-tivity of protozoa in chalk streams.Holarctic Ecol.6,238–246. Borror,A.C.,1972.Tidal marsh ciliates(Protozoa):morphology, ecology,systematics.Acta Protozool.10,29–71.

Bütschli,O.,1887–1889.Erster Band Protozoa.III.Abtheilung.

Infusoria und System der Radiolaria.In:Bronn,H.G.(Ed.), Klassen und Ordnungen des Thier-Reichs,vol.1,Winter, Leipzig,pp.1585–2035.

Cairns Jr.,J.,Yongue Jr.,W.H.,1968.The distribution of freshwater protozoa on a relatively homogenous substrate.Hydrobiologia 31,65–72.

Claparède,é.,Lachmann,J.,1859.études sur les infusoires et les rhizopodes.Mém.Inst.natn.genèv.6,261–482.

Corliss,J.O.,1979.The Ciliated Protoza.Characterization Classi-?cation and Guide to the Literature,2nd ed.Pergamon Press, Oxford.

Deroux,G.,https://www.wendangku.net/doc/651076355.html, série“Chlamydonellienne”chez les Chlamydodontidae(holotriches,Cyrtophorina Fauré-Fremiet).

Protistologica6,155–182.

Deroux,G.,1975.Les dispositifs adhésifs ciliaires chez les Cyr-tophorida et la famille des Hypocomidae.Protistologica10, 379–396.

Deroux,G.,1976a.Le plan cortical des Cyrtophorida unitéd’expression et marges de variabilitéI.Le cas des Plesiotrichop-idae,fam.nov.,dans la nouvelle systématique.Protistologica12, 469–481.

Deroux,G.,1976b.Le plan cortical des Cyrtophorida unitéd’expression et marges de variabilitéII.Cyrtophorida a thig-motactisme ventral généralise.Protistologica12,483–500. Deroux,G.,1976c.Plan cortical des Cyrtophorida III–Les structures différenciatrices chez les Dysteriina.Protistologica12,505–538. Deroux,G.,1994.Sous-classe des Cyrtophoria Fauré-Fremiet in Corliss,1956.Traitéde Zoologie2/2,401–431.

Deroux,G.,Dragesco,J.,1968.Nouvelles données sur quelques Cil-iés Holotriches Cyrtophores a Ciliature Ventrale.Protistologica 4,365–404.

Diesing,K.M.,1866.Revision der Prothelminthen.Abtheilung: Amastigen.I.Amastigen ohne Peristom.Sber.Akad.Wiss.

Wien,Math.-naturwiss.Cl.,I.Abt.52,505–579. Dragesco,J.,Dragesco-Kernéis,A.,1986.Ciliés libres de l’Afrique intertropicale.Faune Trop.26,1–559.

Ehrenberg,C.G.,1835.Zus?tze zur Erkenntniss grosser organis-cher Ausbildung in den kleinsten thierischen Organismen.Abh.

Akad.Wiss.Berlin,151–180.

Fauré-Fremiet,E.,1950.Mécanismes de la morphogénèse chez quelques ciliés gymnostomes hypostomiens.Arch.Anat.

Microsc.39,1–14.

Fromentel,E.,de1875.études sur les Microzoaires ou Infu-soires Proprement https://www.wendangku.net/doc/651076355.html,prenant de Nouvelles Recherches sur Leur Organisation,Leur Classi?cation,et la Description des Espèces Nouvelles ou Peu Connues,G.Masson,Paris, pp.89–192.

Gismervik,I.,2004.Podite carrying ciliates dominate the benthic ciliate community in the kelp forest.Aquat.Microb.Ecol.36, 305–310.

Gong,J.,Song,W.,2003.Morphology and infraciliature of two marine benthic ciliates,Dysteria procera Kahl,1931and

Dysteria magna nov.spec.(Protozoa Ciliophora,Cyrtophorida), from China.Eur.J.Protistol.39,301–310.

Gong,J.,Song,W.,2004.Morphology and infraciliature of two marine species of Hartmannula(Protozoa,Ciliophora,Cyr-tophorida),from scallop farming waters off Qingdao(Tsingtao), China.J.Nat.Hist.38,1327–1337.

Gong,J.,Song,W.,2006.Redescriptions of three cyrtophorid cil-iates from marine bio?lm,with establishment of a new genus, Wilbertella nov.gen(Ciliophora:Crytophorida:Lynchellidae).

Acta Protozool.45,153–165.

Gong,J.,Song,W.,Warren,A.,2002.Redescriptions of two marine cyrtophorid ciliates,Dysteria cristata(Gourret and Roeser,1888) and Dysteria monostyla(Ehrenberg1838)Kahl,1931(Proto-zoa,Ciliophora,Cyrtophorida),from China.Eur.J.Protistol.

38,213–222.

Gong,J.,Lin,X.,Song,W.,2003.Redescription of a poorly-known marine cyrtophorid ciliate,Dysteria pusilla(Claparède et Lachmann1859)(Protozoa:Ciliophora:Cyrtophorida)from Qingdao,China.Acta Protozool.42,215–221.

Gong,J.,Song,W.,Warren,A.,2005.Updating the ciliate genus Chlamydodon Ehrenberg1835,with redescriptions of three species(Ciliophora,Cyrtophorida).Acta Protozool.44,19–32. Gong,J.,Gao,S.,Roberts,D.McL.,Al-Rasheid,K.A.S.,Song,W., 2008.Morphological description of Trichopodiella faurei n.sp.

(Ciliophoria,Phyllopharyngea,Cyrtophoria)and phylogenetic analyses based on SSU rRNA and group I intron sequences.J.

Eukaryot.Microbiol.55,492–500.

Gourret,P.,Roeser,P.,1888.Contributionàl’étude des protozoaires de la Corse.Archs Biol.8,139–204.

Hu,X.,Suzuki,T.,2005.Light microscopical observations on two marine dysteriid ciliates from Japan,including a description of Dysteria yagiui nov.spec(Ciliophora,Cyrtophorida).Eur.J.

Protistol.41,29–36.

ICZN[The International Commission on Zoological Nomencla-ture],1999.International Code of Zoological Nomenclature,4th ed.International Trust for Zoological Nomenclature,London (Adopted by the International Union of Biological Sciences). Jankowski,A.W.,1980.Conspectus of a new system of the phylum Ciliophora.Trudy Zool.Inst.Leningr.94,103–121(in Russian). Jankowski,A.W.,2007.Ciliophora Do?ein,1901.In:Alimov,A.F.

(Ed.),Protista.Part II.Nauka.St.Petersburg,pp.371–993(in Russian).

Kahl,A.,1931.Urtiere oder Protozoa I:Wimpertiere oder Cili-ata(Infusoria)2Holotricha ausser den im1.Teil behandelten Prostomata.Tierwelt Dtl.21,181–398.

Kurth,T.,Bardele,C.F.,2001.Fine structure of the cyrtophorid cili-ate Chlamydodon mnemosyne Ehrenberg,1837.Acta Protozool.

40,33–47.

Li,L.,Song,W.,2006.Phylogenetic positions of two cyrtophorid ciliates Dysteria procera and Hartmannula derouxi(Ciliophora: Phyllopharyngea:Dysteriida)inferred from the complete small subunit ribosomal RNA gene sequences.Acta Protozool.45, 265–270.

Li,L.,Huang,J.,Song,W.,Shin,M.K.,Al-Rasheid,K.A.S.,Berger,

H.,2010.Apogastrostyla rigescens(Kahl,1932)gen.nov.,comb.

nov.(Ciliophora Hypotricha):morphology,notes on cell divi-sion,SSU rRNA gene sequence data,and neotypi?cation.Acta Protozool.49,195–212.

Lynn,D.H.,2008.The Ciliated Protozoa.Characterization,Clas-si?cation and Guide to the Literature,3rd ed.Springer Press, Dordrecht.

10X.Chen et al./European Journal of Protistology xxx(2011)xxx–xxx

Pan,H.,Hu,X.,Gong,J.,Lin,X.,Al-Rasheid,K.A.S.,Al-Farraj, S.A.,Warren,A.,2011.Morphological redescriptions of four marine ciliates(Ciliophora:Cyrtophorida:Dysteriidae)from Qingdao,China.Eur.J.Protistol.47,197–207.

Petz,W.,Song,W.,Wilbert,N.,1995.Taxonomy and ecology of the ciliate fauna(Protoza,Ciliophora)in the endopagial and pelagial of the Weddell Sea,Antarctica.Stap?a40,1–233.

Shao,C.,Song,W.,Gong,J.,2008.Morphology and morphogenesis of a new marine cyrtophorid ciliate,Hartmannula sinica nov.

spec.(Protozoa,Ciliophora Cyrtophorida)from China.Eur.J.

Protistol.44,1–12.

Song,W.,Packroff,G.,1997.Taxonomische Untersuchungen an marinen Ciliaten aus China mit Beschreibungen von zwei

neuen Arten,Strombidium globosaneum nov.spec.und S.pla-tum nov.spec.(Protozoa Ciliophora).Arch.Protistenkd.147, 331–360.

Song,W.,Wilbert,N.,1995.Benthische Ciliaten des Süsswassers.

In:R?ttger,R.(Ed.),Praktikum der Protozoologie.Gustav Fis-cher,Stuttgart,pp.156–168.

Song,W.,Wilbert,N.,2002.Faunistic studies on marine ciliates from the Antarctic benthic area,including descriptions of one epizoic form,6new species and,2new genera(Protozoa:Cilio-phora).Acta Protozool.41,23–61.

Song,W.,Warren,A.,Hu,X.,2009.Free-living Ciliates in the Bohai and Yellow Seas China.Science Press,Beijing.