Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 pbot a on e hipseie.Tedsrbto fteeiin pce ntesraeof of individuals surface colonized the different on the on species density epibiont the the of ensifera calculation of allow distribution C. to The recorded, was specimen. basibiont shrimp the per found was epibionts ( haptorid nlsssoe that showed analysis intermedium ahfntoa ru,btwt rn eutn neulbimaogseisadgroups, the and as species well among as epibionts, equilibrium the in of resulting adaptations possible trend the The discussed. a density. are throughout patterns and with of epibionts colonization requirements diversity but body, particular the for certain the with a group, substratum, of of compensating follows available end colonization functional the densities posterior Their occupying the species. the species each to different epibiont the anterior each the of behaviour, the of of from of genera distribution analysis pattern gradient different patterns a The significantly epibiont showed distribution a shrimp shrimp. and the of the of the between axis pairs difference on longitudinal significant the species a and epibiont was basibiont, There distribution. the on Podophrya distributed widely polypoides hrceitc n aooi oiin ee iitdpooon n oie eeeaie:three ( examined: were species biometrical rotifer and a protozoan morphological and their protozoans ciliate on ciliated based Seven suctorian time position. first taxonomic the and for characteristics analysed were Indonesia), (Sulawesi, History Natural of Journal O:10.1080/00222930701787871 online University, DOI: 1464-5262 Complutense print/ISSN 9, 0022-2933 Pnta 2000b). ISSN Biology, [email protected] 2000a, of Email: Faculty Tato-Porto Spain. Zoology, Madrid, of and 28040 Department Fernandez-Leborans Fernandez-Leborans, 1995; (Dawson Gregorio 1999; Small Crustacea Correspondence: and al. in Morado et 1993; epibiosis Key al. to et 1998; dedicated Gili al. 1992; were et Uriz reports and Dick Maldonado of 1962; number Eldred a 1957; years, recent In Introduction Keywords: shrimp freshwater the of communities epibiont The Abstract 1 FERNANDEZ-LEBORANS GREGORIO Indonesia) (Sulawesi, Poso Lake shrimp ensifera freshwater Caridina the on communities Epibiontic ( fu cetd5Nvme 2007) November 5 Accepted rNtrud,Humboldt-Universita Naturkunde, ¨r eateto olg,Fclyo ilg,CmltneUiest,Mdi,San and Spain, Madrid, University, Complutense Biology, of Faculty Zoology, of Department ,treprtih ( peritrichs three ), – mhlpu fusidens Amphileptus Acineta n nec ntmclui ftesrm.Tems bnatspecies, abundant most The shrimp. the of unit anatomical each on and and aiiaesfr,claepooon,eiiss aePs,rtfr Sulawesi rotifer, Poso, Lake epibiosis, protozoans, ciliate ensifera, Caridina cnt sulawesiensis Acineta , Spelaeophrya Zoothamnium 07 14–8:2891–2917 41(45–48): 2007; , otanu intermedium Zoothamnium ,adtertfrspecies rotifer the and ), Cutca eaoa tia)from Atyidae) Decapoda, (Crustacea, – Amphileptus , Acineta tBri,Bri,Germany Berlin, Berlin, ¨t eeas h nsms ieydsrbtd h statistical The distributed. widely most ones the also were , # cnt sulawesiensis Acineta 07Tyo Francis & Taylor 2007 , and , Podophrya Cothurnia 1 , and , otclaglobosa Vorticella RSIAVNRINTELEN VON KRISTINA & maalaticeps Embata aiiaensifera Caridina , – Embata oohy maupasi Podophrya Vorticella eeteeiin eeamost genera epibiont the were olwdasmlrptenof pattern similar a followed and , ennme f314.6 of number mean A . nei oLk Poso Lake to endemic , ohri compressa Cothurnia Zoothamnium and , Zoothamnium Spelaeophrya 2 – Museum Embata ,a ), 2 , Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 2892 itln(uemo aua itr,Bri,Gray rmtesuhsoeo Lake of shore south the from Germany) Berlin, History, Natural of (Museum Rintelen epibiosis by explained be can Abello taxa 1988; crustacean Ropes basibiont their and of (Bottom ecology biodiversity. and and fields: biology conservation the biological and to many related to fields taxa, relevant those described a are been newly and in epibionts, relationships of evolution considered has epibiotic increase of ecology, been that noticeable physiology, crustaceans association presence recognition a this the only on The has not also involved recently 1997). but etc., has only al. This symbionts although perspective. time, et a new epizoics, long constitutes Wahl a commensals, and are for 1989; hosts cycle, reported as basibiont (Wahl life the known their epibiont while The of substratum, previously the basibiont. phase living the for a sessile and of the support surface epibiont during the the that, to organisms: organisms attached two includes between ‘‘epibiont’’ association term an is Epibiosis h i fti okwst otiuet h ecito n xlnto fthe of of Specimens explanation methods and and description Material the the on communities to epibiont contribute Poso. protozoan Lake the to from of species was distribution shrimp shrimp. of work endemic the of patterns this surface and the on of relationships distribution their aim to as The well as characteristics taxonomical and the for and 2004), (Fernandez-Leborans the Europe Sulawesi, about Northern of on information genus lakes lakes for complementary the case shrimp the ancient in is give the This 1862 the environments. increase and special Loven, may or these of patterns, that biodiversity Europe lakes and evolutionary conservation independent Northern from of of data knowledge comparable lakes provide the could Small Indonesia, lakes, and African Lynn East 1979; (Corliss heterotrichs apostomatids, and groups: following 2000). peritrichs, Nematoda, the of suctorians, Rotifera, representative invertebrate are chonotrichids, Ascidiacea, epibionts Nemertea, and Bryozoa, Protozoan Phoronida, protozoan others). Platyhelminthes, Bivalvia, and Gastropoda, for of Cnidaria, Decapoda, number basibionts a Cirripedia, (Porifera, to decapods, Polychaeta, belong as that and phyla forms found include amphipods, epibionts different been invertebrate isopods, The have 1983). cirripeds, (Ross epibionts which copepods, species cladocerans, include as such groups, aeadotncletdb oa ihre K o itln esnlfedobservation). species field personal the riverine Rintelen, in one von abundant and (K. is fishermen It lacustrine local 1). (Figure three by 1997) collected currently Chace often 1902; of and (Schenkel lake flock Poso species Lake from endemic The described faunas. an not respective of are their systems part for lake The 2004, conditions 2006). shrimp, environmental Glaubrecht al. freshwater similar and et Herder provide Rintelen shrimps molluscs 2004; but 2004; Soeroto 2006), example connected, and al. Cai (Parenti et for and fishes Rintelen and Zitzler flocks, 2006) 2003; 1997; of species Glaubrecht Chace hotspots former and 1937b; are endemic (the Poso, (Korniushin 1937a, several Lake Woltereck Sulawesi and harbour 1902; of system (Schenkel lake and island Malili biodiversity Indonesian the systems, the aquatic lake On ancient two 2006a). Celebes), al. et (Fernandez-Leborans ubro rtza iit pce aebe ecie seiinso crustaceans, on epibionts as described been have species ciliate protozoan of number A h td feiin omnte nbsbot rmdfeetlk ytm,eg the e.g. systems, lake different from basibionts on communities epibiont of study The h pbotcmuiison communities epibiont The .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. aiiaensifera Caridina Caridina aiiaensifera Caridina in-dad,13 rmteMll ytmo Sulawesi of system Malili the from 1837 Milne-Edwards, eecletdb atisGabeh n hmsvon Thomas and Glaubrecht Matthias by collected were ta.19;Abello 1990; al. et ´ .ensifera C. cekl 92(rsae,Dcpd,Ayde,is Atyidae), Decapoda, (Crustacea, 1902 Schenkel, eeaaye ihrsett hi biological their to respect with analysed were n apesn19) Crustacean 1992). Macpherson and ´ yi relicta Mysis Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ntmcluiswt iia ooiain;()vrac opnns(vlainof (evaluation components of variance clusters (4) obtain colonization); (to similar analysis conglomerate with species hierarchical epibiont units of (3) groups anatomical (for distribution); analysis (for the right similar component of analysis principal with colonization determine (2) comparison in units); multiple differences to anatomical and (1) left and species, used following: and epibiont Statgraphics the of were the distribution were using between images used differences performed tests the were The programs. analyses from SPSS Statistical features Zeiss) schemes. (KS300 morphological species Analysis epibiont Image diverse using ocular the obtained an were using and personal calculated images with were Australia; microscope solution epibionts Light Adelaide, glycerol/water the micrometer. of of 10% Measurements (University in hypochlorite. Shiel placed sodium analysed 2.5% were J. specimens, trophi R. isolated the treating by rotifers, the communication), were indicated identify and slides procedure to Permanent isolated order the red. In neutral were using ciliates. Castro and stained they and green the methyl Fernandez-Leborans epibionts, from with by obtained protozoan also described and the (1986), technique as Zaldumbide identify carbonate indicated de silver to were the order species with colonial In treated of zooids. Numbers of microscopes. number light and 2006a). stereoscopic al. (software: under et Microscope with Fernandez-Leborans Electron (see stubs Scanning Sputter kV 1450VP 640 specimen 10 LEO SC7 at aluminium a Polaron V02.03) on a on 32 taken using mounted were gold–palladium critical- with 030, Pictures were coated Coater. CPD they and BAL-TEC pads Afterwards, adhesive min. a 30 standard for with ethanol dried 100% in point stereoscopic dehydrated a were ethanol laboratory, under 95% the observed In was microscopy. unit light of females. anatomical for specimens ovigerous each colonized ethanol and 95% Forty 75% in microscope. dissected fixed to were were transferred Samples 1). shrimps (Figure then 2004 and March in ethanol Indonesia, Sulawesi, Central Poso, pbot ntesraeo h hipaaoia nt eeosre n counted and observed were units anatomical shrimp the of surface the on Epibionts in fixed specimens shrimp epibionts, the of (SEM) microscopy electron scanning For iue1 egahclae ftestudy. the of area Geographical 1. Figure pbotccmuiison communities Epibiontic .ensifera C. eeaaye,o hc iewere nine which of analysed, were aiiaesfr 2893 ensifera Caridina Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ae fteMll ytm hyhv ogrsak n oe ubro etce per tentacles of number lower a and stalk, three longer other a the have in they previously observed system, been actinophore. Malili have the that of species this lakes of ciliates to comparison etr fteeclae a h oiapoogdatrol rudtetnals The tentacles. the around anteriorly prolonged species lorica the the to was belong ciliates ciliates these of feature eae.Teeiinson epibionts The females. ie ondtelrc i u-ieepnin(al ;Fgrs2,3 4). 3, 2a, Figures I; (Table expansion cup-like a via lorica the joined wide) oytikes1.6–1.615.36 31.00 17.28 17.28 9.60 15.36 13.00 17.28 15.50 26.88 9.60 7.68 5.76 6.63 5.76 – 17.28 61.44 – 19.20 26.57 13.2 15.36 49.92 15.36 78.72 5.28 3.84 1.45 3.84 17.28 61.44 3.08 12.07 0.78 7.68 26.88 7.13 1.62 7.68 19.20 28.80 1.30 21.44 14.20 19.20 6.53 10.03 2.61 6.34 9.72 14.36 2.40 43.61 11.92 10.61 49.78 35.95 11.20 39.55 lorica tentacles of of part Number anterior free of Length thickness Body thickness Lorica length Tentacles width vacuole Contractile length vacuole Contractile width Stalk length Stalk width Macronucleus length Macronucleus width Lorica length Lorica width Body length Body ot ooie pcmn of specimens colonized Forty the analyse to order Results in treated the colonization. along statistically on Epibiosis gradient was activity basibionts). longitudinal basibiont of the width the of and influence of length axis in variation anterioposterior versus species epibiont 2894 al .Boercfaue of features Biometric I. Table and suctorians tme,13,teperitrichs the 1935, Stammer, rai h pclsraeo h oywsdpesd h arnceswsruddin (19.2–26.9 rounded stalk was The macronucleus macronucleus. The the depressed. near central was located The body (7.6–15.3 micronucleus tentacles. the located capitate of centrally 13–31 and surface shape with apical each the corners, in area the on protruding actinophores h iitswr raglri uln rbl-hpd oiae,adpdnuae The pedunculate. and loricated, (28.8–78.7 bell-shaped, body or cellular outline the surrounded in completely triangular lorica were ciliates The genus the of Epibionts genus the of epibionts rotifer were there addition, h oiahdafe neirpr vrtebd neoigtetnals(13.2–15.5 tentacles the enveloping body (19.2–61.4 the over body part The anterior free long). a had lorica The hs iitsbln otegenus the to belong ciliates These Cothurnia .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. Acineta hebr,13,adtehaptorid the and 1831, Ehrenberg, hebr,1833, Ehrenberg, Acineta cnt sulawesiensis Acineta aiiaensifera Caridina m cnt sulawesiensis Acineta Zoothamnium .ensifera C. og 19.2–49.9 long, m Acineta m og 7.6–17.3 long, m Podophrya maueet in (measurements eeaaye,o hc iewr ovigerous were nine which of analysed, were Ln n ml 00.Tems distinctive most The 2000). Small and (Lynn enS iiu Maximum Minimum SD Mean hebr,1838, Ehrenberg, ee mn h rtza iits the ciliates, protozoan the among were, Embata m enne-eoase l,20.In 2006. al., et Fernandez-Leborans hebr,13,and 1833, Ehrenberg, ie a w neirlobular anterior two had wide) m Amphileptus m m; m ie.Teewsaspherical a was There wide). m rc,1910. Bryce, m n og 26.8–61.4 long, m 5 80). Vorticella hebr,13.In 1830. Ehrenberg, m og 5.8–9.6 long, m ines 1767, Linnaeus, Spelaeophrya m wide). m m m m m Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 iue2 h pbotseis (a) species. epibiont The 2. Figure otanu intermedium Zoothamnium laticeps arnces i irnces y ynm;r otu;s tl;s,soah ,tnals d rca ic;to, discs; trochal td, tentacles; t, stomach; st, stalk; vitellarium. s, rostrum; v, r, toes; myoneme; my, micronucleus; mi, macronucleus; (i) ; maalaticeps Embata h rpi d aa ic l aa oia ,coc;c,cnrcievcoe ,lrc;ma, lorica; l, vacuole; contractile cv, cloaca; c, lorica; basal bl, disc; basal bd, trophi. the , (e) ; otclaglobosa Vorticella cnt sulawesiensis Acineta pbotccmuiison communities Epibiontic (f) ; ohri compressa Cothurnia (b) ; oohy maupasi Podophrya (g) ; aiiaesfr 2895 ensifera Caridina mhlpu fusidens Amphileptus (c) ; pleprapolypoides Spelaeophrya (h) ; Embata (d) ; Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 2896 iue –0 3 4) (3, 3–10. Figures neirpr riuaeo h oy(ivrcarbonate, (silver body the carbonate, of articulate (silver part macronucleus inferior elongate the showing specimen a (5) tentacles. the with actinophores the (4) (6–9) lorica; the of prolongation aul arw (SEM, (arrow) vacuole tl n h hp ftezod sle carbonate, (silver zooids the of shape the and stalk pleprapolypoides Spelaeophrya .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. cnt sulawesiensis Acineta 6 8) 7 eea pcmn nterlcto ntebsbotsrae(SEM, surface basibiont the on location their in specimens several (7) 280); 6 niiulsoigtebslds,tetnalsadtepr ftecontractile the of pore the and tentacles the disc, basal the showing individual (6) : sle carbonate, (silver 6 6 420). 6) (10) 360). 6 00:()temcoulu n h anterior the and macronucleus the (3) 1070): otanu intermedium Zoothamnium 6 5) 9 h arncesirglradthe and irregular macronucleus the (9) 450); oohy maupasi Podophrya mty green, (methyl ooysoigthe showing colony a , 6 3) (8) 230); 6 792). Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 egh h arnceswseogtdadarne lnsd h ogtdnlai of axis longitudinal the alongside arranged and (24.9–53.7 elongated ciliate the was macronucleus The length. oywt otro oiae ra h ufc fteclaewscniuu rmthe from continuous was 6–13). ciliate 2c, Figures the III; (Table of the disk surface of basal division be The the the can area. show to which not loricated the surface of did apical pores posterior images In the SEM a macronucleus. The vacuoles, with ciliate. contractile several the the body three in of near to macronucleus surface micronuclei one the the were in spherical of distinguished there rest six body the the macronucleus to with of The four middle angle widened. were right appear a There can forming specimens. part section anterior anterior an its had and flattened, considerably was oae vrti ufc;n ciohrswr rsn.Ec etcews18–30.9 was tentacle Each present. were actinophores This no of basibiont. surface; end the of this posterior surface The over the indicated. located 14.9–32.2 to have attachment was for (1945) area disc disc Lu anterior circular and basal a the in Nie with expanded as articulate was angle, to body an the seemed forming body body, the the of of zone posterior this specimens some 19.2–25.7 aa unlsae ra salrc,wt egho 25.7–37.3 of length a by a replaced with sometimes lorica, was a which stalk, as developed area, a funnel-shaped without basal substrate the joined body h iitswr el rtuptsae,wt h oyfatndadcvrdb thick a by 157.4 covered and and 48 flattened between body fluctuated the length with body trumpet-shaped, The or pellicle. bell- were ciliates The and genus freshwater vegetation the to of spherical aquatic Epibionts attached to a and epibionts attached end, were and they their free-living study our at were in crustaceans. ciliates description whereas trumpet-shaped the the objects, In zooid. slightly inanimate (1986) the to layer Curds tentacles external thick of a lorica, body, and cellular positioned the of centrally of macronucleus form II; (Table spherical absence the habitat, macronucleus the freshwater was their ciliates dimensions, the their these common to near similar most placed species was micronucleus 5). oval 2b, Figures An excentrically. h ellrbd a netra ae.Tecptt etce 1–5 eesra over spread were (3.8–15.4 (10–15) macronucleus (15–192 rounded tentacles length The capitate body. considerable The the a layer. of reach surface external entire can an the stalk had the body cellular body, The the to comparison In tl it .613 .47.68 15.0 192.00 32.64 13.44 3.84 53.76 15.36 10.0 15.36 1.36 7.68 3.84 7.68 3.84 1.6 95.19 5.76 6.35 3.62 122.50 12.00 11.05 3.75 21.60 7.68 26.50 8.45 tentacles of Number width Stalk length Stalk width Macronucleus length Macronucleus width Body length Body h niiul a hrceitcshri oy(7.6–53.7 body spheroid characteristic a had individuals The genus the of Epibionts al I imti etrsof features Biometric II. Table h utrasbln otegenus the to belong suctorians The m ) hc nsvrlseiesjie h eto h oyvaacntito.In constriction. a via body the of rest the joined specimens several in which m), Spelaeophrya Podophrya m og 3.8–11.5 long, m oohy maupasi Podophrya m ndaee.I h pclae,teewr 22 tentacles 12–20 were there area, apical the In diameter. in m enS iiu Maximum Minimum SD Mean Podophrya pbotccmuiison communities Epibiontic m maueet in (measurements ie.I eea niiul,temacronucleus the individuals, several In wide). m oohy maupasi Podophrya Crs18;Ln n ml 02.The 2002). Small and Lynn 1986; (Curds m m; n m 5 (13.4–28.8 m m ushi 89 hyhv in have they 1889; Butschli, 80). og 7.6–32.6 long, m aiiaesfr 2897 ensifera Caridina m mxmmwidth, (maximum m m og a located was long) m m ie.The wide). m m m wide). m long). m m min Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 h oi iesos n h -hpdmcoulu VloeiadGgilo1988). Guglielmo description); and the (Valbonesi in macronucleus than studied C-shaped ciliates the the and in dimensions; lower zooid was the colony per individuals of number of of description those area), to (suprastylar similar 15). structure were 14, cup-shaped 2d, a The Figures basibiont. by IV; the zooid (Table the to the striation of attached of longitudinal was width body a it The cellular with where the stalk. colony, to the the joined of inside stalk base the located towards was stalk increased The (spasmoneme) stalk width. myoneme body contractile maximum the The of (3.8–28.8 half broad approximately short, was was disc peristomal The 3.8–23 ubro etce 55 .61 20 20 30.9 6.00 32.24 12 37.33 12 11.52 18.02 4.00 25.75 53.76 2.46 157.44 14.98 2.77 28.80 5.89 25.75 0.83 3.84 15.54 5.92 16.33 19.22 23.90 48.00 24.96 5.04 4.87 21.30 13.44 2.87 2.81 31.42 22.69 9.10 8.23 4.61 22.74 103.01 40.05 19.68 tentacles of Number tentacles of Number micronuclei of Number length Tentacle diameter disc Basal width lorica basal Maximum length lorica Basal width Macronucleus length body Macronucleus the of width Apical length Body a rme-hpdbd ihtnalsdsrbtdoe h itled hr r two are There end. distal the over distributed tentacles of with species body trumpet-shaped a had otie otatl ynm.Tebd ftezodwsoa,eogtd(28.8– elongated oval, was which zooid stalk, the ramified a of by body linked The zooids myoneme. 2–18 76.8 contractile with colonial, a were contained ciliates peritrich The genus the of Epibionts both summary, In 7). described, belong. 6, first (Figures observed show the SEM ciliates not in by the did observed pellicle which included when the be of body could posterior observation ciliate joins species external a the which and The of of bend surface. division to presence basibiont ciliate any the the the allows and with that size, body macronucleus, mechanism body a the specimens, of zone be our the types to hyaline In to seems a species. both area respect two as pseudoloricate with the with or individuals between 1945), differences micronuclei were Lu significant of there and number no (Nie or are tentacles, lorica There of a number 1988). as al. considered been et has (Matthes body the of zone in bent often a in apically into part (divided shape anterior body another the troglocaridis in and differ part species two hyaline these basal (1988) al. et Matthes to According al I.Boercfaue of features Biometric III. Table 2898 hs pbot eogt h genus the to belong epibionts These hs iitsbln otegenus the to belong ciliates These m og 23–38.4 long, m m ie.Temcoulu a peia n ipsdnx otemacronucleus. the to next disposed and spherical was micronucleus The wide). m .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. Spelaeophrya eg ftefne ihasa)adtemcoulu eogtdo wound, or (elongated macronucleus the and seam) a with funnel the of (edge .intermedium Z. Zoothamnium m .polypoides S. ie.Ti tl a eydahnu iharuddcontour. rounded a with diaphanous very was stalk This wide). m : m otanu intermedium Zoothamnium ie.Temcoulu a -hpd(3.8–30.7 C-shaped was macronucleus The wide). m .polypoides S. pleprapolypoides Spelaeophrya n h ihtmul rnhn oois(lhuhthe (although colonies branching dichotomously the in: adlk,aial hcesin thickens apically band-like, , Spelaeophrya aa,11 and 1910 Daday, Zoothamnium enS iiu Maximum Minimum SD Mean maueet in (measurements .polypoides S. pleprapolypoides Spelaeophrya Ln n ml 02.Tetrophonts The 2002). Small and (Lynn rct 95 hycicddwt the with coincided They 1935. Precht, Ln n ml 00.Teciliates The 2000). Small and (Lynn .troglocaridis S. m pclyfne-hpdin funnel-shaped apically , m; n 5 .troglocaridis S. 80). aa,11,to 1910, Daday, tme,1935. Stammer, .Tebasal The ). m long, m S. Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 (13) carbonate, (silver iue 11.(11) 11–19. Figures oysaeadtearneeto h ieis(ivrcarbonate, (silver kineties the of arrangement the and shape body niiulsoigtetohldss otu,adte mty green, (methyl toes and rostrum, discs, trochal the showing individual 1,1)tetoh ftemsa,soigteuc n h et ( teeth the and unci the showing mastax, the green, of (methyl trophi rostrum the the 19) and (18, discs trochal the with ohri compressa Cothurnia 6 0) (12) 400). otanu intermedium Zoothamnium niiulsoigtelrc sle carbonate, (silver lorica the showing individual , otclaglobosa Vorticella rao tl ofunefratcmn otesrm surface shrimp the to attachment for confluence stalk of area , pbotccmuiison communities Epibiontic pcmnwt h otatdsak(ivrcarbonate, (silver stalk contracted the with specimen a , 6 1) 1)dti ftersrm(ehlgreen, (methyl rostrum the of detail (17) 410); 6 6 7) (15–19) 770). 520). 6 0) 1)teatro rao h body the of area anterior the (16) 200); 6 2) (14) 820). aiiaesfr 2899 ensifera Caridina maalaticeps Embata mhlpu fusidens Amphileptus 1)teentire the (15) : 6 6 710). 410); the , Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 lothl ftelrc.Temcoulu a vi n oae nteatro afo the of half anterior the in located and ovoid was macronucleus The lorica. the of half almost notl ihteclua oy h poieedo h oiacnandteapical the in contained width lorica the (15.4–40.3 than the body wider of retracted generally an above, The through end from zone. connected opposite viewed middle was when The lorica the elliptical the body. was end, which cellular posterior aperture, the the In with stalk. endostyle the by aborally attached h iitswr oiae,adsakd h oiawsnro n lnae,cylindrical elongated, and narrow was (32.6–65.3 lorica The posteriorly stalked. rounded and and loricated, were ciliates The (Warren genus epibiotic the being of their Epibionts and environment; freshwater macronucleus; the the of stalk; disposition and the C-shape 1986). the of body; the length was of the ciliates shape and these size the to common similar most species ln t nielnt n a ewe w n iebns(al ;Fgrs2,16). 2e, Figures V; (Table bends five and two between was had stalk and The length peristome. width entire the the on its than elevated along shorter and and convex narrow (13.4–153.6 macronucleus. was was disc the lip elongated peristomial to peristomial The next the body. body, placed the the was of of micronucleus part anterior spherical the A On extremes. its at widening (13.4–21.1 ubro ed e tl .514 .08.00 28.80 9.60 163.20 7.68 18.00 38.40 3.80 3.00 23.04 76.80 30.72 24.96 5.76 5.76 2.00 7.46 1.46 23.04 3.84 49.50 1.89 28.80 17.28 3.84 4.85 1.11 4.69 84.00 5.54 8.00 7.47 13.72 6.72 8.00 9.27 30.52 14.21 47.15 15.36 stalk per bends of colony Number per individuals of Number width vacuole Contractile length vacuole Contractile width Stalk length Stalk width Macronucleus length Macronucleus width Body length Body h iitswr oiayadsakd h oywsgouos n oeo esovoid less or more and globulous, was body The stalked. (23–46 and contracted solitary when were ciliates The genus the of Epibionts of features Biometric IV. Table 2900 al .Boercfaue of features Biometric V. Table tl it 08 .157 19.20 153.60 10.00 34.56 17.28 5.76 46.08 21.12 13.44 2.00 5.11 17.28 5.76 23.04 35.42 13.44 10.88 1.92 4.06 43.65 4.42 5.52 5.47 3.15 27.69 12.67 35.67 17.66 stalk per bends of Number width Stalk length Stalk width Macronucleus length Macronucleus width Body length Body hs iitsbln otegenus the to belong ciliates These .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. m og 5.8–17.3 long, m Cothurnia Vorticella m og 5.8–19.2 long, m m otclaglobosa Vorticella otanu intermedium Zoothamnium og 17.3–34.6 long, m m ie n adtasesl costecne ftezooid the of center the across transversely laid and wide) m m Vorticella og 17.3–28.9 long, m maueet in (measurements enS iiu Maximum Minimum SD Mean m otclaglobosa Vorticella enS iiu Maximum Minimum SD Mean ie,cnandacnrciemyoneme contractile a contained wide), m m Wre 96 ynadSal20) The 2000). Small and Lynn 1986; (Warren ie.Temcoulu a C-shaped was macronucleus The wide). m maueet in (measurements m og 13.4–21.1 long, m m m; m n 5 ie.Teidvdaswere individuals The wide). m 80). hs,12.Te aein have They 1922. Ghosh, m m; n 5 80). m ie occupied wide) m Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 egho h oy h one adled n h ubro ih n etsomatic left and right of number the 2005). al. and et end, Lin caudal 1989; Wilbert rounded and the (Song body, kineties the of was length ciliates these to similar 2rgtsmtcknte n n ofu etsmtcknte.Acnrcievcoewas vacuole the over contractile located A extrusomes numerous kineties. were somatic There 2.9–8.1 body. left each the four of surface, end to posterior one the and at located kineties somatic right 12 15.4 ubro etknte .310 .04.00 8.16 14.00 49.92 11.52 1.00 2.86 15.36 103.68 17.28 10.00 24.96 1.06 19.20 2.63 5.76 1.77 42.24 5.76 2.63 15.36 5.51 8.34 13.44 11.5 2.48 16.52 4.54 1.11 31.49 5.20 8.64 71.42 9.12 16.32 17.28 kineties left of Number kineties right of Number length Extrusome width macronucleus Second length macronucleus Second width macronucleus First length macronucleus First width Body length Body tl it .809 .87.68 3.84 13.44 19.2–49.9 15.36 28.80 65.28 21.12 5.28 40.32 in higher 1.46 are were lorica 7.68 the there of However, width 1991). and 5.76 length short lorica Paynter the was 17.28 and The dimensions: endostyle compressa the (Warren the above. to absent short, respect from 32.64 was was 0.92 in stalk 13.44 viewed differences mesostyle external when and The broad, 0.86 aperture, clefts. of 15.36 deep and 2.15 lorica those two had elliptical to border the similar aperture 3.78 7.68 3.08 in were 1858 found Lachmann, 3.84 9.49 ciliates 2.42 10.82 The 2000). reduced 7.48 12.80 considerably 23.90 specimens several in and 50.88 short 17.58 was (1.5–3.9 stalk The macronucleus. 28.80 the near width Macronucleus length Macronucleus width Stalk length Stalk width Lorica length Lorica width Body length Body of features Biometric VI. Table al I.Boercfaue of features Biometric VII. Table (5.8–15.4 body h iitshdalnelt-hpdbd,ltrlycmrse (42.2–103.7 compressed laterally body, lanceolate-shaped a had ciliates The genus the of Epibionts environments. marine hs iitsbln otegenus the to belong ciliates These hs iitsbln otegenus the to belong ciliates These m ie.Teewsashrclmcoulu oae ewe h arnce,10– macronuclei, the between located micronucleus spherical a was There wide). m m si h ieo h otatdzod naddition, In zooid. contracted the of size the is as og TbeV;Fgrs2,17). 2f, Figures VI; (Table long) m m ie.Teewr w vlmcouli(ah13.4–25 (each macronuclei oval two were There wide). m m og 7.7–13.4 long, m Amphileptus m og(al I;Fgrs2,18). 2g, Figures VII; (Table long m ohri compressa Cothurnia mhlpu fusidens Amphileptus mhlpu fusidens Amphileptus m ie.Temcoulu a peia n situated and spherical was micronucleus The wide). m enS iiu Maximum Minimum SD Mean Cothurnia Amphileptus pbotccmuiison communities Epibiontic enS iiu Maximum Minimum SD Mean maueet in (measurements maueet in (measurements Wre n ane 91 ynadSmall and Lynn 1991; Paynter and (Warren Ln n ml 00.Teseismost species The 2000). Small and (Lynn Kh,12) hyhv ncmo the common in have They 1926). (Kahl, ohri compressa Cothurnia m m m; m; .compressa C. n n 5 5 aiiaesfr 2901 ensifera Caridina 80). 80). a enfudin found been has m Clapare og 5.8– long, m m long, m and `de C. Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ubr feiins al Iicue h ubr fpooonseiso the different on the on species species protozoan epibiont each of of numbers mean numbers highest The the of the shrimp. units includes with the anatomical XI units of Table units the epibionts. anatomical were uropods of and numbers maxillipeds, antennae, Antennulae, te iit pce cone o .3 ftema pbotdniyo h shrimps the on density epibiont mean the of 5.33% for X). accounted (Table species ciliate ( density other epibiont mean the of 94.2% intermedium nteaaoia nt ftesrm hwd ntepo ftetofrtprincipal first two the of plot the in showed, shrimp the species of epibiont the units of anatomical numbers mean the the using on performed Analysis Component Principal the with species the epibionts, ciliate of the the of 0.45% of Among proportion Only number IX). highest were (Table 314.6). The density the epibionts highest (mean representing females. of protists 1114 ovigerous ciliate density and the mean were 14 rotifers, were shrimps between epibionts fluctuated infested these basibiont the per of epibionts cent per Twenty-five epibionts the of 508– Distribution were and 2/2 were teeth The segments. five and foot 635 long a integument, transparent laticeps hip h oie,atog curn nlwnmes a ieydsrbtdo the on distributed widely of was numbers, low in occurring although basibiont. rotifer, The shrimp. 11.2–14.4 (each teeth two having 26.4–33 uncus was each with body the from protruding ytocnpcosclae ic,ec f39.6–79.2 of each discs, ciliated conspicuous two by 31.9–45.5 ihanro tece otwt orte iil hnte eentrtatd h head The retracted. not were they when visible toes four 59.4–85.6 with was foot stretched narrow a with et egh1.109 11 14.35 12.75 41.45 11.16 7.97 85.77 148.44 31.88 0.95 1.64 215.25 115.45 12.31 3.28 59.37 8.84 512.50 79.17 33.54 10.43 141.00 11.29 32.98 271.75 125.61 69.13 39.58 15.34 28.80 121.98 26.39 11.27 287.00 57.78 2.20 28.54 length Teeth width Uncus body the length protruding Uncus rostrum the of Length width Head length Head disc ciliated of Diameter width Body length Body h oyo hs oies si te dlod,hdtreprs ed rn,adfo.The foot. and trunk, head, parts: three had 271.8–512.5 bdelloids, other were in as individuals rotifers, these of body The genus the of Epibionts 2902 al II imti etrsof features Biometric VIII. Table al Isostenmeso pbot nec ntmclui of unit anatomical each on epibionts of numbers the shows XI Table h ttsia oprsnbtentedsrbtoso h pbotseiso h body the on species epibiont the of distributions the between comparison statistical The genus the to belong rotifers The .ensifera C. m og thsbe on ncutcas(ot n he 1986). Shiel and (Koste crustaceans on found been has It long. m Mra,10) hshssrih pr,drcl otrol,asedrbd iha with body slender a posteriorly, directly spurs, straight has This 1905). (Murray, .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. m ,adawdho 8–12.8 of width a and m, and niae infcn ifrnebtenteseis( species the between difference significant a indicated m ogad115.5–148.4 and long m .sulawesiensis A. .ensifera C. otanu intermedium Zoothamnium Embata maalaticeps Embata m eeas h otwdl itiue ntesraeo the of surface the on distributed widely most the also were , a ese nTbeXI h otaudn species, abundant most The XII. Table in seen be can ogad141–215.3 and long m Acineta Embata m TbeVI;Fgrs2,19–23). 2h, Figures VIII; (Table m maueet in (measurements m m og h atxhdwl-eeoe trochi, well-developed had mastax The long. m ie nteha,tecrn a composed was corona the head, the In wide. m hwdtehgetpooto,5.4) The 59.94%). proportion, highest the showed Sgr 02.Te r otsmlrto similar most are They 2002). (Segers and enS iiu Maximum Minimum SD Mean cnt sulawesiensis Acineta m m og.Teucshdalnt of length a had uncus The long). m imtr h ato h rostrum the of part The diameter. m m m m; ie h oywselongate was body The wide. m n 5 80). F 11.05; , hc represented which , P ( .ensifera C. .5.The 0.05). E. Z. . Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ubro oiesprsrm .317 .06.00 1114.00 5.00 0.00 6.00 14.00 1114.00 5.00 33.00 1.75 1110.00 306.07 430.00 2.00 1.00 314.62 29.00 1.43 12.00 266.59 13.00 4.00 0.78 665.60 2.30 305.16 1110.00 4.65 19.00 3.90 0.55 313.19 3.40 24.19 429.00 3.91 4.60 266.05 25.40 662.20 shrimp ovigerous per epibionts of shrimp Number per shrimp epibionts female of ovigerous Number per rotifers of shrimp Number per shrimp rotifers female of ovigerous Number per protozoans of shrimp Number per protozoans (mm) of shrimp Number female ovigerous (mm) of shrimp Width female ovigerous of (mm) Length shrimp the of (mm) Width shrimp the of Length rsaen( crustacean est enS iiu Maximum Minimum SD Mean laticeps E. laticeps Embata fusidens A. fusidens Amphileptus polypoides S. polypoides Spelaeophrya globosa V. globosa Vorticella compressa C. compressa Cothurnia maupasi P. maupasi Podophrya sulawesiensis A. sulawesiensis Acineta intermedium Z. intermedium Zoothamnium Density of specimens the of width and Length IX. Table pbot ma .7prui) ntefut lse eetefrtadlf second left and first of the number were moderate on cluster a epibionts different fourth had of Number the which X. Table In (first, units) pleopods unit). the of per rest of the 8.27 18.75% of (mean epibionts consisted pleopods, of cluster epibionts number third third lowest The and the unit). showed second per units the epibionts these pereiopods, of telson); 2.87 fifth 46.88% and and (mean pleopods, fourth included third, fifth cluster number pereiopod, and second high right fourth The a second unit). eyes, had antennulae, (rostrum, per units the units These epibionts anatomical to units). 15.80 corresponded anatomical (mean units the cluster epibionts The of A of shrimp. (18.75% the 21). (Figure uropods of and clusters units antennae, anatomical five the in on grouped species appeared ciliate epibiont 20). (Figure different The the densities of scarce. low numbers in more body were basibiont the that of species end contained the on cluster first compressa the Cothurnia basibiont; the on laticeps Embata polypoides with one clusters, two components, h irrhclCnlmrt nlsspoue edormuigtemean the using dendrogram a produced Analysis Conglomerate Hierarchical The e vgru eaesrm .023 6 1 2.30 3.40 shrimp female ovigerous per e vgru eaesrm .014 3 0 1.41 1.00 shrimp female ovigerous per e vgru eaesrm .056 13 0 5.61 3.00 shrimp female ovigerous per e vgru eaesrm .075 18 0 7.57 6.40 shrimp female ovigerous per n nte including another and , n e vgru eaesrm .015 3 0 1.52 1.40 shrimp female ovigerous per e vgru eaesrm 18 97 49 3 19.78 31.80 shrimp female ovigerous per 5 e vgru eaesrm 8.0206 9691 79 240.67 284.60 shrimp female ovigerous per 40). e vgru eaesrm 3.02.927362 297 26.49 333.80 shrimp female ovigerous per hsscn lse nlddteeiin pce htwr otcommon most were that species epibiont the included cluster second This . a eaaefo ohcutr.Ti a eepandb t presence its by explained be may This clusters. both from separate was .intermedium Z. aiiaensifera Caridina mhlpu fusidens Amphileptus pbotccmuiison communities Epibiontic aiiaensifera Caridina , 8.2172 585 691 1 0 187.22 188.62 154.66 107.81 ( n 11 64 49 0 16.40 11.14 .sulawesiensis A. 5 .317 6 13 4 0 22 0 1.75 0 18 2.82 0 1.43 1.20 1.14 4.80 0 0.67 1.33 4.61 2.48 enS iiu Maximum Minimum SD Mean nlsdaddsrbto fteeiinso the on epibionts the of distribution and analysed 40). , otclaglobosa Vorticella aiiaesfr 2903 ensifera Caridina , oohy maupasi Podophrya and , Spelaeophrya ,and Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 otu .21.4046 0 Maximum 12.14 Minimum 4.62 SD Mean Rostrum unit Anatomical etfut loo .978 29 60 58 0 116 116 0 47 0 55 7.84 7 15.80 0 4.19 0 13.56 16 7.90 0 6.95 0 15 25.76 25.07 36 0 13.48 10.71 26 0 8.33 14.29 14 9.19 pleopod fourth Left 0 6.81 1.53 pleopod third Right 35 0 4.61 pleopod 32 third Left 0.38 0 48 pleopod 4.19 2.05 second 0 Right 8.01 70 pleopod second Left 2.19 204 0 7.74 pleopod first Right 2.86 211 0 3.72 pleopod first 84 Left 4.05 0 79 pereiopod fifth 9.12 2.62 Right 0 pereiopod 8.11 1 fifth Left 56 4.81 0 12.46 pereiopod fourth 68 Right 6.67 0 19.61 pereiopod fourth 22 Left 8.67 0 57.83 5 pereiopod third Right 55.67 12.57 49.14 pereiopod 0 20.27 third Left 50.24 0 21.09 pereiopod second Right 15.76 0 pereiopod second Left 16.29 18.73 0 pereiopod first 20.82 Right 15.19 pereiopod first 5.27 Left 17.29 1.22 maxilliped Right 2.24 maxilliped Left 0.76 antenna Right antenna Left antennule Right antennule Left orbit ocular Right orbit ocular Left al I itiuino h pbot ntedfeetaaoia nt of units anatomical different the ( on below) epibionts given is the species of protozoan Distribution XI. Table 2904 .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. n 5 40). 06 57 116 0 25.79 10.62 70 204 211 84 0 79 1 56 0 19.52 68 0 57.83 12.48 0 55.57 49.05 0 20.19 50.14 0 21.02 15.62 18.73 16.19 20.82 15.19 17.29 .978 29 60 58 0 116 0 47 0 7.84 53 7 15.68 4.19 0 16 13.55 7.86 0 15 6.90 0 36 25.07 0 26 13.48 0 8.33 14 13.95 0 9.19 1.53 35 0 6.71 4.61 31 0 0.38 4.19 48 0 2.05 8.01 0 2.19 7.76 0 2.86 3.76 4.00 0 9.12 2.52 7.95 4.81 12.46 6.62 8.67 22 5 46 0 0 0 5.27 1.23 2.24 12.16 0.71 4.57 aiiaensifera Caridina tenme of number (the Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ih orhpepd39 .0039 species, two to corresponding distribution: each of clusters, pattern four similar showed a followed shrimp which the of units Maximum 49.6 cluster anatomical (mean fifth infestation The highest unit. the per with epibionts units unit). 0 9.26 the per of (6.25%), epibionts mean maxillipeds a the with to Minimum units), corresponds the of (9.38% pereiopods 8.80 SD 3.90 Mean pleopod fourth Right unit Anatomical pbot.As,i a on tahdt h eepd 1.% n h maxillipeds antennae, the rostrum, (43.2%), and maxillipeds the (18.3%) on pereipods located were the epibionts the the maxillipeds; of to of 50.48% 73.90% attached represented eyes; and it found where and contrast, was pleopods In (15.4%). the it antennulae, on were Also, especially units antennae, body, epibionts. the Anatomical of (rostrum, shrimp. half telson). posterior the and groups epibiont uropods total of pleopods; five the axis pereiopods; and anteroposterior species in epibiont the different considered along of proportion proportion mean mean the shows 23 Figure of axis longitudinal the throughout Distribution 61 13 52 18 0 0 17 0 19.29 4.24 17.51 0 13.90 0 3.90 16.76 5.12 3.97 2.81 2.10 uropod Right uropod Left Telson pleopod fifth Right pleopod fifth Left Continued. XI. Table nlssbtenteuiso h etsd ftesrm niae infcn difference significant a Comparison indicated Multiple shrimp a the ( showed The of side these side right appendages. left and ( the right them uropods, of among and units and the left pleopods, between fourth Analysis between fifth and correlation and third third hand, significant second, other second, first, the maxillipeds, On first, left antennae, pairs. antennulae, pereipods, as the in considered similar to when is shrimps. distribution units corresponded the these of between width variance difference of in significant variation maximum length no total in the the variation of width, 60.53% total represented the the contribution was of to Its shrimp 75% antennula. respect was the represented variance maximum With contribution of the with Its shrimps. width contributing antennula. most and unit right the length epibiosis, the the the to relation of in that, variance total the sulawesiensis to most contributing sulawesiensis h edormotie ihtema ubr fteeiin pce nthe on species epibiont the of numbers mean the with obtained dendrogram The h oprsnAayi fteeiisso etadrgtuisidctdta hr was there that indicated units right and left on epibiosis the of Analysis Comparison The ihrsett h u fnmeso h ntmcluisof units anatomical the on numbers of sum the to respect With F h u feiinso ahaaoia nto h ifrn hip showed shrimps different the of unit anatomical each on epibionts of sum The . , 7.40; , .polypoides S. F 8.14; , P ( .sulawesiensis A. P 0.05). ( – .5.As,teewsasgiiatdfeec ihrsett the to respect with difference significant a was there Also, 0.05). .fusidens A. a oeaudn nteatro ato h shrimp, the of part anterior the on abundant more was and , 38 91 60 52 0 0 19.17 17.36 13.86 16.67 pbotccmuiison communities Epibiontic .042 13 17 17 39 0 0 0 4.24 0 5.00 3.90 3.97 2.71 8.82 2.10 3.86 .maupasi C. .intermedium Z. aiiaensifera Caridina .intermedium Z. – .globosa V. a itiue anyo the on mainly distributed was aiiaesfr 2905 ensifera Caridina – .laticeps E. Fgr 22). (Figure .ensifera C. , .maupasi P. h species the , – A. A. ih orhpripd1.52 pereiopod fourth Right etfut eeoo 1.62 pereiopod fourth Left ih hr eeoo 0.86 pereiopod third Right ih nen 12.48 antenna Right ettidpripd1.95 pereiopod third Left ih eodpripd2.14 pereiopod second Right etscn eeoo 4.10 5.38 pereiopod second Left pereiopod first Right etatna13.05 antenna Left etfrtpripd6.57 pereiopod first Left ih nenl 13.86 antennule Right ih ailpd41.29 maxilliped Right etatnue15.19 0.29 antennule Left orbit ocular Right etmxlie 40.10 maxilliped Left etoua ri 0.10 orbit ocular Left al I.Nme fec pbotseiso h ifrn ntmcluisof unit units Anatomical anatomical different the on species epibiont each of Number XII. Table otu 3.05 Rostrum

Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 sulawesiensis 024 04 04)(–)(0–2) (0–1) (0–45) (0–4) (0–204) 014 01)(–7 (0–2) (0–67) (0–18) (0–144) Acineta 01)(–)(0–11) (0–1) (0–10) 01)(–)(0–8) (0–9) (0–18) 01)(–)(–6 (0–1) (0–26) (0–1) (0–11) 01)(–)(0–2) (0–6) (0–13) 07)(–)(–8 01 01 (0–1) (0–1) (0–1) (0–18) (0–5) (0–74) 01)(–)(0–20) (0–2) (0–19) 02)(–5 (0–1) (0–15) (0–21) 05)(–8 01 01 (0–1) (0–1) (0–1) (0–19) (0–48) (0–4) (0–39) (0–53) 04)(–)(–)(0–2) (0–4) (0–1) (0–47) 06)(–)(–)(–1 04 (0–1) (0–4) (0–11) (0–2) (0–2) (0–68) 07)(–)(–1 03 02 (0–2) (0–2) (0–3) (0–11) (0–9) (0–72) 03)(–)(–)(–7 01 (0–1) (0–1) (0–27) (0–1) (0–2) (0–35) 02 01 01 05 (0–1) (0–22) (0–5) (0–1) (0–1) (0–3) (0–2) ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ 10 0.24 – 11.01 24 5.81 – – – 12.43 .00.05 2.80 .50.43 4.25 .1–0.05 – 2.41 70 0.76 – – 17.07 .4–––0.57 – – – 3.34 .3–0.10 – 4.53 .4–––2.52 – – – 5.74 88 0.76 – – 18.80 67 0.05 16.72 40 0.33 – 54.08 .8–––1.95 – – – 0.10 19.78 0.78 45 0.86 – 44.51 .40.05 0.44 .30.10 8.43 Vorticella globosa ¡ ¡ ¡ ¡ ¡ ¡ .2––0.62 – – 0.22 .6––0.76 – – 1.96 .2–0.29 – 0.22 .4–0.14 – 0.44 .20.05 0.22 .4–0.05 – 0.44 Podophrya maupasi ¡ ¡ ¡ ¡ ¡ ¡ .2–3.05 – 0.22 .4–2.57 – 0.44 .9–3.05 – 0.89 .6–7.33 – 1.06 .3–9.10 – 3.93 .2–0.52 – 0.22 aiiaensifera Caridina Cothurnia compressa ¡ ¡ ¡ ¡ ¡ .02.10 2.30 .12.14 1.41 .60.10 0.96 .81.43 0.48 .21.29 0.22 (mean Zoothamnium intermedium ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ 25 0.05 12.50 31 0.10 – 13.19 68 0.10 – – 16.86 D iiu–aiu)( minimum–maximum) SD; .0––– – – 2.40 .5––– – – 1.95 .3––0.05 – – 7.33 .6––0.10 – – – – – 1.36 5.74 .20.10 4.62 .7––0.05 – – – – – 4.47 5.30 .4––– – – 0.44 .40.29 2.94 .6––– – – 5.06 .1––0.05 – – 1.21 .20.14 3.62 .9–0.10 – 5.89 Spealophrya polypoides ¡ ¡ ¡ ¡ .00.05 0.30 .20.05 0.22 .00.14 0.90 .50.14 0.65 n Amphileptus 5 40). fusidens ¡ ¡ ¡ ¡ ¡ ¡ .20.10 0.22 .20.14 0.22 .6– 0.36 .00.10 0.30 .80.14 0.48 .00.05 0.30 Emabata laticeps ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡

0.22 0.44 0.30 0.22 0.36 0.44 0.44 0.22 0.48 0.22

2906 .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. al I.Continued. XII. Table ih rpd9.67 uropod Right etffhpepd0.62 pleopod fifth Left esn0.33 Telson etuoo 8.14 uropod Left ih it loo 0.10 pleopod fifth Right ntmclunit Anatomical etffhpripd0.38 pereiopod fifth Left ih it eeoo 0.38 pereiopod fifth Right ih is loo 1.43 pleopod first Right etfut loo 0.71 pleopod fourth Left etfrtpepd0.57 pleopod first Left etscn loo 0.57 pleopod second Left ettidpepd0.86 pleopod third Left ih hr loo 0.33 pleopod third Right ih orhpepd0.48 pleopod fourth Right ih eodpepd0.52 pleopod second Right

Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 sulawesiensis Acineta 06)(–)(–)(–1 02)(0–1) (0–23) (0–21) (0–2) (0–5) (0–60) 04)(–3 01)(0–1) (0–16) (0–33) (0–44) 01)(–)(–4 (0–1) (0–44) (0–2) (0–11) 07 02 03)(0–1) (0–38) (0–2) (0–7) 01 03 01)(–)(0–1) (0–1) (0–17) (0–3) (0–1) 05 06 06 01)(0–8) (0–12) (0–6) (0–6) (0–5) 08 09 06 (0–1) (0–6) (0–9) (0–8) 07 (0–15) (0–7) 06 05)(–)(0–2) (0–1) (0–53) (0–6) 03 01 06)(0–1) (0–60) (0–1) (0–3) (0–7) 07 07 02 019 (0–2) (0–109) (0–2) (0–7) (0–7) 08 02 05)(0–1) (0–58) (0–2) (0–8) 07 (0–29) (0–7) 05 02 014 (0–1) (0–114) (0–2) (0–5) ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ 63 0.24 16.37 17 5.33 – – 11.78 .3––0.76 – – 1.83 .10.29 1.11 .0–0.14 – 0.30 .3–––1.67 – – – 1.53 .3––––––– – – – – – – 1.53 .0––0.10 – – 2.60 .1–––3.48 – – – 0.71 .0–––6.10 – – – 1.40 .6––0.14 – – 1.36 .0––0.14 – – 1.90 .10.05 0.91 .4––0.19 – – 1.54 .3–0.33 – 1.63 Vorticella globosa ¡ ¡ ¡ .90.10 1.09 .10.29 1.31 .2––7.52 – – 0.22 Podophrya maupasi ¡ ¡ ¡ ¡ .42.33 0.44 .1–2.62 – 1.31 .5–2.33 – 0.65 .30.14 1.53 Cothurnia compressa ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ 09 3.19 10.91 .51.52 5.15 .00.67 2.30 .47.62 0.44 .87.48 0.48 .89.62 0.48 .85.90 0.48 .13.19 0.51 Zoothamnium intermedium ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡ 39 0.05 13.92 30 0.05 – 13.08 57 0.05 – – 15.74 48 0.05 – 24.84 45 0.10 – – 24.57 34 0.05 – – 13.46 .2––0.05 – – 5.02 .4––0.10 – – 5.14 .3–0.38 – 3.83 .3––0.05 – – 8.63 .5–0.10 – 0.05 4.85 1.56 .0––– – – 3.90 .8––– – – 7.98 Spealophrya polypoides ¡ ¡ .2–– – 0.22 .2–0.10 – 0.22 Amphileptus fusidens ¡ ¡ ¡ ¡ .5– 1.75 .00.10 0.30 .2– 0.22 .2– 0.22 Emabata laticeps ¡ ¡ ¡ ¡ ¡ ¡ ¡ ¡

0.22 0.30 0.22 0.30 0.44 0.22 0.44 0.22

pbotccmuiison communities Epibiontic aiiaesfr 2907 ensifera Caridina Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 hwn itntv itiuinaogtebsbotbd.Teewsn significant no ( species was them each There among difference with body. significant ensemble, a basibiont but of the species, behaviour epibiont along the with distribution among places correlation distinctive in the pattern a a occupied following showing epibionts, species distributed was the of community epibiont which 39% The pleopods. and and pereiopods, 68.8 fusidens for A. accounted they where body, the although of respectively, mean end the posterior using eyes. performed and analysed. Analysis shrimps antennulae, Component different the Principal of the unit anatomical of each components on principal epibionts of first density Two 20. Figure 2908 este feiinso h ifrn ntmcluiso h hip nlsd(ercdsac:Ct Block City distance: mean (metric analysed the shrimps using the performed of units) units (anatomical anatomical Analysis different Ward). Cluster the method: on Hierarchical (Manhattan); epibionts the of of densities Dendrogram 21. Figure between fluctuated areas the colonization; in difference remarkable a showed shrimp P nene nenle n ys (77.9%). eyes) and antennulae, antennae, epibionts). ( .5.Tettlcutdt niae htn oeo h neootro xso the of axis anteroposterior the on zone no that indicated data count total The 0.05). .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. .polypoides S. h nso h oy while body, the of ends the .globosa V. .compressa C. a oeaudn nteatro ato h oy(rostrum, body the of part anterior the on abundant more was a eakbynmru nteprios(53 fthe of (35.3% pereipods the on numerous remarkably was and .laticeps E. .maupasi P. .globosa V. a itiue nteatro end, anterior the on distributed was anyclnzdtemxlies and maxillipeds, the colonized mainly eemr ueostwrsthe towards numerous more were F 5.98; , Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 NeadL 95,on desmaresti 1945), Lu and (Nie polypoides Spelaeophrya and ( found polypoides species Spelaeophrya epibiont protozoan The females. h pbotseisfudi h pce of species the in found species epibiont the genera the of species although a rvosybe bevda pbot ncutcas(oaoadSal1995; Small The and 2004). genera (Morado Fernandez-Leborans the crustaceans to 2000b; belong on found 2000a, species epibionts ciliate Tato-Porto as and observed Fernandez-Leborans been previously had xsi iegop faaoia nt ftesrm,hr hr a infcn difference significant ( a species was epibiont there different here the shrimp, the of of distribution units the anatomical between of groups five the in axis of 45.63% of axis for accounted telson the and of 54.37% uropods, for colonization. accounted pleopods, basibiont (rostrum, region the Pereiopods, anterior of The colonization. maxillipeds) maxillipeds. Block and the eyes, City on antennulae, 31.67% distance: mean and antennae, (metric end the posterior analysed the using shrimps on 11.02% performed the of species) units (epibiont anatomical Analysis different Cluster Ward). the method: Hierarchical on (Manhattan); epibionts the of of densities Dendrogram 22. Figure ot ooie pcmn of specimens colonized Forty Discussion several Podophrya FradzLbrn ta.2006a). al. et (Fernandez-Leborans coarctata 1963, Biernacka, h itiuino h pbotseiso h ifrn oyaesaogtelongitudinal the along areas body different the on species epibiont the of distribution The mhlpu fusidens Amphileptus .ensifera C. Caridina (Clapare Mle,13)(uoe,adon and (Europe), 1831) (Millet, n aeol eetybe bevda pbot on epibionts as observed been recently only have and pce rmSlws FradzLbrn ta.20b.TbeXI lists XIII Table 2006b). al. et (Fernandez-Leborans Sulawesi from species d n ahan 88,and 1858), Lachmann, and `de ssoni iue2.A ntedsrbto hogottelongitudinal the throughout distribution the in As 24. Figure in shown is ohri compressa Cothurnia , a enfudpeiul on previously found been has aentbe eoddfrel seiinson epibionts as formerly recorded been not have ) otanu intermedium Zoothamnium aamntsantennarius Palaemonetes Acineta .ensifera C. cnt tuberosa Acineta pbotccmuiison communities Epibiontic , , Podophrya otclacampanulata Vorticella Acineta eeaaye,o hc iewr ovigerous were nine which of analysed, were Xiphocaridina Caridina , cnt sulawesiensis Acineta , MleEwrs 87 and 1837) Edwards, (Milne , Cothurnia Zoothamnium Zoothamnium otclaglobosa Vorticella rmSlws.Tescoinspecies suctorian The Sulawesi. from Caridina hebr,1833, Ehrenberg, ove,10 Arc n Asia) and (Africa 1909 Bouvier, F 5.07; , , Zoothamnium aiiaesfr 2909 ensifera Caridina Kh,1933), (Kahl, p nCugig(China) Chungking in sp. , sp Vorticella P . , ( , aebe on on found been have ohri compressa Cothurnia oohy maupasi Podophrya aiialanceolata Caridina 0.05). aiiaensifera Caridina hrcl elegans Thuricola ,and , Vorticella Opercularia Atyaephrya Cothurnia ,and , , , Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 rsae,adVrert.Tertfrseisfudi hsstudy, on 1989). this in previously (May found species found involved rotifer Mollusca, The relationships Echinodermata, Vertebrata. associated Bryozoa, of and Mollusca, species Crustacea, Annelida, types parasitic Rotifera, and the Porifera, epizoic Protozoa, of include with known Bdelloidea associated and fauna is Monogononta, the little Seisonidae, of but ciliates organisms, as considered other typical be not should are they they with although therefore, crustaceans, and on epibionts ciliates as sessile found previously been never have 78.Ohrrtfr r pzi ndvrefehae rsae,as crustacea, freshwater diverse on Asellus epizoic are rotifers Other 1758). 2910 Ddy11;Nzw 98 Hadz 1938; Nozawa the 1910; along epibionts, of (Daday density mean total and densities) (mean species of epibiont axis each anterioposterior of Distribution 23. Figure Samr13;Matjas 1935; (Stammer troglocaridis .ensifera C. efry 1762, Geoffroy, .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. aebe ecie rmtecv shrimp cave the from described been have oiesaewl-nw seiinso slvn ncoeascainwith association close in living as or epibionts as well-known are Rotifers . aiiaensifera Caridina slu aquaticus Asellus ˇic ˇ 96 ate ta.18) h pce ftegenus the of species The 1988). al. et Matthes 1956; Gammarus ntmcluisaecniee nfv groups. five in considered are units Anatomical . i14) pce iia to similar Species 1940). ˇi Lnau,15)and 1758) (Linnaeus, arcu,17 My18) rdecapods or 1989), (May 1775 Fabricius, rgoai schmidti Troglocaris amrspulex Gammarus maalaticeps Embata Daphnia .polypoides S. omte,1853 Dormitzer, ulr 1785, Muller, Amphileptus (Linnaeus, a been has , and S. Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 iue2.Dsrbto fec pbotseis(endniis ln h neipseirai of axis anterioposterior the ( along densities) (mean species epibiont each of ensifera Distribution 24. Figure rsneo oie pce ncutca Excepting crustacea. on species rotifer of presence Garzo Chasmagnathus iits oee,i eea ae,te xii euircaatrsis hc may which characteristics, peculiar exhibit they cases, several as in considered and However, maxillipeds generally the ciliates. as are on such and locations feed on could basibiont especially they host, their Also, the 1989). to by uropods. (May treated harm association matter organic the any the from cause transport to obtaining epizoans, appear the from of not shrimp case the the along in do present As higher scarcely uropods. relative was the its It to to epibionts. antennae due other perhaps to abundance, comparison maximum in lowest biomass the with species epibiont the Austropotamobius h pbotseisosre r iia oohr on seiinso sfree-living as or epibionts as found others to similar are observed species epibiont The nadMneo17;Fnaeoe l 04.TbeXVsmaie h recorded the summarizes XIV Table 2004). al. et Fontaneto 1973; Montero and ´n ntmcluisaecniee niiuly e,pripd l,pepd r,uropod. uro, pleopod; ple, pereiopod; per, individually. considered are units Anatomical . eHa,13)(ae 96 99 aln13;Wlet15;Man 1957; Wulfert 1939; Carlin 1959; 1926, (Hauer 1835) Haan, De krkw 1908, Skorikow, Astacus pbotccmuiison communities Epibiontic amrspulex Gammarus arcu,1775, Fabricius, pleprapolypoides Spelaeophrya and slu aquaticus Asellus aiiaesfr 2911 ensifera Caridina Potamon , ain,1816, Savigny, .laticeps E. h rotifers the , Caridina was ˜e ´- Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 oie epibiont Rotifer ao Locality epibionts 2006b). Ciliate 2006a, al. et Fernandez-Leborans from revised Taxon been have names species some study; on this found from epibionts of List XIII. Table 2912 otclacampanulata Vorticella globosa Vorticella pleprapolypoides Spelaeophrya elegans Thuricola maupasi Podophrya Zoothamnium n ahan 1858) Lachmann, and coarctata Opercularia maalaticeps Embata otanu intermedium Zoothamnium psyi coronata Epistylis compressa Cothurnia fusidens Amphileptus tuberosa Acineta ahan 1858 Lachmann, eoas ize,adGblno 2006 Gabilondo, and Zitzler, Leborans, sulawesiensis Acineta .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. p aiilks aeTwt,Nshore N Towuti, Lake lakes, Malili sp. Mra,10)Lk oo shore S Poso, Lake 1905) (Murray, hebr,13 aiilks aeTwt,Nshore N Towuti, Lake lakes, Malili 1833 Ehrenberg, hs,12 aePs,Sshore S Poso, Lake 1922 Ghosh, irak,16 aiilks aeMhln,N shore NW Mahalona, Lake lakes, Malili 1963 Biernacka, uc,17 aiilks aeMtn,Sshore S Matano, Lake lakes, Malili 1970 Nusch, ushi 89Lk oo shore S Poso, Lake 1889 Butschli, Clapare Kh,12)Lk oo shore S Poso, Lake 1926) (Kahl, Fernandez- (Clapare Kh,13)Mll ae,Lk aao shore S Matano, Lake lakes, Malili 1933) (Kahl, aa,11 aePs,Sshore S Poso, Lake 1910 Daday, rct 1935 Precht, d and `de `de Caridina aiilks aeTwt,LeaIln,Sshore S Island, Loeha Towuti, Lake shore lakes, NW Malili Mahalona, Lake lakes, Malili aiilks ee River Petea lakes, Malili shore NW Mahalona, Lake lakes, Bantimurung Malili karst, Maros Sulawesi, South River Petea lakes, Malili aiilks aeMhln,N shore NW Mahalona, Lake shore lakes, S Malili Matano, Lake shore lakes, SW Malili Towuti, Lake lakes, Malili River Tarrak Sulawesi, South Masapi Lake lakes, Malili River Petea shore lakes, W Malili Towuti, Lake shore lakes, E Malili shore Towuti, S Lake Island, lakes, Loeha Malili Towuti, Lake lakes, Malili aePs,Sshore S Poso, Lake aiilks aeMhln,N shore NW Mahalona, Lake lakes, Malili ot uaei arkRiver Tarrak Sulawesi, South River Petea lakes, Malili aiilks aeMhln,N shore NW Mahalona, Lake shore lakes, S Malili Matano, Lake shore lakes, SW Malili Towuti, Lake lakes, Bantimurung Malili karst, Maros Sulawesi, South Masapi Lake lakes, Malili River Petea shore lakes, W Malili Towuti, Lake shore lakes, E Malili shore Towuti, S Lake Island, lakes, Loeha Malili Towuti, Lake lakes, Malili shore S Poso, Lake Bantimurung karst, Maros Sulawesi, South River Tarrak Sulawesi, Masapi South Lake system, lake Malili River Petea shore lakes, W Malili Towuti, Lake shore lakes, E Malili shore Towuti, S Lake Island, lakes, Loeha Malili Towuti, Lake lakes, Malili shore S shore Poso, NW Lake Mahalona, Lake shore lakes, S Malili Matano, Lake shore lakes, SW Malili Towuti, Lake lakes, Malili rmSlws FradzLbrn ta.20a 06,addata and 2006b, 2006a, al. et (Fernandez-Leborans Sulawesi from .ensifera C. ensifera C. .lanceolata C. lanceolata C. lanceolata C. .loehae C. ensifera C. lanceolata C. typus C. loehae C. .lanceolata C. .lanceolata C. lanceolata C. lanceolata C. C. masapi C. loehae C. lingkonae C. lanceolata C. lanceolata C. .ensifera C. C. .ensifera C. .lanceolata C. .lanceolata C. C. loehae C. .lanceolata C. lanceolata C. lanceolata C. typus C. masapi C. loehae C. lingkonae C. lanceolata C. lanceolata C. ensifera C. ensifera C. typus C. C. masapi C. loehae C. lingkonae C. lanceolata C. lanceolata C. ensifera C. lanceolata C. lanceolata C. lanceolata C. C. Caridina p 2 sp. p 1 sp. p 2 sp. p 2 sp. p 1 sp. basibiont Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 hldn convergens Philodina parasitica Embata o nypoet h tl,btas h ooy(enne-eoase l 06) The 2006a). al. is et life (Fernandez-Leborans epibiont the colony to the adaptation also obvious most but the stalk, showing genus the protects only not stalk. .truncata T. eael astacicola Lepadella grande Encentrum esnanltsNebalia magna Daphnia excisum Diaphanosoma sociales Rotaria nudus Paraseison annulatus Seison rubens Brachionus sessilis Brachionus and Settele 1989; May 1986; Shiel Epibiont and Koste from (data crustacea study). on epibiont present species 2003; rotifer Thalhofer of List XIV. Table maaltcp aiiaensifera Caridina example, For life. epibiont the to adaptations constitute laticeps Embata rotatoria rotatoria Rotaria ol ecreae otebhvoro h hip(enne-eoase l 06) Like 2006a). of al. et communities appendages. (Fernandez-Leborans epibiont shrimp right the lanceolata the of and behaviour C. left in the to the observed correlated between be also could anterior differences was the significant to phenomenon corresponded without colonization This body, maximum the the and of body, areas the of crustaceans. end shrimp. posterior on the the squash- epibionts of a as appendages of the found of ciliate been most its surface the only the of are and have with adoption linkage suctorians ciliates flattened, the the These these by to considerably friction due where the was probably antennae, reducing widened, macronucleus shape the appear the can can of body part individuals, surface the anterior area several the posterior the to In in next and abundant. flattened, resting is folded, body the be lorica, conspicuous a lacking is as such Something ciliates, basibiont. the Other of surface the basibiont. to close in the settle occurs to of ciliate similar the allows movement which stalk, the short from them by protecting possibly tentacles, caused Cothurnia the enveloping damage body the over possible part anterior free a as eael borealis Lepadella crassipes Cephalodella rcinsnveelniePedmnalemnae Pseudomonia novaezealandiae Brachionus volutata Collotheca rcinsvraii ahi longispina Daphnia curvirostri Acantholeberis Daphnia variabilis Brachionus epicopta Testudinella daphnicola Proales aePs n fe on nplgcsam K o itln esnlfedobservation). field personal Rintelen, von (K. swarms pelagic in found often and Poso Lake rae auiPgrsbernhardus Pagurus granulat Chasmagnathus tardigrada Rotaria paguri Proales chasmagnathi Anomopus cancrophila Macrotrachela h itiuino pbotseison species epibiont of distribution The Zoothamnium , .caeca T. r rtce yalrc opeeysronigtebd,adwt noticeably a with and body, the surrounding completely lorica a by protected are , , .magna-calcarata R. , , nteMll aesystem, lake Malili the in , .proboscideus P. .astacicola L. .laticeps E. , var. , , .branchiola L. maaparasitica Embata , irnpou hauerianus Dicranophorus irnpou hauerianus Dicranophorus Podophrya eslsMnsiu dispar Monospilus sessilis a ra tl,wihi oiebywdron wider noticeably is which stalk, broad a has , nmpstlhseTlhs fiuviatilis Telphusa telphusae Anomopus , .hamaca E. , .lata L. , , , .clau Nebalia ciliatus P. etdnlaelliptica Testudinella .parasitica L. hc,i otatt h relvn pce,hsavr short very a has species, free-living the to contrast in which, , .laticeps E. , .hurau abrsaffinis Cambarus hauerianus D. , rae daphnicola Proales , var. , pbotccmuiison communities Epibiontic .boreales L. .commensalis E. .ensifera C. siedleckii .ensifera C. , , , , slu aquaticus Asellus amrspulex Gammarus Ceriodaphnia Crustaceans Astacus rectirostris rsae bacilchambers) (branchial Crustacea olwdagain rmteatro to anterior the from gradient a followed saudn n ieydsrbtdin distributed widely and abundant is cnt sulawesiensis Acineta sp. sp. sp. , .brachiata M. sp. aiiaesfr 2913 ensifera Caridina , .pulex D. Spelaeophrya , aiialanceolata Caridina , Basibiont eidpnalongispina Ceriodaphnia ahi carinata Daphnia , , .sarsii D. oyhmspediculus Polyphemus , .longispina D. .lanceolata C. , a h lorica the has lhuhit Although . .brachyurum D. , , Moina This . and Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 2914 n hsmyifunetedsrbto n bnac fbt oe n ihrorganisms web, higher food and lower aquatic very both the of abundance of a and components distribution community are the other the common influence in a affect Protozoa may changes are key significantly thus The they and 1985). ecosystems. may have and al. levels protozoa they et freshwater trophic (Porter of and higher larvae structure in the fish web to and food cycling crustaceans energy lake for of limnic of nutrient and transfer the Protozoa the flow in in shrimp. link the link important energy of major ciliates in activities a Peritrich the as other functions ciliates. movement from considered on other free arising on are or have nutrients feed environments community, (that the suctorians on host epibiotic example, depend the the for species could to as, in several associated basibiont), present and community the are relations), around the other that to or groups, protozoans belonging loop other (microbial organisms and environments, on feed-back free energy in feed species happens of can as sort among therefore, a and, is links equilibrium there trophic diverse towards are there community trend functional general density. each species and of a The diversity requirements for pattern. with particular compensating colonization with their but substratum, varied by available which group, of verified the pattern basibiont be occupy certain the to can a of show tend which to axis ensemble, seems longitudinal by community the the behaviour data However, along shrimp, basibiont, statistical species. the distribution the between of The units on of significantly anatomical 1%. distribution pattern the on distinctive than the species a epibiont less and of had was presence species differential epibionts the epibiont involving different other the of that the density showed the sexes, Abello 1997). and while al. two 1988; protozoans, burying et Fernandez-Leborans Ropes the areas, 1993; al. and geographical between et Gili (Bottom different pattern indicate 1992; etc. of Macpherson can and moult basibiont density behaviours, populations of the The between feeding of basibiont. differences units moult the anatomical seasonal of asynchronic different biology moult, the the the on and terminal of to epibionts species, and aspects contribute epibiont of physiology, may important the distribution Epibiosis dynamics, of of communities. structure, discernment pattern invertebrate and the colonization the protozoan the therefore, the reflect and, of organic basibiont development 1993) of the al. passage of wide et important ecology their (Gili an to adhere with shrimp. they due the body of possibly the movement and uropods, in digestion Other the position the function. from were their their material to and epibiosis due surface, high possibly particles exposed nutrient with these which the of of sites characteristics location presence and protected the morphological three frequent the surface, of epibionts the showed available and surface wide of maxillipeds, appendages, the densities the epibiosis: the settlement high remarkable body, with example the the units explain the for Other for colonization. of epibionts, the substrata part facilitates of appendages anterior provide these of the antennae movement In and colonization. antennulae for important were 2006a). al. et (Fernandez-Leborans lakes Malili 90,aogwt t ihmblt anybig h neirpr ie h feeding the substrates hard (i.e. and part in soft observed anterior of similarly kinds the was different This brings with macrophytes). mainly contact sand, into mobility wood, shrimp (rocks, high the its of for described appendages) with as behaviour, along feeding 1960), characteristic and rapid Its h rtza iit pbot rbbyd o amtebsbot ihnteepibiont the Within basibiont. the harm not do probably epibionts ciliate protozoan The ciliate mainly species, epibiont of diversity and density high a showed Poso Lake which to subtratum living the of characteristics the upon depend organisms sessile Many morphology their and surfaces basibiont the of characteristics physical the addition, In .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. ta.19;Abello 1990; al. et ´ Caridina .lanceolata C. ngnrl(Fryer general in rmthe from and ´ Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ikM,DnlsnW,Vnn W 98 pbot ftetne crab tanner the of Epibionts 1998. IW. Vining WE, Donaldson MH, Dick M, Corn In: environments. marine U in quality 1939. water of B. indicators Carlin as Protists 1993. PV. McCormick Jr, J Cairns LIPI Soroako. in Inco T. P. Bogor, References to Bogoriense, and permits. Indonesia research Zoologicum to provided in grateful Sciences) (Museum support further of Institute are Marwoto logistic (Indonesian We their Ristiyanti Poso. History, for Lake Natural and Indonesia) from of material Wowor (Museum the Rintelen Daisy collecting von for Thomas Germany) and Berlin, Glaubrecht Matthias thank We Acknowledgements similar a established strategy. has particular species a each species cases, following all community distribution. located in of epibiont and were The pattern species the species present 2006). the which The of Gabilondo pattern. in Independently and colonization environment a (Fernandez-Leborans dynamic acquire reflect a environment could represents which the community, basibiont of epibiotic an et biodiversity to (Patterson small transferred the a be diversity On could trophic 2002). conditions important Fernandez-Fernandez these an and benthic scale, and Fernandez-Leborans in 1990; rates Fenchel especially growth 1989; environments, high al. free show 1992). they in Fahnenstiel where significance and areas, ecological Carrick 1993; important McCormick have and Ciliates Cairns 1989; Crisman and (Beaver otmM,RpsJ.18.A nietmto o siaiglneiyo h ossoca ( horseshoecrab the of longevity estimating for method indirect An 1988. Ecology Microbial JW. ecosystems. freshwater Ropes pelagic ML, in protozoa Bottom ciliated of role The 1989. TL. Crisman JR, Beaver Abello Abello ece .19.Terl fpooo nntr ntrso ucinlpoete eae osz.Zoological size. to on related epibionts properties protozoan functional of of distribution terms in Comparative nature 2004. in G. protozoa Fernandez-Leborans of role The 1990. T. Fenchel barnacle the of attachment The 1962. B. Eldred 4. Kinetofragminophora). (Ciliophora, Suctoria 1957. the CE. u Dawson Untersuchungen of 1910. EV. revision Daday New A ed. 2nd literature. 1986. the to CR. guide and Curds classification characterization, 1907– Protozoa: ciliated Expedition, The Phillipine 1979. Albatross JO. Corliss the of Decapoda) (Crustacea: situ shrimps In caridean Michigan. The Lake 1997. in FA. protozoa Chace planktonic of production and Growth 1992. GL. Fahnenstiel HJ, Carrick ,Vlauv ,Gl M 90 pboi nde-e rbppltosa niao fbooia and biological of indicator as populations crab deep-sea in Epibiosis 1990. reproductive JM. the to Gili relation R, in Villanueva patterns P, infestation rhizocephalan ´ and Epibiosis 1992. E. Macpherson P, ´ dtr adoko aadu aeil.NwYr:Aaei rs.p627–638. p Press. Academic York: New materials. hazardous of Handbook editor. polyphemus eaiua hrceitc ftehs.Junlo h aieBooia soito fteUie Kingdom United 17:111–136. the of Association Biological Marine the of Journal 70:687–695. host. the of characteristics behavioural of biology Msdca rmtrelksi otenErp.Cutcaa76:1037–1054. Crustaceana Europe. Northern in lakes three from (Mysidacea) cec 7:51–58. Science shrimp, rock the 18:519–528. Biology Crustacean of Journal Alaska. Island, Kodiak 50:59–91. Series Zoology History) (Natural Museum British the of Bulletin relatives. morphological Press. Oceanography Pergamon York: and Processidae, 587:1–106. Limnology Bathypalaemonellidae, Zoology to Rhynchocinetidea, dynamics. Contributions Eugonatonotidae, Smithsonian Atyidae, web Hyppolytidae. Families and food 7: Part to 1910, importance and comparison vitro 37:1221–1235. in versus 2:3–68. Institution Limnologiska ae neianlsiprshells slipper epifaunal on based ) ihdsferox Lithodes ¨ Balanus e i oaoinengrSe e nbd.Mdeudnfa ud Universitets Lunds fran Meddelunden Aneboda. bei Seen einiger Rotatorien die ber iynadorsalis Sicyonia oln fsrm.Sine126:1068. Science shrimp. of fouling ihl 85(nmr:Ltoia) ora fCutca ilg 12:561–570. Biology Crustacean of Journal Lithodidae). (Anomura: 1885 Filhol, brdeSu die ¨ber igly rsaen 3:203–206. Crustaceana Kingsley. ßasrMkoan etc-sarks olgc 23:1–314. Zoologica Deutsch-Ostafrikas. ¨ßwasser-Mikrofauna pbotccmuiison communities Epibiontic aau mhieniveus amphrite Balanus rpdl fornicata). Crepidula ora fSelihRsac 7:407–412. Research Shellfish of Journal awn n te oln raim to organisms fouling other and Darwin, hooctsbairdi Chionoecetes aiiaesfr 2915 ensifera Caridina yi relicta Mysis Podophrya ntergo of region the in oe,1869 Loven, n its and Limulus Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 ae .15.Ruprstsh oaoinasdrKiemenho der aus Rotatorien Raumparasitische 1959. J. Hauer edrF ot W fedrJ cwre ,HdayR,ShiwnU.20.Aatv aito and radiation Adaptive 2006. UK. Schliewen RK, Hadiaty J, Schwarzer J, Pfaender AW, Nolte F, Herder e M iso E op W efisW,VrsH.19.Byza oln ftebu crab blue the of fouling Bryozoan 1999. HK. Voris WB, Jeffries JW, Volpe JE, Winston MM, Key onuhnA,Gabeh .20.Nvlrpoutv oe nfehae lm:bodn n larval and brooding clams: freshwater in modes reproductive Novel 2003. M. Glaubrecht AV, Korniushin adnd ,Ui J 92 eainhp ewe pne n rb:pten feiisson epibiosis of patterns Man crabs: and guide sponges illustrated between An Relationships editors. 1992. P, MJ. Bradbury Uriz GF, M, Leedale Maldonado JJ, Lee In: Ciliophora. Phylum 2000. EB. Small DH, Lynn China, from ciliates pleurostomatid marine new Two Australian 2005. Digononta). A. (Rotifera: Warren Bdelliodea W, I. Song X, waters. Lin inland Australian from Rotifera 1986. RJ. Shiel W, Koste ae .12.De eeLpdlaAtnasdnKiemenho den aus Lepadella-Arten neue Drei 1926. J. Hauer Suktorien: crab symphorische the Neue in 1940. duration J. intermolt Hadzi and Epibionts 1993. R. Villanueva P, Abello JM, Gili ate ,Gh ,Hie .18.Scoi Suifsre) n utraudUcoaide Protozoenfauna Urceolariidae. und Suctoria In: (Sauginfusorien). Suctoria 1988. G. Haider W, Guhl D, Matthes Matjas re .16.Tefeigmcaim fsm ti rwso h genus the gill of the prawns atyid from some Bdelloidea) of Monogononta, mechanisms (Rotifera: feeding The rotifers 1960. Epizoic G. 2004. Fryer G. Melone shrimp H, freshwater Segers the on D, epibionts Fontaneto ciliate Protozoan 2006b. R. Gabilondo K, Zitzler G, Fernandez-Leborans enne-eoasG ize ,GblnoR 06.Eiin rtza omnte on communities protozoan Epibiont 2006a. R. Gabilondo K, Zitzler G, Fernandez-Leborans enne-eoasG aoProM.20b eiwo h pce fpoooneiinso rsaen.I. crustaceans. on Co epibionts Herrero protozoan G, of Fernandez-Leborans species the of I. review crustaceans. A on 2000b. epibionts ML. protozoan Tato-Porto on of G, Fernandez-Leborans species community the of epibiotic review the A 2000a. estuarine of ML. Tato-Porto sublittoral variability G, Fernandez-Leborans Inter-annual a 2006. in R. groups Gabilondo functional G, Protist Fernandez-Leborans 2002. D. Fernandez-Fernandez G, Fernandez-Leborans enne-eoasG atod admieM 96 h opooyof morphology The 1986. M. Zaldumbide de Castro G, Fernandez-Leborans 2916 oaoJ,SalE.19.Claeprstsadrltddsae fCutca eiw eiw nFisheries in Reviews review. a on Crustacea: Studies of 1945. KSY. diseases Lu related D, Nie and parasites Ciliate 1995. 186/187:59–67. EG. Hydrobiologia Small rotifers. parasitic JF, and Morado Epizoic 1989. L. May ˜e ´-Garzo 16:459–464. crnk.Beitra Schranck). yrdzto nWlae’ rapns vdnefo ali ivrie nteMll ae fSulawesi. of Lakes 273:2209–2217. Malili B the London in of Society silversides Royal sailfin the from of evidence Proceedings Dreamponds: Wallaces’s in hybridization sapidus otePooo.Lwec K) le rs.p371–656. p Press. Allen (KS): Lawrence Protozoa. the to 37:765–792. Research Freshwater and Marine of Journal of taxa Asian 84:293–315. Southeast in morphology Dcpd,Mjde.Mrn ilg 113:281–286. Biology Marine Majidae). (Decapoda, 41:163–173. and karamani 64:217–244. Edinburgh of Society clg rgesSre 98:107–113. Series Progress Ecology 5:71–73. eit eBiologı de Revista aia rnua e ageod estuario de cangrejo del branquial cavidad hmesof chambers 40:1983–2000. History Natural Caridina Cutca eaoa rmteMll ae fSlws Idnsa.Zooice negr245:167–191. Anzeiger Zoologischer (Indonesia). Sulawesi of lakes Malili the from Decapoda) (Crustacea, otndcrab portunid 73:643–684. Crustaceana ciliates. Peritrich 73:1205–1237. Crustaceana ciliates. Suctorian bernhardus 25:382–392. Estuaries area. epibenthic 20:713–721. History Natural of Journal Scuticociliatida). cec 3:275–354. Science rs-ae shrimp, fresh-water 7:1–226. ˇic ˇ .15.Osrain ntecv Suctorium cave the on Observations 1956. J. mhlpu yuianus Amphileptus nF otr .17.Uaneasei eRtfr Bdelloidea, Rotifera de specie nueva Una 1973. R. Montero F, ´n .FradzLbrn .vnRintelen von K. & Fernandez-Leborans G. tBafr,NrhCrln.Blei fMrn cec 64:513–533. Science Marine of Bulletin Carolina. North Beaufort, at Cutca eaoa tia)fo h aiilk ytmo uaei(noei) ora of Journal (Indonesia). Sulawesi on system lake Malili the from Atyidae) Decapoda, (Crustacea, .s.uddrnEtikigkes.Blei fteAaeyo cecso egae6B:45–56. Belgrade of Sciences of Academy the of Bulletin Entwicklingskreise. deren und sp. n. rmSoln.EtaieCatladSefSine66:35–54. Science Shelf and Coastal Estuarine Scotland. from oao fluviatile Potamon icriu depurator Liocarcinus g e auknlceFrcuggmishf Su Forschungsgemeinschaft Naturkundliche der ¨ge adlUuuy1:139–144. Uruguay del ´a Caridina o.se.(iipoa luotmtd) uoenJunlo Protistology of Journal European Pleurostomatida). (Ciliophora, spec. nov. ptoytu caridina Spathocyathus p iesa16:57–64. Sinensia sp. roaM,GmzdlAc .19.Dsrbto fclaeeiinso the on epibionts ciliate of Distribution 1997. P. Arco del Gomez MJ, ´rdoba Hrs,18) ora fNtrlHsoy38:1225–1232. History Natural of Journal 1785). (Herbst, Dcpd:Bahua.Ivrert ilg 116:171–177. Biology Invertebrate Brachyura). (Decapoda: pleprapolypoides Spelaeophrya Corbicula hsantu granulata Chasmagnathus e.e p o. utra tahdt h neneo a of antennae the to attached suctorian a nov., sp. et gen. plepratroglocaridis Spelaeophrya Mluc,Bvli,Criuia) caZoologica Acta Corbiculidae). Bivalvia, (Mollusca, he e lskess rhvfu Archiv Fluszkrebses. des ¨hlen hedsSenrbe ( Steinkrebses des ¨hle .sp., n. detetclns18:92–105. ¨dwestdeutschlands aa 81(eaoa Brachiura). (Decapoda, 1851 Dana, nprsarenicola Anophrys Caridina plepralacustris Spelaeophrya nmpschasmagnathi Anomopus ahnce piperitus Bathynectes tme.Book Vestnik Bioloski Stammer. rnatoso h Royal the of Transactions . mhlpu gui Amphileptus oaoistorrentium Potamobius aiialanceolata Caridina .s.(Ciliophora, sp. n. rHydrobiologie ¨r ncu aguiarii Inachus p n., sp. .s. ela de sp., n. o.spec. nov. Callinectes Marine . Pagurus Acineta Downloaded By: [Fernandez-Leborans, Gregorio] At: 08:02 8 January 2008 oaaK 98 oenwfehwtrscoi.Anttoe olgceJpnne 17:247–259. Japonenses 2004. Zoologicae B. Annotationes Soeroto suctoria. LR, fresh-water new Parenti Some 1938. K. Nozawa eesH 02 h oecaueo h oiea noae hcls fvldfml n eu-ru names. genus-group and family Kleingewa episodischer valid Untersuchungen of 2003. A. checklist Thalhofer annotated A, Settele Rotifera: the York: of New nomenclature 7. The Naturforschenden Volume 2002. der H. Crustacea. Verhandlungen Segers Celebes. of von biology Dekapodenfauna der The Kenntnis zur editor. Beitrag DE, 1902. E. Bliss Schenkel In: adaptive relations. drive specialization Symbiotic trophic 1983. and DM. Escalation Ross 2004. M. Glaubrecht A, Meyer bivalve freshwater AB, the of Wilson the flock Tv, species in endemic Rintelen an strategy in sessility reproductive of evolution unique of Rapid 2006. a Journal M. radiation: Glaubrecht webs. Tv, adaptive Rintelen food an planktonic of in Protozoa Anatomy 1985. 2004. RW. M. marine Sanders Glaubrecht in M, Tv, living Pace ciliates Rintelen BF, and Sherr flagellates heterotrophic EB, of Sherr ecology KG, The Porter 1989. JO. Corliss J, Larsen DJ, Patterson ize ,CiY 2006. Y. Cai K, Zitzler ogW ibr .18.Txnmsh nescugna uwcsiitn(rtza iipoa im Ciliophora) (Protozoa, Aufwuchsciliaten Protozoen: an troglobionte neue Untersuchungen Zwei 1935. HJ. Taxonomische Stammer 1989. N. Wilbert W, Song ufr .15.EnnusRa neues der Ein Revue Internationale 1957. Caridinen. K. der Verbreitung Wulfert geographischen und Systematik Zur 1937b. E. Woltereck of revision Hydrobiologia A interactions. 1991. J. consumer–prey Paynter on genus A, the epibiosis Warren of of revision Effects A 1997. 1986. A. P. Warren Enderlein Series ME, Progress Ecology Hay Marine peritrich aspects. epizoic M, basic the antifouling-some Wahl and with Fouling community I. zooplanktonic epibiosis. Marine lagoon 1989. a M. of Wahl Infestation 1988. L. Guglielmo A, Valbonesi otrc .13a ytmtshvrainaayiceUtruhne u Untersuchungen Systematisch-variationsanalytische 1937a. E. Woltereck ora fNtrlHsoy36:631–640. History Natural of Journal 13:485–585. Basel in Gesellschaft 163–212. p Press. Society Academic Royal the of Proceedings Indonesia. Sulawesi, 271:2541–2549. on B lakes London ancient of in gastropods freshwater of radiation 85:513–542. Corbicula Society Linnean the of Journal Biological implications. biogeographical gastropod freshwater endemic 32:409–415. Protozoology 3:185–277. Protistology in 51:10–19. Progress Research sediments. Ichtyological Indonesia. Sulawesi, Poso, Lake from Adrianichthyidae) Beloniformes: h nin aiilk ytmo uaei noei.RflsBlei fZooy54:271–276. Zoology of Bulletin Raffles Indonesia. Sulawesi, of system lake Malili ancient the 158:26–30. Ho 3:1–222. Lauterbonia Bonn. Weiher, Poppelsdorfer Universita (Germany): Ulm dissertation]. nat. rer. [Dr. 1820 Jurine tempora eatnHdoilgeudHdorpi 34:294–330. Hydrographie und Hydrobiologie Gesamten 34:208–262. Hydrographie und Hydrobiologie 50:1–57. Series Zoology History) (Natural 355:49–59. 58:175–189. intermedium Zoothamnium Ho der Su ultno h rts uem(aua itr)ZooySre 57:17–59. Series Zoology History) (Natural Museum British the of Bulletin ßasranlndrGattung der ¨ßwassergarnelen ¨hlengarnele ¨hlenasselgattung rnEiissdsRa des Epibiosis ¨ren rmacetlkso uaei noei.Booia etr 2:73–77. Letters Biological Indonesia. Sulawesi, on lakes ancient from rgoai schmidti Troglocaris aiiaspongicola Caridina Minolistra dincty roseni Adrianichthys rct(eirci,Zohmide.Bletn iZooi 3:179–183. Zoologia di Bollettino Zoothamniidae). (Peritrichia, Precht ¨dertieres drirasdrKiemenho der aus ¨dertier Tylomelania rhvfu Archiv . om und Dorm. Caridina e pce,afehae hip(rsae:Dcpd:Ayde from Atyidae) Decapoda: (Crustacea: shrimp freshwater a species, new , Vorticella rcinsrubens Brachionus Cothurnia pbotccmuiison communities Epibiontic rPoitnud 84:518–527. Protistenkunde ¨r Crtiie:Pcyhlde nSlws,Idnsaadits and Indonesia Sulawesi, on Pachychilidae) (Cerithioidea: Dcpd,Ayde.Itrainl eu e Gesamten der Revue Internationale Atyidae). (Decapoda, and aeohy monolistrae Lagenophrys Clohr:Prtihd) ultno h rts Museum British the of Bulletin Peritrichida). (Ciliophora: Clohr:Prtihd)adismrhlgclrelatives. morphological its and Peritrichida) (Ciliophora: plepratroglocaridis Spelaeophrya rza nebulosus Oryzias hevon ¨hle hebr 88afdrCladocere der auf 1838 Ehrenberg se ne eodrrBeru besonderer unter ¨sser tUm Fakulta Ulm, ¨t abrsaffinis Cambarus w e ieihs(Atherinomorpha: ricefishes new two , brdeRse-udAtidn bei Artbildung und Rassen- die ¨ber .s.vndnKenn(Pleopoden) Kiernen den von sp. n. aiiaesfr 2917 ensifera Caridina .g,n p o e nennder Antennen den von sp. n. g., n. tfu ¨t olgshrAnzeiger Zoologischer . rNaturwissenschaften. ¨r csctgn der ¨cksichtigung on brachiata Moina