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Freshwater nematodes from Loch Ness, Scotland. Part 1. The orders Tylenchida Thorne, 1949 and Rhabditida Chitwood, 1933 (Nematoda, Secernentea) PDF

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Preview Freshwater nematodes from Loch Ness, Scotland. Part 1. The orders Tylenchida Thorne, 1949 and Rhabditida Chitwood, 1933 (Nematoda, Secernentea)

) K*£33*U"&-0 Bull. nat. Hist.Mus. bond. (Zool.)67(1): 1-24 Issued28June2001 Freshwater nematodes from Loch Ness, Scotland Part The orders Tylenchida JUN 2001 I. D Thorne, 1949 and Rhabditida Chitwood, 1933 siL^rWTY (Nematoda, Secernentea). F. R. WANLESS AND R. HUNTER* DepartmentofZoology, TheNaturalHistoryMuseum, CromwellRoad, London SW75BD, UK CONTENTS Introduction 1 Nematodefauna 2 Materia] andMethods 2 Abbreviations 4 Specieslist 4 Descriptionsofspecies 5 Acknowledgements 22 References 22 SYNOPSIS. DuringasurveyofthenematodesofLochNess.Scotland(Project Urquhart 1993-94)94nematodespecieswere identifiedfrombenthiccoresamplescollectedfromtheprofunda!(90-211 m)andfromrivermouthareas(<0.5m).Thispaper presentsataxonomicreviewofthenematodesofthetwoordersTylenchidaThorne. 1949andRhabditidaChitwood, 1933.Thirty five species are described and figured; some morphological features are illustrated by Normarski interference-contrast microscopy. INTRODUCTION and sand (63-125 urn. 36.4%) (Smith B. D. etal, 1981). Water levels remain relatively constant throughout the year al- thoughstream levelsvarybecauseofsnowmeltinspring,butmore This study isbasedon nematodescollected in LochNess, Scotland importantly because of fluctuations in rainfall. The loch has a during Project Urquhart (1993-94). The first part presented here catchment areaof 1775 km: providing 84 m3 S"1 mean inflow with dealswithspeciesthatbelongintheordersTylenchidaThorne. 1949 themajorityofthecatchmentbeingroughgroundorforest,andonly and RhabditidaChitwood, 1933. The second part, now in prepara- 7.5 km: urbanized. Surface water is slightly acidic with an annual tion, will cover species of the remaining orders. The results of mean pH 6.7; it has low conductivity 30 uS cm-1 at 20°C and is ecological studies(R. David,1998-Ph.D. thesis)will be published stainedwithhumics.Lowconductivitylevelsinlakewaterareoften elsewhere. The following briefprofile ofLoch Ness is essentially associatedwithdarkcolourationandahighconcentrationoforganic basedontheliteratureMaitland(1981)andShine&Martin(1987). compoundsbecausetheyhavebeendrainedthroughswampy/peaty LochNess is the largest body offreshwater in the United King- regions.Thetotalorganiccarbonconcentrationsofthesurfacewater dom. It runs in a north-east/ south-west direction and is 39 km in is 3 mg Cl_l Maitland (1981). In many systems oflake classifica- length,butonly 2.4kmat itswidestpointwithanaveragedepthof tion. Loch Ness falls within the nutrient-poor Spence (1967) and 132 m. A new maximum depth of 239.5 m was found during a oligotrophiccategoryVollenweider(1968). However,Baileyetal., Project Urquhart sonar survey. The loch basin has in most places 1981 debate whether Loch Ness along with Loch Awe should be very steep, sometimesprecipitous shelvingsides,thus ithas avery described as dystrophic (acid), a category considered by Jarnefelt smallshorezoneforitssurfaceareaMaitland(1981).Theflatbedof (1958)assupplementaryto.ratherthancomparablewitholigotropic the loch is divided into two basins of 220 m depth as a result of andeutrophiccategories. fluvial deposits of material opposite the entrance to the River The dissolved oxygen content of the Loch Ness water column Foyers. The floor is extensively covered by black lake sediments was, along with other lochs studied, consistently found to be over whichoccupiesover50%oftheareaofLochNessgreaterthan 150 80% saturated (often approaching 100%) even during periods of m depth. Shine & Martin (1987) found that a 4 m core onlyjust stratification Smith, I. R. etal, (1981 penetratedtoclay,suggestingthattheorganicsedimentisatleastof UndertheclassificationsystemofYoshimura(1936),LochNess thatdepth. Sedimentsanalysedfromasinglecoreobtainedfromthe isawarmmonomicticlakewhichmeansthatthetemperaturenever profundal area of Loch Ness were found to consist of fine size falls below 4°C at any depth, there is one circulation ofthe water fractions withclay (< 39 urn, 45.8%), fine silt (39-63 urn, 18.1%) column every year in winter and the loch is stratified during the 'R.HunterneeR.David. ©TheNaturalHistoryMuseum,2001 F.R.WANLESSANDR. HUNTER summerMurray&Pullar(1910).Fromlateautumntothespringthe MATERIAL AND METHODS lochisthesametemperaturethroughout,withprogressivedevelop- mentofstratifiedconditionsfromMaytothefollowingautumn.The temperaturerangeinmidsummeristypically 14.5°Catthesurfaceto Samplingand fixation 5.5°C at220m (Mackal & Love, 1969; Murray& Pullar, 1910). Profundal sampling: coresweretakenwithaBowersandConnolly Maitland (1981) gives a detailed account of human activities Multicorer deployed from the research vessel Seol Mara. This whichinfluencetheenvironmentalparametersinherentinscientific devicetakesfourcoretubes (eachof5.9cmdiametergivingacore studieseffectingLochNess.Thelochisrelativelyunpolluteddespite surface area of 27.34 cm2), allowing sediments to be recovered arable, tourist and urban influences. The Scottish Environmental withoutdisturbingthe sedimentwaterinterface. Protection Agencyhas supplieddetailsofsewerage inputstoLoch Sediments were sampled at four stations: Fort Augustus Bay, Ness which indicate that there are seven direct septic tank inputs DeepSouthBasin,FoyersPlateauandDeepNorthBasincoveringa fromsmallcommunitiesorhotelsandtouristattractions,onlytwoof range ofdepths between 91 and 211 m (Fig. 1, Table 1). At each which have monitored flows. Another septic tank inflow enters station samplingwasconcentrated within approximately 10-20m: LochNessindirectlyviatheRiverMoriston.Therearetwoactivated between 10—13th July 1993, allowing for boat drift. Eight cores sludge plants for full biological treatment, one at Drumnadrochit were collected from each station except for the Deep North Basin and another at Fort Augustus. When maximum flow is reached, whereonlyfourcoresweretakenbecauseofpoorweatherconditi- sewerage by-passes the treatment plant and discharges untreated ons.Oftheeightcorescollectedsixwereusedforfaunalanalysis;of intothewatercourse. Seasonalinformationontheloadingvariation thesefivewere sectionedintotwo, onecentimeterlayers. One was wasnotavailable. However, DrumnadrochitandFortAugustusare sectioned into five onecentimeterlayers i.e. 0-1 cm, 1-2cm, 2-3 knowntofrequentlybreachtheirconsentlimits.Thewaterregimeof cm, 3—4 and 4—5 cm. The seventh core was used to take redox Loch Ness is affected directly by an hydro-electric pump storage potentialreadingsimmediatelyonrecovery.Thetoptwocentimeters schemebasedatFoyersandindirectlybyfourmoreinthecatchment ofthe remaining core was placed in a cold box forcarbon content area.Theseinfluencesmayhaveaneffectonthefloraandfaunaof analysis. Loch Ness. Riversampling: sedimentswerecollectedbyhand withacoring tube ofthe same dimensions as usedforcollecting profundal sam- ples. The cores were sectioned as above. Three stations i.e. Rivers NEMATODE FAUNA Oich, FoyersandMoristonwere selectedby surveyingtheareafor soft sediment similarin grain size to the profundal samples where So far as is known the only study ofnematodes from freshwater coretubescouldusedwithease.Sampleswerethentakenrandomly lakes ofthe United Kingdom is that ofMartin et al., (1993) who within an areaof 10m2. All samples were sublittoral rangingfrom publishedafulllistingofmacrobenthicandmeiofaunalspeciesfrom 0.08-0.39 m depth. Eighteen cores, six from each station were profundal sediments of both Loch Ness and Morar. Their list of collectedfrom7-8thAugust 1994. nematodesfromLochNessprofundalcores(>200m)althoughnot After slicing, all samples were immediately fixed using 8% exhaustiveincludesIronustemiicaudatusdeMan, Tripylaglomerans formaldehyde,atambienttemperature,givingafinaldilutionof4% grp, Tobrilus gracilis grp, Tobrilus cf. pellucidus and Dorylaimid after mixing. A trace of rose bengal was added to the fixative to 'A'. Additional species listed (Eumonhystera filiformis grp, temporarily stain the nematodes, thus making them easier to see Eumonhystera cf. longicaudatula, Ethmolaimus pratensis grp, whencounting and sorting. Aphanolaimus sp. andDorylaimus cf. stagnalis) were collected at In addition to the stations listed in Table 1, four sub-littoral lesserdepths such as 50-170m. qualitativecoresweretakenfromasandybeachatBorlumBay,Fort InadditionColes(1996)recordsChromadoritatenuisSchneider, Augustus (NH085 385)on 22July 1992fromdepthsof0.10-0.38 1906 andAphanolaimus ? aquaticus Daday, 1894 from 'trickle of m. Some ofthe nematode species founddid notreoccurduringthe freshwater spring from beside footpath from Goyes Hotel, Foyers 1992-93surveys,sowehavetakenthisopportunitytoincludethem Harbour, Loch Ness, Scotland'. inthispaper.They areidentifiedinthedescriptionsasBorlumBay Table 1 Summaryofsamplingstations,coredepthsandsedimentcharacteristics. Core Depth Station No's Position Date (m) Sediment FortAugustusBay(FA) 2-7 57"08.80'N4"39.45'W 10/7/93 91-105 Organicmatter(OM)floe,occasionallyferrugineousmixedwith greysiliceoussand W DeepSouthBasin(DSB) 8-13 57°12.55'N4°34.55- 11+12/7/93 194-210 Asabovebutwithslightsulphuroussmell W FoyersPlateau(FP) 14-19 57"15.80'N4°29.60" 12+13/7/93 150-161 Asabove W DeepNorthBasin(DNB) 20,22 57°16.40'N4°29.10' 13/7/93 211 211 Asabove RiverOich(RO) 31.32 NH094382 7/8/94 0.31 Sublittoral.Rivermouth.AlgaeandOMonsurface,coarsesandbelow 33,34 NH094382 7/8/94 0.20 Smallstonesandgravel 35 NH094382 7/8/94 0.33 Coarsesandandsmallpebbles 36 NH094382 7/8/94 0.17 Coarsesand RiverFoyers(RF) 40^15 NH211 495 7/8/94 0.14-0.39 Stagnantsublittoral.50mfromrivermouth.OMandgreysiliceous sandcontainingblackstreaks. Strongsulphuroussmell RiverMoriston(RM) 50 NH211 495 8/8/94 0.15 100mfromrivermouthSandbetweenboulders (accountoffulldataset 51,52 NH211 495 8/8/94 0.100.08 Coarsesandsmallpebbles inprep.) 53 NH211 495 8/8/94 0.20 Sandandgravel,somepebbles 54 NH211 495 8/8/94 0.15 Coarsesand 55 NH211 495 8/8/94 0.16 Sandandgravel . FRESHWATERNEMATODESFROMLOCHNESS R. Ness 200m contour Farigaig R. Foyers 200m contour 1 RiverOich (RO) 2. FortAugustus Bay (FA) 3. RiverMoriston (RM) 4. DeepSouth Basin (DSB) Direction offlow 5. RiverFoyers (RF) 6. Foyers Plateau (FP) 7. Deep North Basin (DNB) ** Activated sludgetreatment * Significantseptictank input R. Tarff Fig. 1 MapofLochNessshowingsamplingstations. (1992collection).Also,theyhavenotbeenincludedintheecologi- amounts of organic matter of very similar density to that of the cal part of this study which is based entirely on the 1992-93 nematodes.Inordertoreducesamplesortingtime,itwasdecidedto samples. subsample the remaining 1 cm core slices usingthe 'Asko sample splitter' Elmgren (1973). Twohundrednematodeswerepickedout Extractionandslidepreparation fromeachcore; iflessthan200nematodeswerepresentinthefirst Nematodes were extracted by the Ludox centrifugation-floatation subsample, further subsamples were extracted until the required technique(Persmarketal., 1992).However,afterextractingthefirst number was found. Extracted nematodes were then dehydrated to core it was found that the nematodes were still obscured by large glycerineby the Seinhorst method (Seinhorst 1962). 1 F.R. WANLESSANDR. HUNTER Figures, photographs and measurements MiculenchussalvusAndrassy, 1959 + Specimens were examined, figured and photographed with an Ol- Tylenchus rexAndrassy, 1979 - + ympus BH-2 interference contrast microscope fitted with camera Tylenchinae sp. + + lucida. The drawings areoriginal andbasedentirely on Loch Ness specimens. Allabsolutemeasurementsofnematodesaregiven um. FamilyANGUINIDAENicoll, 1935 Sedimenthorizon Thiscanbetakenas0-1 cmunlessastatementismadetothecontary Ditylencluts sp. + — in the descriptions. Abbreviations used Family BELONOLAIMIDAEWhitehead, 1960 a = body length-fgreatestbody width b = body length-fdistancefromanteriorendtojunctionof Geocenamuscf. nanus (Allen, 1955) + esophagus and intestine Tylenchorhynchus sp. - + b' = body length -f distance from anterior end to posterior endofoesophageal glands c = body length -ftail length Family HOPLOLAIMIDAE Filipjev, 1934 c' = tail length -fbody widthatanus L = totalbody length Helicotylenchuspseudorobustus (Steiner, 1914) + + MB = distance ofmedian bulb from anteriorend -r oesopha- geal length x 100 R = total numberofbody annules Family HETERODERIDAEFilipjev, 1934 Roes = numberofannules in oesophagealregion. Rex = number of annules between anterior end ofbody and Meloidogyneardenensis Santos, 1968 + MeloidogynekralliJepson, 1983 - + excretory pore Rv = numberofannules between anteriorend andvulva Rvan = numberofannules between vulvaandanus Ran = numberofannules ontail Family CRICONEMATIDAETaylor, 1936 tail/V-a = tail length-fdistance betweenvulvaandanus Criconemaannuliferum (de Man, 1921) - + VV = distanceofvulvafromanteriorend-rbodylengthx 100 Criconemademani (Micoletzky, 1915) + - = distance ofvulva from anterior end -f distance ofanus Criconema sp. - + from anteriorend x 100 Discocriconemella sp. - + VL/VB = distance between vulva and posterior end of body -r Hemicydiophora sp. - + body width atvulva Ogma sp. + - SPECIES LIST FamilyTYLENCHULIDAE Skarbilovich, 1947 Paratylenchussp. - ThesystematicclassificationoftheorderTylenchidaisbasedonthat of Fortuner, Geraert, Luc, Maggenti and Raski (1987-1988) 'A reappraisalofTylenchina(Nemata)'.ThatoftheorderRhabditidais FamilyAPHELENCHOIDIDAE Skarbilovich, 1947 afterLorenzen (1981). Aphelenchoides sp. 1 + Aphelenchoides sp. 2 + OrderTYLENCHIDAThome, 1949 S* Aphelenchoides sp. 3 + FamilyTYLENCHIDAEOrley, 1880 Aglenchusagricola (de Man, 1884) + OrderRHABDITIDA Chitwood, 1933 CoslenchuspolonicusBrzeski, 1982 + Family RHABDITIDAEChitwood, 1933 CCoosslleenncchhuuss sspp.. 21 -+ cf. Caenorhabditissp. + Filenchuscf. discrepans (Andrassy, 1954) + Filenchusfaeultativus(Szczgiel, 1969) Filenchusthornei(Andrassy, 1954) + FamilyDIPLOGASTERIDAE Micoletzky, 1922 Filenchussp. 1 - Diplogasteritusnudicapitatus(Steiner, 1914) + Filenchussp. 2 — Filenchussp. 3 - Lelenchusleptosoma (de Man, 1880) Family CEPHALOBIDAEFilipjev, 1934 Lelenchus sp. - MalenchusacarayensisAndrassy, 1968 + Eucephalobusoxyuroides (de Man, 1876) Malenchusbryophilus (Steiner, 1914) + Eucephalobussp. Malenchuspachycephalus Andrassy, 198 Malenchuspressulus (Kazachenko, 1975) * S =. Sublittoral, P= Profundal. FRESHWATERNEMATODES FROM LOCHNESS DESCRIPTIONS OF SPECIES genitalbranch 123-146umlong;spermathecapartlyoffset,usually poorly defined, ovoid to elongate-ovoid, sometimes appearing double, length variable 20-25.6 um; postuterine sac lacking. Aglenchus agricola (de Man, 1884) Andrassy, 1954 Males, (n=6). L=591.5 |jm (469-670); a= 38.9 (33^3); b (n (Fig. 2) - 4) = 6.2 (5.6-6.7); c = 3.1 (2.9-3.3); c' = 17.5 (15.3-19); Materialexamined. FortAugustus Bay: 92, 105 m;cores2, 3; gubernaculum(n=5)=6.4um(6.3-6.4);spicules(n=5)= 14.7um 29$. Foyers Plateau: 150-161 m; cores 14-17; 29946*6*. River (14-15.3); tail = 193.8 urn (153-219). Foyers: 0.14-0.39 m; cores40, 43,45: 299, 266. Similartofemale.Annules 1.6-1.96umwideatmid-body. Spear 12-12.8umlong,withroundedknobsabout2.6umwide.Oesopha- Horizon. Core 2 (2-3 cm), core 14(3^4cm), core40(2-3 cm). gus 81.6-101 um long; median bulb ovoid with usually distinct F65.6e)6m;(a6Vl.e3s-=6,.797).;(6nc(==7653)...53-L(738=..246-)53;0.4tu)a;rinlc/'(V6=-2a51-4=.65617.(9)1:2(.1a7.-=71-235.1.08)():;26tV.a4i=l-3=545..1491)4;(5\b3i=m- aovsaclycvmuempseytiartnig4c6a2l^4;-—82b%u5r%osfaoofaedbsaoondpayhlal3gee2nga-tl3h8;l.ec4nlgoutahmc.alTleolsnitgpisswtiu1tb4eh4--ls1imk5oe3,o.ts6houmomerwlvhoeanrtgy finely crenate margins. (188-200). Headcontinuous, notannulated. Body annules 1.6-1.8 urn wide Distributionandhabitat. Cosmopolitan,invarioussoils,lives at mid-body. Lateral fields with three lines; outer lines sometimes on ornearplant roots, in mossesandfreshwater. appear double, whereas the inner line which is usually weakly expressed,couldbeinterpretedastwolinesverycloselysetasthey REMARKS. Morphometric measurements of this population fall midwaybetweenthoseoftwovery similarspecies,A. agricolaand aSpppeeaarrmaosdeeritahteerlygulrloybuosrt,ri1d1g.e0--li1k2e.8(duerpnenlodnegn,tboansaolptkincaolbssedtitsitnigns)c.t A. muktii Phukan & Sanwal, 1980. A. tnuktii, according to the about3.2 urn wide. Oesophagus92.7-101.7 um long; median bulb originalauthors,isdistinguishedfromA. agricola 'inhavinganon- ovoidwithusuallydistinctvalvesat44-50%ofoesophageallength. annulated lipregion, 3 incisures in the lateral field,andavery long Vulva with flaps about 5.0 um long covering about three annules; filiform tail. (A. agricola has 3 annules in the lip region, 1 lateral vagina curved anteriorly with swollen pyriform walls; anterior incisuresandatailwhichisnotasfiliformasinA.muktii).'Ofthese charactersthefirsttwoaremistakenbecauseinA.agricolathehead is not annulated and the lateral field is comprisedofthree lines. In practiceA. muktiiisdistinguishedfromA.agricolaby itsverylong filiform tail. Tail length inA. agricola rangesfrom 134to 179 um, whereas in A. muktii the range is 178-276 um (Geraert & Raski. 1988). Femaletaillengthofthispopulationvariesbetween 153-197 um. Br/eski (pers. comm.) notestail lengthsofA. agricola varying from 134-208um.Inmalesofbothspeciestherewouldappeartobe adistinction between tail lengths and ratioc'. Tail length 140-169 um inA. agricola vs 178-330um inA. muktii:ratioc' 13-15 vs22- 25 in A. muktii. However the Loch Ness males bridge the gap in respect of tail length 153-219 urn and fall between the range of values in the caseofratioc', c'= 15.3-19. Theoverlapintaillengthsandothermorphometricmeasurements suggest that the specific status ofA. muktii is doubtful. This popu- lation is therefore identifiedasA. agricola. Coslenchuspolonicus Brzeski, 1982 (Fig. 3) Material examined. Deep South Basin: 204 m; core 9; 1 9. RiverFoyers: 0.22 m; core40; 1 9. Horizon. Core40, (1-2 cm) Females, (n=2). L=635, 744um; a=41.5,46.5; b=5.4,6.2;c =5.8,5.4;c'=ll,12;V=65.8,64;V'=79.6,80;Rex=47,52;Roes =62,63;Rv=203;Rvan-44.49; Ran=55,65;tail/V-a= 1.0;tail = 110, 137.6 um. Head more or less continuous with body contour, 6.4-6.7 um wide with 4 very fine annules. Annules 2.3, 2.8 um at mid-body. Lateral fields with4 lines, the two inner lines closely spaced, thus sometimesappearingasasingleline.Cuticlewith 18-20longitudi- nal ridges (difficult tocount) excluding lateral fields. Spearrobust 15, 13.4 um long; knobs rounded, 3.8, 3.2 um wide. Oesophagus Fig.2 Aglenchusagricola(deMan, 1884).A-E,female.A,oesophageal 118, 120 um long; median bulb ovoid, valve at about 45% of region;B,posteriorendofreproductivesystem;C.tail;D,lateralfield; oesophageal length. Vulva, with lateral vulval membranes, 6.4 um E,habitus:F,malespicularregion.Scalebarsa=20|jm,b=40|jm,c= longorabout3annuleswide;vaginainclinedanteriorlywiththick, 100um. somewhat pyriform, walls; postuterine sac small; spermatheca F.R. WANLESSANDR. HUNTER Fig.3 CoslenchuspolonicusBrzeski, 1982.A-F,female.A,oesophageal region;B,lateralfield;C,posteriorendofreproductivesystem;D, habitus;E,tailtip;F,tail. Scalebarsa=20um,b= 100um. Fig.4 Coslenchussp. 1. A-E.female.A.oesophagealregion;B.lateral field;C,vulvalregion; D.habitus:E,tail. Scalebarsa=20um,b= 100 elongate, 38, 41 jam long, packed with sperms about 2.0 um in um. diameter. Distance fromvulvatoanus 107-128 um. Tail tip finely roundedwithannulesinonespecimen;roundedwithoutdiscernible sloping posteriorly, 2.6, 3.0 pm wide. Oesophagus 96.3, 94.7 um annules inother. long; median bulb ovoid; valve well developed at about 46% of Distribution and habitat. 'Collected in several places in Po- oesophageal length. Vulva with lateral vulval membranes 7.0, 7.6 urn or 3 adjacent annules long; vagina perpendicular or slightly landandinAndreevskoe,Karelia,USSR,alwaysinlowpeatsoil,in inclined anteriorly, with thick walls; postuterine sac not evident; small numberofspecimens' (Brzeski, 1987). spermatheca without sperm. Distance from vulvato anus 100, 120 Remarks. These nematodes were not originally identified as C. um. Tail tip finely rounded. polonicusbecausetheyhavefewercuticularridges(18-20vs26in Remarks. UsingBrzeski's (1987) andGeraert & Raski's (1988) C. polonicus). However, Brzeski (pers. comm.) recognized these fareemalneos,otahnedrssuigggneisfticeadntthedyifafeprpeenacreesdbteotbweeeCn.ptohleosneicfuse.maAlessthaenrde ktheeysm,itdh-ebsoednyemanantuoldeessacroemmearcglionsaelltoyCt.ooanwdirdaess2.y6i,B2r.z7esvksi2,.11-928.75,buumt in C. andrassyi, alsothe numberofannules betweenthe vulvaand dtheesclroipwteironnsuomfbeC.rpooflocuntiiccuusla(rBrrziedsgkeis,l1i9es82w,it1h9i8n7)thietirsaansgesuomfevdartihaa-t theanteriorendofthebody aretoolow, 136, 138 vs 157-186in C. andrassyi. tionthatmightbe expected inthis species. Coslenchus sp. 2 Coslenchus sp. 1 (Fig. 5) (Fig. 4) Material EXAMINED. Fort Augustus Bay: 92 m; core 4; 19. Materialexamined. RiverFoyers: 0.22m; core40; 1 9. Foyers Plateau: 157 m; core 16; 19. Horizon. Core40, (1-2 cm). FEMALES, (n=2).L=520,527umV; a=23.2,24.7;b=5.5,5.4;c FVemale. (n= l).L=546pm;a=24;b=4;c=5.4;c'=7;V=65; -5.6,6.1;c'=7.2,6.9;V=64,65; =77,79;Rex=37,42;Roes =79.5;Rex=40;Roes=63;Rv= 152;Rvan=41;Ran=66;tail/ =42, 46; Rv = 136, 138; Rvan = 40, 37; Ran = 59, 53; tail/V-a= V-a= 1.1; tail = 102 pm. 0.92,0.93; tail = 93.4, 86.4 um. Head narrower than adjacent body; 7.7 um wide with 5 Head offset in one specimen; 7.0, 7.5 um wide with 4 annules. annules.Annules 2.7 pm wide at mid-body. Lateral fields with 4 Annules about 2.6, 2.7 urn at mid-body. Lateral fields with four lines. Cuticle with longitudinal ridges and additional short ridges lines. Cuticlewith 14longitudinal ridges (excluding lateral fields). distributed irregularly, giving an estimated total of 15-16 ridges Spearmoderatelyrobust11.5, 12.8umlong,knobsrounded,slightly excluding lateralfields. Spearslender, 13.5 pm long; knobs small, FRESHWATERNEMATODES FROM LOCHNESS A/1 1 ill Fig.5 Coslenchussp.2.A-E,female.A,oesophagealregion: B. lateral field;C,vulvalregion;D.tail;E.habitus. Scalebarsa=20pm.b= 100 Fig.6 Filenchuscfdiscrepans(Andrassy, 1954). A-E,female. A. pm. oesophagealregion: B. lateralfield;C,reproductivesystem;D.tail;E, habitus. F, malespicularregion.Scalebarsa=20pm.b= 10pm,c= roundedabout2.6umwide.Oesophagus 136umlong;medianbulb 50pm. ovoid,valvewelldevelopedat45.5%ofoesophageal length. Vulva withlateralvulvalmembranes,7.6umlongorabout3annuleswide; mid-body. Lateral fields with two lines. Spearslender, 6.0-7.6 pm vaginaslightlyinclinedanteriorlywiththickenedwalls;postuterine sac short; spermathecaempty. Distance from vulvato anus 91 um. longwithsmallbasalknobs,about 1.2pmwide.Oesophagus69-92 Tail damaged, but annules upto finely rounded terminus. pmlong;medianbulbsomewhatfusiform,valveweakat35-42%of oesophageal length. Anterior genital branch 68-156 pm long. Remarks. Although the tail of this specimen is damaged, it is Spermatheca axial, oblong with rounded sperms about 1.3 pm in clearlydifferentfromtheotherspeciesofCoslenchusfoundinLoch diameter. Postuterine sac 7.0-13.4 pm long. Vagina more or less Ness. Ratios effected by this damage are considered to be close to perpendiculartobody axis. measurementsthatwouldhavebeenobtainedhadthespecimenbeen undamaged. It keys out close to C. pastorAndrassy, 1982, but the MALE, (n = 1). L = 453 pm; a = 47; b = 5.0; c = 4.5; c' = 14.5; morphometries are apoorfit. gubernaculum = 3.0pm; spicules = 13.8 pm; tail =101 pm. Similartofemale,exceptbodymoreslenderwithnarrowerlateral Filenchus cf. discrepans (Andrassy, 1954) Raski & fields. Testis 152 pm long, occupying 33% ofbody length. Bursa Geraert, 1987 adanal, 19 pm long. (Fig. 6) Distribution and habitat. Widely distributed in Europe with additional recordsfromMexico,Russia,USA(terrestrial soils)and Materialexamined. Fort AugustusBay: 98, 102m;cores5,6; New Guinea (terrestrial andfresh water). 299. Deep South Basin: 196, 204 m; cores 8. 10; 299. Foyers Plateau: 158,-160m;cores 14, 15, 18;399.DeepNorthBasin:cores REMARKS. This population keys out to F. discrepans. The mor- 20,22;211 m.299.RiverFoyers:0.25-0.39m;cores41,42,44; 16, phology ofthe female reproductive system closely resembles that 399. figured forthe type specimen, especially in respect ofthe oblong Horizon. Core 14, (1-2 cm). spermathecainfemales.However,thecephalicregionisnotmakedly narrower than the adjacent body region as figured by Andrassy Females, (n= 10). L=429 um(337-526);a=35 (27.0-45.8);b (1954) anddescribedbyTroccoli & Geraert (1995). Brzeski (pers. =5.3V(5.0-6.1);c=3.8(3.4-4.5);c'= 14.1 (12.8-17.6);V=60(57- comm..)suggeststhatthesenematodesshowaffinitiestoF.misellus 64); = 82.3 (80.5-84); tail/V-a = 2.1 (1.7-2.5); tail = 109 urn (Andrassy, 1958),butinFmisellustaillength(28-71 pm)andratio (91.4-140). tail/V-a (0.7-1.6) are different (Brzeski, 1979); tail shape also Head3.8-5.0pmwideatbase.Annules 1.2um(0.8-1.6)wideat differs. F.R. WANLESSANDR. HUNTER Filenchusfacultativus (Szczygiel, 1969) Raski & Geraert, species of Filenchus, the same range is also given by Karegar & 1987 Geraert (1988) in their summary of characters distinguishing F. facultativus from other very similar species, whereas Torres & (Fig. 7) Geraert (1996) record spearlengths of7.5-9.0 um in apopulation Material examined. Fort Augustus Bay: 91, m; cores 2, 5; describedfromArgentina. 3$?. Horizon. Core 2, (2-3 cm). Filenchus thornei (Andrassy, 1954) Andrassy, 1963 Females. (n=3).L=515um(434-557);a=36(35-37.8);b=6.1 (Fig. 8) V(6.0-6.3);c=4.0(3.9-4.1);c'= 13(11.6-14.5);V=63.2(62-64.6); Material examined. Deep South Basin: 161 m; core 19, 19. =84.8.(83.7-85.6);tail/V-a=2.1 (2.0-2.2);tail= 127um(105- RiverFoyers: 0.22m;core40, 12. FoyersPlateau: 159m;core 14, 142). 1<J. Head about 4.6 um wide at base with 2-3 annules. Amphids sinuous. Annules 1.7 um(1.5-1.9) atmidbody. Lateralfieldswith Horizon. Core 14, (2-3 cm); core 19, (1-2 cm); core 40, (2-3 twolines.Spearslender9.0um(n=2)longwithsmallbasalknobs, cm). baublobutov1o.id2,umvawlivdeew.eaOeksaotp3h9a^g1u2s%8o3f.8oeusomph(a7g3e.a6l-9l1e)ngltho.ngE;xcmreedtioarny F=e5m.a0,le4.s1,;c'(=n=142.)5..L18=; 9V97=,519,00536;uVm;=a=733,87,34.45.;7t;abil=/V6-.a9,=60..59;5,c pore62.4,68.7um(n=2)fromanterior.Anteriorgenitalbranch 145 1.16; tail = 131.5, 191.5 um. um (127-159) long. Spermatheca axial or slightly offset, elongate Head about 7.6, 8.0 um wide at base with 3-4 annules; height 23,26umlongwithroundedsperms(n=2).Postuterinesac7.7-12 about3.5 um. Amphidsnotobserved. Annules 1.3-1.6 um wideat um long. Vagina 7.0-8.0 urn long, more or less perpendicular to mid-body. Lateral fields with four lines, inner lines very faint and body axis. Tailtippointedtofinely rounded. notalwaysevident. Speardistinct,moderatelyslender, 13.4, 12 um Distribution and habitat. A widespread species found inter- long with smallbasal knobs about 2.0 urn wide. Oesophagus 144- restrial soils. 155 um long; median bulb more or less ovoid, valves moderately distinct at 39.7, 42% of oesophageal length. Excretory pore Remarks. F. facultativus is a variable species and these nema- sclerotized at 112, 122 um from anterior. Anterior genital branch tdoedsecsriabreediwnitthheinlittehreatruaren.geHoowfevmeera,sutrheemsepneatrsatof9.o0tuhemrlpoonpgulisatatiotnhse 4la3r2tourbno(dny=a1xi);ssapbeorumtat1h3e.5cau4m2luomngl.onTagi,loftfispeft.inVealygipnoainpteerdp.endicu- extreem range recorded for this species (6.0-9.0 um.). A range of 6.0-8.0 um is given by Raski & Geraert (1987) in their key to - F Fig.7 Filenchusfacultativus(Szczygiel, 1969).A-G,female.A. Fig.8 Filenchusthornei(Andrassy, 1954).A-E,female.A,habitus;B, oesophagealregion;B,headshowingamphid;C,posteriorendof posteriorendofreproductivesystem;C,tail;D,headregion;E, reproductivesystem;D,lateralfield;E,reproductivesystem;F,tail oesophagealregion.F,malespicularregion. Scalebarsa=200urn,b= terminus;G,habitus.Scalebarsa= 10urn,b=30um,c= 100um. 30urn,c=20um. ) FRESHWATERNEMATODES FROM LOCH NESS 9 Male, (n= 1). L=960pm; a=44.2;b= 7.1; c =5.1; c'= 12.8; (56-73) from anterior. Oesophagus 85 pm (74-94.2) long; median gubernaculum =6.0 pm; spicules = 22.6 urn; tail = 188 pm. bulb with poorlydevelopedvalvesat40.4% (35^15.6)ofoesopha- Similartofemale;bodyslightlysinuousandcurveddistally,also geal length. Anterior genital branch 163 pm (115-198) long. narrowed in region ofcloaca- possibly an artefact as the body is Spermathecaoffset,27.7pm(22.4-32)long.Postuterinesac8.9pm slightly twisted in this region. Bursa 38 urn long, with crenate (8.3-9.6) long. Vagina7.0-7.7 pm long, gently inclined anteriorly borders; cloacal lips protruding. Tail tip finely rounded. with thickened walls. Tail terminuspointedtofinely rounded. Distribution and habitat. Cosmopolitan, found in damp soils Remarks. The specific identity of these females is uncertain. and freshwaterhabitats. They show affinities with F.facultativus described above and may REMARKS. ThisspecieswasrecentlyredescribedbyBrzeski(1997 bleoncgoenrsapencdiftihce.waHlolwsevofert,heivnatghiensaeanreemsaltioghdtelsy tthhiecksepneeadr.is slightly and these specimens fall well within the range of measurements presented forF. thornei. The heightofthe head is perhaps slightly Filenchus sp. 2 lower, butthis is notthought tobe ofany significance. (Fig. 10) Filenchus sp. 1 Materialexamined. Foyers Plateau: 157 m;core 14, 1 9. Fig. 9 Female, (n= 1). L=625 pm; a=40.1; b=5.2: c=5.5;c'=9.0; Materialexamined. FoyersPlateau: 159m;core 14.299.Deep V =65.8; V'= 80; tail/V-a= 1.0; tail = 108 pm; North Basin: 211 m; core 22, 1 9. Head5.7 pm wideatbase. Amphidsnotobserved. Annules fine, 1.3 pm wide at mid-body. Lateral field with four lines, inner two Horizon. Core 14, (1-2 cm; 3-4cm). faint.Spearindistinct,9.3pmlongwithsmallbasal knobsabout 1.3 Females, (n=3). L=531 pm(439-615);a=40.6(39.7—41.8);b pm wide. Excretory pore 82.5 pm from anterior. Oesophagus 119 = 6.0 (5.5-6.5); c = 3.4 (3.2-3.6); c' = 18.5 (15.8-20.2); V = 58.6 pm long; medianbulbovoid, valvesweakat40.7%ofoesophageal V (57-60); =83.3 (82.6-84.5); tail/V-a=2.5 (2.2-2.7); tail = 157 length. Anterior genital branch 175 pm long; spermatheca axial, urn (132-191.5). about 22 pm long, with rounded sperms about 1.9 pm in diameter. Head 5.0-5.7 urn wide at base with two annules. Amphids Postuterinesac6.4pmlong,occupying41%ofcorrespondingbody sinuous.Annulesabout 1.6urnwideatmid-body.Lateral field(only diameter.Vaginaabout5.8pinlong,perpendiculartobodyaxis.Tail evidentintwospecimens)withfourlaterallines,theinnertwobeing terminus finely rounded. veryfaintandclosetoouters. Spearindistinct,slender,9.6urn long withsmallroundedknobsabout 1.3urnwide.Excretorypore64urn Fig.9 Filenchussp. 1.A-F,female.A.oesophagealregion;B.head Fig. 10 Filenchussp.2.A-E,female.A,oesophagealregion:B.habitus; showingamphid;C,lateralfield;D,tailtip;E,habitus;F.posteriorend C.lateralfield:D,tail;E,posteriorendofreproductivesystem.Scale ofreproductivesystem. Scalebarsa= 10pm,b= 100pm. barsa=20pm,b= 100pm. 10 F.R. WANLESSANDR. HUNTER Remarks. ThisnematodeisclosetoF. butteus(Thorne&Malek, 1968) Brzeski, 1997,butthetail is longerand differs in shape. Filenchus sp. 3 (Fig. 11) Materialexamined. FortAugustusBay: 92 m; core 3, 16. MALE, (n = 1). L=434 urn; a= 35.7; b = 6.4; c =4.4; c'= 10.3; gubernaculum = 5.8 urn; spicules = 15 um; tail = 99 urn. Head 5.0 um wide at base with 3-4 annules. Amphids not observed.Annulesfineabout0.9umatmid-body.Lateralfieldwith fourlaterallines,hardlyevident.Spearmoderatelyslender, 11.2um long with small basal knobs 1.9 um wide. Excretory pore 61.4 um from anterior. Oesophagus poorly preserved, about 68 um long; median bulb ovoid, valves not observed, centre at about 59% of oesophageal length. Testis 138 um long, occupying 32% ofbody length,bursaadanal,about 19umlong,withfinecrenatedmargins. Tail tippointed. Remarks. Noconclusion canbe reachedon the specific identity ofthis specimen. It is close to F.thornei, but differs by its shorter body length, spicules andtail. Lelenchus leptosoma (de Man, 1880) Raski & Geraert, 1985 (Figs 12, 36B) Materialexamined. FortAugustusBay:92m;core3; 1 9.Deep SouthBasin: 194m;core 13; 1 9. FoyersPlateau: 159, 160m;cores Fig. 12 Lelenchusleptosoma(deMan. 1880).A-E.female.A, 14, 15; 399. oesophagealregion: B.headshowingamphid;C.posteriorregion:D. Females, (n = 5). L = 546 (487-583) um; a = 46.6 (40-49.8); habitus; E.reproductivesystem. Scalebarsa=20pm.b= 10pm.c= b=5.5 (4.9-6.3); c = 3.1 (2.7-3.35); c'=24.8 (22-27.9); V =52 60um,d= 100um. V (48-54.8); =77.6(75.5-78); tail/V-a=2.2 (1.9-2.5);tail= 180 um (156-215). Cephalic region narrowed dorso-ventrally. Cuticle smooth by light microscopy. Amphids sinuous. Lateral field absent. Spear slender, 7.7-9.3 um long with small rounded knobs about 1.4 um wide. Oesophagus 88-108 um long; median bulb spindle-shaped with valves at 41-43% of oesophageal length. Anterior genital branch 96.7-121.5 um long. Spermatheca more orless bispherical with rounded sperm (absent in one specimen) about 1.3 um in diameter. Postuterine sac not evident in any specimen, possibly collapsed. Vagina about 5.5 um long with thin walls, curved anteriorly. Distributionandhabitat. Cosmopolitan;soilsandfreshwater. Remarks. These specimens, for the most part, agree with the description andmorphometriespresentedinthe literature(Raski & Geraert, 1985). Minor differences slightly extending the range of severalratios are notconsideredtobe ofany significance. Lelenchus sp. (Figs 13. 36A) Materialexamined. DeepSouthBasin: 195,204m;cores9, 11; 299. FoyersPlateau: 157, 159 m; cores 14, 16; lcJ, 1 9. FEMALES, (n=3).L=620(512-763)um;a=47.4(42-54.5);b= Fig. 11 Filenchussp.3.A-E,male.A,oesophagealregion:B,lateral 6.0(V5.5-6.5); c=3.5 (3.3-3.8); c'=22.8 (21.4-24); V=57.6(57- field;C,spicularregion: D.habitus;E,tailandspicularregion.Scale 58); =81 (80-82);tail/V-a=2.1 (1.8-2.4);tail= 173 (154-200) barsa=20pm,b= 100pm. um.

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