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Baujardia Mirabilis Gen. N., Sp. N. from Pitcher Plants and Its

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Baujardia Mirabilis Gen. N., Sp. N. from Pitcher Plants and Its Nematology,2003,V ol.5(3), 405-420 Baujardia mirabilis gen. n., sp. n.from pitcher plants andits phylogenetic position withinPanagrolaimidae (Nematoda: Rhabditida) 1; 2 1 Wim BERT ¤, Irma TANDINGAN DE LEY , Rita VAN DRIESSCHE , Hendrik SEGERS 3 and Paul DE LEY 2 1 Departmentof Biology,Ghent University, Ledeganckstraat 35, 9000 Gent, Belgium 2 Departmentof Nematology,University of California, Riverside, CA 92521,USA 3 FreshwaterLaboratory, Royal Belgium Institute for Natural Sciences, V autierstraat29, 1000 Brussels, Belgium Received:16 September2002; revised: 13 January2003 Acceptedfor publication:13 January2003 Summary – Measurements,line drawings and scanning electromicrographs are provided of Baujardiamirabilis gen.n., sp. n., isolated frompitcher uidof Nepenthesmirabilis fromThailand. The new genus differs from all known nematodes in having two opposing andoffset spermatheca-like pouches at the junction of oviduct and uterus. It also differs from most known Rhabditida in having four cephalicsetae instead of papillae.Phylogenetic analysis of small subunit rDNA sequencedata robustly places the new genus within Panagrolaimidaeas asistertaxon to Panagrellus .Theseunusual nematodes resemble Panagrellus inbodysize (1.8-2.7 mm infemales, 1.3-1.9mm inmales) and in the monodelphic, prodelphic female reproductive system with thickened vaginal walls and prominent postvulvalsac. However, they differ from Panagrellus inthecharacters mentioned above, in their comparatively longer stegostom and intheshape of themale spicules. Because of its aberrant characters, inclusion of this new genus in Panagrolaimidae requires changes tothefamily diagnosis. Keywords – Nepenthes,newgenus, new species, Panagrellus ,phylogeny,taxonomy, Thailand. Duringa visitto Tung-Kai Botanical Garden in Trang Materials andmethods Province,near the Isthmus of Kra, SouthernThailand, oneof us (Hendrik Segers) sampledthe liquid con- SAMPLING tainedwithin pitchers of thecarnivorous plant Nepenthes mirabilis (Lour.)Druce, in order to obtain some of the Sampleswere collectedby lteringthe content of peculiarrotifers recorded from suchhabitats. Except for some50 pitchers through a 50 ¹mplanktonnet and someunidenti able bdelloids, no rotiferswere found,but were immediately xedby adding one tenth of the thesamples contained abundant material of aremarkable samplevolume of concentratedformalin (36%) at ambient newnematode genus together with some diplogastrids. temperature. Nepenthesmirabilis was encounterednear Someof the unique morphological characters of this theend of thetrail through the primary swamp forest of newgenus make its placement dif cult. In order to ob- TungKai Botanical garden. A singlebulk sample was tainadditional information about the identity of the ne- collectedon 18 July 1999. Several additional samples, matodeand its position in relation to known taxa, we includinga samplepreserved in 70% ethanol, were conductedmolecular phylogenetic analyses based on the collectedon 13January2002. smallsubunit (SSU) ribosomalDNA. Inthis paper, we describethis new genus and species from Nepenthes and MORPHOLOGICAL OBSERVATIONS presentan emended diagnosis of thefamily Panagrolaim- For lightmicroscopy, nematodes were processedto an- idae. hydrousglycerol (Seinhorst, 1959) and mounted on alu- miniumslides with double cover slips. Measurements and * Correspondingauthor, e-mail: [email protected] © KoninklijkeBrill NV ,Leiden,2003 405 Alsoavailable online - www.brill.nl W. Bert et al. illustrationswere preparedfrom cameralucida line draw- mony(MP) andmaximum likelihood (ML) algorithms ingsusing Olympus BX50 and BX51 DIC microscopes implementedin P AUP* 4b10(Swofford, 2002).During andthe drawings were preparedusing Illustrator ® 8.0 theanalyses, trees were rootedusing Plectus. Gaps were software(Adobe Systems, Mountain View ,CA,USA). alwaystreated as missing data. T oestimateoverall sup- For scanningelectron microscopic (SEM) observation, portfor monophyly,NJ distancecalculations were per- glycerine-embeddednematodeswere rst transferredinto formedwith 3000 bootstrap replications with corrections adropof glycerine,gradually transferred to distilled wa- for multiplesubstitutions using Log Determinant distance terover a periodof 4h,thendehydrated in a 25,50, 75, measure(Lockhart et al.,1994).MP heuristic analysis 95,and 100% ethanol series at 2hintervals,followed by was performedwith 3000 bootstrap replicates and without anovernight dehydration in 100% ethanol. Dehydrated bootstrappingwith 100 replicates of randombranch addi- nematodeswere critical-pointdried with CO 2 (Wergin, tion.Finally, a non-bootstrapheuristic ML searchwas also 1981),sputter coated with 20 nmgold,and examined with performedwith 100 replicates of random branch addi- aJEOL JSM-840at 15kV. tion.The trees were thencompared statistically using the Inaddition to slide voucher specimens, morphology Kishinoand Hasegawa(1989) pairwise tests with ML and ofthe new genus was recordedas video clips that MPoptimalitycriteria, and T empletontest with MP opti- mimicmultifocal observation through a lightmicro- malitycriterion. Cladograms were examinedwith TREE- scopefollowing the V ideoCapture and Editing pro- VIEW 1.6(Page, 1996) and converted into graphic les ceduresdeveloped by De Leyand Bert (2002). The for AdobeIllustrator ® 9.0. resultingvirtual specimens are available on the web (http://faculty.ucr.edu/ pdeley/vce/Nep/nep.html). » * Terminologyon stoma morphology was adaptedfrom Baujardiamirabilis gen.n., sp. n. De Ley et al. (1995).The papilla formula follows De Ley (Figs 1-4) et al.(1999):pre- andpostcloacal papillae are separated by‘/’,withprecloacal papillae listed rst,and the number MEASUREMENTS ofpairs in each cluster of papillae separated from the adjacentcluster with a ‘+’. The‘ p’for phasmidpair is See Table 1. omitted,as we didnot observe any phasmids. DESCRIPTION MOLECULAR ANALYSIS Female Thenematodes were rehydratedin distilled water for Bodylarge (1.8-2.7 mm), almoststraight after xa- 1htoremove ethanol. DNA extractionand sequencing tion.Cuticle nelyand faintly annulated, 0.8 ¹m wide were asdescribedin T andinganDe Ley et al. (2002). atmidbody. Lateral eldnarrow (2.5 ¹mwideat mid- TheDNA sequenceof Baujardia gen.n. (GenBankac- body),starting at end of procorpusand becoming incon- cessionnumber: AF547385) was alignedwith the pub- spicuousat anal region. At midbody,lateral eldform- lishedsequences of 21othertaxa. A secondarystructure inga singleprotruding ridge ankedby ne,faint, indis- alignmentwas constructedusing Dedicated Comparative tinctlongitudinal lines. Mouth opening partly covered by SequenceEditor (DCSE v.3.4)(De Rijk& De Wachter, sixliplets each bearing a labialpapilla. Liplets continu- 1993),based on theunweighted SSU rRNA modelof V an ouswith six discrete lips separated by shallow grooves. de Peer et al.(1998)derived from base-pairingsdetected Fourshort cephalic setae, 3-4 ¹mlong,placed at base of ascompensating mutations, with equal weighting to stems subventraland subdorsal lips. Amphid opening slit like, andloops. Using Modeltest (Posada & Crandall,1988) encircledby a roundshallow incisure and situated two thisalignment was analysedto determine the appropri- discontinuousannuli posterior to setae. Stoma triradiate, atemodel of DNA substitution(GTR +I+ 0, 6 substi- relativelywide along full length (on average one third as tutiontypes) and to estimate the substitution rates, rela- wideas headdiam.). Lengths of cheilostom,gymnostom tivebase frequencies, gamma distribution shape parame- ter,and proportion of invariablesites. * Thegenus name is given in honor of Dr Pierre Baujard for his Usingthe parameter estimates obtained from Mod- outstandingcontributions to nematology. It istobe regarded as eltest,molecular phylogenetic relationships were recon- femininein gender.The speci c epithetrefers to the biotope this structedwith neighbour joining (NJ), maximumparsi- nematodeinhabits, i.e.,thepitcher uidof Nepenthesmirabilis. 406 Nematology Baujardiamirabilis gen.n., sp. n. from pitcher plants Fig. 1. Baujardiamirabilis gen.n., sp. n., female (holotype). A: Neckregion; B: Reproductivesystem showing granulated sperm cells (arrows);C: Headregion (super cial view); D: Stoma;E: Endof ovary;oviduct, spermatheca with two offset pouches (arrows) and beginningof uterus (arrowheads); F: Vulva-tailregion; G: Entirefemale. (Scale bars: B, G 100 ¹m; A, F 50 ¹m; C, D, E D D D 10 ¹m.) Vol.5(3), 2003 407 W. Bert et al. Fig. 2. Baujardiamirabilis gen.n., sp. n., male. A: Neckregion; B: Cloacalregion with copulatory papillae; C: Entiremale. (Scale bars: C 100 ¹m; A 50 ¹m; B 10 ¹m.) D D D 408 Nematology Baujardiamirabilis gen.n., sp. n. from pitcher plants Fig. 3. Baujardiamirabilis gen.n., sp. n., SEM studiesof female.A: En face view;B: Lateralview head; C: Longitudinalsection of stomaregion; D: Lateral eld(at middle of body); E: Longitudinalsection at the vulval region; F: Vulvalateral view; G: Anusventral view;H: Middleof tail.Arrowheads: cephalic setae. Arrows: digitate stoma projections. (Scale bars: C, E,F 10 ¹m;A, B,D,G,H D 1 ¹m.) D Vol.5(3), 2003 409 W. Bert et al. Fig. 4. Baujardiamirabilis gen.n., sp. n.,SEM studiesof male.A: Longitudinalsection of neck region; B: Longitudinalsection of stoma regionshowing the subventral sectors; C: Longitudinalsection of stomaregion showing the dorsal sector ,blackarrow projectionof subventralsector ,whitearrows basallamina; D: Cloacaland tail region, lateral view; E: Cloacalregion, ventral view; D F: Lateral D eld(at middle of body).(Scale bars: A, B,C,D,E
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