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TWO NEW SPECIES OF AINURDRILUS (CLITELLATA TUBIFICIDAE) FROM SOUTH-WESTERN AUSTRALIA, WITH NOTES ON AINUDRILUS NHARNA PINDER AND BRINKHURST PDF

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Preview TWO NEW SPECIES OF AINURDRILUS (CLITELLATA TUBIFICIDAE) FROM SOUTH-WESTERN AUSTRALIA, WITH NOTES ON AINUDRILUS NHARNA PINDER AND BRINKHURST

Recordsofthe WesternAustralian Museum 21: 1-7 (2002). Two new species ofAinudrilus (Clitellata: Tubificidae) from south-western Australia, with notes onAinudrilus nhama Finder and Brinkhurst AdrianM.FinderandStuartA.Halse DepartmentofConservationandLandManagement,P.O. Box51,Wanneroo,6946WesternAustralia,Australia Abtsract-Twonew species ofAinudrilus are described fromsouth-western Australiaand thedescriptionofA. nharnaisamended.Ainudrilusangustivasa sp. nov., from Lake Logue near Eneabba, is characterised by simple spermathecal ducts, thinvasa deferentia and lackofhairchaetae.Ainudrilus ngopitchup sp. nov., from a swamp near Kojonup, has long spermathecal ducts,broadvasadeferentiaandhairchaetae. INTRODUCTION SYSTEMATICS The freshwater representatives of the tubificid genusAinudriluswererecentlyreviewedbyFinder SubfamilyRhyacodrilinae and Brinkhurst (2000). Fourfreshwaterspecies, all GenusAinudrilusFinogenova,1982 fromAustraliaandnumerousmarinespecies,from Australia and elsewhere in the Pacific and Typespecies Caribbean, are currently recognised (Erseus, 1990, Ainudrilus oceanicus Finogenova, 1982 by original 1997; Finogenova, 1982). Two new species have designation. recently been collected from freshwater wetlands sampled during a biological survey of the Diagnosis wheatbelt and adjacent coastal areas of south- Hairchaetae presentorabsent. Chaetae ofpenial western Australia. Wetlands in this region are segmentmodified,chaetaeofspermathecalsegment threatened by secondary salinisation and usually unmodified. Vasa deferentia usuallybroad waterlogging resulting from rising groundwater and glandular, entering atria medially to caused by replacement of perennial native subapically.Atriavariablyshapedbutusuallymore vegetation by annual crops that use less water or less erect or directed posteriad, consisting of an (George et al., 1995). The biological survey is ampulla, often constricted medially and with designedtoprovide aframeworkforplarming the spacious lumen, often containing sperm, usually conservationofregionalbiodiversityin theface of leading to well developed ejaculatory ducts. this threat. Non-marine tubificids are particularly Prostate absent. Penes absent, though ejaculatory sensitive to salinity and discovery of two new ducts often eversible. Spermathecae with distinct, species in the south-west, which otherwise has and often complex ducts. Sperm loose in only one endemic tubificid {Ainudrilus nharna spermathecal ampullae. Coelomocytes large and Finder and Brinkhurst, 2000), is of conservation generallyabundant. significanceinrelationtotheaboveenvironmental Included Australian freshwater species: concerns. Ainudrilusfultoni (Brinkhurst, 1982), A. billabongus (Brinkhurst, 1984), A. stagnalis Erseus, 1997, A. MATERIALSANDMETHODS nhama Finder and Brinkhurst, 2000, A. angustivasa sp.nov.andA.ngopitchupsp.nov. Thetwonewspecieswerecollectedinsamplesof benthic invertebrates takenusingaD-frame sweep netwithameshporesizeof250pmandpreserved AinudrilusnhamaFinderandBrinkhurst,2000 in ethanol. Specimens were stained with Figure1 Grenacher's borax carmine, cleared with methyl salicylate and slide mounted in Permount®, either Ainudrilus nharna Finder and Brinkhurst, 2000: 55, wholeorwiththegerutalsegmentsdissected.Type Figure3. material is deposited with the Western Australian Materialexamined Museum (WAM) with other material held by the Department of Conservation and Land HolotypeandParatypes Management(CALM). Franikland River at Roe Road ford, 34 km NNW 2 A.M.Finder,S.A.Halse Figure1 Ainudrilus niiarna Finder and Brinkhurst: A, male genitalia of holotype; B, spermatheca of holotype; C, WAM WAM foldedvasadeferentiaof V4159andV4166;D,malegenitaliafrom 37-98;E,sectionthroughvas WAM deferensof V4156,showingglandularnatureofliningcells;F,ventral chaeta;G,dorsalchaeta.Scale lines;A-D50pm;E-G,20pm. of Walpole, Western Australia, 34°41’02''S SE of Corrigin, 32°27'29"S 117°59'53"E, 22 October 116°51'13"E,9September1996(WAM1-99to4-99). 1997; Hamersley River, Fitzgerald River National Park, 33°53'25"S 119°5r39"E, 12 September 1998; OtherMaterial Jimperding Brook at Lover's Lane, 31°35'32"S Not previously listed. All specimens from the 116°2r43"E, 28 October 1997; Lake Bryde, 30 km south-west of Western Australia, collected by the SW of Newdegate, 33°21’14"S 118'’49’26"E, 3 authors with D.J. Gale or J.M. McRae and held by October 1997; Lake Coomelberrup, 10 km SE of CALM. Calyerup Creek, west of Fitzgerald River EXimbleyung, 33°24'36"S 117°47'01"E, 05 Nov 1998; NationalPark,33°56’11"S119°03'56"E,12September Lake Dulbining, 40 km E of Narrogin, 32°54'24"S 1998; Dale River at Lupton's Bridge, 32°18'08"S 117°36'49"E, 24 October 1997; Melaleuca swamp 30 116°41'16"E, 29 October 1997; Dam on Gorge Rock, km NW of Hopetoun, 33‘='49'40"S 120°24‘20"E, 11 NewAustralianAinudrilus(Clitellata:Tubificidae) 3 September 1998; Melaleuca swamp in Paperbark records listed above show that the species is Nature Reserve, 21 km SE of Corrigin, 32°24'58"S widespreadinthesouth-west,includingthecentral 118°05'51"E, 25 October 1999; Peenebup Creek, 10 andsouthernwheatbeltandsouth-coastandoccurs km S of Ongerup, 34°06’02"S 118°32'12"E, 27 in a wide variety of wetlands. Pinder and September 1998; Lake Pleasant View, Manypeaks, Brinkhurst (2000) noted thatA. nhama was present 34°49'51’'S118°10'59"E,29September1998. atLakeWalbyringwhenthiswetlandhadasalinity Material listed in Pinder and Brinkhurst (2000) of 2.8 parts per thousand (ppt) (Halse et ah, 2000) newly deposited with WAM. Thomas Spring, but was apparently absent at a later date when south-west of Augusta, 34°21'00"S 115°09’35"E, 17 salinity was 20 ppt. However, the species was September 1996 (WAM V4159); northern tributary subsequently collected at Lake Coomelberrup and of Collier Creek on Cemetary Road, Walpole, Hamersely River at salinities of 22 and 20 ppt 34°58'30"S 116°45T2"E, 11 September 1996 (WAM respectively and is evidently tolerant of moderate 4160);FranklandRiveratRoeRoadFord,34°41'02"S salinity. 116°51'13"E,9September1996(WAMV4156-4158). Remarks Ainudrilusangustivasasp.nov. This species was described and illustrated in Figure2 Pinderand Brinkhurst(2000)butre-examinationof new and previously documented material has Materialexamined revealed some new features and intra-specific Holotype variation for some characters. Pinder and Dissected specimen. Lake Logue, 12 km SSW of Brinkhurst (2000) noted 4-7 penial chaetae per Eneabbain Lake LogueNature Reserve, 29°59'20"S bundle but up to 10 per bundle have now been 115°08'50"E, Western Australia, 27 October 1999, boobstehrvoefd.thAessnpoetremdatfohrectahle tawmopunlelwaespoefciAes.,nohnaemoar S.A.HalseandD.J.Gale(WAMV4145). sometimes lie in IX, though still with pores anteriorly on X. The vasa deferentia ofthis species Paratypes are ciliated, with the lining tissue consisting of a Fiveonslides(1 dissected,4whole-mounted)and sIiEn)gliendliacyaetrinogfgtallalncdeulllsarfialclteidviwtiyt.hTvhaeculoulmeesn(Foifgutrhee csoelvleercatilonmdaattuaraessforahnodlotiympmea(tWurAeMsV4i1n47a-l4c1o5h3o)l., vasa deferentia is generally difficult to follow and waswrongly interpreted asbeingfolded vertically Etymology in the holotype (partly because of corrfusion with Fromthe latinangustus (narrow), referring to the theovary,whichisnormallycloselyassociatedwith thinvasadeferentia. thevasdeferens)butisactuallyonlyslightlyfolded horizontally (Figure lA). The vasa deferentia vary Description from fairly straight (Figure ID) to moderately Dimensions of preserved and slide-mounted folded(Figure1C)indissectedspecimens.Theatria specimens: length 7.5-9 mm, width 0.45-0.5 mm. ofthesectionedholotypeappearfairlyerect(Figure Number of segments 42-54. Prostomium rounded. lA), whereas in dissected and slide mounted Pharynx in II and III, pharyngeal gland cells specimens (e.g. Figure IE), including specimens abundant, mostly on pharynx and oesophagus collected with the holotype, the atria appear more rather than post-pharyngeal septa. Oesophagus squashed. This is probably due to compression from pharynx to IX, stomach in X-XII (type under thecoverslip, althoughthe lowerpartofthe specimenswithnumerousparasiticciliatesattached atrium and the ejaculatory duct appear to be tointeriorofstomach wall). Coelomocytespatchily eversible (seen protruding from the male pore in distributed, particularly abundant in some pre- some specimens) so the shape of the atria will clitellar segments, ovoid (10-15 pm) and depend on the degree of contraction. Despite the granulated. Clitellum coveringposterior third ofX real or apparent plasticity of the shape and and all ofXI and XII, least developed ventrally on arrangement of the atrium and vas deferens, all XI. specimens attributed to A. nharna have the Hair chaetae absent, all chaetaebifid with upper characteristic short spermathecal duct with the teeth slightly shorter than lower, nodulus distal. constrictionclose to thepore (Figure IB),andhave Ventral and dorsal chaetae from II, 65-75 pm, 6-9 chaetae of the same size and form (Figure IG, H) per bundle anteriorly, 3-5 per bundle posteriorly. andsoareconsideredtobeconspecific. Penial chaetae of XI 4-8 per bundle, 65-87 pm, ArecordofthisspeciesfromLakeWalbyringnear with simple bent ectal ends (som.e bifid on semi- Narrogin (Pinder and Brinkhurst, 2000) was mature specimens) and indistinct distal nodulus. considered to be a northern outlier from other Penialchaetae lying in aline parallel toeach other records listed in the samepublication,butthenew and protruding through the body wall ventral to 4 A.M.Finder,S.A.Halse Figure2 Ainudrilusangustivasasp. nov.: A, malegenitaliaofholotype; B,spermathecaofholotype;C,penialchaetae; D,ventralchaeta;E,dorsalchaeta.Scalelines:A-B50pm;C-E,20pm. male pores. Chaetae on spermathecal segment ejaculatory ducts narrowing between atria and presentbutnotmodified. pores (everted in some specimens). Prostate tissue Genitalia paired. Genital pores in line with absent.Femaleporesnotobserved. somatic ventral chaetae, spermathecal pores anterioronX,maleporesmedialonXI.Malepores Remarks each a narrow slit within longitudinal depressions Thelackofprostatetissue,medialentryofthevas either side of penial chaetae. Testes and ovaries into the atria and the bipartite atria are typical of antero-ventralinXandXIrespectively.Spermsacs the genus Ainudrilus and the numerous parallel to VIII and XVII, egg sacs to XV. Spermathecal penial chaetae are at least similar to the other ampullae ovoid in IX and/or X, 210-225 pm long, freshwater forms (marine species generally have containingloosesperm,cormectedtoporesonXvia fewer, exceptforthe typespeciesA. oceanicus). The muscular ducts (130-150 x 65-80 pm). Ratio of narrowvasadeferentiaaretmusualbutalsoknown length of spermathecal ampulla to duct length inthemarineAinudriluslutulentus (Erseus, 1984)of about T.0.75. Ducts with lining tissue surrormded southernChina,whichwasattributedtothisgenus by thick circular muscle, with sparse covering of by Erseus (1990)because it is 'otherwise similar to peritoneal cells. Sperm funnels on 10/11 feed the type species' (i.e. itlacks prostate and thevas/ narrow (14-19 pm) non-glandular ciliated vasa atrial union is not apical). Rhyacodrilus simplex deferentiawhichenteratriamedially.Atria115-145 (Benham, 1903) of New Zealand also lacks pm wide, in two parts separatedby a ringofatrial prostates, which are normally present as a diffuse muscle around the middle, upper part with thick coveringovertheatriainRhyacodrilus,andhasthin, lining tissue aroimd broad ciliated lumen, lower non-glmdular vasa deferentia but the latter curl part with thick lining tissue around narrower around the atrium and enter it more apically. ciliated lumen, except for a more open unciliated Ainudrilus angustivasa differs from the remaining areaectallynearejaculatoryducts.Atriallumennot two species from Western Australia in the lack of containing sperm in type material. Eversible hair chaetae and can be distinguished from other NewAustralianAinudrilus(Clitellata:Tubificidae) 5 Australian freshwater species, which all lack hairs, muscular ducts (245-330 x 50-75 pm), ratio of by the thin vasa deferentia and the morphology of ampulla to duct about 1:1.3. Duct with narrow the other chaetae, see Brinkhurst (1982, 1984) and lumen, surrounded by thick lining tissue then Erseus(1997). circularmuscleandacontinuouslayerofperitoneal Lake Logue is afre.sh tobrackish lake fringed by cells. Junction of duct and spermathecal ampulla Melaleuca and Acacia trees. It usually dries particularlymuscularwithcircularmuscleforming seasonally,althoughoccasionallycontainswaterfor a sphincter just ectal to duct-ampulla union. several years. At the time of sampling the water Spermathecal ducts of paratype penetrating septa was moderately coloured at280TCLf, pH was 7.83 9/10and 10/11, withoneampullainIXand onein to8.18andsalinitywas1.1ppt. XI. Holotype with at least one ampulla in XI prior to dissection. Sperm funnels on 10/11 feed broad (up to 50 pm) ciliated, possibly glandular, loosely Ainudrilusngopitchupsp.nov. folded vasa deferentia. Exact position of atrium/ Figure3 vas deferens union not visible but presumably medial. Atria 85-120 pm wide, in two parts Material examined separatedbyaringofmuscletissue,entalpartwith Holotype thick lining tissue aroimd a broad ciliated lumen Dissected specimen, Ngopitchup Swamp, in and ectal part with upper half filled with lining Water Reserve 2184, 21.5 km south-west of tissuearoundnarrowciliatedlumenandlowerhalf Kojonup,33°57'27"S117°20’32"E,WesternAustralia, forming a broad unciliated lumen. Atria without coll.A.M.FinderandJ.M.McRae(WAMV4154). sperm in type specimens. Ejaculatory ducts narrowing towards pores. Prostate tissue absent. Femaleporesnotobserved. Paratype Whole-mounted specimen, collection details as forholotype(WAMV4155). Remarks Thelack ofprostate tissue,broad vasa deferentia Etymology and the bipartite atria are all typical of the genus Namedforthetypelocality. Acihnauedtraielusa.reThAe.onnlhyamotahearnAdinutdhrrieleusmtaoriponsesesspsechiaeisr {Ainudrilus brendae Erseus, 1997,Ainudriluspiliferus Description Erseus, 1997 and Ainudrilus taitamensis Erseus, Both specimens with post-clitellar segments 1990). The marine species differ from both A. mmiosusnmitnemgd ssopecliemnegntshatunIkXn0o.3w4n.mmWi(dhtolhotyopfe)slaindde tngwoopistmcahlulpastnrdaiAg.htnhpaenmiaalincthhaaettaAe. pbreerndbauenhdalseoannldy 0.46 (paratype).Prostomiumrounded.Pharynx has ventral chaetae with upper teeth much longer in II and III, pharyngeal gland cells mostly on thanthelower,andtheothertwohavelongtubular pharynx and oesophagus in IV-VI. Oesophagus atria. The long spermathecal ducts readily frompharynxtoIX,stomachinX-XII.Coelomocytes distinguish the new species from A. nhama. The sparse, ovoid (about 10 pm long) and granulated. chaetae of A. ngopitchup are also slightly shorter ClitellumfromViXtoendofXII. thanthoseofA.nhama. Anterior ventral chaetae 4-6 per bundle from II, The type locality, Ngopitchup Swamp, is a 85-100 pm, bifid with upper tooth thirmer and shallowperchedseasonalsedgeswampwithsandy slightlyshorterthanupper,nodulusdistal.Anterior clay sediment. At the time of sampling the water dorsal bundles each with 4-5 long (235-330 pm) wasmoderatelycoloured at280TCU,pHwas8.47 thinhairs and anequalnumberofcrotchetchaetae andsalinitywas0.58ppt. of similar dimensions and form to ventral chaetae but with fine pectinations on some. Penial chaetae of XI 3-5 per bvmdle with simple bent ectal ends, DISCUSSION aboutsamelengthassomaticventralchaetae,lying ApomorphiesunitingspeciesofAinudrilusarethe inparalleltoeachotherandprotrudingthroughthe lack ofprostate (presumably lost independently in body wall ventral to the male pores. Chaetae on Rhyacodrilussimplexandothergenera)andthenon- spermathecalsegmentpresentbutnotmodified. apical entry of the vasa deferentia into the atria. Genitalia paired. Genital pores in ventral chaetal The type species, A. oceanicus, has vasa deferentia line. Spermathecalpores anterior on X, male pores lining cells that are clearly glandular, as in A. medial on XL Male pores each a narrow slit in nhama, which suggested to Baker (1982) that this depression lateral to penial chaetae. Testes and specieswasnotcloselyrelatedtoA.fultoni(thenin ovaries antero-ventral in X and XI respectively. Rhyacodrilus) and Rhyacodrilus simplex, which he Spermathecal ampullae ovoid, 210-260 pm long observed to have non-glandular vasa deferentia. with loose sperm, connected to pores via long Rhyacodrilusfultoniwastransferred toAinudrilusby 6 A.M.Finder,S.A.Halse Figure3 Ainudrilus ngopitchupsp. nov.: A, malegenitalia ofholotype; B,spermathecaofholotype;C,penialchaetae; D,ventralchaeta;E,dorsalchaeta.Scalelines:A-D50pm;E-G,20pm. Erseus(1990)onthebasisofitslackofprostateand west of Western Australia, where a large non-apical atrial vas/union. Other species {A. proportion of freshwater wetlands are lutulentus and A. gibsoni) with these features, but threatened or already affected by secondary also with vasa deferentia that are not clearly salinisation and hydrological disturbance. The glandular, have now also been included in the salinity tolerances of the new species are genus (Erseus, 1990). In fact, A. fultoni has vasa unknown but both appear to be uncommon deferentia with an amorphous appearance, due to compared toA. nhama, which has been found in the thick lining tissue, and thus appears many wetlands and rivers throughout the south- intermediateinformbetweenA.angustivasaandA. westand isknowntooccurinwaterup to22ppt. nharna/A. oceanicus, whether glandular or not. Neither of the localities for the new species BakersuggestedthatA. oceanicushasatriawiththin (Ngopitchup Swamp and Lake Logue) is lining tissue because the glandular role normally threatened by salinity, as the former is perched performed by the atrial cells was being performed above the surrounding landscape and the latter by the vasa deferentia. However, A. nhama, which lies in a coastal sandplain with low salinity also has clearly glandular vasa deferentia (Figure groundwater.However,thenewspecieshighlight IF),hasatrialliningtissueaswelldevelopedasthat theconservationvalueofwetlands,suchas these, of A. angustivasa which has a thin, non-glandular that are likely to maintain a diverse freshwater vasa deferentia. This suggests that there is no faunainaregionbroadlyaffectedbysalinity. correlation between the glandular development of theatriaandvasadeferentia. Both ofthenew species are known fromsingle ACKNOWLEDGEMENTS freshwater localities in the agricultural south- The biological survey of the wheatbelt is part of NewAustralianAimidrilus(Clitellata;Tubificidae) 7 the State Salinity Strategy. Field work was Erseus, C. (1997). The marine Tubificidae (Oligochaeta) undertaken with Jane McRae and David Cale and of Darwin Harbour, Northern Territory, Australia, samples were sorted by Jane McRae, Melitta with descriptions of fifteen new species. In Hanley, Pennifold and Edyta Jasinska. We thank Ralph J.R., Caswell, G., Megirian, D. and Larson, H.K. The BrinkhurstandChristerErseusforadviceaboutthe Marine Flora and Fauna ofDarwin Harbour, Northern Territory,Australia.ProceedingsoftheSixthInternational gutprotozoainA.angustivasa. MarineBiological Workshop99-132. Museumsand Art GalleriesoftheNorthernTerritoryandtheAustralian MarineSciencesAssociation,Darwin. REFERENCES Finogenova,N.P. (1982).Aimidrilusoceanicus,novyirodi Baker,H.R. (1982). Vadicolaaprostatusgen.nov.,sp.nov., vidsemeistvaTubificidae(Oligochaeta).Zoologicheskii a marine oligochaete (Tubificidae; Rhyacodrilinae) Zhurnal61:1255-1258. fromBritishColumbia.CanadianJournalofZoology60; George, R.J., McFarlane, D.J. and Speed, R.J. (1995). The 3232-3236. consequencesofa changinghydrologicenvironment Benham, W.B. (1903). On some new species of aquatic for native vegetation in southwestern Australia. In Oligochaeta from New Zealand. Proceedings of the Saunders,D.A.,Craig,J.L.andMattiske,E.M. (eds) ZoologicalSocietyofLondon2:202-232. Nature Conservation 4: The Role of Networks 9-22. Brinkhurst, R.O. (1982). Additional aquatic Oligochaeta Surrey-Beatty,Sydney. fromAustraliaandNewZealand.RecordsoftheQueen Halse, S.A., Pearson, G.P. McRae, J.M. and Shiel, R.J. VictoriaMuseum78:1-13. (2000). Monitoring aquatic invertebrates and Brinkhurst, R.O. (1984). Two new speciesofTubificidae waterbirds at LakeToolibin and Walbyring Lakes in (Oligochaeta) from the Northern Territory of theWesternAustralianwheatbelt.JournaloftheRoyal Australia. Proceedings of the Biological Society of SocietyofWesternAustralia83;17-28. Washington97:142-147. Pinder, A.M. and Brinkhurst, R.O. (2000). A review of Erseus, C. (1984). The marine Tubificidae (Oligochaeta) the Tubificidae (Annelida: Oligochaeta) from of Hong Kong and Southern China. Asian Marine Australian inland waters. Memoirs ofthe Museum of Biology1:135-175. Victoria58:39-75. Erseus, C. (1990). MarineOligochaeta ofHong Kong. In Morton, B. (ed.) The Marine Flora and Fauna ofHong Kong and Southern China. Proceedings of the Second Manuscriptreceived3May2001;accepted1August2001. InternationalMarineBiologicalWorkshop259-334.Hong KongUniversityPress,HongKong.

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