Downloaded By: [University of California] At: 16:05 23 May 2007 Dpea hrnmde revealed ) (Diptera: of identity The 114 – 103 29(2): 2007; June Aquatic SN06-44pitIS 7445 online 1744-4152 USA. 95616, print/ISSN CA 0165-0424 Davis, ISSN California, of University Entomology, of [email protected] Department E-mail: Cranston, S. Peter Correspondence: our description, and report first the Since a of observation. (1923) Tonnoir rare of by 1993; understanding a description the (Tokeshi remains and fish unusual Blephariceridae organisms are and chironomid, Host hosts Hemiptera Dipteran ectoparasitism. commensal 2004). and involve al. and Megaloptera cases et common Roque some Ephemeroptera, very is least Odonata, insects at include aquatic evidently other widespread: and globally larvae chironomid involving Commensalism Introduction phoresy Keywords: immature aquatic the of genus affinity not blepharicerid has commensal/phoretic common taxon The the strong the as discovery. to of apparent first belonging species its described the since with enigmatic Blephariceridae despite stages remained has restudied on Zealand been New commensal from 1923 species Tonnoir, chironomid The Abstract USA CA, Davis, California, of University CRANSTON S. PETER O:10.1080/01650420701308844 DOI: of light the in and of Courtney, genera Greg hemisphere and southern clear. obscure’ been Zwick never ‘more Peter has some by taxon of commensal descriptions collected this recent material of relationships recent and With identity the and genotype) Zealand New the genus, sister putative the from substantially differs is it and since it Naonella description, n.), that original (comb. collected suggest (Tonnoir) the recently namely analyses to from Phylogenetic Orthocladiinae, stages 1999, recognised larvae. history Edward, austral is & blepharicerid life species certain Cranston all torrenticole the to of with Here morphology close association dubium. of nomen tight congruence a the the as by treated material is and identity Dactylocladius 94 e genus, new A 1994. . hrnmde rhcaine e eln,Behrcrde commensalism, Blephariceridae, Zealand, New Orthocladiinae, Chironomidae, ife,10 sn ogrmaigu eas h yeseisi otado uncertain of and lost is species type the because meaningful longer no is 1906 Kieffer, hudsoni Dactylocladius Tonnoirocladius ope nwihocrec a ec 0%o l ouain.Tegenus The populations. all of 100% reach can occurrence which in complex atlcaiscommensalis Dactylocladius Pirara atlcaiscommensalis Dactylocladius otry rntn 95 n especially and 1995, Cranston, & Boothroyd ife a lee t oiadbai h bec fa of absence the in dubia nomina (to altered has Kieffer e.n setbihdfrtespecies the for established is n. gen. ª 07Tyo Francis & Taylor 2007 Kaniwhaniwhanus Neocurupira oni,o e eln larval Zealand New on Tonnoir, otry,1999, Boothroyd, ab 93 seilyspecies especially 1913, Lamb, onioldu commensalis Tonnoirocladius atlcaiscommensalis Dactylocladius Naonella Botryocladius Boothroyd, Downloaded By: [University of California] At: 16:05 23 May 2007 0 .S Cranston S. P. 104 cotcasi ih vlae nmdsuu,tba pr flrd ihsalt strong to hooked/scalpellate small dorsalis, small with crista or 2 ‘flared’ – 1 virga spurs point, eyes, anal tibial without volsella. mid-scutum, bare claws, inferior curved in pectinate rectangular has stout terminally with area short, with adult oval male three denticles, light The with transverse in lobe with fringe. VI anal acrostichals – IV spines, without sternites anteriorly-directed hooked), macrosetae, of (not band transverse bands spines posterior and anteriorly-directed conjunctival spinules II, of of tergite bands conjunctiva on transverse anal hooklets anterior on the and with of horn each one VIII thoracic – with a III fine mola, lacks tergites by of and pupa flanked tooth beard The innermost tooth antero-medially. dense between protruding median with gap tubules broad plate(s), large with with ventromental mandible mentum to setae, branch, equating simple inner perhaps weak layers with sclerotised bifid double SI has larva The Diagnosis oni,dsoee fti nsa ao,ad– and taxon, unusual this of discoverer Tonnoir, Tonnoirocladius Germany Munich, Staatssammlung, Zoologische Research Landcare Division, Entomology Collection, ZSM Zealand New Road, Cromwell Museum, Australia History Entomology, Natural NZAC CSIRO The Collection, Department, Entomology National Australian and BMNH (1980) Sæther ANIC following in terminology limited Standard thus and features. pharates used. on some is based (1994) interpret are Cranston free No descriptions to exuviae). adult pupal their for ability studied—all within DNA both been retained been ringed, developed, have for had subsequently (i.e. adults mountant, adults Euparal isopropanol Blephariceridae pharate Hoyers either and which or high-strength using pupae neutralising in larvae, prepared to and were vials clearing slides or transferred in after Microscope Zwick mountant found were sequencing. (P. or molecular specimens collector and author, extraction freshest the the by by The either or stored. Blephariceridae Courtney), larval W. from G. removed were Specimens to methods generic and undertaken its Material thus been and have relationships, analyses its of molecular assessment preliminary end. allow this and to Morphological features seek placement. and species 1999; the Edward, & Cranston (e.g. Etymology. commensalis Dactylocladius species. Type h olwn coysaeue o h ntttosweemtra slocated: is material where institutions the for used are acronyms following The Kaniwhaniwhanus e eln t. rvt a 27,Acln,NwZealand New Auckland, 92170, Bag Private Ltd., Zealand New UK London, From rntn e.n. gen. Cranston, Tonnoir otry,1999; Boothroyd, h ega-on nioenaotdDpeitAndre Dipterist antipodean-adopted Belgian-born, the – Pirara otry rntn 95 ti ieyt redescribe to timely is it 1995) Cranston, & Boothroyd oni,12,b ooyyadpeetdesignation. present and monotypy by 1923, Tonnoir, Fgrs1–16) – 1 (Figures Naonella cladius rnh si clade. in as branch, a , otry,1994; Boothroyd, Botryocladius ´Le ´on Downloaded By: [University of California] At: 16:05 23 May 2007 iue 4. – 1 Figures onioldu commensalis Tonnoirocladius atlcaiscommensalis Dactylocladius Tnor.Lra 1 nen.()Lbu.()Mnu.()Mandible. (4) Mentum. (3) Labrum. (2) Antenna. (1) Larva. (Tonnoir). Dpea hrnmde eeld105 revealed Chironomidae) (Diptera: Downloaded By: [University of California] At: 16:05 23 May 2007 eiai a ml) 8 nllb,gntlasc(female). sac genitalia lobe, Anal (8) (male). sac genitalia L f P,Oao on siigNP,Mkrr ie,‘ai lt,1.x20,40m s. .W. G. asl., m. 490 16.ix.2004, Flat’, ‘Davis River, Makarora ex- N.P., Aspiring Courtney, ex- Mount Courtney, Otago, W. 2Pm, elev G. Pf, asl., 22.ix.2004, 6L, m. crossing, 490 above 15.ix.2004, River, Creek hudsoni Hollyford Bakers N.P. Fjordland N.P., 1Pm, ex- Paparoa 4Pf, Courtney, s. 9L, W. rd G. coast m.asl., Westland, 5 3Pm, 1Pf, 2P, 8L, Cranston S. P. 106 orhisa av eimszd . mln,ha asl 6 400 – 360 capsule head long, mm 3.6 medium-sized, larva Fourth-instar Larva Hokitika of trib. Greymouth, nr Island, ex- South Zwick, ZEALAND, P. NEW 1972, 2Pm, Feb 1Pm, R, 3Pf, 1P, 1Pe, 1L(P), 2L, Material commensalis Dactylocladius commensalis Tonnoirocladius 8. – 5 Figures eann ea ipe I n IItiknd I ige eysot hea short, chaetae short; without and very teeth single, inner no SIV teeth, apical thickened, 2 of SIII 1. indication and weak segment with of SII stout, 1/4 Lauterborn premandible simple, at simple; segment, organ setae 3rd Ring S 0.5. than than remaining c shorter longer ratio little slightly Antennal a style 3. branch segment apex; inner to bifid, antennal subequal blade to organs 4; extending segment than not shorter outer, slightly or to S5) fragmentation. subequal S4, bearing anterolateral plate some of with margin sclerite, anterior on labral (S3 complete apotome and frontoclypeal delimited with head of arm(iue2 ihS id ihinrbac eywa,otrbac stout; branch outer weak, very branch inner with bifid, SI with 2) 3 segment (Figure subequal, Labrum less or more segments 3 apical segmented, 5 1) (Figure Antenna (NZAC). onioldu commensalis Tonnoirocladius ecrpr hudsoni Neocurupira oni,12.Tp-aeilntfud rsmdlost. presumed found, not Type-material 1923. Tonnoir, ecrpr hudsoni Neocurupira ecrpr hudsoni Neocurupira (ANIC). Tnor.Pp.()Cpai ra 6 neirtoa.()Aa lobe, Anal (7) thorax. Anterior (6) area. Cephalic (5) Pupa. (Tonnoir). Tnor 93 ob n. comb. 1923) (Tonnoir, (ZSM). (NZAC). Fgrs1–16) – 1 (Figures m .Dra surface Dorsal m. Neocurupira Downloaded By: [University of California] At: 16:05 23 May 2007 etoetlpae u opie tlat2slrtsdlyr otrltrlt aeo outermost of base ‘conventional’ to a posterolateral include layers not sclerotised inner 2 may lateral- least its or mid-brown at may to than comprises and but anterior paler complex plate, laterals protruding highly ventromental pairs outermost mentum two the Posterolateral first pair. teeth, and most lateral mentum of smaller median pair pairs Broad median neighbours. with 4 teeth recessed; shallow slightly separated of indistinctly and four pair comprising short perhaps one few or spatulate. margin, with with sclerite squat scales; Basal and unfused chaetae. short pectinate Ungula 3 or pectinate. of simple weakly and epipharyngis other scale-like, Pecten laterales absent. chaetulae lamellae Labral brush. iue 10. – 9 Figures etm(iue3 ihmda ot ra n ihrsml n ihceuaeanterior crenulate with and simple either and broad tooth median with 3) (Figure Mentum onioldu commensalis Tonnoirocladius atlcaiscommensalis Dactylocladius Tnor.Pp.()Admnltrie.(0 boia sternites. Abdominal (10) tergites. Abdominal (9) Pupa. (Tonnoir). Dpea hrnmde eeld107 revealed Chironomidae) (Diptera: Downloaded By: [University of California] At: 16:05 23 May 2007 0 .S Cranston S. P. 108 rna ptm Fgr )wt ekfotlste(0–35 – (30 setae frontal weak with 5) (Figure apotome Frontal Pupa tubules anal all 10), Figure between extends (1923, and Tonnoir antero-ventrally length. base to in common contrast protrudes subequal a In from are position. one elongate-oval arises postero-ventral apparently more that of partner a in paired in such pair conventional its is one and modifed (as parapods comprising parapods posterior pair and to tubules setae, posterior second claws. short area Anal a anal Chironomidae); brown laterally from setae. simple bearing simple posterolaterally anal of spurs, projecting and without tubules group medium-length high, claws apical as 4 yellow wide with as apically pectinate separate, pale, short, highly parapods very Posterior shorter Procercus claws. slightly interna pale spines, setae. Seta shorter simple yellow short. elongate simple, subdentalis relatively inner Seta elongate, bearing of squat, mola. line Maxilla beyond and branches. protruding tooth simple scarcely few innermost teeth, with between narrow inner gap 3 Large stout. combined of setae) teeth. than laterad cephalic to shorter central mentum much other basal tooth (and from submenti extending Setae setae tooth. simple mental fine outermost of of beard apex dense with tooth; lateral neoeilyo I I I ihbodadde neirtases ado n spinules, fine of band transverse anterior deep and broad with VII – III II, on anteromedially aeas eldvlpd n rcrel,sbqa nlnt,smtmswt hr,fine short, with sometimes length, in subequal dc precorneals, precorneal; fine 4th developed, asymmetrical well 3 frons laterals; pearls; lack strong. sheaths seta pedicel Ocular tubercles, contiguous creases. raised transverse slightly with to anterior just sheaths dc oywtotayeiec fltrlste neirprpd eaae ihconof crown with separate, parapods Anterior setae. lateral of evidence any without Body Apical rounded. unevenly margin inner smooth, margin outer with 4) (Figure Mandible boe Fgrs7–1)wt egtsIadI ae rwt io n spinulation fine minor with or bare, II and I tergites with 10) – 7 (Figures Abdomen hrx(iue6 ekyrgls oslyadatrol;2dra nerntl,0 antepronotals, dorsal 2 anteriorly; and dorsally rugulose weakly 6) (Figure Thorax 4 raa eamyb rsn.Toai onabsent. horn Thoracic present. be may seta Prealar . iue11. Figure onioldu commensalis Tonnoirocladius 1 n dc and 2 ls,dsatfo lsl prxmtddc approximated closely from distant close, Tnor.Ml genitalia. Male (Tonnoir). m og yn ewe pedicel between lying long) m 3 and Downloaded By: [University of California] At: 16:05 23 May 2007 n 2coeyapoiae;sgetVI ih2,2,1.AlLstesml n fine. and simple setae L All 1L. 2V, 2D, with VIII segment with approximated; VI closely – absent. IV L2 B and conjunctives and A corres- spinules; spurii on of Pedes conjunctives. than row tergal stronger transverse on absent. as mostly Apophyses with spines, but anterior-directed fine VIII – long, with VI slender, anteriorly-directed, corresponding (V) long, pattern tergites, of tergites; spinulation ponding posterior band with more VIII transverse – on V multiserial size multiserial 30 in to becoming with up increasing spines, V, and VI on – small tubercles III and toward few conjunctives spines, tending of band width transverse and posterior with VIII – V riorly; mle (ca. smaller 16. – 12 Figures iwpplpstos 1)Ltrlve,pplposition. pupal view, Lateral (16) positions. pupal view eain emn ih5,I,2 ea;sget I–VIwt D Vad3 ea,L1 setae, 3L and 5V 5D, with VII – II segments setae; 2L IV, 5D, with 1 segment Setation: 5 onioldu commensalis Tonnoirocladius 5 m m og trie Fgr 0 I ihu pnlso pnlto,anterior spinulation, or spinules without III – I 10) (Figure Sternites long. m )adcutrdatrol,lne (ca. longer anteriorly, clustered and m) atlcaiscommensalis Dactylocladius Tnor.Lra 1 4 eta iwlra oiin.(5 Ventral (15) positions. larval view Ventral 14) – (12 Larva. (Tonnoir). Dpea hrnmde eeld109 revealed Chironomidae) (Diptera: 4 5 m )admr eaaeposte- separate more and m) Downloaded By: [University of California] At: 16:05 23 May 2007 oyodRvr(7 8cprgtG .Cute;rpoue ihpermission). with reproduced Courtney; W. G. copyright 18 (17, River Holyford 1 .S Cranston S. P. 110 iue 7–1.NwZaad ot sad lpaiei / blepharicerid Island, South Zealand, New 18. – 17 Figures ra htmyecmasmr hn1st;3–4pelr;7–9uieilsuelr.Lateral scutellars. uniserial 9 – 7 prealars; 4 – oval 3 pale seta; from 1 arising setae. than dorsocentrals without more uniserial thorax encompass hooked/scalpellate 9 uniserial small may mid-scutum; 2 in that – 1 area less areas antepronotals; oval lateral light or 3 in – 2 acrostichals chaetica. setation: more sensilla Thoracic or medially. 7 consecutive pit contact – 6 each without segments, segment to Third 5 previous. with than restricted Palps longer setose. segment setation moderately quadrate, Clypeus Temporal postorbitals. 13 extension. on sub-apical several dorsomedial and 3 flagellomere on chaetica extending sensilla groove 0.22. 1 seta; flagello- 13; A.R. subapical with 13 to strong elongate, lacking with 3 apex, increasingly Antenna flagellomere to becoming from mm. extending segments 1.0 plume – successive complete 0.9 globular, but length near sparse (sheath) 3rd wing and mm, 2nd anal 3.4 meres, c. of length body apex Small, as only) far specimens (pharate as male Adult extending not sheaths lobe genital anal beyond female 8). well 7), extending (Figure and lobe (Figure long sheaths macrosetae genital Male and macrosetae. stout short 3 with uvliwl-eeoe,hl egho emnlypciaeclaws. absent. terminally-pectinate divergent apparently strongly of chaetica with length each Sensilla denticles; half spur, flared absent. well-developed, longer with pseudospurs of Pulvilli basally biserial; length mid-legs half irregularly and than comb more fore- denticles; spur of shorter spurs with spurs; tibia subequal hind short two with tibia nllb one,ntpoue.Csawt lgtetnin C itlt - (ratios r-m to distal FCu setae. extension; 10 with slight Squama with setae. Costa without veins produced. remaining Cu not 6, incalculable) – rounded, 5 with lobe R Anal seta, 1 with Brachiolum igmmrn ihfiepntto,wtotmcorci.R macrotrichia. without punctation, fine with membrane Wing in narrowed, medially not lobes developed, without well ommatidia, Antepronotum of brown. medium margin Thorax mesal to close microtrichia with bare, eye with Head spinules, fine of patch circular median and margin crenulate with rounded, lobe Anal e aisiclual,fr iilsu oeaeyln,sotrta iilae,medium apex, tibial than shorter long, moderately spur tibial fore incalculable, ratios Leg 1 straight. Tonnoirocladius oain.(7 pe tr ie.(18) River. Otira Upper (17) locations. 4 þ 5 nigaoeM above ending 3 þ 4 . . Downloaded By: [University of California] At: 16:05 23 May 2007 ek or weak, whanus rntne l 18) ae ntebr igadee n h rsneo iegn lateral divergent of spurs. presence leg the hind and and eye, mid- and the wing of bare shaft the the on on spines based (1989), al. et Cranston Eukiefferiella ‘ae’ iilsu tutr skyda fuiu to unique if in as absent are keyed and (both area) is pale distinct structure mid-scutal are spur a virga to tibial and restricted ‘‘flared’’ point and scalpellate anal being to the than character (rather contrast phylogenetically-informative antepronotum (putative) (in following the share states species included all but aetesm pce sTnordsrbdfo iia oain n otblepharicerids we host that and reasons locations morphological similar or from biological described on Tonnoir doubt 1923). as no (Tonnoir species is same there the material, have historic of lack of non-retention) (or loss seeming the given type: Orthocladius sIhv eni ntopo odto o et fe eecitos’(rea 99 p. 1959, (Freeman redescriptions’’ to offer Zealand to species me Chironomidae, the New for material transferred condition Zealand the ‘‘such a Freeman New that poor in this, species too found, the in the Despite they of been is for 418). seen noted study not are have types, the his I mentioning has nor and as specifically in In Nelson, material not Canberra, 1999). Institute, while excellent original Cawthron Freeman, (Crosby Collection, was the The Collection of Insect stages. description collections Arthropod are the National history to original now of as life that search Australian clues Tonnoir’s personal few orthoclads all gives although by non-descript of allocation including identity, dark generic details original small (modern) the (or providing of Thus, genus true genera. variety the of period a the protracted diversity for a a for to used However, allocated was details). for subgenus) 1983 as Ashe (see dubium nomen genus The spermathecal setose. on convoluted quite cerci discussion opening highly developed, Taxonomic common moderately with cerci to neck, and prior X or Tergite bulb bare. microtrichia common hyaline, without labia in eminence; dark. capsules, ending with and undivided, broad ducts seminal lobe apparently apodeme spermathecal oval VIII lobe; Gonapophysis dorsomesal pale, of setae. sign Two short any 6 obscuring – lobe 4 ventrolateral long, large pointed. 7 – simple, 6 claws with long. protruding, than broader Clypeus 0.68. AR except 5-segmented, male Antenna as only), specimens projecting. (pharate female obviously Adult not dorsalis. corner, inferior crista without absent; distomedial and volsella hyaline megaseta with superior rounded simple, virga; Gonostylus with without very rectangular, placed Hypopygium projections volsella oral weak. weak very phallapodeme with rounded, lateral; Sternapodeme point. anal without setae; segment. posterior more Tokunagaia ekn oeniett,teautml a ekydto keyed be can male adult the identity, modern a Seeking egt Xbod niie,eel u preysts;gncxt Xqiestrongly quite IX gonocoxite setose; sparsely but evenly undivided, broad, IX Tergite apical or medial without successively setae, long each anterolateral few on with IX sparser Tergite 11). becoming (Figure Genitalia setae, long scattered with unicolorous, Tergites goigti etr,tetxnky to keys taxon the feature, this Ignoring . Psectrocladius Dactylocladius æhr 93adt oeetn in extent some to and 1973 Sæther, up 84 with 1874, Wulp, heean 96adi h uta genera austral the in and 1926 Thienemann, ife,10 fcniee og u nalohrfaue relationship a features other all on but long, considered if 1906 Kieffer, atlcaiscommensalis Dactylocladius Tonnoirocladius ife,10 a rce sasbeu ftewl established well the of subgenus a as erected was 1906 Kieffer, rhcaiskervillei Orthocladius :tedsic nerntlstesata the at start setae antepronotal distinct the ): Dpea hrnmde eeld111 revealed Chironomidae) (Diptera: Bryophaenocladius Eukiefferiella Chaetocladius ife,19 eintdsbeunl as subsequently designated 1899 Kieffer, .kervillei O. Orthocladius Chaetocladius Tonnoirocladius Botryocladius Chaetocladius fteplil r considered are pulvilli the if h aei eadda a as regarded is name the , sahmgnosgrouping, homogeneous a is heean 94and 1934 Thienemann, owtsadn the Notwithstanding . tocr loin also occurs it , and .Atog the Although ). ife,11 in 1911 Kieffer, Kaniwhani- Downloaded By: [University of California] At: 16:05 23 May 2007 1 .S Cranston S. to P. 112 iue1.Rltosisof entomology.ucdavis.edu/chiropage/tonnoirocladius.pdf Relationships 19. Figure rnia ifrnele ntesaeo h neirvlel hti hrceitcly‘‘hooked’’ taxa, characteristically is candidate that volsella all inferior the of and of species shape both genes in the in all lies difference how for principal to of as species incomplete arises both then question is from between The unpublished). relationship CAD) Cranston close and and (Morse a indicates CO1 analysis 28s, preliminary (18s, genes with 19) Cardiocladius (Figure clade resolved Kaniwhaniwhanus a reveals Naonella matrix for this 2004), parsimony resemblances, (1994, phenetic with Boothroyd Botryocladius taxa austral from the and added for ‘‘ were derived the matrix morphology with this amongst To relationships 2000a). investigate (Cranston of to set relationships data subsequently to The testing genera. constructed expanded for Orthocladiinae been of originally has set matrix designed global data a was amongst a placement Thus, phylogenetic resemblance. the phenetic assess any even or and beard relationship, ventromental the of density mandible. and the extent the of mentum, structure outer sternites the the of and shape tergites the in the differs no of any any bands lacking However, with spine distribution horn, structure. and or thoracic lobe spinulation structure a anal in the lacking disagree and the in spines in resemblance conjunctival and superficial strong tergites a on bears it hooklets which couplets. to multichoice incompatible, (1986), internally and ambiguity much involves this (1977), oresemble do pr.os,cnr otry 94 n er eea eebac to resemblance general a bears of and pupa 1994) The Boothroyd 1994). contra Boothroyd obs., contra (pers. obs., (pers. acrostichals located oekonb oenmsi rntn20c n hlgntcifrnefo such from anal inference developed phylogenetic better and 2000c) has (including Cranston orthoclads but in austral several names sternites), somewhat to code having common the and by is B, type on known and pupal A some a (including spurii such pedes However patterns II, macrosetae. tergite spinule on hooklets similar horn, thoracic a lacking vdnl h oaci est ahlf tgspouen osseteiec for evidence consistent no produce stages life each to keys Holarctic the Evidently to (1983) al. et Cranston in keys larva The to unambiguously and readily keys pupa The Psectrocladius ihbosrpspoto 8.Ti ruigfrsa neovdtihtm with trichotomy unresolved an forms grouping This 68%. of support bootstrap with ife,11 tscesv etrmv.Atog oeua apigfrfour for sampling molecular Although remove. next successive at 1912 Kieffer, Tonnoirocladius rmseiesadfo rntnadEwr 19) nlsdunder Analysed (1999). Edward and Cranston from and specimens from suraitc lhuhtefml a ekydto keyed be can female the Although unrealistic. is Naonella þ Pirara Naonella Tonnoirocladius n ute,in further, and , and Naonella nlcigsnil heiaadhvn iial omdand formed similarly having and chaetica sensilla lacking in mlctywehri eevsgnrcsau.I h ae the male, the In status. generic deserves it whether implicitly , Echinocladius siae ymxmmprioyo aaarx vial thttp:// at Available datamatrix. on parsimony maximum by estimated rmeaiaino pcmn Caso collection) (Cranston specimens of examination from Naonella Kaniwhaniwhanus Metriocnemus ,with Metriocnemus Paracladius h aecasaesimple. are claws male the Parapsectrocladius Eukiefferiella . Tonnoirocladius ivnj,17,fo hc it which from 1973, Hirvenoja, up 84i ofane al. et Coffmann in 1874 Wulp, rmBohod(99,for (1999), Boothroyd from Brillia Tonnoirocladius Metriocnemus and Caso 00)and 2000b) (Cranston Eukiefferiella Tonnoirocladius Naonella ’gopo genera of group ’’ Tonnoirocladius Tokunagaia and Naonella Botryocladius a frontals has aesuch have nSæther in itrto sister differs adults and in Downloaded By: [University of California] At: 16:05 23 May 2007 eebac suwratd h av of of that larva to construction The different of plate unwarranted. ventromental is resemblance seP 93 aaou fciooi eeaadsbeeao h ol nldn yoys(Diptera: synonyms including world the of subgenera and of Description genera 1994. chironomid IKG. of Boothroyd catalogue A 1983. P. Ashe all ‘‘nearly that observed Courtney chiltoni specimens. the many involve to on appear phoretic associations were collected who chironomids Courtney, Greg to According Biology otry K,Caso S 95 w rhcaine(hrnmde eeacmo oNwZaadand Zealand New to common genera (Chironomidae) Orthocladiinae Two 1995. PS. Cranston IKG, Boothroyd of species new A 2004. IKG. Boothroyd of Description 1999. IKG. Boothroyd References systematics. Diptera in endowment host Schlinger studies Evert support sought the to of support who Davis, the California, with Courtney of place took University (e.g. Greg study to Zealand cold- This New 18). by and Island, and collected South 17 Zwick throughout specimens Figures conditions the arduous Peter under without larvae and place entomologists Blephariceridae taken aquatic have locomotion not stenothermic could their study commensal revisionary This with numerous of interfere ‘‘load’’ must Acknowledgements the surely rocks, of pupae surface general. and/or in the hydrodynamics larvae across frequently. specimens quite progress chironomid multiple observed were larvae, to 4 – orientated with 3 suckers larvae: the were As with blepharicerid always 16). 15), large (Figure one pupae (Figure host with pupae: support the associated of be with lateral body could co-occur the providing small to suckers could appressed on surface ventral Larvae found ventral be host. their could with large transversely larvae single instar a Late 0%’’. on was 13). extreme (Figure other suckers The Neocurupira ventral 100%. the to around close wound probably were 4) & 3 instars (on populations of species other on chironomids found ofa P rntnP,Oie R æhrO.18.Ky n igoe fteppeo h subfamily the of 2000a. pupae PS. and Biology Cranston the Chironomidae: editors. of LCV, diagnoses Pinder PS, and Cranston Keys PD, Armitage 1986. In: Morphology. OA. 1994. Sæther PS. Cranston DR, Oliver PS, Cranston WP, Coffman ie rtgrs(02 bevtoso h s ybehrcrdlra fterventral their of larvae blepharicerid by use the on observations (2002) Frutiger’s Given commensalis Tonnoirocladius hrnmde.Etmlgc cniaiaSplmn 71–68. – 17:1 Supplement Scandinavica Entomologica Chironomidae). Australia: 15. – 27:11 349. Entomologist – 33:341 Zealand Research New Freshwater and Marine of Journal Zealand New Zealand. 315. – 21:309 Zoology of Journal clg fNnbtn igs odn lso,NwYr,Tko ebun,Mda:CamnadHall. and Chapman Madras: Melbourne, Tokyo, 30. York, – New 11 Glasgow, p London, Midges. Non-biting of Ecology 296. – 28:147 Supplement Scandinavica Entomologica Chironomidae). (Diptera, Orthocladiinae 408. – 389 p CSIRO. Melbourne: Ecosystems. 311–111. – 23:101 , ohhri micrognathia Nothohoraia ave(iue1) n eea ifrn ntr of instars different several and 14), (Figure larvae Pirara .gn and gen. n. Austrobrillia atlcaiscommensalis Dactylocladius Eukiefferiella rea:imtr tgs n e pce rmteNorpc.Spixiana Neotropics. the from species new a and stages, immature Freeman: avewr oae ihrtasesl Fgr 2 rsometimes or 12) (Figure transversely either located were larvae Kaniwhaniwhanus Naonella Naonella and ecrpr hudsoni Neocurupira heean n rntnP,eio.Cioois rmGnsto Genes from Chironomids: editor. PS, Cranston In: Thienemann. Peritheates Neocurupira .gn Dpea hrnmde rhcaine.NwZealand New Orthocladiinae). Chironomidae: (Diptera: gen. n. otry Ciooia:Otoldia)fo e Zealand. New from Orthocladiinae) (Chironomidae: Boothroyd Neocurupira e.n Dpea hrnmde rhcaine rmNew from Orthocladiinae) Chironomidae: (Diptera: n. gen. Naonella p.Lvl fpoeyi some in phoresy of Levels spp. Dpea hrnmde eeld113 revealed Chironomidae) (Diptera: on ociooison chironomids no found I . Tonnoirocladius pce o ytmtcsuis larval studies, systematic for species ope,atog occasionally I although complex, a ea eet distinctive a beneath setae has .commensalis T. . ol co-occur could Neocurupira .hudsoni N. Downloaded By: [University of California] At: 16:05 23 May 2007 oeh .19.O h vlto fcmesls nteCiooia.Fehae ilg 941–489. – 29:481 Biology Freshwater Chironomidae. the Scandinavica e in cycle commensalism Entomologica Le of 1923. Chironomidae). evolution A. the (Diptera: Tonnoir On terminology 1993. other chironomid M. on Tokeshi of living glossary larvae Chironomidae A of 1980. Records OA. 2004. Freshwater EN. Sæther Blephariceridae). Fragoso (Diptera: M, midges Jancso net-winged S, larval Trivinho-Strixino of FdeO, suckers Roque the of function Museum The British the 2002. of A. Bulletin ). Frutiger (Diptera, Chironomidae Zealand New Available: the Diptera. of study (NZAC): A 1959. Collection P. Arthropod Freeman Zealand New Orthocladiinae the subfamily Orthocladiinae in the subfamily of specimens males the type adult Primary of the 1999. of larvae T. diagnoses the and Crosby Keys of 1989. diagnoses OA. and Sæther DR, Keys Oliver 1983. PS, Cranston OA. Sæther DR, Oliver PS, Cranston 1999. DHD. edu/ Edward http://entomology.ucdavis. Available: PS, Australia. Cranston of Chironomidae the to Guide Electronic 2000c. PS. Cranston Cranston 2000b. S. PS. P. Cranston 114 upeet1: 51. – 14:1 Supplement 9. – 4:1 Neotropica Biota Brazil. in aquatic 302. – 47:293 Biology 437. – 393 7: Entomology History Natural 2006. Dec 26 Accessed http://www.landcareresearch.co.nz/research/biodiversity/invertebratesprog/nzac/dipt_intro.asp. 352. – 34:165 Supplement Scandinavica 291. Entomologica – 19:149 Chironomidae). Supplement (Diptera, Scandinavica Entomologica Region. Holarctic the of Chironomidae) (Diptera, 333. – 24:305 Entomology Systematic Chironomidae). (Diptera: 2006. Dec 26 Accessed chiropage/index.html. 120. – 31:103 Evolution and Systematics Insect Andes. southern Parapsectrocladius vltfde ´volutif atlcaiscommensalis Dactylocladius Botryocladius e eu fotoldieCiooia Dpea rmPtgna the Patagonia, from (Diptera) Chironomidae orthocladiine of genus new a : e.nv:anwtasnaci eu fotoldiemidge orthocladiine of genus transantarctic new a nov.: gen. p o.Anlsd ilgeLcsr 129–291. – 21:279 Lacustre Biologie de Annales nov. sp.