Aquatic Insects

This article was downloaded by:[Umea University Library] On: 23 August 2007 Access Details: [subscription number 781079123] Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Aquatic Insects International Journal of Freshwater Entomology Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713817864 Morphological and molecular species delimitation within the Holarctic Ilybius angustior complex with a focus on Beringia (Coleoptera: Dytiscidae)

Online Publication Date: 01 September 2007 To cite this Article: Nilsson, Anders N. and Ribera, Ignacio (2007) 'Morphological and molecular species delimitation within the Holarctic Ilybius angustior complex with a focus on Beringia (Coleoptera: Dytiscidae)', Aquatic Insects, 29:3, 159 - 171 To link to this article: DOI: 10.1080/01650420701383136 URL: http://dx.doi.org/10.1080/01650420701383136

PLEASE SCROLL DOWN FOR ARTICLE

Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article maybe used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 Downloaded By: [Umea University Library] At: 08:31 23 August 2007 egahclrne,telwgntcvrainwti hsseiscmlxsgetseito within speciation suggest complex species this within variation Beringia. including genetic that refugia low suggest Pleistocene genes the the mitochondrial I ranges, subunit within geographical oxidase c events cytochrome and speciation ribosomal large studied the within eiiain Sakhalin delimitation, ot lsaadteCnda o rtc h ml n arwbd hrceitco hs w species widespread two the these of of form characteristic climate body cold narrow and the small of The extreme arctic. an low represents Canadian the and Alaska north eswl-nw eg iso omn19) ncnrs,mn atPlaci species Palearctic East so-called many the or contrast, revised more in In 1999) are (1994, 1995). species occurring Nilsson Palearctic Holmen species known. West & The poorly the Nilsson 2001). and remain (e.g. (1987), (Nilsson Larson well-known Holarctic by less five revised were and America Nearctic North nine Palearctic, are Dytiscidae) (Coleoptera: Beringia within Holarctic delimitation the species molecular and Morphological 171 – 159 29(3): 2007; September Insects Aquatic SN06-44pitIS 7445 online 1744-4152 Umea print/ISSN Science, 0165-0424 Environmental ISSN and Ecology of [email protected] Department E-mail: Nilsson, Sweden. N. Anders Correspondence: The Introduction Keywords: Holarctic the of species fourth A Abstract 2 1 NILSSON N. ANDERS O:10.1080/01650420701383136 DOI: complex, species this within diagnostic from are separated be genitalia cannot Male species new east. the far shape and Russian size body the from in although Sakhalin of Island ard Spain Madrid, opee,adw ilhr ecieanwseisi h Holarctic the in species new a describe here will we and complexes, ta.20,rpeetdby represented 2004, al. et h obnto frde eaasmrsadacrnt n uoesenm6i males partly in the in the 6 and in sternum size, species body rugose the smaller the and of in antennomeres carinate found are distal a differences infuscate main and two metatarsomeres The 1999). ridged (Nilsson of combination the eatmnod idvria Biologı y Umea Biodiversidad Science, de Environmental Departamento and Ecology of Department The .subaeneus I. .angustior I. iigbeetles, Diving gopo h genus the of -group cmlxi eysmlrt h lsl related closely the to similar very is -complex lbu angustior Ilybius lbu angustior Ilybius .angustior I. 1 .angustior I. lbu angustior Ilybius GAI RIBERA IGNACIO & ª 07Tyo Francis & Taylor 2007 Ilybius aEouia ue ainld inisNaturales, Ciencias de Nacional Museo Evolutiva, á Glehl and (Gyllenhal) cmlxaefil eet ncmiainwt known with combination In recent. fairly are -complex e pce,Hlrtc iohnra N,species DNA, mitochondrial Holarctic, species, new , rcsncretyicue 2seiso hc 18 which of species 32 includes currently Erichson complex, ope ihafcson focus a with complex .angustior I. nvriy Umea University, ˚ lbu minakawai Ilybius .anjae I. 2 .angustior I. -complex. iso,rsetvl) sharing respectively), Nilsson, .poppiusi I. .crassus I. .s. sdsrbdfo the from described is sp., n. ,See,and Sweden, ˚, Glehl.Lwvariation Low (Gyllenhal). .churchillensis I. .angustior I. nvriy 0 7Umea 87 901 - S University, ˚ cmlx(Ribera -complex and - -complex. .poppiusi I. alsfrom Wallis ˚, - Downloaded By: [Umea University Library] At: 08:31 23 August 2007 e.SK hln A;Kmhta erpvos,53 Petropavlovsk, Kamchatka, CAN; Kholin, S.K. leg. 8v.91 1 18.vi.1991, 1 19.vii.1970, – 18 106 < 69 ..Koi,CN acak,pnsiln rmtebybtenCp hly suh n Cape and (south) Zheltyi Cape between bay the from 51 inland (north), ponds Ilya Kamchatka, CAN; Kholin, S.K. ot uis aauhr nadfo aeo ai’eaPnnua 50 Peninsula, Vasil’yeva of base W from inland Paramushir, Kurils, North eedissected: were of material following The localities. mapped maximum Statistica all software the for using as run given were measured are was analysis was latitude the statistical length width All and of view. Penis penis Longitude apex lateral bead. and in 6.0. the shaft 15), marginal to penis Figure the inside capsule of (1999, measured head width Nilsson the was in of (WS) as margin metasternum measured anterior of the Width from elytra. measured was length Body analyses Morphological the methods concerning and hypotheses Material some advance also data. We sequence DNA the Minakawa. on based complex, N. the of species the by of sent origin larger temporal recently author and a Sakhalin geographical and to senior expeditions 1995, 2003 in and two the Kholin 2002 the S.K. the to of from by species material collected the six was in the species included of new was series four the for of data male single DNA known mitochondrial A provide complexes. three and The east, far morphology. Russian analysed on the separate. been based to have (2000) difficult complex and al. differentiated et this poorly Larson within are and limits species (1987) as Species Larson 1939). (1837) by Nearctic The (Wallis chiefly Kirby the 1985). arctic (Persson complex, by distribution low Holarctic the America Canadian a with of North species species valid from a third as described restored recently was until not complex the picipes Colymbetes of species second .Ags A;Sbra akl y rsosi,1 mSJlns,52 Jelantsy, S km 18 Krestovskij, Mys Baikal, Siberia, CAN; Angus, R. 69 bJnsjEp 86F rbm HS iadosiisl,70 insula, Nikandrovski NHRS: Trybom, F. 1876 Exp. Ob-Jenisej Ribera I. & Nilsson N. A. 160 wdn rvneHa province Sweden, 8 8 ZN olgclIsiue cdm fSine,S.Petersburg St. Sciences, of Academy Stockholm Institute, History, Zoological Natural (ZIN) of Madrid Museum Science, Swedish Natural (NHRS) of Vladivostok Museum Pedology, National and (MNCN) Biology Helsinki of History, Institute Natural (IBPV) of Museum Finnish Berlin (FMNH) Hendrich, Lars Coll. Plymouth (CLH) Bilton, David Umea Coll. Nilsson, (CDB) N. Anders Coll. (CAN) the of species fourth a describe here will We h oaci itiuinof distribution Holarctic The 2 45 04 8 53 , 0 0 e.Kas .Ja O. & Krause leg. , 160 N N83 0 ,2 0v.99 1 30.vi.1999, – 20 E, 8 8 40 57 < L;Skai,Oh itit eeoy agy 53 Langry, Beregovye district, Okha Sakhalin, CLH; , 0 ,2 E, 8 0 ,19,1 1994, E, 33 ib’ pce a yoyie with synonymised was species Kirby’s . < 0 ,157 N, e.R nu,CN rus,Tbli 51 Tibelti, Irkutsk, CAN; Angus, R. leg. , reae,Fare,2vi20,5 2.vii.2001, Flatruet, ¨rjedalen, < 2 , 8 gr A;Brai,Kahadsrc,5k OKrnvlae 50 village, Kiran NO km 5 district, Kiakhta Buryatia, CAN; ¨ger, < uik 69 Dudinka ; 43 .angustior I. < 1 0 ,2.i.99 5 27.vii.1999, E, e.I enk L;Ykta oyadla ohdky Edoma, Pokhodskaya delta, Kolyma Yakutia, CLH; Melnik, I. leg. , , e.A oak D;Ykta rdeoys,67 Srednekolymsk, Yakutia, CDB; Hobaek, A. leg. , .angustior I. a tde opoerclyadgntlao l males all of genitalia and morphometrically studied was 8 25 0 ˚ N86 a enrcgie led ySap(82.A (1882). Sharp by already recognised been has < .churchillensis I. .angustior I. 8 5 10 < , 0 e.N iaaa&KL uosi CAN; Kurowski, K.L. & Minakawa N. leg. , ,1 E, 5 , e.J egtn A.-Rsi:Siberia, Russia: - CAN. Bergsten, J. leg. , 8 02 < 8 cmlxfo h ahlnIln in Island Sakhalin the from -complex aui,Olene Yakutia, ; .angustior I. 46 0 158 N 0 103 N als a ecie rmthe from described was Wallis, 8 40 8 02 0 8 N82 8 38 8 0 39 141 N 14 0 ,1.i.97 1 10.vii.1997, E, 0 yBadn(85,and (1885), Branden by 106 N 0 ,8–1.i17,1 15.vi.1970, – 8 E, 8 50 k 68 ¨k, 8 0 55 ,1 E, 8 0 23 ,3.x19,1 30.ix.1995, E, 8 8 0 01 < 8 ,2.i19,2 25.vi.1993, E, 32 27 0 1 155 N 0 0 112 N 153 N , Saostrov, ; < , 8 8 8 8 leg. , 24 40 17 20 leg. , 0 0 0 0 E, < E, E, N , Downloaded By: [Umea University Library] At: 08:31 23 August 2007 .aenescens I. Outgroups eune F029t F032fo ieae l 20) sqec f16S of *sequence (2001). al. et Ribera from AF309352 Methods). to AF309289 Sequences vittiger I. fraterculus I. discedens I. albarracinensis I. .picipes2 I. .picipes1 I. .minakawai I. .angustior4 I. .anjae I. angustior5 I. angustior3 I. .angustior2 I. .angustior1 I. Species opoercdt nNearctic on data Morphometric 49 Bay, Nemo from 1 16.viii.1997, eoi N a bandtruhasadr hnlclrfr xrcinusing extraction genes mitochondrial phenol-chloroform of fragments standard ( Two unit ethanol. ribosomal a large in the collected through sequenced, were specimens obtained of tissue was abdominal DNA Genomic Larson. D.J. Dr. methods from Molecular received were (1987) Larson in 74 Figure to corresponding odtoso h mlfiain.A diinlfamn a eune o oeo the of some for sequenced was fragment additional 3 the An spanning I), amplification). (Table specimens the of conditions ihacsinnmesa nTbeI oceso h eune pcmn r eti the in using kept are edited (Madrid). specimens Naturales and sequenced Ciencias de the GeneBank Nacional assembled of Museo to the Vouchers and were I. submitted (London) Table Museum were which History in Natural sequences as fragment, numbers New accession each Corporation). with (GeneCodes for 4.2 obtained Sequencher were sequences xds uui ( I subunit oxidase c TTTGAATATATCCT-5 rgetadtervrepie 23(io ta.19)(3 1994) al. et (Simon M223 primer small the reverse as primer the forward same and the with (ND1), fragment 1 subunit dehydrogenase NADH gene < Q801 Q802MC-I0 wdnJA Sweden MNCN-AI102 DQ285002 DQ285011* Q802NMI28See B idl district, Vindeln VB: Sweden NHM-IR268 DQ285012 Q800 Q805MC-I0 usaSkai,Vlriver Val Sakhalin, Russia MNCN-AI101 DQ285005 DQ285010* Q808 Q803MC-I0 wdnA Sweden MNCN-AI103 DQ285003 DQ285008* Q809 F031NMI29Rsi uieIslands, Kurile Russia NHM-IR279 AF309351 DQ285009* AF309295 Q803D250 H-R8 aaaA:Longview, AB: Canada NHM-IR283 DQ285007 DQ285013 F029A394 H-R7See A Sweden NHM-ER17 AF309345 AF309289 rLcox1 rrnL 1 , 8 e.N iaaa A;NrhKrl,Oeoa,pnso h lta inland plateau the on ponds Onekotan, Kurils, North CAN; Minakawa, N. leg. , h Holarctic The 36 cox1 0 ,154 N, c.80b)(e ieae l 03frdtiso h rmr and primers the of details for 2003 al. et Ribera (see bp) 800 (ca. ) 0 ,apiyn oa fc.80b.Bt owr n reverse and forward Both bp. 800 ca. of total a amplifying ), Q804MC-I5 wdnJA Sweden lakes MNCN-AI156 Waterton AB: DQ285004 Canada NHM-IR284 DQ285006 F036Sweden Islands Kurile Canada Russia Portugal Finland AF309336 AF309337 AF309352 AF309334 AF309350 al .Mtra sdfrtemlclrstudy. molecular the for used Material I. Table 8 49 0 ,2.i.99 1 24.vii.1999, E, .angustior I. lbu angustior Ilybius 0 n fthe of end oce e.CutyLclt Leg. Locality Country ref. Voucher rrnL (48 c.50b)adte3 the and bp) 500 (ca. ) rrnL < e.N iaaa&KL uosi CAN. Kurowski, K.L. & Minakawa N. leg. , ope Clotr:Dtsia)161 Dytiscidae) (Coleoptera: complex < ee h RALuadte5 the and tRNA-Leu the gene, 45 , and ) ˚ ˚ ¨ ¨ :Ja N: :Rundvik, N: 28.6.2002 13.5.2000 ot,28.7.2002 mouth, uahr 17.8.1999 Kunashir, 28.6.2002 28.6.2000 NP, 28.6.2000 27.7.1999 :O :O .churchillensis I. ¨ ¨ truddistrict, stersund truddistrict, stersund ¨rna rrnL 0 s ..02AN Nilsson A.N. 8.8.2002 ¨s, -GGTCCCTTACGAA 0 n ftecytochrome the of end RAluadN1(see ND1 and tRNA-leu , (44 0 .Bergsten J. ..Nilsson A.N. .Minakawa N. .Minakawa N. .Bergsten J. & Ribera I. .Rbr & Ribera I. ..Nilsson A.N. n fthe of end .Cieslak A. .Cieslak A. < 51 rrnL , ) Downloaded By: [Umea University Library] At: 08:31 23 August 2007 4vi.03 e.N iaaa 3 Minakawa; N. leg. 14.viii.2003, hc tdfes oee,i h hp fteml genitalia. minakawai male Ilybius the of shape the in however, differs, it which lbu angustior Ilybius .Mnkw;1 Minakawa; N. ooa ie,Eo endv,49 Leonidovo, of E River, Poronay 0vi.91 e.A aaui,CH 1 CLH; Basarukin, A. leg. 10.viii.1991, 6 .N iso .Ribera I. & Nilsson N. A. 162 neirmttra lwi aewt umda iain(iue n ) nfemale in 6); subapical and and 5 view lateral (Figures in knob dilation apical with submedian robust 9) with (Figure male Penis apically. in curved stronger weakly claw punctures metatarsal of Anterior rows Longitudinal angustior reticulation. female. I. in dorsal than in fine incised in stronger with strongly than wide reticulation less relatively and present; Head larger meshes sheen. meshes with metallic reticulation faint Elytral a with black surface 2 e pce.Tenwseisi xenlyvr iia oteNearctic the to similar very externally is species new The species. new ahln uahv,48 Pugachevo, Sakhalin, 24 esrmnsadrto si al II. Table in as ratios and Measurements penis. Description robust more a and wing, metasternal narrow body, narrow Diagnosis. arw(iue1 ihdra ufc oehtls rhdta in than arched less somewhat surface dorsal with 1) (Figure narrow p ooyedpstdi I;4prtpsi BV nCH nFN,2i HS in NHRS, in 2 FMNH, in 2 locality. CLH, Type in CAN. 3 in IBPV, 79 in and paratypes MNCN-AI101), (ref. 4 MNCN ZIN; in deposited Holotype material. Type eune aigmlil re.Otrusicue eea pce fthe of addition the species random of several 1000 example included with one Outgroups plus searches group, trees. heuristic multiple TBR saving using sequences 2002), (Swofford 4.0b10 version stenwsmldSkai ouain ipa nqecmiaino hrces relative the characters, of of species combination recognised unique already a three display the populations to Sakhalin sampled now the As (2004). al. et Ribera from obtained Results were I) (Table sequences Outgroup trees. the root to itit 0k fNgii y’rvr 51 river, Tym’ Nogliki, of S km 20 14 Paratypes: district, label. holotype our and Minakawa’’ 143 5 , h nesxa ifrne nbd eghadwdhaesaitclysgicn ( significant statistically are width and length body in differences intersexual The hlgntcaaye eecnutduigprioyudreulwihsi PAUP* in weights equal under parsimony using conducted were analyses Phylogenetic kadsrc,1 mEo itn 52 Piltun, of E km 15 district, Okha , , 8 kadsrc,Smd eisl,Tyzny ae 54 lake, Tayezhnoye peninsula, Shmidt district, Okha , .5,weestedfeecsi h aisT-/WadW/Saent Body not. are WC/WS and TL-h/MW ratios the in differences the whereas 0.05), 06.942 .angustior I. eaii ihpntto nvnrlfc oehtwae hnin than weaker somewhat face ventral on punctation with Metatibia . 0 ihnthe Within E. Holotype usa ahln olk itit ermuho a ie,52 River, Val of mouth near district, Nogliki Sakhalin, Russia, Glehl 88:Nlsne l 99 .12(npr,misident.). part, (in 122 p. 1999, al. et Nilsson 1808): (Gyllenhal, < kadsrc,7k Wo aorvrbig,53 bridge, river Sabo of SW km 7 district, Okha , eatra ignro Fgr ) aeadfml tru sin as 6 sternum female and Male 4). (Figure narrow wing Metasternal . p n. sp. 8 11 .angustior I. < 0 142 N aeld‘R:NSkai 3,bgn a ie ot,28/7-02, mouth, River Val nr bog #34, Sakhalin N ‘‘RU: labelled 8 8 lbu chalconatus Ilybius 30 14.692 , kadsrc,na itnBy 52 Bay, Piltun near district, Okha , 0 cmlxrcgie ntecmiaino ml and small a of combination the on recognised -complex ,2.x19,lg ..Kholin. S.K. leg. 22.ix.1995, E, 8 44.400 , 0 8 143 N 38.994 eta ahln ooas itit nioso lower of environs district, Poronaysk Sakhalin, Central , 0 143 N < .angustior I. 8 0 00.483 142 N 24 , ru Rbr ta.20) hc a used was which 2004), al. et (Ribera group 8 rmsm oaiya ooye 4 holotype; as locality same from 16.793 8 0 ,4vi.02 e.N iaaa 1 Minakawa; N. leg. 4.viii.2002, E, 58.633 8 00.778 cmlx hyaehr ecie sa as described here are they -complex, 0 ,7vi.03 e.N iaaa 15 Minakawa; N. leg. 7.viii.2003, E, 0 ,3vi.03 e.N iaaa 1 Minakawa; N. leg. 3.viii.2003, E, 0 142 N 8 45.136 8 8 52 04.556 .churchillensis I. 0 143 N .angustior I. .angustior I. 0 ,1.ii20,leg. 10.viii.2003, E, 0 lbu subaeneus Ilybius 143 N 8 16 < .angustior I. 0 4 ,3.i.– 30.vii. E, 8 8 21.338 primary ; , 05.622 Dorsal . < Nogliki , South , ,from t -test, 0 0 E, N < < . Downloaded By: [Umea University Library] At: 08:31 23 August 2007 churchillensis angustior olco ftehltp,M.Nbr Minakawa. Noburu Mr. holotype, the of collector Locality angustior angustior angustior Sex minakawai minakawai minakawai Species of specimens selected from ratios and Measurements II. Table Etymology. in than shorter somewhat Paramere 8.0. in – 7.1 ratio angustior length/width I. view; dorsal in constriction 5 includes sample Each .angustior I. n ihstto ndsa atsasr(iue 0ad1) eaegncx as gonocoxa Female 11). and 10 (Figures sparser part distal in setation with and h pcfi pte sanu ntegntv ae eie rmtenm fthe of name the from derived case, genitive the in noun a is epithet specific The . iue1. Figure ohKmhta1 .101 .00.08 0.07 5.00 3.99 0.16 0.20 9.81 7.96 10 10 Churchill Manitoba, Kamchatka both both ohSkai 080 .439 0.17 0.08 0.09 0.24 3.93 0.20 3.88 4.55 3.98 0.24 4.57 0.16 0.18 0.28 8.01 0.30 7.92 8.92 8.10 10 8.72 5 10 5 10 Sweden ASL m 940 Sweden Churchill Manitoba, Sakhalin both both Sakhalin Sakhalin both both female male h Holarctic The < n 5 and lbu minakawai Ilybius , n h w ee r ie losprtl for separately also given are sexes two the and , 5 lbu angustior Ilybius 0 S 098 .250 0.12 5.09 0.22 9.80 10 ASL m 100 .s. oslhbtso ooye cl a mm. 1 bar Scale holotype. of habitus dorsal sp., n. ope Clotr:Dtsia)163 Dytiscidae) (Coleoptera: complex N minakawai Ilybius enS enSD Mean SD Mean oylength Body (mm) , .angustior I. .minakawai I. and , . .churchillensis I. oywidth Body (mm) . Downloaded By: [Umea University Library] At: 08:31 23 August 2007 oecmo ntenrhpr fteiln hni h ot n eta at rmwhich from parts central and history. south Natural the 12). in (Figure than known island are the of specimens part single north only the in common more Ribera I. & Nilsson N. A. 164 Distribution. 9. – 7 Figures ae ihsnybtos h pcmnfo uahv nSuhSkai a olce na in collected was shallow Sakhalin and South ponds in mossy Pugachevo from including specimen also The and bottoms. habitats, sandy similar with lakes from are records Other tundra. oaiy cl a mm. 1 bar Scale locality. wdn ltut ifrn cl asfr2–4(et m n rgt . mm). 0.1 (right, 6 – 5 and mm) 1 (left, 4 – 2 for bars scale Different Flatruet. Sweden, S24 (4) 2.4; WS 6. – 2 Figures .minakawai I. Ilybius Ilybius h e pce ss a nw nyfo h ahlnIln.I sseemingly is It Island. Sakhalin the from only known far so is species new The ei nltrlve.( 8) – (7 view. lateral in penis , 2–4 eatra ig 2–3) – (2 wing; Metasternal 4) – (2 . ttetp locality, type the At CW .;( )Ml eaaslcas (5) claws; metatarsal Male 6) – (5 3.4; WC/WS , .minakawai I. .angustior I. .angustior I. wdn 7 Umea (7) Sweden; , a olce nbgwt hc osi the in moss thick with bog in collected was wdn 2 Umea (2) Sweden; , .minakawai I. ;()Fare;(9) Flatruet; (8) ˚; ,W/S24 3 ltut WC/ Flatruet, (3) 2.4; WC/WS ˚, yelclt;(6) locality; type , .minakawai I. .angustior I. type , , Downloaded By: [Umea University Library] At: 08:31 23 August 2007 (squares). ae ntefis afo uut hs hnlgcldt r otlkl isdfo h fact the from biased likely most periods. are certain data to phenological most concentrated These with was August. September, collecting of late to half that July first late the from in collected taken were Specimens pond. forest man-made of material Palearctic East examined of location Geographical 12. Figure 11. – 10 Figures cl a mm. 1 bar Scale Ilybius aaeei nenlve.(10) view. internal in paramere , h Holarctic The lbu angustior Ilybius .angustior I. ope Clotr:Dtsia)165 Dytiscidae) (Coleoptera: complex wdn ltut (11) Flatruet; Sweden, , lbu angustior Ilybius .minakawai I. dt)and (dots) yelocality. type , .minakawai I. Downloaded By: [Umea University Library] At: 08:31 23 August 2007 iue1.Rltosi ewe eaiewdho eatra igadbd eghi eetdseiesof specimens selected in length body and wing metasternal of width churchillensis relative between Relationship 13. Figure rm 1 aioa hrhl,()See,Fare,90mal 3 wdn Umea Sweden, (3) asl, m 940 Flatruet, Sweden, Kamchatka. (2) Russia, Churchill, (4) Manitoba, (1) from: ssalr arwr n a arwrmtsenlwnsthan wings metasternal narrower has and narrower, smaller, is Externally, Ribera I. & Morphometrics Nilsson N. A. 166 3 al I.Within II). Table 13, agradhv rae eatra ig hntoefo h aainlwaci or significant arctic statistically low are Canadian differences the These from II). those Table than 13, 5 wings (Figure (ANOVA, region metasternal alpine broader Swedish have and larger 4.Telnt/it ai ftepnsi esta . in 8.1 than less is penis the of ratio angustior I. length/width The 14). of penis the 171), Figure ( groups hoc hr a ovrainaogthe among variation no was There analyses Molecular .angustior I. hra h ei of penis the Whereas oprsn hwta oylnt n CW ifrbtenbtntwti hs three these within not but between differ WC/WS and length body that show comparisons .churchillensis I. , .minakawai I. n rae hn95in 9.5 than greater and , and .minakawai I. < þ .poppiusi I. 5 , , and , p 5 þ .angustior I. .churchillensis I. .minakawai I. .minakawai I. 0.001; .angustior I. sietclt h Nearctic the to identical is opee,adtevrainaogthe among variation the and complexes, F oln ouain rmSee n acak are Kamchatka and Sweden from populations lowland , ifrn ybl sdfrdfeetseisand species different for used symbols Different . aus L191;T/W2.6 CW 175.00). WC/WS 21.16; TL/MW 139.13; TL values: .churchillensis. I. rrnL-tRNALeu-ND1 smr outta nbt te pce Fgrs7–9, – 7 (Figures species other both in than robust more is lowland , smr lne hnin than slender more is .angustior I. .churchillensis I. n arctic/alpine and , eune fteseiso the of species the of sequences .angustior I. .angustior I. .minakawai I. cox1 n hsseispair species this and , district, ˚ Lro ta.2000, al. et (Larson eune a very was sequences Fgrs2–4and 4 – 2 (Figures .angustior I. .angustior I. 5 . . in 9.6 – 8.0 , 0 s,and asl, m 100 specimens Ilybius Post ). Downloaded By: [Umea University Library] At: 08:31 23 August 2007 hrdbtenoeo hs and these of of one specimens between two shared the of each in n oe htol eaaino ae a ‘aryrlal’.Externally, the reliable’’. ‘‘fairly of was males of species separation only four to that noted referred the and 384) p. genitalia, (1987, Larson male differentiated. diagnostic highly ifrnitdfrom differentiated the of species two the Whereas Discussion the of one pce fthe of species specimens two the between 0.004, was group the of of specimens among distance p uncorrected Kamchatka), teltrwt n pcmnof specimen p one with between later were (the differences maximum the small: of males selected in width penis and length penis between Relationship 14. Figure h otdansi hrce eaaigteetreseisi,hwvr aegenitalia male however, is, species three these separating character diagnostic most The churchillensis I. ¼ h hlgntcaayi eutdi igete f35ses with steps, 305 of tree single a in resulted analysis phylogenetic The .angustior I. .1) ihnthe Within 0.011). .churchillensis I. .angustior I. .angustior I. rmAbra(al I). (Table Alberta from pcmn from specimens h Holarctic The .minakawai I. and , pcmn rmAbrawith Alberta from specimens .angustior I. ope,wihfre aa oyoywt igend linking node single a with polytomy basal a formed which complex, .minakawai I. .anjae I. .angustior I. .angustior I. .angustior I. n asn(97 on tpsil osprt most separate to possible it found (1987) Larson and , ope,teol aito a n uaoopi change autapomorphic one was variation only the complex, lbu angustior Ilybius .minakawai I. . ope r elsprtdmrhlgcly including morphologically, separated well are complex rmAbra orsodn oa uncorrected an to corresponding Alberta, from nbd ieadsaei ra fsympatry. of areas in shape and size body on rmAbra n n yaoopi change synapomorphic one and Alberta, from .anjae I. .angustior I. TbeIadFgr 5.Temaximum The 15). Figure and I (Table ope Clotr:Dtsia)167 Dytiscidae) (Coleoptera: complex .minakawai I. n h et agn rm6t bp 8 to 6 from ranging rest, the and and .angustior I. .picipes I. lbu angustior Ilybius Fgr 15). (Figure .churchillensis I. .anjae I. s‘sbigspecies’’ ‘‘sibling as ope r less are complex fo wdnand Sweden (from itrt the to sister snot is Downloaded By: [Umea University Library] At: 08:31 23 August 2007 dnicto,a a enpitdotb ueosatos(..Mire l 06and 2006 al. et those Meier for the (e.g. of inconvenience authors species clear numerous the by a of out however, 2002; fragment pointed Penkrot is, therein). reduced been & references This has a Wiens as reviews). e.g. of identification, (see for use under definitions the 2004 assumable species perfectly advocating not Marshall for situation and a & criteria 2003), sorting, or Omland Sites incomplete to concepts & due species (Funk likely species many most is recent this among below), uncommon (see group the of species paraphyletic a within nested ‘oel’mrhtp.Ormlclraayi Fgr 5 ugssthat suggests 15) (Figure analysis molecular Our morphotype. ‘‘boreal’’ bipustulatus ugssta rtcadapn ouain ifrfo oeloe ntesm ieto sthe as direction same the from in differ ones boreal species from two differ populations other alpine and arctic that suggests ooooso a vle neednl rmteacsrlsoki egahclisolation geographical in stock within Variation ancestral climate. the similar from the a independently within under species evolved the has of or relationships homologous the present angustior At 13). I. (Figure wing metasternal narrower from way same eaaeseisi ae nisuiu obnto fbd ie oysaeadpenis and shape body size, with body sympatric of is combination it unique that fact its the on with based together is shape, species separate a Ribera I. & Nilsson N. A. 168 a eni iue1) n eae r oepolmtc u eonto of recognition Our problematic. more are females and 14), Figure in seen the (as on search heuristic an with found tree parsimonious most single the of Phylogram 15. Figure ubr bv oe otta upr aus e al o h oaiyo h specimens. the of locality the for I Table See values. support bootstrap node, above Numbers h eei iest mn h eune iohnra ee ftefu sampled four the of genes mitochondrial sequenced the among diversity genetic The ti oebethat noteable is It nSadnva(rt ta.20) ugsigtemlil needn rgno a of origin independent multiple the suggesting 2001), al. et (Drotz Scandinavia in ope r nnw,adtu ti o osbet a fti iiaiyis similarity this if say to possible not is it thus and unknown, are complex .angustior I. .angustior I. .churchillensis I. and nldn mle n arwrbd ncmiainwt a with combination in body narrower and smaller a including , .angustior I. .angustior. I. .poppiusi I. and u otelwgnrlgntcdvrec mn the among divergence genetic general low the to Due . .minakawai I. .angustior I. opee srmral o,epcal ihnthe within especially low, remarkably is complexes iia rn a endcmne in documented been has trend similar A ohi h ertcadtePalearctic, the and Nearctic the in both , r opooial ifrnitdi the in differentiated morphologically are cox1 .angustior I. eefrseisrcgiinand recognition species for gene . .minakawai I. .minakawai I. cox1 sequence. Agabus as is Downloaded By: [Umea University Library] At: 08:31 23 August 2007 yohoecoiae1 n bu afi ecnie h hl iegneo h combined the of divergence whole the the for consider divergence we if 1% half approximately about within and species 1, region morphological oxidase recognised Holarctic c currently cytochrome northern six be whole will the there spanning divergence, sampled the of range angustior I. data. unpublished and the (1999), of Nilsson species (2000), al. the et of Larson limits Range 16. Figure ope.I h w isn pce ( species missing two the If complex. h Holarctic The lbu angustior Ilybius angustior and - .churchillensis I. poppiusi ope Clotr:Dtsia)169 Dytiscidae) (Coleoptera: complex cmlxs ae nmp nLro (1987), Larson in maps on Based -complexes. and .poppiusi I. alwti the within fall ) Downloaded By: [Umea University Library] At: 08:31 23 August 2007 References CGL2004- project Noboru the and through financed Kholin was the K. work in IR. obtained Molecular (MNCN, Sergey to were Izquierdo Vogler. 00028 specimens specimens. NHM Ana Alfried of collected. of of Sequences they work. laboratory Johannes loan sequencing specimens for data. sending thanked the is for raw Madrid) for thanked morphometric both thanked his are sharing Minakawa all for are thanked Stockholm, is Umea John’s, St. Bergsten, Larson, J. David Acknowledgements locally. exist Yamal the and Tyumen well from may known CLH) are Plutenko, nearctic records The A. Kodar 2004). Siberian Petrov the leg. & West from (Andrejeva 1.vi.1995, females Peninsula species. environment, two this Additionally, village Krasnoyarsk. represent (Tschara from Mts. female a Range on reported (1995) e ieysatrdlclte ntewsenaci anad agn rmChurchill, Russian from the ranging in Sakhalin mainland, of Island arctic the also western to restricted the east, Seemingly far Alaska. on northern localities to Manitoba, scattered widely few aentbe bet ofimtepeec of presence Mountains. the Rocky confirm the to along able extension been 1999). southern not a (Nilsson have of with that range encloses States, extended range United This more the of a parts suggests northern Yakutia from angustior record Ilybius northern single the of anjae I. atr ta.20;Buae ta.20) dst h yohsso h xsec fspeciation refugia. of northern existence the isolated of multiple, hypothesis in the to 2000; complex the al. adds Nearctic the et 2005), of Abbott within al. western evidence 1994; et processes increasing (Elias and Brubaker area 2004; the Palaearctic Beringian al. with the et eastern in together Waltari the refugia This, the of Pleistocene Beringia. northern species around of Pleistocene the existence centred the of in i.e. be distribution speciated 16), to The have (Figure 2004). seems to Vogler considered complexes & to distributions two (Ribera comparable restricted refugia and with observations), southern (e.g. species unpublished Pleistocene close and family very 2003 same al. between et that the Ribera of 2003; Drotz species fragment; other or of (Linnaeus) variability intraspecific Ribera I. with & Nilsson N. rrnL-tRNALeu-ND1 A. 170 rt K 03 ouaingntc n hlgorpyo h etPalearctic and west trees the boreal of for phylogeography refugium and glacial genetics a Population as 2003. Beringia MK. 2005. Drotz AV. the Lozhkin and ME, Peninsula Yamal Edwards southern PM, of Anderson migration (Coleoptera) LB, plant beetles Brubaker of Adephaga analysis aquatic Molecular The 2000. 2004. J. PN. Balfour Petrov K, TR, Wolff Andrejeva RMM, Crawford RL, Milne LC, Smith RJ, Abbott lbu anjae Ilybius .poppiusi I. hus e esetvsfo apdple aa ora fBoegah 283–848. – 32:833 Biogeography of Journal data. pollen mapped 20. from – 109(3):9 perspectives Ser.) new (Biol. shrubs: Naturalists of Society Moscow of Bulletin Urals. Polar 1346. – 289:1343 Science Arctic. the in refugia and ope.PDdsetto,Dprmn fMlclrBooy Umea Biology, Molecular of Department dissertation, PhD complex. .angustior I. skonfo ot hn,JpnadteRsinfres Fgr 6.Ms records Most 16). (Figure east far Russian the and Japan China, North from known is .minakawai I. r rmnrhr oglaadtesuhrms ato atSbra although Siberia, East of part southernmost the and Mongolia northern from are and eaeacoriacea Meladema a ierneta oesms fNrhErp,Sbra aaa n the and Canada, Siberia, Europe, North of most covers that range wide a has ,DvdBlo,Pyot,Lr edih eln n etViklund, Bert and Berlin, Hendrich, Lars Plymouth, Bilton, David ˚, skonfo h otenotpr fteiln,tetoseismyco- may species two the island, the of part northernmost the from known is .poppiusi I. and a h otrsrce ag fteseiso h w opee.As complexes. two the of species the of range restricted most the has cox1 .picipes I. r emnl loarcoe uho atSbra hra only whereas Siberia, East of much over allopatric seemingly are ee.Ti ag fvraiiyi eylwee fcompared if even low very is variability of range This genes. aot,wt a %o iegnei h aegene same the in divergence of 2% ca. with Laporte, htseigyi oersrce nms ietos We directions. most in restricted more is seemingly that , .picipes I. lbu churchillensis Ilybius nEs iei.NlsnadHolmen and Nilsson Siberia. East in University. ˚ gbsbipustulatus Agabus skonfo nya only from known is gbsbipustulatus Agabus iigbeetle diving Downloaded By: [Umea University Library] At: 08:31 23 August 2007 asnD.18.Rvso fNrhAeia pce of species American North of Revision 1987. DJ. Larson alsJ.13.Tegenus The 1939. JB. Wallis cole des Catalogue 1885. C. Branden den van in J eko A 02 eiiigseisuigDAadmrhlgclvrainaddsodn species discordant and variation morphological and DNA the of using biogeography species historical Delimiting the 2002. on perspective TA. molecular Penkrot A JJ, 2004. Wiens JA. Cook DR, Klein JR, Demboski E, Waltari eso .1985. S. Persson Palearctic the of intraspecific high revision of tale A a Diptera: 1994. in taxonomy AN. and Barcoding Nilsson DNA 2006. PKL. Ng Region, G, Nearctic Vaidya the K, of Shiyang Dytiscidae) R, (Coleoptera: Meier beetles diving Predaceous 2000. RE. Roughley Y, Alarie DJ, Larson ib .13.Teiscs n ihrsnJ dtr an oel mrcn,o h olg ftenrhr parts northern the of zoology the with or Americana, consequences, – Boreali and Fauna editor. causes, J, Richardson frequency, In: polyphyly: insects. The and 1837. W. paraphyly Kirby Species-level 2003. KE. Omland DJ, Funk wfodD.20.PU* hlgntcaayi sn asmn *adohrmtos,Vrin4.0b10. Version Society methods), Dublin Royal other the of and Transactions Scientific Dytiscidae. (* or Coleoptera parsimony carnivorous aquatic using On 1882. D. analysis Sharp Phylogenetic PAUP*. Systematics and 2002. Ecology of DL. Review Annual Swofford species. delimiting for criteria Operational 2004. JC. of Marshall utility Jr, phylogenetic JW Dytiscidae). and Sites weighting, (Coleoptera, Evolution, 1994. refugia P. Flook Pleistocene H, beetles in Liu B, beetles diving Crespi AT, diving Beckenbach Agabinae F, Iberian Frati of of C, Simon Speciation biogeography 2004. historical AP. and Vogler Phylogeny I, Ribera 2004. AP. Vogler AN, Meladema Nilsson of history I, population Ribera and phylogeography in DNA movements Mitochondrial range and 2003. rates AP. speciation Vogler on DT, type Bilton habitat I, of effect Ribera The 2001. AP. Vogler TG, Barraclough I, Ribera iso N 99 ecito fanwEs aertcseisof species Palearctic East new a of Description 1999. AN. Nilsson iso N hlnS,Mnkw .19.Tedvn ete fKmhta ihadtoa eod rmSakhalin from records additional with Kamchatka, of beetles diving Dytiscidae. The II. 1999. Denmark. N. Minakawa and SK, Fennoscandia Kholin of AN, (Coleoptera) Nilsson Adephaga 395. aquatic – 3:1 The Insects 1995. of M. Catalogue Holmen World AN, (Coleoptera). Nilsson Dytiscidae 2001. AN. Nilsson pp Press. Institution Smithsonian DC: Washington, environments. their and insects Quaternary the 1994. to SA. applied problem Elias delimitation species The 2001. AN. Nilsson A, Saura MK, Drotz yicde.Anlsd aSocie la de Annales Dytiscidae). iisi pn iad Seoou) ytmtcBooy5:9–91. – 51:69 Biology Systematic (Sceloporus). lizards spiny in limits 600. – 85:591 Mammalogy of Journal latitudes. high northern 199. – 71:192 aiblt n o dnicto ucs.Sseai ilg 575–728. – 55:715 982. Biology p Systematic Press. success. Research identification NRC low Ottawa: and Alaska. variability and Canada of 413. fauna – 95:341 the Society on Entomological emphasis York with New the of Journal species. Palearctic on notes fBiihAeia at4 odn oga.p xxxix pp Longman. London: 423. 4. – 34:397 Part Systematics America. and British Ecology of of Review Annual DNA. mitochondrial animal from insights 2219–1003 – (2)2:179 Associates. and Sinauer MA: Sunderland, 227. – 35:199 the of Annals primers. 701. – reaction 87:651 chain America polymerase of conserved of Society compilation Entomological a and sequences gene mitochondrial 193. – 13:179 Ecology – 30:545 Molecular Evolution and Phylogenetics Molecular Molecular sequences. Dytiscidae). DNA (Coleoptera, 562. mitochondrial basin from Mediterranean inferred (Coleoptera) the in and Islands 167. Atlantic – 12:153 the Ecology on 735. – beetles 10:721 Ecology diving Molecular phylogenies. level – species from inferences beetles: aquatic 268. – 16:265 Scandinavica Entomologica Dytiscidae). isrf 1:5–61. – 115:55 Tidskrift n h ui sad Clotr:Dtsia) Beitra Dytiscidae). (Coleoptera: Islands Kuril the and 192. – 32:1 Scandinavica Entomologica Fauna poppiusi I. 22. – 73:11 Society Linnean the xiii of Journal Biological Scandinavia. north in Dytiscidae) (Coleoptera, þ 284. ate Clotr:Dtsia) oepeooiceRnshu6:3–40. – 69:33 Rundschau Koleopterologische Dytiscidae). (Coleoptera: Zaitzev lbu angustior Ilybius þ l.7–18. – 7 pls. h Holarctic The Ilybius Glehl and (Gyllenhal) ´te r nNrhAeia(oepea yicde.TeCnda Entomologist Canadian The Dytiscidae). (Coleoptera, America North in Er. noooiu eBliu 91: 116. – 29(1):5 Belgique de Entomologique ´ lbu angustior Ilybius ópte rscrasesautqe Hlpia,Apiode eoideet Pelobiidae Amphizoidae, (Haliplidae, aquatiques carnassiers `res lbu crassus Ilybius .picipes I. Ilybius g u noooi 917–131. – 49:107 Entomologie zur ¨ge þ ope Clotr:Dtsia)171 Dytiscidae) (Coleoptera: complex Kry sdsic,hlrtcseis(Coleoptera: species holarctic distinct, as (Kirby) 329 ope Clotr,Dtsia) Entomologisk Dytiscidae). (Coleoptera, complex – rcsn(oepea yicde,wt systematic with Dytiscidae), (Coleoptera: Erichson þ 1pl. Ilybius rcsnpeiul ie pwith up mixed previously Erichson gbsbipustulatus Agabus complex