EcologyEcologyEcology andandand PhylogenyPhylogenyPhylogeny ofofof DamselfliesDamselfliesDamselflies (Zygoptera)(Zygoptera)(Zygoptera) ininin FrenchFrenchFrench PolynesiaPolynesiaPolynesia

RonaldRonald A.A. EnglundEnglund1,, SteveSteve JordanJordan2,, DanDan A.A. PolhemusPolhemus1 1BishopBishop MuseumMuseum Hawaii,Hawaii, USAUSA 2BucknellBucknell UniversityUniversity OverallOverall Objectives:Objectives: PolynesiePolynesie FranFranççaiseaise OdonataOdonata SurveysSurveys

1) Survey previously unsampled aquatic and terrestrial habitats for Odonata 2) Assess origins and dispersal patterns of FP Odonata via molecular systematics 3) Assess the conservation status and Bora Bora: Ischnura cardinalis impacts of introduced aquatic species on native Odonata, 4) Provide museum specimens and an ecological information baseline for future researchers 5) Assess ecological shifts previously only found in Hawaii (Aquatic to Terrestrial) Raiatea: new Ischnura sacrebleu WhatWhat waswas knownknown forfor EndemicEndemic ZygopteraZygoptera inin PolynesiePolynesie FranFranççaiseaise (5(5 sp.)sp.) ??

Societies Genus Ischnura: • 2 species described in 1930’s - Bora Bora/Raiatea (1) - Tahiti (1)

Marquesas (Genus Bedfordia) Raiatea: Trois Cascades • 2 species described in the 1930’s - Ua Pou (1) - Nuku Hiva (1) Australs (Genus Ischnura) • Rapa (1) described in 1966

Bora Bora damselfly stream habitat AustralAustral DamselfliesDamselflies -- DamselflyDamselfly MorphologyMorphology

(Cook Islands) Rarotonga Rimatara Rurutu Tubuai Raivavae

•• 33 totaltotal endemicendemic speciesspecies fromfrom Rurutu,Rurutu, Raivavae,Raivavae, RapaRapa Rurutu: Puputa Cascade •• WorldWorld’’ss largestlargest IschnuraIschnura fromfrom RapaRapa (45(45 mm)mm) •• TubuaiTubuai lackinglacking endemicendemic damselfly,damselfly, butbut limitedlimited surveysurvey time/invasivetime/invasive speciesspecies possiblepossible reason?reason? •• NewNew genusgenus fromfrom Raivavae,Raivavae, currentlycurrently inin Rapa IschnuraIschnura Raivavae: new Ischnura jeanyvesmeyeri SocietySociety IslandsIslands -- DamselflyDamselfly MorphologyMorphology

•• BoraBora Bora:Bora: 11 describeddescribed IschnuraIschnura speciesspecies •• Tahaa:Tahaa: 22 IschnuraIschnura speciesspecies (2(2 new)new) •• Raiatea:Raiatea: 33 IschnuraIschnura speciesspecies (2(2 new)new) Huahine: Ischnura new sp. •• Huahine:Huahine: 2?2? IschnuraIschnura (2(2 new)new) •• Moorea:Moorea: 11 IschnuraIschnura (1(1 new)new) •• Tahiti:Tahiti: 4?4? IschnuraIschnura speciesspecies soso farfar (3(3 new)new) •• IncompleteIncomplete surveys,surveys, FredFred JacqJacq hashas severalseveral newnew speciesspecies Bora Bora: Ischnura cardinalis MarquesasMarquesas -- MorphologicalMorphological WorkWork DamselfliesDamselflies •• Eiao:Eiao: 11 permanentpermanent streamstream andand 11 newnew speciesspecies collectedcollected by Fred Jacq/Jean-François Butaud •• NukuNuku Hiva:Hiva: 33 speciesspecies total,total, 22 newnew BedfordiaBedfordia sp.sp. •• UaUa Huka:Huka: 11 newnew BedfordiaBedfordia sp.sp. Fatu Hiva: Bedfordia new sp. •• UaUa Pou:Pou: 11 sp.sp. ((BedfordiaBedfordia halecarpenterihalecarpenteri)) •• HivaHiva Oa:Oa: 11 newnew BedfordiaBedfordia speciesspecies •• Tahuata:Tahuata: SShareshares 11 BedfordiaBedfordia withwith HivaHiva Oa?Oa? •• Mohotane:Mohotane: 00 damselflydamselfly sp.sp. (1(1 newnew SimuliidSimuliid)) •• FatuFatu Hiva:Hiva: 11 newnew BedfordiaBedfordia sp.sp. Ua Pou:Bedfordia new sp. IschnuraIschnura thelmaethelmae LieftinckLieftinck 19661966:: RapaRapa

Rarotonga

(Cook Islands) Rurutu Tubuai Rimatara Raivavae

male Bichromatic female

Rapa

Marotiri Rocks IschnuraIschnura jeanyvesmeyerijeanyvesmeyeri EnglundEnglund && PolhemusPolhemus (2009)(2009) RaivavaeRaivavae (also(also aa newnew genusgenus toto bebe describeddescribed later)later)

Rarotonga

(Cook Islands) Rurutu Tubuai Rimatara Raivavae

Rapa

Marotiri Rocks IschnuraIschnura rurutanarurutana EnglundEnglund && PolhemusPolhemus (2009)(2009) RurutuRurutu

Rarotonga

(Cook Islands) Rurutu Tubuai Rimatara Raivavae

Rapa

Marotiri Rocks AUSTRALAUSTRAL ANDAND SOCIETYSOCIETY ISLANDISLAND DAMSELSDAMSELS

Rarotonga

(Cook Islands) Rurutu Tubuai Rimatara Raivavae

Tahiti- Ischnura nov. sp. Ischnura thelmae

Rurutu Raivavae Rapa Ischnura rurutana nov. sp. New genus Ischnura jeanyvesmeyeri nov. sp. Rapa Marotiri Rocks Methods:Methods: mtmt DNADNA COIICOII andand A6A6 andand A8A8

mtDNA

1186 bp

tRNA-Asx COII tRNA-Lys A8 A6

C2-J-3102 TK-N-3785

C2-J-3711 A6-N-4379

Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: Marquesas mtDNA, Mark Olaf: Methods http://www.genomenewsnetwork.org/articles/04_02/mito_dna.shtml http://bioweb.wku.edu/courses/biol22000/6StructureNA/images/F04-10A.JPG Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: Marquesas mtDNA, Mark Olaf: Methods

• 27 specimens – 22 ingroup taxa • From most major Marquesas populations

– Outgroups from 2 genera • Ischnura – widespread • Megalagrion orbates – Hawaii Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: Marquesas mtDNA, Mark Olaf: Results

97 Ua Huka 1 100 Ua Huka 1 Ua Huka 2 100 Ua Huka 2 98 100 100 ML Bootstrap Ua Poa1 100 100 Ua Pou 1 100 97 Bayesian Ua Pou 5 Posterior 100 Ua Poa5 100 ML Bootstrap Nuku Hiva BB 2 Hiva Oa 10 82 Nuku Hiva 22 Bayesian Hiva Oa 8 100 Posterior 61 Hiva Oa 12 Nuku Hiva 25 74 Tahuata1 Hiva Oa 10 73 73 Hiva Oa 3 80 70 Hiva Oa 3 Fatu Hiva 8 99 100 Hiva Oa 12 55 Nuku Hiva 13 68 73 99 Nuku Hiva 1 83 Fatu Hiva8 74 99 68 Nuku Hiva 5 100 Nuku Hiva 13 55 96 87 Nuku Hiva 7 100 Nuku Hiva 1 97 72 Nuku Hiva 15 Nuku Hiva 4 Nuku Hiva 2 92 100 73 Nuku Hiva 26 Nuku Hiva 26 100 85 Nuku Hiva 4 Nuku Hiva 2

100 Nuku Hiva BB1 Nuku Hiva 15 85 56 Nuku Hiva 7 100 Nuku Hiva 25 100 91 61 Nuku Hiva 22 Nuku Hiva 5 90 Nuku Hiva BB 2 100 Tahiti 8 100 Tah iti 8 72 Tahiti 9 84 100 Ta h iti 9 100 50 70 Ischnura aurora 38 (Fiji) 98 Ischnura aurora 38 (Fiji) 58 96 Megalagrio n Megalagrion orbates orobates (Kauai) I. heterosticta 23 (Fiji) (Kauai) I. heterosticta 23 (Fiji) 0.1 0.1 Analysis I: All Analysis II: 1st and 2nd Characters Codon Positions Only

Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: Marquesas Nuclear DNA – Jenn Mrozek

• 30 individuals NH13 BB2 NH17 • 29 alleles 60 NH8 •1055 bp of nuclear Elongation Factor-1α gene NH2 NH1 UP5 • 86 variable characters UP1 UP3 NH22 • 31 parsimony informative characters NH5 NH15 NH7 • K80+gamma Maximum Likelihood analysis NH25 HO3 • 1000 bootstrap pseudoreplicates HO10 TA1 68 HO8 HO1 100 HO14 86 HO11, 21 FH6 FH2 58 FH1 FH8 NH BB1 UP6 SiTahiti8 EF-1alpha SiTahiti9 0.01 Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: Society/Austral mtDNA: Sherri Liang

SITahaa7 665 bp SITahaa8 mtDNA SIBo1, 10, 6, Tahaa 1, 14, 5 81 SITahaa22, 23, 29 Societies/Australs mtDNA SIBo5 SIBo14, 2, 9, Tahaa 2 • 665 bp of mt COII gene 92 SIHu1, 5 • 167 parsimony informative characters SIHu11 • GTR + I + gamma model 87 SIHu10 • 39 individuals SIHu12, 2, 6 • 22 haplotypes Tahiti Iti • 500 bootstrap pseudoreplicates 100 Rurutu504, 510, 516 Rurutu512 79 SIBo16 Rapa505, 506 90 Rapa509 Rapa507 SITahaa28 89 Raivavae508 Raivave511 Raivavae513, 515 SITahaa30 0.1 Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: All Together

Nuku Hiva 13 Nuku Hiva 5 • 665 bp mt COII gene Nuku Hiva 15 99 Nuku Hiva 26,7 Nuku Hiva 1 • GTR+I+G model ML analysis 88 Nuku Hiva 4 Fatu Hiva 8 • 476 bootstrap pseudoreplicates Nuku Hiva 25 Nuku Hiva Big Blue 1 Hiva Oa 10 Hiva Oa 12 Hiva Oa 3 Marquesas 91 Nuku Hiva 2 85 Tahuata 1 Hiva Oa 8 97 Nuku Hiva 22 100 Ua Pou 1 Ua Pou 5 Ua Huka 1, 2 Nuku Hiva Big Blue 2 100 Raivavae 508 92 Raivave 511 Raivavae Raivavae 513, 515 Megalagrion orobates Hawaii Bora Bora 1,10,6; Tahaa 1,14,15,22,23,29 Bora & 94 Tahaa 7, 8 Bora Bora 5 Tahaa 90 Bora Bora 2,9,14;Tahaa 2 Huahine 1, 5 100 99 Huahine 11 Huahine 10, 12, 2, 6 Huahine 89 Tahiti 8 Tahiti 9 Tahiti Iti 514 100 Ischnura aurora 38 83 Bora Bora 16 100 Rurutu 504, 510, 516 Rurutu 512 Rurutu Tahaa 28 99 Rapa 505,506 Rapa 509 Rapa 507 Rapa Ishnura heterosticta - Fiji 0.1 Englund, Jordan et al. – Molecular Systematics of the Damselflies of French Polynesia: Conclusions Preliminary Conclusions 1. Archipelago monophyly • Endemic Marquesan Zygoptera are likely monophyletic. • Society Island damselflies may not be. • Austral Zygoptera probably are not. 2. Zygoptera of each individual Austral Island are likely monophyletic. 3. mtDNA data suggest that Raivavae damselflies may represent a different genus from others in the Australs and even FP. 4. mtDNA suggest complex biogeography for Marquesan damselflies, while nuclear data show a much simpler pattern. More data are needed. 5. Ua Huka, Ua Pou, Hiva Oa/Tahuata and Fatu Hiva each host one endemic damselfly species. Nuku Hiva may be home to multiple species, though nuclear data put this into question. 6. Bora Bora/Tahaa and Huahine probably each host a single species. PolynesiePolynesie FranFranççaiseaise NativeNative DamselfliesDamselflies SummarySummary •• ExcellentExcellent indicatorindicator speciesspecies forfor aquaticaquatic ecosystemecosystem health,health, foundfound inin undisturbedundisturbed habitatshabitats •• EachEach highhigh islandisland withwith waterwater foundfound toto havehave atat leastleast 11--22 endemic,endemic, largerlarger islandsislands withwith moremore (most(most newnew spp.)spp.) •• TerrestrialTerrestrial damselfliesdamselflies foundfound onon RaiateaRaiatea •• 55 sp.sp. inin 19301930’’s,s, upup toto ~24~24 speciesspecies todaytoday (19(19 new)new) •• DispersalDispersal patternspatterns stillstill unclear,unclear, andand datadata areare incompleteincomplete esp.esp. fromfrom thethe SocietiesSocieties GapsGaps toto fillfill forfor thethe FrenchFrench PolynesiaPolynesia AquaticAquatic InsectInsect BiodiversityBiodiversity AssessmentAssessment Societies: • Highest Priority is Tahiti Nui and Tahiti Iti, particularly high elevation forest and wetland areas • Moorea: Upland stream/wetland areas • Tahaa: Upland native forest areas for terrestrial damselflies • Raiatea: Upland native forest/wetland areas Mt. Toomaru damselfly habitat Marquesas • Eiao: Stream/wetland areas • Nuku Hiva: High Elevation stream and wetland areas • Tahuata: upland stream areas Australs Raiatea: new terrestrial • Tubuai: Upland stream areas Ischnura from Mt. Toomaru Mahalo! •• Team Team Members:Members: JeanJean-Yves-Yves Meyer,Meyer, BenoBenoîtît Fontaine,Fontaine, OlivierOlivier Gargominy,Gargominy, JeanJean-François-François Butaud,Butaud, ElinElin Claridge,Claridge, DianaDiana Percy,Percy, MatthewMatthew Prebble,Prebble, JacquesJacques Florence,Florence, JeanJean-Claude-Claude Thibault,Thibault, RosieRosie Gillespie,Gillespie, FredFred JacqJacq,, ElieElie PoroiPoroi •• Jean Jean-Yves-Yves Meyer,Meyer, DDélégationélégation àà lala RechercheRecherche (Minist(Ministèreère dede lala Culture,Culture, dede ll’Enseignement’Enseignement SupSupérieurérieur etet dede lala Recherche,Recherche, GouvernementGouvernement dede lala PolynPolynésieésie franfrançaise)çaise) •• Bishop Bishop Museum:Museum: NealNeal Evenhuis,Evenhuis, AllenAllen Allison,Allison, DavidDavid Preston,Preston, AlAl Samuelson,Samuelson, AlistairAlistair Ramsdale,Ramsdale, LaVonneLaVonne EnglundEnglund

Raivavae: 2002 Tubuai: 2003 Rimatara: 2004