Overview of the Hymenoptera AToL

Orussidae

PRIMARY OBJECTIVE

To generate a robust phylogeny of the phylogeny of the Hymenoptera at the superfamily level, using molecular, behavioral, and morphological data.

SECONDARY OBJECTIVES

• Robust phylogenies of selected grades and superfamilies at finer taxonomic levels

• Interactive key to the families of Hymenoptera

• Fully vouchered matrices with SEM and light images

• Integration of fossil data and estimations of ages of major clade splits in the Hymenoptera

• Detailed phylogenetic analyses of selected morphological character suites, i.e., male and female genitalia, and internal musculature of the thorax

• Production of a website and publication of two books to disseminate results

APOCRITAN RELATIONSHIPS

APOCRITAN RELATIONSHIPS

CARPENTER: ACULEATA, SYMPHYTA

ENGEL: APOIDEA, FOSSILS

HERATY: CHALCIDOIDEA, MYMAROMMATOIDEA

RONQUIST: CYNIPOIDEA, PROCTOTRUPOIDEA, CERAPHRONOIDEA

SHARKEY: ICHNEUMONOIDEA, PLATYGASTROIDEA, TRIGONALYOIDEA, STEPHANOIDEA, MEGALYROIDEA, EVANIOIDEA

PRINCIPAL INVESTIGATORS

COLLABORATORS• Andy Bennett Ichneumonidae• Andy Boring Female genitalia• Denis Brothers Aculeata• Matt Buffington Cynipoidea, databasing• Andy Deans Evanioidea• Michael Engel Fossils, Apoidea• David Gaitros Databasing• Jeremiah George Chalcidoidea• David Hawks Chalcidoidea• John Heraty Chalcidoidea• Johan Liljeblad Chalcidoidea• Chrissy Romero Chalcidoidea• Neelima Jammigumpula Databasing• Lynn Kimsey Aculeata: Chrysidoidea• Vasudevan Mahalingam Databasing• Lubomir Masner Proctotrupoidea, Platygastroidea• Debra Murray Molecules• Michael Ohl Apoidea s.l.• Laurence Packer Apoidea• James Pitts Aculeata• Alex Rasnitsyn Fossils• Susanne Schulmeister Symphyta, Male genitalia• Katja Seltmann Web page, Interactive keys• Barb Sharanowski Braconidae• Lars Vilhelmsen Symphyta, Thorax, including internal musculature• Matthew J. Yoder Proctotrupoidea

AND MANY OTHERS…..

COLLABORATORS• Andy Bennett Ichneumonidae• Andy Boring Female genitalia• Denis Brothers Aculeata• Matt Buffington Cynipoidea, databasing• Andy Deans Evanioidea• Michael Engel Fossils, Apoidea• David Gaitros Databasing• Jeremiah George Chalcidoidea• David Hawks Chalcidoidea• John Heraty Chalcidoidea• Johan Liljeblad Chalcidoidea• Chrissy Romero Chalcidoidea• Neelima Jammigumpula Databasing• Lynn Kimsey Aculeata: Chrysidoidea• Vasudevan Mahalingam Databasing• Lubomir Masner Proctotrupoidea, Platygastroidea• Debra Murray Molecules• Michael Ohl Apoidea s.l.• Laurence Packer Apoidea• James Pitts Aculeata• Alex Rasnitsyn Fossils• Susanne Schulmeister Symphyta, Male genitalia• Katja Seltmann Web page, Interactive keys• Barb Sharanowski Braconidae• Lars Vilhelmsen Symphyta, Thorax, including internal musculature• Matthew J. Yoder Proctotrupoidea

AND MANY OTHERS…..

Chris DarlingGerard DelvareMichael GatesGary GibsonChrister HanssonJohn HeratySteve HeydonJohn HuberJohn LaSalleJohn PintoAndy PolaszekJean-Yves RasplusMichael SchauffSimon van NoortGreg ZolnerowichJohn NoyesChris BurwellGerhard PrinslooJo BerryPaul HansonEric GrissellAlex GumovskyHans BauerStefan SchmidtDino ReinaJung-Wook KimJames MunroAlbert OwenRoger BurksJeremiah GeorgeAndrew CarmichaelMatt BuffingtonChris DesjardinsLars KrogmannSerguei Triapitsyn

ChalcidoideaTWiG

Taxon Objectives

Skeleton Matrix• superfamily group representation across Hymenoptera (102 species)• taxa that can be commonly collected for multiple purposes• taxa that are well referenced in the literature• taxa that best represent the variety of form, behavior and ecology• sampled for the greatest proportion of morphology and molecules

Multiple TWiGs• subfamily group representation across TWIG (Chalcidoidea, etc.)• taxa sampled for in depth relationships across taxonomic groups• initial preference for taxa that can be sequenced (later to include all)• taxa that best represent the variety of relationships within clades• sampled for a subset of relevant morphology and molecules

LIST OF EXEMPLAR TAXA

Morphology project• Taxon Sampling

– Skeleton and TWiG taxa– Mainly at subfamily level (multiple genera per subfamily if possible)– Initially guided by molecular analysis– Later addition taxa that cannot be sequenced

• Morphological information– Digitized images from SEM, slides– Scans and PDF’s from literature– Mounted specimens

• Character coding– Initial list from literature (600+ characters and growing [all levels])– Reworked as coding progresses (200-300? characters [all levels])

Thorax/Mesosoma: Lars Vilhelmsen

Head: John Heraty, Johan Liljeblad, Jeremiah George

Legs: Fredrik Ronquist and Matt Buffington

Wings: Sharkey

Abdomen/metasoma: Sharkey

Female genitalia: Andy Boring

Male genitalia: Susanne Schulmeister

Immature Stages: David Wahl

Behavior and life history: Jim Carpenter

Morphology Based Research Units

MorphoNetimage management

• proprietary (proprietary (www.canto.comwww.canto.com))• includes XML metabridge for includes XML metabridge for

queries by XML applicationsqueries by XML applications• ODBC compliant for alternate ODBC compliant for alternate

storage in Oracle or SQL.storage in Oracle or SQL.

MorphBankimage management

Character list, taxon images and annotated matrix

Molecular Data

Skeleton Matrix• Ribosomal genes (18S, 28S D2-D5)• Nuclear genes (fkh, wng, CAD)• Mitochondrial gene (COI)• 4000-5000 kb of sequence data (9 regions/18 reactions)

TWiGs (choices dependent on relevance within taxa)• Ribosomal genes (18S E17-35, 28S D2-D5)• Nuclear genes (fkh)• Mitochondrial gene (COI?)• 1700-2200 kb of sequence (5 regions/10 reactions)

DNA and Specimen Vouchering

Agaonidae 1 2 7 6[5] 1 2 76 757Aphelinidae 5 5 7 2 15 16 30 1165Chalcididae 4 4 5 1 12 19 87 1464Encyrtidae 2 2 2 11 12 460 3735Eucharitidae 3 3 3 40 42 55 423Eulophidae 4 4 4 12 17 297 4472Eupelmidae 1 3 3 2[0] 2 6 45 907Eurytomidae 3 3 3 4 5 88 1424Leucospidae 1 1 1 1 1 4 134Mymaridae 2 2 2 5 8 103 1424Ormyridae 1 1 1 1 1 3 125Perilampidae 2 2 2 9 9 13 262Pteromalidae 7 20 31 14[11] 47 88 588 3506Rotoitidae 0 0 1 1 2 2Signiphoridae 1 1 1 4 4 4 76Tanaostigmatidae 1 1 1 2 2 9 92Tetracampidae 3 3 3 4 5 12 44Torymidae 2 2 2 3 4 68 986Trichogrammatidae 2 2 2 47 47 83 839 Akapalinae 1 1 1 1 1 1 2 Calesinae 1 1 1 1 1 1 3 Philomidinae 1 1 1 3 3 3 15Sum: 19+3 48 64 84 26[20] 224 232 2031 21857

Mymarommatidae 1 1 1 1 1 1 9

chalcidoidmolecular

taxa

possibleTaxon # subfamilies missing # genera # species

SUBFAMILIES69% done19% in queue

GENERA11% done>8% in queue

Eucharitidae

Perilampidae

Trichogrammatidae

Azotinae

Signiphoridae

Coccophaginae

Aphelininae

Eulophidae

Pteromalinae

Gillespie et al. submitted

228S [and 18S] secondary structure for alignment

3h' RAA REC 3q RAA 3q' REC RAA 3g' (23) (4) (24) (4') (25) - - ))))) )))) )Aprosto cetus Ôsp. 3 Õ |GGCUC| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Chrysoc haris pubi corni s |GGCUC| [UAAAA----] [ UUC------- ----- --] [ GAAA------] [--- ----- ----- -GGA] [---- ----- ----] |UUACC|Cirrosp ilus sp. |GGCUC| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Deutere uloph us sp. |GGCUC| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Henryan a mag nific a |GGCUC| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Horisme nus sp. |GGCUC| [UACAA----] [ UUC------- ----- --] [ GAAA------] [--- ----- ----- -GGA] [---- ----- ----] |UUACC|Melitto bia d igita ta |GGCUC| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Polston ia pe lagoc oryph a |GGCUC| [UUAC-----] [AUUCGUG------- ---] [UUUA------] [--- ----- --CACGGAU] [---- ----- ----] |AUACC|Gastran cistr us sp. |GGCUC| [UUGC-----] [AUUC------- ----- -] [G AAA------] [--- ----- ----- GGAU] [---- ----- ----] |AUACC|Scutell ista cyane a |GGCUC| [UUAU-----] [AUUC------- ----- -] [G AAC------] [--- ----- ----- GGAU] [---- ----- ----] |AUACC|Eunotus sp. |GGCUC| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Rileya sp. (t -comp lex) |GGCUC| [GUUAC----] [ AUUC------- ----- -] [G AAC------] [--- ----- ----- GGAU] [---- ----- ----] |GUACC|Rileya sp. 1 |GGCUC| [AGAC-----] [AUCC------- ----- -] [ AAUC------] [--- ----- ----- GGAU] [---- ----- ----] |AUACC|Eurytom a sp. 1(a ) |GGCUC| [---- ----- ] [UCU------- ----- --] [ CGAAA-----] [---- ----- ----- GGG] [---- ----- ----] |AUACC|Eurytom a sp. 1(b ) |GGCUC| [---- ----- ] [UCU------- ----- --] [ CGAAU-----] [---- ----- ----- AGG] [---- ----- ----] |AUACC|Brasema sp. |GGCUC| [UU------- ] [UGAU------- ----- -] [ AAUAC-----] [---- ----- ----- AUA] [---- ----- ----] |AUACC|Eupelmi s sp. |GGCUC| [UUAC-----] [AUUC------- ----- -] [G AAC------] [--- ----- ----- GAAU] [---- ----- ----] |AUACC|Cheilon eurus fulv escen s |GGCUC| [---- ----- ] [UAAA------- ----- -] [ CGCA------] [--- ----- ----- UAUA] [---- ----- ----] |UUACC|Aceroph agus sp. |GGCUC| [GUU------] [GCCGC------- ----- ] [CUCGC-----] [---- ----- ---GCGGC] [---- ----- ----] |AUACC|Metaphy cus h elvol us |GGCUC| [GUU------] [GCCGCG------- ----] [CUCG------ ] [-- ----- ---- CGCGGC] [---- ----- ----] |AUACC|Neodusm etia sangw ani |GGCUC| [GU------- ] [UAUUAUUU------- --] [ UCUUUU----] [ ----- ---- AAAUAAUA] [---- ----- ---U] |CUACC|Leptoma stide a sp. |GGCUC| [AUU------] [CCAUA------- ----- ] [UUCAU-----] [---- ----- ---UAUGG] [---- ----- -AAA] |UUACC|Anagyru s sp. |GGCUC| [GU------- ] [UAUUA------- ----- ] [AUUU------] [--- ----- ---- UAAUA] [---- ----- ---U] |AUACC|Anagyru s ?pseudoc occi |GGCUC| [---- ----- ] [GUUAU------- ----- ] [UAUUC-----] [---- ----- ---AUAAU] [---- ----- --AU] |AUACC|Plagiom erus ?diaspid is |GGCUC| [U------- -] [ GGGCGAA------- ---] [CUCG------ ] [-- ----- ---UUCGCUC] [---- ---- UCUAA] |GUACC|Comperi ella bifas ciata |GGCUC| [U------- -] [ GUGGGGAGA------- -] [G CAA------] [--- ----- UCUCUUCAC] [---- GUUAAAAAA ] |AUACC|Trechni tes sp. |GGCUC| [U------- -] [ GUUU------- ----- -] [ UUU------- ] [-- ----- ----- -GUAC] [---- ----- ---U] |CUA-C|Copidos oma f lorid anum |GGCUC| [G--- ----- ] [GAAAAGC------- ---] [CUAGC-----] [---- ----- -GCUUUUC] [---- -AAUUAAAA] |UUACC|Copidos oma t runca tellu m |GGCUC| [G--- ----- ] [A------- ----- ----] [CUA------- ] [-- ----- ----- ---- U] [---- ----- ----] |UUACC|Ooencyr tus j ohnso ni |GGCUC| [UCG------ ] [GCGCGUGAG------- -] [ UUCG------ ] [-- ----- -CUCGCGCGU] [---- ----- ----] |CGACC|Ablerus clis iocam pae |GGCUC| [U------- -] [ GGCUGCU------- ---] [UUCG------ ] [-- ----- ---GGCAGCC] [---- ---- UCCGA] |UUACC|Ablerus sp. 2 |GGCUC| [C------- -] [ GGCUGCUCC------- -] [ CUCG------ ] [-- ----- -GGGGCAGUC] [---- UCAUUCGAA] |UUACC|Ablerus sp. 3 |GGCUC| [U------- -] [ GGCUGCU------- ---] [UUCG------ ] [-- ----- ---GGCAGCC] [---- ---- UCCGA] |UUACC|Ablerus sp. 4 |GGCUC| [C------- -] [ GGUUGCUUU------- -] [ CUCG------ ] [-- ----- -AAAGCAACU] [---- ----- CGUA] |UUACC|Ablerus sp. 5 |GGCUC| [---- ----- ] [GGGGCUGCU------- -] [ UUCG------ ] [-- ----- -GGCAGUCUC] [---- -UCUAACGA] |UUACC|Ablerus sp. 6 |GGCUC| [U------- -] [ GUUGC------- ----- ] [UCUG------ ] [-- ----- ----- GCAAU] [---- ----- --CA] |UUACC|Ablerus sp. 7 |GGCUC| [G--- ----- ] [AAUUGCU------- ---] [CUCG------ ] [-- ----- ---GGCAAUU] [---- -AUUAGAAA] |AUACC|Ablerus sp. 8 |GGCUC| [C------- -] [ AAUUGCU------- ---] [CUCG------ ] [-- ----- ---GGCAAUU] [---- ---AUCGAA] |UUACC|Ablerus sp. 10 |GGCUC| [UG------ -] [ UUGCUCU------- ---] [CGG----- --] [ ----- ----- GCAAUUA] [---- ----- ----] |UUACC|Chartoc erus sp. 2 |GGCUC| [UCUCGA---] [ CAGCUUAU------- --] [ AUCA------] [--- ----- --AUAAGCG] [---- ----- ---A] |AUACC|Chartoc erus sp. 6 |GGCUC| [UUUCGA---] [ CAGCUUA------- ---] [UUCG------ ] [-- ----- ---- UAAGCG] [---- ----- ---U] |AUACC|Signiph ora sp. 1 |GGCUC| [UCCCAA---] [ GUCC------- ----- -] [ UUCAUU----] [ ----- ----- ---GGAC] [---- ----- ----] |AUGCA|Signiph ora sp. 2 |GGCUC| [UCGAAA---] [ GUUCGC------- ----] [CUCG------ ] [-- ----- ---- GCGGAC] [---- ----- ----] |AUGCA|Signiph ora sp. 3 |GGCUC| [UAUAAUGC-] [GUUCGUCCCC------- ] [UCG------ -] [- ----- -GGGGGCGGAC] [---- ----- ----] |AUGCA|Signiph ora sp. 5 |GGCUC| [UCUUAU---] [ GUUCAAUCCCC------] [UCAC------] [--- ---GGGGAUGGGAC] [---- ----- ----] |GCGCA|Signiph ora sp. 7 |GGCUC| [UCUUAU---] [ GUUCG------- ----- ] [UUUA------] [--- ----- ---- CGGAC] [---- ----- ----] |AUGCA|Signiph ora sp. 8 |GGCUC| [UCUUAU---] [ GUUCAAUCCCC------] [UCAC------] [--- ---GGGGAUAGGAC] [---- ----- ----] |GCGCA|Clytina sp. |GGCCU| [CGUUU----] [ GUUCGCCG------- --] [ UUUA------] [--- ----- -CGGCGGAC] [---- ----- ----] |GUACC|Thysanu s sp. 4 |GGCCU| [CGUUU----] [ GUUCGCCCU------- -] [ UCGC------] [--- ----- GGGGCGGAC] [---- ----- ----] |GUACC|Aphytis ling nanen sis |GGCUA| [---- ----- ] [-- ----- ----- -----] [--- ----- --] [ ----- ----- ----- --] [ ----- ----- ---] |UUACC|Aphelin inae new g enus |GGCUC| [UUUCAA---] [ GUUUCU------- ----] [CUUG------ ] [-- ----- ---- AGAAAC] [---- ----- ---A] |UUACC|Coccobi us sp. 1( b) |GGCCU| [UAAAU----] [ AUUCGA------- ----] [ACGC------] [--- ----- ---UCGGAU] [---- ----- ----] |AUACC|Coccobi us sp. 2 |GGCCU| [CCGA-AU--] [AUUC------- ----- -] [G AAC------] [--- ----- ----- GGAU] [---- ----- ----] |AUACC|Coccoph agus sp. 3 |GGCUC| [GUCAAC---] [ GUUCGCCUCU------- ] [CACAA-----] [---- ---GGAGGCGGAC] [---- ----- ----] |CUACC|Coccoph agus rusti |GGCUC| [UCGAAUG--] [ GUUCGCCCCCU------] [CACA------] [--- --AGGGGGGCGGAU] [---- ----- ----] |UUACC|Coccoph agoid es fuscipe nnis |GGCUC| [---- ----- ] [-- ----- ----- -----] [UCUA------] [--- ----- ----- ----] [---- ----- ----] |UUACC|Encarsi a nig ricep hala |GGCUC| [UUAAC----] [ GUUCGACCC------- -] [ UUCGC-----] [---- ---- GGGUCGGAC] [---- ----- ----] |AUACC|Mirufen s sp. 1 |GGCUC| [UGUCGAAC-] [AUUCGUCG------- --] [ CUCG------ ] [-- ----- --CGACGGAU] [---- ----- ----] |AUACC|Mirufen s sp. 2 |GGCUC| [UG------ -] [ GCGCGCGGGGCGCGUUC] [GCAA------] [GGGCGCGAUCCGUGUGC] [AAAUAAAAUGAAA] |AUACC|

Summary

• Skeleton taxa and TWiGs (both altered through reciprocal illumination)

• Layered molecular approach (most detailed for skeleton taxa)

• Collaborator Networks

• Databases for sequence and image data (Filemaker, MorphoNet, MorphBank)

• Web page development (glossaries, keys, images)

• Independent analysis (parsimony to Bayesian)

Problems

• Interoperability (different software preference)

• difficulties of meshing skeleton and TWiG focus

But we do have a logo and t-shirt!!!

Where are the *@$#& chalcids?

Acknowledgements

• National Science Foundation• Sharkey lab• Ronquist lab• Heraty lab• Carpenter lab

• Katja Seltman• Susanne Schulmeister• Johan Liljeblad• David Hawks• Matt Buffington• etc., etc., etc……