Finding the nearest relatives of nasonia hymenoptera pteromalidae
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Finding the nearest relatives of Nasonia (Hymenoptera: Pteromalidae). Roger Burks University of California, Riverside Department of Entomology. What is Nasonia ?. Gregarious puparial parasitoids of calyptrate flies in bird nests and refuse

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Finding the nearest relatives of Nasonia (Hymenoptera: Pteromalidae)

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Finding the nearest relatives of nasonia hymenoptera pteromalidae

Finding the nearest relatives of Nasonia (Hymenoptera: Pteromalidae)

Roger Burks

University of California, Riverside

Department of Entomology


What is nasonia

What is Nasonia?

  • Gregarious puparial parasitoids of calyptrate flies in bird nests and refuse

  • Model system, better known than any other species of Chalcidoidea—genome project ongoing

  • Three species, each infected by two unique strains of Wolbachia


The three species of nasonia

The three species of Nasonia

  • Females almost identical (Darling & Werren 1990)

  • Males differ in degree of wing reduction

  • Nasonia vitripennis worldwide, synanthropic

  • N. giraulti in eastern North America, N. longicornis in western North America

    • specialized on flies in bird nests


Wolbachia basic background

Wolbachia basic background

  • Bacteria infecting arthropods and filarial nematodes

  • Transmitted vertically from mother to offspring (Binnington & Hoffmann 1989)

  • Cause crossing incompatibility in Nasonia (Breeuwer & Werren 1990)

  • Phylogenetic congruence between bacteria and host usually absent– horizontal transmission?

  • May cause rapid speciation in arthropods (Laven 1959, 1967; Breeuwer & Werren 1990)


How wolbachia affects nasonia

How Wolbachia affects Nasonia

  • Cytoplasmic Incompatibility (Breeuwer & Werren 1990)

    • Causes death of offspring of mothers that do not have same Wolbachia strains as the father

  • Incompatible crosses:

    • Uninfected female x infected male

    • Infected female x male infected by at least one different strain

  • Infection rate near 100% in wild Nasonia

    • “Cured” colonies used to study Wolbachia effects in lab


Why nasonia s relationships still need studying

Why Nasonia’srelationships still need studying

  • Nasonia is a model system for evolutionary biology studies, yet…

  • Ancestral states cannot be inferred with only three analyzed species!

  • No agreement in classification of wasps in its family (Pteromalidae)

  • Needed: means to reject some pteromalids as close Nasonia relatives


Pteromalidae is a scary taxon

Pteromalidae is a scary taxon

  • 587 genera in 31 subfamilies

  • Pteromalinae with only 283 genera

  • Parasitoids of various terrestrial arthropods

  • No previous phylogenetic analysis using more than 10 pteromaline genera

  • Previous analyses with either morphology only or 28S ribosomal sequences only


Pteromalinae molecular vs morphological rates of evolution

Pteromalinae molecular vs. morphological rates of evolution

  • 283 genera of Pteromalinae, but...

  • 28S D2 sequence divergence equal to that of the genus Aphelinus (Heraty 2004)

  • Rapid morphological evolution or ribosomal constraints?

  • Rapid evolution due to Wolbachia?


Tools for the search

Tools for the search

  • Morphology

    • 105 morphological characters (work in progress)

  • 28S D2-D5 ribosomal DNA, Wingless

    • Secondary structure alignment for 28S (Gillespie et al. 2005) to be compared with POY results

  • Analysis with parsimony (PAUP, TNT, POY), maximum likelihood, Mr. Bayes

  • Hypothesis testing with ML using CONSEL


Outgroup selection

Outgroup selection

  • Based on Heraty lab matrix of Chalcidoidea

    • 28S D2-D5, 18S E17-E35 ribosomal DNA

    • 471 taxa (including outgroups)

    • All families, 84 total subfamilies represented

  • Subfamilies Diparinae, Ormocerinae are legitimate outgroups for Pteromalinae


Finding the nearest relatives of nasonia hymenoptera pteromalidae

Combined 28S and Wingless molecular results, Parsimony (PAUP)

black = Pteromalinae

red = other Pteromalids

* = Wolbachia positive

Numbers indicate

bootstrap support

(1000 replicates)

Agrees with simple POY

run in topology

1176 steps in PAUP

rci = 0.209

ri = 0.403


Combined 28s and wingless molecular results mr bayes 3 1

Combined 28S and Wingless molecular results, Mr. Bayes 3.1

black = Pteromalinae

red = other Pteromalids

* = Wolbachia positive

6 parameters, 4 chains, partitioned

by gene region, 1 million

generations

Numbers indicate

posterior probability


Combined 28s and wingless molecular results likelihood

Combined 28S and Wingless molecular results, Likelihood

black = Pteromalinae

red = other Pteromalids

* = Wolbachia positive

model: GTR+I+G

program: PAUP


Finding the nearest relatives of nasonia hymenoptera pteromalidae

Testing hypotheses not present in the optimum maximum likelihood tree (500 total sampled trees for test)

au = approximately unbiased test (Shimodaira 2002)

sh = Shimodaira-Hasegawa test (Shimodaira & Hasegawa 1999)


Finding the nearest relatives of nasonia hymenoptera pteromalidae

Problem: Not enough variation to have statistical power

Solution: Add a more rapidly evolving gene

Candidates:

Long-wavelength Rhodopsin—multiple copies?

Pten—contains intron, but short

Cytochrome Oxidase I & II—AT richness


Perspective

Perspective

  • Trichomalopsis sarcophagae 28S sequence (>1100 base pairs) differs from that of Nasonia vitripennis by only 1 base pair

  • Sampling remains incomplete

    • Nasonia not well-surveyed in Palearctic region

    • Trichomalopsis with 54 species!

Trichomalopsis microptera male


They differ by only one base pair in 28s

They differ by only one base pair in 28S??

Trichomalopsis sarcophagae

Nasonia vitripennis


Further goals

Further goals

  • Sequence from more species of Trichomalopsis, other genera near Nasonia (>120 specimens to be sequenced)

  • Finish morphological analysis

  • Wolbachia survey across Pteromalinae, comparing bacteria and wasp phylogenies


Acknowledgments

Acknowledgments

Heraty lab:

Dave Hawks

Johan Liljeblad

James Munro

Jeremiah George

Jason Mottern

Chrissy Romero

Adena Why

Jutta Burger

Matt Buffington

Funded by: NSF FIBR: 0328363

Advisory committee:

John Heraty

Richard Stouthamer

Bob Luck

Cheryl Hayashi

Jack Werren

Matt Yoder

Doug Yanega

Serguei Triapitsyn

Lara Baldo

James Russell

Genet Tulgetske

Danel Vickerman


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