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Finding the nearest relatives of Nasonia (Hymenoptera: Pteromalidae) Roger Burks University of...
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![Page 1: Finding the nearest relatives of Nasonia (Hymenoptera: Pteromalidae) Roger Burks University of California, Riverside Department of Entomology.](https://reader036.fdocuments.us/reader036/viewer/2022081519/56649d375503460f94a0f8aa/html5/thumbnails/1.jpg)
Finding the nearest relatives of Nasonia (Hymenoptera: Pteromalidae)
Roger Burks
University of California, Riverside
Department of Entomology
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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
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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
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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)
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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
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Why Nasonia’s relationships 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
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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
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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?
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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
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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
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Combined 28S and Wingless molecular results, Parsimony (PAUP)
Numbers indicate bootstrap support (1000 replicates)
Agrees with simple POY run in topology
1176 steps in PAUPrci = 0.209ri = 0.403
black = Pteromalinaered = other Pteromalids* = Wolbachia positive
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Combined 28S and Wingless molecular results, Mr. Bayes 3.1
Numbers indicate posterior probability
black = Pteromalinaered = other Pteromalids* = Wolbachia positive
6 parameters, 4 chains, partitioned by gene region, 1 million generations
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Combined 28S and Wingless molecular results, Likelihood
black = Pteromalinaered = other Pteromalids* = Wolbachia positive
model: GTR+I+Gprogram: PAUP
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Testing hypotheses not present in the optimum maximum likelihood tree (500 total sampled trees for test)
constraint tree with: au test p value sh test p value
Nasonia+ Urolepis clade 0.81 1.00
Nasonia + Trichmalopsis + Urolepis paraphyly
0.50 0.85
monophyletic Pteromalinae
0.38 0.80
Nasonia + Trichomalopsis clade
0.23 0.79
monophyletic Trichomalopsis
0.07 0.70
paraphyletic Nasonia 0.01** 0.50
au = approximately unbiased test (Shimodaira 2002)sh = Shimodaira-Hasegawa test (Shimodaira & Hasegawa 1999)
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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 shortCytochrome Oxidase I & II—AT richness
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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
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They differ by only one base pair in 28S??
Trichomalopsis sarcophagae
Nasonia vitripennis
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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
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AcknowledgmentsAdvisory 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
Heraty lab:Dave HawksJohan LiljebladJames MunroJeremiah GeorgeJason MotternChrissy RomeroAdena Why
Jutta BurgerMatt Buffington
Funded by: NSF FIBR: 0328363