PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS

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Acknowledgements I thank first my advisors and other graduate committee members for the guidance, funding, and patience which made this dissertation possible. Supplementary funds for travel and field work were provided by the University of Maryland Center for Biodiversity, the Williston Diptera Research fund of the Smithsonian Institution, and the U.S. Department of Agriculture. Thanks also to the following for hosting visits for collection of specimens and/or access to bulk samples: Charles Godfray and the Centre for Population Biology of Imperial College, London; Greg Zolnerowich and Bob Kula (Kansas State University); and Riley Nelson (Brigham Young University). Saulius Pakalniškis (Institute for Ecology, Vilnius, Lithuania) and Charles Godfray (Oxford University) sent important specimens which were much appreciated. Conrad Labandeira (US National Museum) provided access to important agromyzid fossils. Special appreciation is extended to Matthew Lewis (USDA Systematic Entomology Laboratory), who gave valuable technical assistance in the laboratory and provided most of the sequences for PGD. Leo Shapiro provided additional sequences for holly leaf miners. Akito Kawahara translated an important reference from Japanese. Alexei Drummond provided useful advice regarding his BEAST program. Thanks also to Scott Wing, Conrad Labandeira, and Vicki Funk (U.S. National Museum) for helpful discussions and for reading over drafts of the final chapter. Finally, I thank Joelynn Winkler for assistance in field work, and for her patient support of my studies. iv
Table of Contents Foreword....................................................................................................................... ii Acknowledgements...................................................................................................... iv Table of Contents.......................................................................................................... v List of Tables .............................................................................................................. vii List of Figures............................................................................................................viii Chapter 1: Introduction................................................................................................. 1 Chapter 2: The Phylogenetic Dimension of Insect-Plant Interactions: A Review of Recent Evidence........................................................................................................................ 6 Conservatism of Host-Plant Use..................................................................................... 8 Conservation of Host-Taxon Associations.................................................................. 9 Variation in Rates of Major Host Shift ..................................................................... 12 Other Conserved Aspects of Host Use...................................................................... 15 Parallelism, Reversal, and Genetic Constraints on Host Shift................................. 17 Conservatism, Host Shifts, and Speciation ............................................................... 19 Phylogenesis of Host Range...................................................................................... 22 Signatures of Long-Term History in Extant Insect-Plant Interactions ......................... 29 Diversification of Phytophagous Insects ...................................................................... 39 Synopsis and Conclusions............................................................................................. 50 Chapter 3: Phylogeny of the Leaf-mining Fly Genus Phytomyza Fallén s.lat. (Diptera: Agromyzidae), with Comments on Life History Traits and on the Status of Chromatomyia Hardy.................................................................................................. 55 Introduction................................................................................................................... 55 Methods......................................................................................................................... 62 Taxon sampling......................................................................................................... 62 Extraction and sequencing........................................................................................ 69 Phylogenetic analysis................................................................................................ 72 Results........................................................................................................................... 74 Discussion..................................................................................................................... 83 Delimitation of Phytomyza ....................................................................................... 83 Species groups, major clades and infra-generic classification of Phytomyza ......... 89 The status of Chromatomyia ................................................................................... 105 Evolution of life history and host use...................................................................... 109 Chapter 4: The History of a Temperate Adaptive Radiation: Diversification and Host Use Evolution in Phytomyza Leaf-mining Flies (Diptera: Agromyzidae)....................... 117 Introduction................................................................................................................. 117 Phylogenetic patterns in insect/plant evolution...................................................... 117 Earth history, niche conservatism, and “biome tracking”..................................... 119 Study group: leaf-mining flies................................................................................. 121 Fossil history........................................................................................................... 126 Study aims ............................................................................................................... 131 Methods....................................................................................................................... 131 Data sets and divergence time estimation .............................................................. 131 v
Page 1 and 2: ABSTRACT Title of Document: PATTERN
Page 3 and 4: PATTERNS OF DIVERSIFICATION IN PHYT
Page 5: Note: This dissertation should not
Page 9 and 10: List of Tables Table 2.1. Summary o
Page 11 and 12: Chapter 1: Introduction With nearly
Page 13 and 14: Determining the historical timing o
Page 15 and 16: compared to the history of the host
Page 17 and 18: The “escape and radiation” mode
Page 19 and 20: Conservation of Host-Taxon Associat
Page 21 and 22: Fig. 2.1. Frequency of host shifts
Page 23 and 24: analysis with better control for ph
Page 25 and 26: feeding lineage, there have been re
Page 27 and 28: as the phylogenetic evidence cannot
Page 29 and 30: Conservatism, Host Shifts, and Spec
Page 31 and 32: Table 2.1. Summary of host and dist
Page 33 and 34: The study of host range evolution i
Page 35 and 36: Table 2.2. Synopsis of nine recent
Page 37 and 38: of transition to and from specializ
Page 39 and 40: Several of the foregoing hypotheses
Page 41 and 42: (reviews in Magallón 2004, Welch a
Page 43 and 44: Table 2.3. Synopsis of 18 recent st
Page 45 and 46: Datings based on fossils, molecular
Page 47 and 48: the most basal seed plants. The fiv
Page 49 and 50: Aphids, agromyzids and other groups
Page 51 and 52: apidly multiplying lineage” (pg.
Page 53 and 54: diversification effects known (Coyn
Page 55 and 56: sound detectors allowing adults to
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insect diversification and correlat
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comparisons (Table 2.4). The only p
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We have focused on phylogenies at t
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6. Because phylogenetic studies dir
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Chapter 3: Phylogeny of the Leaf-mi
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phylogenetic relationships between
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Chromatomyia alone is monophyletic,
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Fig. 3.1. Phylogeny of host plant f
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Table 3.1. Species sampled for this
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LocColl- GenBank Accession # Specie
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LocColl- GenBank Accession # Specie
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newly obtained for this study, exce
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Table 3.2. Primers sequences used f
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esults. The tree of highest likelih
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elow 50% for many deeper nodes of t
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Phytomyza s. nov. Phytomyza main li
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Within Phytomyza sensu novo, the tw
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was monophyletic with strong suppor
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monophyly of the entities Phytomyza
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Fig. 3.5. Nucleotide sequence of va
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accommodate the incongruous charact
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Table 3.4. Revised species groups o
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excluded from Phytomyza group min.
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particularly primitive characterist
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including species feeding on either
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eduction of the paired tubules of t
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For reasons including the need to c
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are herbaceous. The inclusion of P.
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(Spencer 1969), is surprising, but
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) a slipper-shaped, usually lightly
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apodeme, and possibly also by a red
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Hering), both possibly associated w
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pupate internally in the host plant
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The phylogenetic position of P. gym
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Although a detailed investigation o
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phylogenies between the plant speci
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pronounced expansion of open habita
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of about 750 described species in t
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Region # total described agromyzid
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difficult. Despite this, and althou
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leaf mines in Platanus (Crane and J
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pattern. Most of the Mexican Hat sp
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partitions was not available from o
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independent estimates not based on
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estimation, integrating over the li
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with supplementary results obtained
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diversification rate in these clade
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warming event of approximately 24 M
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strong (83%) support in the bootstr
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Table 4.3. Selected clades of Phyto
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Ancestral host reconstruction Ances
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Fig. 4.4. Clade size vs. clade age
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DT (based on benthic isotope ratios
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intervals (Table 4.2), especially g
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albipennis/ranunculella grps., nota
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clades of Lamiales (e.g. Orobanchac
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emaining three clades (A2, A3, A5),
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event may have also occurred very r
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paleontological data, as well as so
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the Phytomyza group which are not w
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tropical lineages have simply had m
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allow us to reliably estimate the s
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widespread climate change and Phyto
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fauna as “a host of intriguing qu
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Phytomyza cecidonomia Hering, 1937
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Phytomyza conii Hering, 1931 [PA] (
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Phytomyza orientalis Spencer, 1962
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Phytomyza nemopanthi Griffiths & Pi
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Phytomyza thalictricola Hendel, 192
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Phytomyza nigritella Zetterstedt, 1
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Literature Cited Agrawal, A. A., an
Page 200 and 201:
Bokma, F. 2003. Testing for equal r
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Cook, L. G., and P. J. Gullan. 2004
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Drummond, A. J., and A. Rambaut. 20
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of the Compositae. Biol. Skr. 55: 3
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Grimaldi, D. A. 1999. The co-radiat
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Janis, C. M. 1993. Tertiary mammal
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Kumada, T. 1984. K y ju no pisufure
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McAlpine, J. F. 1989. Phylogeny and
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Nee, S., T. G. Barraclough, and P.
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Pellmyr, O. 2003. Yuccas, yucca mot
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Rundle, H. D. and P. Nosil. 2005. E
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adiation: the evolutionary biology
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Strong, D. R., J. H. Lawton, and R.
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Vrba, E. S. 1985. Environment and e
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Specialization, speciation, and rad
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