PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS

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Ancestral host reconstruction ................................................................................. 135 Diversification rate analysis................................................................................... 136 Results......................................................................................................................... 142 Divergence time estimates ...................................................................................... 142 Diversification rate analysis................................................................................... 148 Discussion................................................................................................................... 153 Fossil calibrations and age of the Agromyzidae..................................................... 153 Ancestral host of Phytomyza and early adaptive radiation.................................... 155 Insect/host plant evolution: delayed colonization and co-diversification .............. 156 Evolutionary context: history of north temperate climate and flora ...................... 160 Origin of the Phytomyza group and timing of Cenozoic temperate radiation ....... 164 Biome tracking in Phytomyza?............................................................................... 169 Conclusion .................................................................................................................. 171 Appendix A............................................................................................................... 175 Literature Cited ......................................................................................................... 188 vi
List of Tables Table 2.1. Summary of host and distribution overlap versus non-overlap for 145 sister species pairs from 45 phytophagous insect phylogenies. Table 2.2. Synopsis of nine recent studies bearing on phylogenetic patterns of host range. Table 2.3. Synopsis of 18 recent studies testing for parallel insect/plant phylogenesis at lower taxonomic levels. Table 2.4. Sister group diversity comparisons between endo- and exophytophage lineages Table 3.1. Species sampled for this study, arranged by species group. Table 3.2. Primers used for this study. Table 3.3. Number of parsimony informative sites and average pairwise distances for the three gene partitions used in this study. Table 3.4. Revised species groups of Phytomyza. Table 4.1. Number of Phytomyza species (including Chromatomyia) reported in various regional catalogues, surveys, or revisions, showing latitudinal diversity gradient. Table 4.2. Divergence times of Phytomyza and other clades. Table 4.3. Selected clades of Phytomyza showing minimum ages, diversities, and clade size probabilities. vii
Page 1 and 2: ABSTRACT Title of Document: PATTERN
Page 3 and 4: PATTERNS OF DIVERSIFICATION IN PHYT
Page 5 and 6: Note: This dissertation should not
Page 7: Table of Contents Foreword.........
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
Page 57 and 58: 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
Page 202 and 203:
Cook, L. G., and P. J. Gullan. 2004
Page 204 and 205:
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
Page 212 and 213:
Kumada, T. 1984. K y ju no pisufure
Page 214 and 215:
McAlpine, J. F. 1989. Phylogeny and
Page 216 and 217:
Nee, S., T. G. Barraclough, and P.
Page 218 and 219:
Pellmyr, O. 2003. Yuccas, yucca mot
Page 220 and 221:
Rundle, H. D. and P. Nosil. 2005. E
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adiation: the evolutionary biology
Page 224 and 225:
Strong, D. R., J. H. Lawton, and R.
Page 226 and 227:
Vrba, E. S. 1985. Environment and e
Page 228 and 229:
Specialization, speciation, and rad
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PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS

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