PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS

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Chapter 3 is a phylogenetic study of the genus Phytomyza sensu lato, using over 3,000 nucleotides of DNA sequence data from three genes. Results indicate that the genus Chromatomyia, considered by some as synonymous with Phytomyza, is in fact polyphyletic and nested within Phytomyza. Possible parallelism in a biological trait (internal pupation in leaf tissue) which is one of the defining traits of species in the former Chromatomyia is discussed. In addition, the internal classification of Phytomyza is assessed and revised insofar as the data permit. Divergence times for the Agromyzidae, and also for Phytomyza and related genera, were estimated using a molecular phylogeny calibrated by three agromyzid fossils (Chapter 4). Results suggest that the temperate Phytomyza group of genera originated in the relatively warm Eocene epoch. Ranunculaceae, a primitive plant family, is inferrred as the ancestral host for a clade including most Phytomyza species, but is probably secondary to feeding on more derived plant families (“asterid clade”). Ten clades were identified for comparison of diversification rates between Ranunculaceae- and asterid-feeding lineages, which showed that asterid-feeding clades exhibit higher rates of diversification. Phytomyza originated approximately at the early Oligocene global cooling event, but contrary to expectations, diversification significantly slowed during the Oligocene cool period, when suitable habitats for Phytomyza were presumably widespread.
PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS: PHYLOGENY AND EVOLUTION OF PHYTOMYZA LEAF-MINING FLIES (DIPTERA: AGROMYZIDAE) By Isaac Scott Winkler Dissertation submitted to the Faculty of the Graduate School of the University of Maryland, College Park, in partial fulfillment of the requirements for the degree of Doctor of Philosophy 2008 Advisory Committee: Professor Charles Mitter, Chair Sonja J. Scheffer Wayne N. Mathis David J. Hawthorne Charles F. Delwiche
Page 1: ABSTRACT Title of Document: PATTERN
Page 5 and 6: Note: This dissertation should not
Page 7 and 8: Table of Contents Foreword.........
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.
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diversification effects known (Coyn
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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
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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
Page 228 and 229:
Specialization, speciation, and rad
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PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS

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