PATTERNS OF DIVERSIFICATION IN PHYTOPHAGOUS INSECTS

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significant host range conservation? Source Significant host taxon conservation? forms of directionality reported Criterion for ”specialist” vs. “generalist” #specialists vs. #generalists <strong>IN</strong>SECT TAXON Taxon sample no: P>0.50, PTP, all criteria 8 no: P > 0.18 (PTP) for host subtribe use depends on criterion 1 vs. >1 host genus, or subtribe, or tribe 11 vs. 8, or 14 vs. 5, or 15 vs. 4 19/59 spp.; sampling limited to part of Brazil Diptera: Tephritidae: Tomoplagia host genus conserved, P 1 host genus 14 spp. (17 taxa)/21 spp. (remainder described subsequently) 11 vs. 6 Phasmida: Timematidae: Timema Abbreviations: gen. (generalist); PTP (Permutation Tail Probability test); spec. (specialist); spp. (species); vs. (versus). Sources: 1. Dobler et al. (1996), 2. Kelly and Farrell (1998), 3. Morse and Farrell (2005), 4. Wahlberg (2001), 5. Janz et al. (2001). 6. Weingartner et al. (2006), 7. Rubinoff and Sperling (2002), 8. Yotoko et al. (2005), 9. Crespi and Sandoval (2000). 26
of transition to and from specialization, the rates can most often be closely predicted from just the proportions of specialists and generalists among the terminal taxa (Nosil 2002, Nosil and Mooers 2005). This outcome, intuitively expected if the states are distributed randomly on the tree, might be taken to suggest that phylogenies have little to contribute to the understanding of host range evolution. And indeed, it is possible that much of the variation in host range analyzed so far is in fact phylogenetically “random” in the sense of reflecting idiosyncratic local fluctuation, for example in the availability of, and/or selective advantage of using, particular hosts. This may be especially true when all the species within the study group are specialists in the broad sense of feeding on plants in, for instance, the same family. As several authors have noted, however, it is plausible that larger-scale phylogenetic regularities remain to be discovered, through the elaboration of more detailed, process-oriented models of host range evolution (Stireman 2005). Multiple approaches can be distinguished. Thus, host range might be thought of as a trait phylogenetically ephemeral in itself, but with probabilities of change predictable from the states of other, more conserved features, inviting use of the “comparative method.” For example, distribution of the use of two versus more than two tribes of legumes appears by itself to be random on a phylogeny of the seed beetle genus Stator. Closer inspection, however, shows that independent origins of broader host range are significantly concentrated in lineages which oviposit on pre-dispersal seeds, rather than on intact seed pods or dispersed seeds (Morse and Farrell 2005). 27
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 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: Table 2.2. Synopsis of nine recent
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
Page 59 and 60: comparisons (Table 2.4). The only p
Page 61 and 62: We have focused on phylogenies at t
Page 63 and 64: 6. Because phylogenetic studies dir
Page 65 and 66: Chapter 3: Phylogeny of the Leaf-mi
Page 67 and 68: phylogenetic relationships between
Page 69 and 70: Chromatomyia alone is monophyletic,
Page 71 and 72: Fig. 3.1. Phylogeny of host plant f
Page 73 and 74: Table 3.1. Species sampled for this
Page 75 and 76: LocColl- GenBank Accession # Specie
Page 77 and 78: LocColl- GenBank Accession # Specie
Page 79 and 80: newly obtained for this study, exce
Page 81 and 82: Table 3.2. Primers sequences used f
Page 83 and 84: esults. The tree of highest likelih
Page 85 and 86: elow 50% for many deeper nodes of t
Page 87 and 88:
Phytomyza s. nov. Phytomyza main li
Page 89 and 90:
Within Phytomyza sensu novo, the tw
Page 91:
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
Page 100 and 101:
Table 3.4. Revised species groups o
Page 102 and 103:
excluded from Phytomyza group min.
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particularly primitive characterist
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including species feeding on either
Page 108 and 109:
eduction of the paired tubules of t
Page 110 and 111:
For reasons including the need to c
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are herbaceous. The inclusion of P.
Page 114 and 115:
(Spencer 1969), is surprising, but
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) a slipper-shaped, usually lightly
Page 118 and 119:
apodeme, and possibly also by a red
Page 120 and 121:
Hering), both possibly associated w
Page 122 and 123:
pupate internally in the host plant
Page 124 and 125:
The phylogenetic position of P. gym
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Although a detailed investigation o
Page 128 and 129:
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
Page 134 and 135:
Region # total described agromyzid
Page 136 and 137:
difficult. Despite this, and althou
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leaf mines in Platanus (Crane and J
Page 140 and 141:
pattern. Most of the Mexican Hat sp
Page 142 and 143:
partitions was not available from o
Page 144 and 145:
independent estimates not based on
Page 146 and 147:
estimation, integrating over the li
Page 148 and 149:
with supplementary results obtained
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diversification rate in these clade
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warming event of approximately 24 M
Page 154 and 155:
strong (83%) support in the bootstr
Page 156 and 157:
Table 4.3. Selected clades of Phyto
Page 158 and 159:
Ancestral host reconstruction Ances
Page 160 and 161:
Fig. 4.4. Clade size vs. clade age
Page 162 and 163:
DT (based on benthic isotope ratios
Page 164 and 165:
intervals (Table 4.2), especially g
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albipennis/ranunculella grps., nota
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clades of Lamiales (e.g. Orobanchac
Page 170 and 171:
emaining three clades (A2, A3, A5),
Page 172 and 173:
event may have also occurred very r
Page 174 and 175:
paleontological data, as well as so
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the Phytomyza group which are not w
Page 178 and 179:
tropical lineages have simply had m
Page 180 and 181:
allow us to reliably estimate the s
Page 182 and 183:
widespread climate change and Phyto
Page 184 and 185:
fauna as “a host of intriguing qu
Page 186 and 187:
Phytomyza cecidonomia Hering, 1937
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Phytomyza conii Hering, 1931 [PA] (
Page 190 and 191:
Phytomyza orientalis Spencer, 1962
Page 192 and 193:
Phytomyza nemopanthi Griffiths & Pi
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Phytomyza thalictricola Hendel, 192
Page 196 and 197:
Phytomyza nigritella Zetterstedt, 1
Page 198 and 199:
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
Page 206 and 207:
of the Compositae. Biol. Skr. 55: 3
Page 208 and 209:
Grimaldi, D. A. 1999. The co-radiat
Page 210 and 211:
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
Page 222 and 223:
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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