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286 dropping on ihe MP communication system of its competitor, a system that evolved in ihe coniext of informing self or conspecirics. If this is the cisc. it remains unclear if such eavesdropping constitutes an evolved trait. While it is possible that C. rhodesUmus evolved a sensitivity lo its congener's MP. it is al.so possible that C. macultiius lost a pre-existing sensitivity to its congener's MP (a sensitivity derived perhaps from shared ancestry). In short, it appears thai a respon.se by one species, or loss of response, to another species' MP can readily evolve under selection, particularly when ipecies differ in competitive ability. However, whether a MP itself can evolve strictly to signal the presence of a heterospecitic competitor is uncertain. Given the ubiquitousnessof MPs conveying information about self or conspecitics. MPs mediating di.scnmmation between species probably rarely evolve independently of MPs mediating discrimination within a species. Rather, the same MP may evolve simultaneously to function at both intraspecific and interspecific levels. Phylogenclic perspectives Eci>li>);ical correlates of host-markin g The Rhagnietis data on cross-recogniiion reviewed above suggest that there is something to be gained from a phylogenetic approach to host-marking evolution. The Rliaftoleiis data are intriguing; yet it is noteworthy that data were not collected with phylogenetic analysis in mind. .Not all of the comparisons or even the most informative comparisons have been made, especially in light of revisions of the phylogeny of Ihe genus. In this group and others, there is a need for an organized phylogenetic approach to the evolution of host-marking behavior. In particular, phylogenetic analyses offer a means of evaluating the roles of various ecological factors in the origin and maintenance of host-marking behavior. For example. It is a truism that MPs are found mainly in insects that develop as juveniles on resources that are limited in quantity, such as seeds, fruit, plants or other insects (Roitberg & Prokopy. 1987). Phylogenetic analysis would be useful in quantifying the evolutionary gains and losses in host-marking behavior in relation to resource limitation, as well as in ruling out other, correlated ecological factors. In this regard, a phylogenetic survey of patterns in host-marking in the tephritid fly genus Rhagoletis 35 represents a beginning. Within the North American clade of Rhaifoletts. the pattern of host-marking in one species group, ihe suavis group, differs strikingly from the pattern in other groups (C. Nutio. D. Papaj. and H. Alonso-Pimentel. unpubl. data). Whereas host-marking behavior is present among all members of at least three other species groups as well as one unplaced species, host-marking behavior within the Kiiavis group is spottily distributed. One species in (hat group. Rhagolelis juglanclis. marks vigorously, but at least (hree species wi(hin (hat group mark inconsistently or not at all. Variation among species groups in host-marking may reflect variation in larval ecology. Whereas native fruit for most species m other groups are so small that just one or a few larvae can survive to pupation in a single fruit, the walnut host fruit used by all members of the suavis group frequently yield many pupae. It IS conceivable that some other ecological factor relating (o life on walnu(s. other (han reduced resource limitation, accounts for the variable pattern m host-marking within (he suavis group. Nevertheless, the pa((ern is in(riguing and deserving of further study. Another ecological correlate lo consider that may influence the evolution of marking behavior relates to the detectability of cues to brood presence. One might expect the occurrence of host-marking behavior lo be inversely correlated wi(h Ihe conspicuousness of non-MP cues (o brood presence such as oviposition wounds. In Anasrreplw flies, for example, it has been proposed that the host-marking behavior is correlated with Ihe degree of latex relea.sed by host fruit during oviposition which is in turn correlated with deposition of oviposition within the pulp vs. seed of a fruit (Aluja et al.. 1999). In A. saggira. whose larvae feed on seeds of Fnuieria fruit, females oviposit deeply into the fruit, causing a great deal of late.x to be released, which could serve as a cue of brood presence. In a closelyrelated species. .4. serpentina, which feeds on the pulp of the same fruit species, females oviposit less deeply inio more-mature fruit, resulting in the release of relatively little latex. The difference in degree of laie.x release is associated with a difference in host-marking behavior whereas .4. serpentina marks the Pouteria host. .4. saggila does not. Other ecological correlates worth considering in a comparative coniext include the ephemerality of (he host, an insect's host breadth, the paichiness of the host in space, mobility of the juvenile stages, and cannibalistic tendencies m the young (Roitberg & Prokopy. 1987; Diaz-Reischer et al.. 1999).
Role !t Vehrencanip. IWS. Szamado. I9'W|. In terms ol sheer volume ol el fort focused on t.he mechanism, function and taxonomic distribution, work on MP coinmunicalion would seem to have much to contribute to the held of animal communication. Many of the key systems studied in .\IP communication (fruit tlies. bean weevils, panisitoidsi would seem to be ideal for tests ol theory addressing a range ol issues in signal evolution, including signal error, sensory bias, and deception. Finally, our understanding of both the mechanism .ind evolution ol hi>st-inarking communication would benefit greatly trom knowledge of MP chemistry in both parasitoids and phytophagous in.sects. To date, few MPs have been identihed. Lack of knowledge of MP chemistry makes it difficult to construct and evaluate hyp«)theses of sensory bias or to evaluate the basis of phyUigenetic patterns m cross-recognition and even to distinguish between instances in which insects utilize cues associated w ith brood presence and instances in which insects utilize an actively-produced signal. •Acknow ledgcmenls We acknowledge financial suppt^rt from the Interdisciplinary Degree Program in In.sect Science, a .National Science Foundation .Minority Cr;iduate Research Fellowship. and NRICCP Award No. .'.'>.^I);-IW77. We thank Judie Bronstein. .Michael Greenfield. Frank Me.ssina. and Dena Smith for comments on previous drafts. Rererenccs Alphen. J. J- M v;«i H. W. NcU. 19X2. SiipcrparaMii^m and hir>t discnminaliifn hy A sttixirtt tahuhi i .Mvvi- 36
Page 1: INFORMATION TO USERS This manuscrip
Page 4 and 5: UMI Number: 3053915 Copyright 2002
Page 6 and 7: STATEMENT BY AUTHOR This dissertati
Page 8 and 9: DEDICATION This work is dedicated t
Page 10 and 11: ABSTRACT Choosing where offspring w
Page 12 and 13: An explanation of tiie problem CHAP
Page 14 and 15: euse occurs in the field, the impac
Page 16 and 17: Appendix D, "Aggregative behavior i
Page 18 and 19: offspring performance. Counter to t
Page 20 and 21: Lalonde RG, Mangel M, 1994. Seasona
Page 22 and 23: APPENDIX A HOST MARKING BEHAVIOR IN
Page 24 and 25: kluwer the language of science Dr.
Page 26 and 27: 174 belter categorized as cues than
Page 28 and 29: fubfe 2. Kinds ot cvicknce used lo
Page 30 and 31: 278 defend themselves and because i
Page 32 and 33: 280 lion of a signal. Costs associa
Page 34 and 35: 282 than were conspccifics with low
Page 36 and 37: 284 thai (he marker was a mutant fo
Page 40 and 41: 28S inac>. 3 larval partsiloiil of
Page 42 and 43: 290 Hol'svang. T. 1988. Mechanisms
Page 44 and 45: 29Z Roitbcfg. B- D it R- G- Laionde
Page 46 and 47: APPENDIX B HOST UTILIZATION BY THE
Page 48 and 49: Dispersal by Larvae of the Stem Bor
Page 50 and 51: POPULATION ECOUXTY Host Utilization
Page 52 and 53: October 2000 NUFIO ET AL.; HOST UTI
Page 54 and 55: October 2000 Nufio et au: Hoct Unuz
Page 56 and 57: October 2000 NUFIO ET AL4 HOST UIHI
Page 58 and 59: Reuse of larval hosts by the walnut
Page 60 and 61: KEYWORDS Life history" marking pher
Page 62 and 63: Alternatively, a weak correlation b
Page 64 and 65: previously attacked hosts early in
Page 66 and 67: walnut fly oviposition punctures. T
Page 68 and 69: Effects of offspring weight on adul
Page 70 and 71: she deposited over a lifetime and t
Page 72 and 73: RESULTS Levels of host reuse in the
Page 74 and 75: Survival from egg to egg hatch Of t
Page 76 and 77: significantly correlated with pupal
Page 78 and 79: DISCUSSION While the femaie-prefere
Page 80 and 81: discriminate between hosts that hav
Page 82 and 83: will consume fewer resources and th
Page 84 and 85: females gain not by simply reusing
Page 86 and 87: of her offspring but also of the nu
Page 88 and 89:
ACKNOWLEDGEMENTS We thank Henar Alo
Page 90 and 91:
Bjorkman C, Larsson S, Bommarco R,
Page 92 and 93:
Fujiyama N, Katakura H, 2001. Varia
Page 94 and 95:
Mayhew PJ, 1997. Adaptive patterns
Page 96 and 97:
Robertson IC, Roitberg BD, 1998. Du
Page 98 and 99:
Wiklund C, 1981. Generalist vs. spe
Page 100 and 101:
Table 2. The influence of various f
Page 102 and 103:
Table 4. Analysis of variance compa
Page 104 and 105:
nCURE CAPTIONS - Continued Figure 5
Page 106 and 107:
Figure 1. Fruit Cohorts (Number of
Page 108 and 109:
Figure 3. A. B. A > on •m e « 50
Page 110 and 111:
Figure 5. B. es & 1 1 0.8 • 0.6 -
Page 112 and 113:
Figure 7. B. > 09 "S s e s o •Ml
Page 114 and 115:
Figure 9. 400 9t W s 1 t 300 « Of)
Page 116 and 117:
Aggregative behavior is not explain
Page 118 and 119:
that the larval aggregations that f
Page 120 and 121:
2001). The shifting benefits and co
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spatial patterning of clutches depo
Page 124 and 125:
16-fl oz (473-ml) plastic Solo cups
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percentage of hatchlings that emerg
Page 128 and 129:
previously created oviposition punc
Page 130 and 131:
RESULTS Experiment 1. Effect of off
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deposited) again improved the model
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median pupal weight in the lone two
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survival, using two-clutch and thre
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DISCUSSION Absence of an AUee efTec
Page 140 and 141:
simultaneously. It would seem that
Page 142 and 143:
Spatial distribution of clutches In
Page 144 and 145:
If reuse of larval hosts negatively
Page 146 and 147:
Page 148 and 149:
Hausmann SM, Miller JR, 1989. Ovipo
Page 150 and 151:
Papaj DR, Alonso-Pimentel H, 1997.
Page 152 and 153:
Page 154 and 155:
Page 156 and 157:
FIGURE CAPTIONS continued Figure 4.
Page 158 and 159:
Figure 2 MD E DC n Om 3 Qi S V DA E
Page 160 and 161:
Figure 4. B. c. -g 60 S a. a 40 - S
Page 162 and 163:
Host marking beiiavior as a quantit
Page 164 and 165:
encountered either control fruit th
Page 166 and 167:
unparasitized and parasitized hosts
Page 168 and 169:
Finally, it is conceivable that the
Page 170 and 171:
Experiment 1. Relationship between
Page 172 and 173:
or 30 minutes). Fruit were then hun
Page 174 and 175:
After two replicates of the four tr
Page 176 and 177:
Experiment 3. Female responses to h
Page 178 and 179:
RESULTS Experiment 1. Effects of cl
Page 180 and 181:
Experiment 3. Female responses to f
Page 182 and 183:
example, the possibility that ovipo
Page 184 and 185:
field cage assays. What remains to
Page 186 and 187:
Page 188 and 189:
Bauer G, 1986. Life-history strateg
Page 190 and 191:
Landolt PJ, Averill AL, 1999. Fruit
Page 192 and 193:
Quiring DT, McNeil JN, 1984. Intras
Page 194 and 195:
FIGURE CAPTIONS Figure 1. The relat
Page 196 and 197:
Figure 2. w OA 8 k 98 s o a. QH s 8
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