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fubfe 2. Kinds ot cvicknce used lo document the exisience of a marking pheramone 1. Behavtoral assays of response lo NtP. 0. Quaniirication of rejection patterns of females olTered 'marked' hosts. b. Dt'^tinguishing elTects of a mark' from elTecLs of other stimuli associated with an oviposition event. I. QuamiHcation of rejection patterns of females offered hosts treated with extracts of marked hcsLs. eggs, or fecal matenal. II. QuantiHcaiion of rejection paiiems of ferrules offered >urrogate hosts on %bhich females have Jeposiied a putative mark, c Char:u:tenzaiK>n of the chemical nature of the mark I. Structural identihcation of the ;u;tivecompourHl(s) m conjunction with behavioral avsays that establish activity II. Synthesis of the active compound(s) m conjurwtion N^iih behavioral as.says that establish the activity of the synthesized compound(s) 2. Devrnption of a putative hosi^marking behavior « Determmation of mechanisms for MP production and detection. X Identmcation of tis.sue involved in prodiKing and storing the MP or MP precursor^. I Quaniihcaiiun of rejection patterns of females olTercd ho^ts or surrogate hosts treated with extracts ot different tivsues b Identihcation of mode of MP reception. I Ablation of candidate semory %tructures and quantincation of changes in rejection patterrts 4 .\sscssment of ecological consequences ot MP a%e. J- Ouantihcatum ot clutch dislnbution patterns in the tield cffect of infestation on female behavior is chemicallymediated. However, neither line of evidence provides ironclad evidence for a MP. For example, a difference m either case could be due to a chemical change in the host it.self that accompanies infestation tSee section on "Cues versus signals of brood presence" above). In lieu of direct evidence that the insect itself produces the chemical compounds involved, behavioral assays of egg-free surrogate hosts bearing 3 putative mark are sometimes used. Where investigators are able to control the surrogate host stimuli precisely, the effect of a marked egg-free surrogate on host acceptance can be rea.sonably strongly attributed to the insect, and probably something deposited by it. Prokopy et al. (1982b) utilized such marked surrogate hosts to test for the presence of a MP in the fruit fly Anasirepha fraierculus. These re.searcheis created marked surrogate hosts by allowing females to deposit a clutch into an artilicial agar sphere and (hen. following egg deposition, transferring the females onto egg-free surrogate models where they were allowed to deposit the putative MP. Females e.xpcsed to the eggfree marked and umarked models rejected the marked models significantly more, supporting the hypothesis that A. faterrulus produces and deposits a MP which deters funher host reuse (Prokopy et al.. 1982b). The case for the presence of a MP is made more convincing if the insect e.xpresses a 'marking behavior'. such as the brushing or dragging of ovipositors on the host, and more convincing still if the putative 25 marking behavior results in the physical deposition of some substance on or in the host. In Rhagoletis Ries. for example, females drag their ovipositor on the fruit surface and this ovipositor-dragging results in the obvious deposition of a clear substance on the fruit surface (.\verill & Prokopy. 1988). However, caution IS advised as apparent marking behaviors may not necessarily be associated with the deposition of a MP (Cirio. 1971; Prokopy & Koyama, 1982; Fitt. 1984). Support for the hypothesis thai females are utilizing a MP sometimes takes the form of tield censuses that show that clutches are not distributed randomly among available hosts, but rather uniformly among them (Bauer. 1986; Thiery et al.. 1995). However, a uniform distribution of clutches might also result if females arc able to detect changes in hosts that occur when eggs are deposit. .Mcchanisms of host-marking pheramone communication Effects of MPs on female behavior The presence of a MP on or in a host may affect female behavior in multiple ways, both deterring and enhancing cviposition (Corbet. 1973a; Prokopy. 198 la; Paine et al.. I99T). The most often reported effect of MPs on behavior, and the one on which this review focuses, is a reduction in the number of eggs allocated to pre
viously marked and uiilized hosis. The rediicluin in allocation can iKcur m a numher ot W3y>i Frei|iienily. a MP will decrease ihe lendeiicy tor a temalc visiting a marked host to lay esgs .in that host. Where ovipositiiin IS not entirely suppressed by MP. females may still lay smaller clutches (Ikavva & Okabe. I9S5; Papaj et al.. IWO). The extent to which clutch si/e is reduced in a previously utilized host has been shown in gregarious parasitoids to be a function of the number of eggs that were deposited previously in that host iBakkeret al.. 1472; van Dijken l murlwinc pticriMiiniic on the oviptwiiion hohjMor ol ihc V!cUiicrrjntf.in iniii i!v, > iifjiuitu iMn? ie*i (t'f t.ltfl;iiK) sition behavior, namely a tendency for a rty in recent contact with MP to terminate some stationary behavior such a.s resting, grooming or oviposition. and to initiate UKomotion. be it walking or Hying. In the folklore of fruit fly research, contact with MP is said to 'irritate' females. .-X generalized "interruptor'. or irritant. etTect may be a neurally economical way in which a MP can mediate a suite of functional effects on oviposition. Wliv pnultices the murk ami where i.s it plated'.' In entomophagous Hymenoptera. females may use either, or both, internal or external marks to indicate which hosts have been previously exploited (Salt. 1937; revs, van Lenteren. 1976. 1981; Hofsvang. 1990). .According to Bosque & Rabinovich (1979). whether MPs are deposited on the inside or the outside of the host depends on the life stage attacked. Egg parasitoid.s tend to mark hosts externally while parasitoids utilizing other host stages tend to mark hosts internally. Bosque & Rabinovich (1979) contend that this pattern is functional; while an external mark may be more readily detected by females inspecting a potential host than an internal one. shedding of a host larva's cuticle during molting would remove the external mark. Bosque & Rabinovich also argue (hat placement of the mark relates to sen.sory contexts for host examination. Larval and pupal stages are generally le.ss accessible than eggs to antennal examination. Larvae may be Ies.s accessible both because they can
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 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 38 and 39: 286 dropping on ihe MP communicatio
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
Page 122 and 123:
spatial patterning of clutches depo
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16-fl oz (473-ml) plastic Solo cups
Page 126 and 127:
percentage of hatchlings that emerg
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previously created oviposition punc
Page 130 and 131:
RESULTS Experiment 1. Effect of off
Page 132 and 133:
deposited) again improved the model
Page 134 and 135:
median pupal weight in the lone two
Page 136 and 137:
survival, using two-clutch and thre
Page 138 and 139:
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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