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wax threads from 6 rows of dorsal wax pores. About 6 weeks after colonization the aphids began to reproduce parthogenetically. Despite two catastrophes (malfunctioning incubator), we succeeded in rearing the aphids for 19 successive generations on the roots, by transferring l newborn offspring to new cages at 3-day intervals. • B._Next came similar success with Aploneura lentisci Pass. Galls of this species were collected in November 1996 on its host plant Pistacia lentiscus and alates were held in jars. The crawlers ofA. lentisci are light lemonyellow, smaller and more elongated than all species of Fordinae examined so far. (This species is mentioned by Bodenheimer and Swirski (1957) as facultatively holocyclic; it belongs in the same tribe, Baizongiini, as the former species). They started feeding immediately after transfer to the cages, and soon produced dense colonies along the roots. They produce copious amounts of wax threads and honeydew droplets, which become coated with wax and look like small pearls. The reproductive rate of this species seems extremely high, the _ dense wax threads making it difficult to census this species in the cages. At the time of writing, we have the colonies going for 6 months and they seem to be thriving. C.---Attempts to rear the crawlers of Geoica spp. were far less successful. We were particularly interested in rearing the winter generations of Geoica on roots in Figure 1._A photograph of a root cage. The white order to sort c,ut the taxonomic problem of the "G. spots on the roots are accumulations of wax threads utricularia" species complex (Brown and Blackman covering the aphids Baizongia pistaciae. 1994), because we seem to have at least two species (galling P palaestina and P atlantica, respectively) Aphids and Handling which were lumped together in previous work (Wertheim 1954 and earlier references there, Koach and Wool 1977, . GallS of several species were collected in the autumn of Wool 1977). Moreover, we expected to succeed with 1995 and 1996, At this time of year the galls contain these species because anholocyclic populations of alates (fall migrants) ready to disperse. The galls were Geoica were described from North America and Northopened in the laboratory and the aphids were held in jars em Europe (Mordvilko 1928). Our efforts began in • . linedwith filter paper. The alates readily gave rise to their offspring (crawlers) as they would have done in _ 1995, employing the methods of Campbell and Hutchinson (1995) and Royer (1991), but failed. In nature on the ground. In a cool and humid environment 1996, we began again, from galls collected in September , the crawlers may survive several weeks without food and October on the two Pistacia hosts. Each gall was •(Wertheim'1954). The jars were held at 20°C and emptied into a separate jar and the alates readily repr_- - samples of crawlers were periodically transferred to root duced. The crawlers were distinctly different in color: cages. The.cages were held at 20°C and 13 L: 11 D light those from P. palaestina galls were orange, those from P cycle or in an ambient temperature . . and continuous light, atlantica greenish. When introduced to the root cages after a first molt in the holding jars, the crawlers did not RESULTS commence feeding. They remained small and most of them died. In a late attempt to induce feeding, we A.---The first successful colonization of the roots was transferred two holding jars to a 4°C refrigerator for 2 with crawlers of Baizongia pistaciae L., collected on P weeks. The crawlers from these jars did start feeding palaestina in November 1995. The dark green crawlers when introduced to the cages, and some matured and , molted once in the jars and commenced feeding immedi- reproduced. Keeping the root cages in a sheltered ately upon transfer to the roots. After another molt their outdoor porch during the winter (January - March 1997) color changed to light yellow. They tended to settle in and taking extreme care in handling them, we managed groups along the roots. Feeding aphids produced long to get F2 and some F3, which eventually died. 128
Although Geoica belongs in the Baizongiini, the root Until now, aphids ofA. lentisci and B. pistaciae reproforms (from both hosts) do not secrete wax threads, duced continuously on the roots and showed no evidence Neither do they aggregate on the roots but tend to scatter of becoming alate (producing sexuparae). In free-living and feed individually. The mature forms are pale, aphids and in Pemphigus, short day length, low tempera- although those from P palaestina (given the specific : tures and crowding are known to induce sexuparae A name G. wertheimae by Brown and Blackman 1994) formation (Judge 1968, Moran et al. 1993a, b). In 1, tend to a pale orange tint. The reproductive rate of this Pemphigus, sexuparae appear in the fall, while in the form seems to be very low: they reproduce their young Fordinae they fly in spring, so it is unclear if the same in batches of four or five, no more then three such factors are involved in sexuparae induction. Success batches in their lifetime. We hope to have better success with the use of the root cages should provide answers to when we try again next year. these questions. The crawlers of two species of Forda behaved contrary ACKNOWLEDGMENT to expectation. Galls of Fordaformicaria von Heyden and F. ri'ccobonii Stef. were collected in October and I benefitted from the experience of Dr. N.A. Moran, Dr. November 1996, and the alates produced crawlers in the T.A. Royer, and particularly Dr. C.D. Campbell with • hbldingjars. In the cages, the long-legged, greenish rearing Pemphigus aphids on roots. crawlers moved up the seedlings out of the cage through the notch and settled on the leaves. However, most of LITERATURE CITED them were lost in a day or two. The few survivors were found near the seed where they grew and matured to Bodenheimer, F.S.; Swirski, E. 1957. The Aphidoidea of produce F1 offspring, but no further reproduction was the Middle East. Israel: The Weizmann Science Press. obtained. Brown, P.A.; Blackman, R.L. 1994. Morphometric Ants variation in the Geoica ultricularia (Homoptera: • Aphididae) species group on Pistacia (Anacardiaceae), Subterranean stages of Fordinae species are often with descriptions of new species and a key to emigrant reported as associated with ant nests. Shaposhnikov alatae. Systematic Entomology. 19:119-132. (1985) interpreted certain morphological structures in Fordinae as adaptations for these mutualistic interactions. Burstein, M.; Wool, D. 1993. Gall aphids do not select In our laboratory and rearing rooms, "household" ants optimal sites (Smynthurodes betae, Pemphigidae). (Monomorium pharoni) are very abundant (to our Ecological Entomology. 18: 155-164. dismay!). The ants invaded some of the rearing cages with Baizongia pistaciae. When observed under the Burstein, M.; Wool, D.; Eshel, A. 1994. Sink strength and stereomicroscope, these wild ants---which are not known clone size of sympatric gall-forming aphids. European to tend aphids---were seen, in many instances, to perform Journal of Entomology. 91" 57-61. the exact kind of behavior that aphid-tending ants perform: they "tapped" the aphids with their antennae Campbell, C.D.; Hutchinson, W.D. 1995. Rearing until a honeydew droplet was excreted, and devoured, methods and demographic statistics for a subterranean "Milking" the aphids may not require specialized kinds morph of the sugarbeet root aphid (Homoptera, • of ant-aphid species associations. Aphididae). Canadian Entomologist. 127" 65-77. DISCUSSION Inbar, M.; Wool, D. 1995. Phloem-feeding specialis_s ' sharing a host tree: resource partitioning minimizls The root'cages, although still imperfect, seem suitable for interference competition among galling aphid species. - the study of the ecology of subterranean stages of the Oikos. 73:109-119. Fordinae; which was previously unknown. The main unresolved problems are both technical (e.g., how to get Inbar, M.; Eshel, A.; Wool, D. 1995. Interspecific " rid of or prevent thedevelopment of fungi in the cages, competition among phloem-feeding insects mediated which make observations difficult and may destroy the by induced hostplant sinks. Ecology. 76:1506-1515. eol0nies) and biological (i.e., why only two of the species were successfully reared for long periods; solving Judge, ED. 1968. Polymorphism in a subterranean aphid, the difficulties in rearing Geoica and Forda spp.). Pemphigus bursarius. I. Factors affecting the development of sexuparae. Annals of the Entomological The differences among species in adapting to the cages Society of America. 61" 819-827. . suggest that the method should be adjusted to the biology of each species. However, the biology is unknown, and only trial_and_error studies can break the vicious circle. 129
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LJSDA United States i Department Ag
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• , THE BIOLOGY OF GALL-INDUCING
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. TABLE OF CONTENTS - BIOGEOGRAPHY
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Genetic and environmental contribut
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.. GALL INDUCING AND OTHER GALL MID
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Table l. -,Gall midge species assoc
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C e. f. . Figure i'.--Galls of gall
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species and several inquilines may
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XYLOPHILOUS GALL MIDGE SPECIES from
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Rtibsaamen, E.H. 1899. Ueber die Le
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Table ° 1.--A summary of studies o
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• 3 ]5
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Skuhrava, • M.; .. Skuhravy, V. 1
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midge sPecies, several hosts yieldi
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Fedotoval Z,A. 1991. Gall midges (D
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known, nol_having been noted except
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Actilasioptera tumidifoHum GAGNE, N
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° _._. _. ..._..... _._:..--._...,
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Figures 13-16, Postabdomens ofActil
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-7"-% 25 i'.., 27 26 28 30 31 32 I
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Thorax. Wing length, 2.1-2.3 mm (n
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I' . McAlpine, J.F. 1981.2. Morphol
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Table 1.---Comparison between Tephr
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Table 3.raThe Afrotropical Myopitin
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........ _"i!_; _ 2,:_::;_ :L I,_ ,
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anthracina (Doane) (Diptera: Tephri
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o Host Plants Gall-forming insects
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, ENCLOSED 63% Figure 3.--A pie cha
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80% 70% 60% ° 50% - 40% 30% ' 20%
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Table 3. Major galling plant famili
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Appendix 1. Data Sheet for Galls Co
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j " GALLS Ephedra nebrodensis Jt ?W
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IS THERE A COOL-WET TO HOT-DRY GRAD
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10 - These findings suggest that ho
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ROLE OF PERICLISTUS (HYMENOPTERA: C
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° tl A o "!. . I ' Ir Figure l.---
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f . It Figure 2.--Schematic represe
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, ° , ° 3 . _ _ "' . . " Figures
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Table 2._Number of Periclistus larv
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. ° . 12 13 B Figures 12.---Schema
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emain as does the circumscribing la
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. Table 3.--Incidence of inhabitant
Page 85 and 86: the previous two galls, Aprostocetu
Page 87 and 88: Periclistus apparently have many of
Page 89 and 90: O'Brien, T.E; McCully, M.E. 1981. T
Page 91 and 92: indicator trait for male fitness, a
Page 93 and 94: Table 1.---Initial analyses to esta
Page 95 and 96: ., these cells, suggesting that fem
Page 97 and 98: o 8- 6- Krmnm 4- J _1) IP 2- '0- T
Page 99 and 100: Table 1. Mortality ofT. taxi expres
Page 101 and 102: kf) =_ i---. J i _ • i J _-" i i
Page 103 and 104: Table 2.---_-Correlationsfor number
Page 105 and 106: t Table 3. Mean values for total mo
Page 107 and 108: . 100 - ' / .r-! 0 i J0 - ,/O_OI 0
Page 109 and 110: (Figdre 5 continued) -. Tree 63 eve
Page 111 and 112: " Table 6.---Correlations (r) betwe
Page 113 and 114: Cameron,.R.A.D.; Redfem, M. 1978. P
Page 115 and 116: FurIhermore, Osten Sacken (1861 c)
Page 117 and 118: this letter, which was read at the
Page 119 and 120: 9 WOLBA CHIA-INDUCED THELYTOKY IN C
Page 121 and 122: Lancieux Ile Besnard • Erquy / Sa
Page 123 and 124: I Table 3.-mFrequencies offemales h
Page 125 and 126: Table 7.---z-Frequency offemales in
Page 127 and 128: . DISCUSSION considered as independ
Page 129 and 130: Fulton, B.B. 1933. Notes on Habrocy
Page 131 and 132: • . (26,galls), Nassau (30.27S, 1
Page 133 and 134: with competition (Stewart 1996) and
Page 135: " GALL-FORMING APHIDS ON PISTA CIA:
Page 139 and 140: THE LIFE-CYCLE OF THE BLACKCURRANT
Page 141 and 142: known" (1) Common form (E), which i
Page 143 and 144: Table 3._Average number of buds per
Page 145 and 146: Our prediction of a low cost of res
Page 147 and 148: - ? J Table 1.--Mean dry stem weigh
Page 149 and 150: stands; does the variation in resis
Page 151 and 152: Strong, D.R.; Larsson, S.; Gullberg
Page 153 and 154: The HR exhibited by the B. brevipes
Page 155 and 156: HR.is an important mechanism whereb
Page 157 and 158: q, J 10o • • O _80 • '= • O
Page 159 and 160: percent (n= 24) of the plants that
Page 161 and 162: REPROGRAMMING PLANT DEVELOPMENT: TW
Page 163 and 164: harvested and snap frozen. Protein
Page 165 and 166: Table 3.---Expected and observed fr
Page 167 and 168: whi.le it is not possible to discar
Page 169 and 170: . PLANT HORMONES AND GALL FORMATION
Page 171 and 172: As a follow up to Beck's (1947) fin
Page 173 and 174: Table 2.-_--Levels of extractable c
Page 175 and 176: .. J 0 0.16 m 0.14 - . 0.12 - 0.10
Page 177 and 178: probably occurs in the intact gall,
Page 179 and 180: Biddington, N.L.; Thomas, T.H. 1973
Page 181 and 182: .THE EFFECTS OF GALL FORMATION BY L
Page 183 and 184: three.larval instars (Mook 1967, Ru
Page 185 and 186: , + . 12 I I i i 4 .o • 1 10-- "
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, o Figure 5-1 to 5-6.----detailed
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Table 3.1Nutritional composition of
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gro.wing point, it is the latter th
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Byers, J.A+; Brewer, J.W.; Denna, D
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Skuhravy, V_1978. Invertebrates: de
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de.rivative spectra of pigment-prot
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Thylakoid Morphology CONCLUSION Ult
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.I THE DEVELOPMENT OF AN OVAL-SHAPE
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. LITERATURE CITED Rohfritsch, O. 1
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.PATTERNS IN GALL-INDUCING INSECTS
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fact, large Shrubs supported the hi
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We may conclude that the reduced ap
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Price, EW: 1997. Insect ecology. 3d
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of egg insertion. The galls of Euur
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species within each group. Close sp
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, According tO our present knowledg
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saying, that, according to A.K. Skv
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Benson, R.B. 1960a. Studies in Pont
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J Table 1. The Ranges of host plant
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(table 2 continued ) Host plant gen
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(table 4 continued) .. Host plant g
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Table 6.--List of host plants for P
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Table 9.--List of host plants for P
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(table o 9 continued) Host plant ge
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J America north of Mexico and addit
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Table 1.-,Host species and habitat
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this sampling and rearing of insect
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. Eastern USA Host Oaks White 0.8 g
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composed of Q. bicolor and Q. macro
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Q..rubraand Q. coccinea. Furthermor
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Fritz, R.S.; Nichols-Orians, C.M.;
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MOLECULAR PHYLOGENY OF NORTH AMERIC
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. RESULTS The heuristic search usin
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a few genera to consider evidence o
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. MOLECULAR PHYLOGENY OF THE GENUS
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. Table 1._Author names of the taxa
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28 26 24 22 A 20 18 0 q_ 1.4 O. 0 !
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•" _ _ _ o ,. _ _ -.--_.,. _ Ii .
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Among species groups, the topology
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. D. ignota D. nebulosa _!I D. fusi
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. LITERATURE CITED Dalla Torre, K.W
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PATTERNS IN THE EVOLUTION OF GALL S
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Figure 1. Gallstructures induced by
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the s.ection Quercus, and the 'blac
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t SG AG Species Clade " _ _ Name 9a
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non-Andricus cynipine species (fig.
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SG AG Species Clade _ t Name 100 94
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.an asexual-only ancestor. The sexu
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.. "j V 6 '_ _!i..-".'='-,.. " •
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Abe, Y. 1997. Well-developed gall t
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Stone, G.N.; Schrnrogge, K.; Crawle
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Quercus cerris, while a parthenogen
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Table t.----Summary statistics for
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Distance from root . 0.00 0.04 0.08
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Table 3.--Summary statistics for ge
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° . A. 40- -, 30- _ w _ A. kollari
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the alleles they possess at specifi
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e!atively narrow water barrier to l
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. TESTS OF HYPOTHESES REGARDING HYB
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classifieati0nerrors for various ba
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Tabl.e 2. Canonical discriminantfun
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encourage hybrid zone workers to ab
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Strauss, S,Y. 1994. Level of herbiv
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sense heritability of resistance to
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0.25 - Adelges abietis • 0.20 q)
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0 a 1 . W ! 309 A
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" I I_)! o Io Io Io I__1o I I I I_)
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classes for A. abietis suggests tha
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0 POSTER ABSTRACTS . W ! O ,l 315
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A . REFERENCES .. Huntley, B.H.; Bi
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CECIDOMYIID-INDUCED GALLS OF MA CHI
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. GALL MIDGE FAUNAS (DIPTERA: CECID
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GALL-INDUCER AND GALL-INQUILINE CYN
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" INDUCED RESISTANCE IN WILLOW AGAI
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.BROAD OVERVIEW OF A GALL-SYSTEM: F
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TWOSPECIES OF EURYTOMID GALL FORMER
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Csdka, Gyuri; Mattson, William J.;
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