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Tooker and Hanks Influence Plant Community Structure Natural Enemies Pine Needle Scale (Homoptera Diaspididae) Urban Landscapes.pdf

Tooker and Hanks Influence Plant Community Structure Natural Enemies Pine Needle Scale (Homoptera Diaspididae) Urban Landscapes.pdf

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Influence of Plant Community Structure on Natural Enemies of Pine Needle Scale (Homoptera: Diaspididae) inUrban Landscapes
Department of Entomology, University of Illinois at Urbana-Champaign, Urbana, IL 61801
Environ. Entomol. 29(6): 1305Ð1311 (2000)
Pine needle scale,
Chionaspis pinifoliae
(Fitch), is a pest of many species of conifersin urban habitats and Christmas tree farms. We found that the scale was abundant in impoverishedhabitats, such as ornamental landscapes, and scarce in more natural, park-like habitats. Rates of parasitism were highest in impoverished habitats, suggesting that parasitoids were not effective insuppressing scale populations. Generalist predators, however, were more diverse and abundant innatural habitats and appear to be more effective in controlling scales in structurally complex plantcommunities. Total densities of arthropods and densities of plant-feeding species were greatest inimpoverishedhabitats,suggestingthatpopulationswerepoorlyregulated.Outbreaksofpineneedlescale in ornamental landscapes and Christmas tree farms may be discouraged by increasing plantstructural and species diversity to favor natural enemies.
Aphytis, Chionaspis heterophyllae,
habitat effects, natural enemies,conservation biological control
and managed ecosystems, speciesdiversity of arthropod natural enemies is often posi-tively correlated with the diversity of plant species(e.g., Risch 1981, Yue et al. 1994, Dean and Milton1995).Plantssustainpopulationsofnaturalenemiesbyprovidingprey,ßoralresourcesforadults,andsuitablemicroclimates (Powell 1986, van Emden 1990). Thus,diversifying the composition of plant communitiescould encourage natural enemies to suppress popula-tionsofphytophagousinsects.Ornamentallandscapesare amenable to this pest management tactic becausethey are relatively stable ecosystems and are not con-strained by the simple structure and periodic disrup-tioncharacteristicofagronomicsystems(Rauppetal.1992). Diverse and structurally complex plant com-munities in ornamental landscapes support a higherdiversity and abundance of natural enemies, resultingin better regulation of phytophagous insects (e.g.,Hanks and Denno 1993, Shrewsbury 1996).We examined the inßuence of plant communitystructure of urban landscapes on the abundance anddiversity of natural enemies, and, indirectly, on pop-ulationdensitiesof pineneedlescale,
Chionaspis pini- foliae
(Fitch). Pine needle scale is native to NorthAmerica(BurdenandHart1989),butisaseriouspestof many species of introduced evergreens, includingspecies in the Pinaceae (
 Abies, Picea, Pinus, Pseudot-suga,
), Cupressaceae (
), and Tax-aceae (
Taxus, Torreya;
Shour and Schuder 1987). Thescaleisoneofthemostimportantpestsofornamentalpines in the United States, and has been labeled the
white malady
because heavily infested trees appearwhitewashed (Johnson and Lyon 1988).Damaging infestations of pine needle scale are usu-allylimitedtomanagedecosystems,suchasnurseries,tree farms, and ornamental landscapes (Johnson andLyon 1988) that are often characterized by low levelsof plant species diversity and structural complexity(Raupp et al. 1992). The scale is rare in more naturalhabitats, such as forests and wood lots (Ruggles 1931)where complexity and plant diversity are higher (seeRisch 1981, Szentkiralyi and Kozar 1991, Yue et al.1994,DeanandMilton1995).Naturalenemiesofpineneedle scale include coccinellid beetles (Cumming1953, Nielsen and Johnson 1973, Luck and Dahlsten1974, Eliason and McCullough 1997) and at least ninespeciesofaphelinidparasitoids(Krombeinetal.1979;Burden and Hart 1989, 1993), but the scale is alsoprobably attacked by more generalist predators of armoredscaleinsects,includingharvestmen,crickets,earwigs, and neuropteran larvae (Clausen 1940, 1978;Crumb et al. 1941; Cloudsley-Thompson 1958; Ebling1978; Hanks and Denno 1993).In this study, we examined the inßuence of plantcommunitystructurewithinurbanlandscapesonratesofpredationandparasitismofpineneedlescale,scalepopulation density, and the abundance and speciesdiversity of the arthropod community as a whole. Inanother portion of this study, we excluded preda-tors from the study trees to better evaluate the im-pact of parasitoids on scale populations. Finally, wetested the ability of natural enemies to disperse shortdistances, which is of interest because of the smallspatialscaleofornamentallandscapeswherethescaleis a pest.
2000 Entomological Society of America 
Materials and Methods
Chionaspis pinifoliae
has long been confused with
Chionaspis heterophyllae
Cooley, the
pine scale,
be-causeofsimilarappearanceandnaturalhistory(Shour1986, Kosztarab 1996). Both species overwinter in theegg stage (Kosztarab 1996) and crawlers of both spe-ciesemergesynchronouslyinspringandsummer.Thetwo species also share parasitoid species (Burden andHart1993).Differencesinphenologyandmorphologyare too subtle to distinguish
C. pinifoliae
C. het-erophyllae
in the Þeld (Shour 1986). The two speciesalso overlap in geographic distribution,
C. pinifoliae
occurring over most of North America while
C. het-erophyllae
is conÞned to the eastern and midwesternstates (Shour and Schuder 1987, Kosztarab 1996). Forthepurposesofthisreport,weacknowledgethepres-ence of the two scale species in the area of our study,butrefertothespeciescomplexas
(see Johnson and Lyon 1988).
Influence of Plant Community Structure on Pop-ulation Density of Pine Needle Scale.
We estimatedpopulation densities on pine trees in three types of urbanhabitats:(1)
ornamen-tal landscape plantings with pines in proximity topavedroadsorparkinglots,andsurroundedbygravelormulch(
ornamentallandscapes with pines surrounded by turf (
24sites); and (3)
wooded habitats,
park-like andwooded primarily with pine species (
24 sites).StudysiteswereinChampaign-Urbana,IL,andateachsite there were at least three of the common hosts of pine needle scale (Shour and Schuder 1987):
Turra (0.25Ð2 m tall),
Pinus sylvestris
L. (1.5Ð4m), or
Pinus nigra
Arnold (2.5Ð5 m).There are apparently no standard protocols for es-timating population densities of scale insects on co-nifers (see Kozar 1990, Jactel et al. 1996). Our initialattempts to estimate density of pine needle scale byrandom sampling proved inaccurate at low densitiesbecause of the patchy distribution of scales withintrees. Instead, we estimated population densities byexaminingneedlesdistributedthroughoutthecanopyand counting female scales for a standard 3-min pe-riod. We counted only scales of the current genera-tion, as evidenced by an intact and pure white cover.This method was reproducible in quantifying densityeven of very lightly infested trees. Similar visual tech-niques have been used to estimate population densi-tiesofavarietyofarthropodspecies(e.g.,WilsonandSimberloff 1969, McCullough and Sadof 1998, Seboltand Landis 1999). We estimated population densitiesof pine needle scale in June 1997 during the Þrstgeneration.
Weinfestedsmallpine trees with pine needle scale to evaluate the im-pact of natural enemies on scale populations. Ourpotted pines were 25Ð40 cm tall Scotch pines (
variety French Blue) in plastic pots (15Ð19cmdiameter)withasterilizedpottingmixtureofequalproportionsofsoil,peat,andperlite.Pineswerepottedin mid-April 1997 and held in a greenhouse underlong-day photoperiod, were watered as needed, andfertilizedat
2-wkintervals(15:30:15;ScottsMiracle-Gro Products, Port Washington, NY). We infestedtrees with pine needle scale 1 mo after potting bylaying scale-infested cuttings on them during crawleremergence; cuttings were taken from infested Scotchpine tree (Hoot Owl Christmas Tree Farm, Urbana,IL).Aftercrawlershadsettled,wedisposedofcuttingsto prevent parasitoids from emerging in the green-house.
Influence of Plant Community Structure on RatesofPredationandParasitism.
Toidentifytheparasitoidspecies that attacked pine needle scale in our studyarea, we collected scale-infested cuttings and rearedparasitoids in lidded cardboard buckets (17 cm diam-eter,18cmhigh)towhichwasattachedaglassvialtrap(15 mm o.d.). Parasitoids were stored in 70% ethanoland 10 specimens of each of three morphospecieswere submitted for identiÞcation to the USDA Sys-tematic Entomology Laboratory in Beltsville, MD. Afourthspecies,discoveredlater,wasidentedbyJ.M.Heraty at the University of California, Riverside.We caged predaceous arthropods with scale-in-fested needles of Scotch pine to conÞrm that theywould eat pine needle scale: only species that weremost likely to be predators were tested, includingharvestmen (Opiliones), snowy tree crickets (
sp.),larvalgreen lacewings (Chrysopidae), and twicestabbedlady beetles [
Chilocorus stigma
(Say); see Clausen1940, 1978; Crumb et al. 1941; Cloudsley-Thompson1958; Ebling 1978; Hanks and Denno 1993]. Predatorswere caged individually (
3 per species) in plasticcontainers with moistened cotton rolls (to providewater) and pine needles having similar densities of female scales (
10 scales) in the second or thirdinstar. After 24 h, we inspected scales for evidence of predation,suchasdamagetoscalecoverorremovalof the scale body.To determine whether rates of predation and par-asitismwereinßuencedbyhabitat,wepositionedpot-ted Scotch pines infested with scale next to or at thebase of study trees in impoverished turf and woodedhabitats (
10, 10, 15, respectively) on 16Ð25 June1997. Scales on both potted and resident trees wereeithersecondorthirdinstars,thelifestagesvulnerableto parasitization (unpublished data). Potted pineswere subsequently watered as needed.After 5 wk, we returned the potted trees to thelaboratory where they were held for
2 wk to allowparasitoids to develop. To estimate the density of scalesperunitofneedlelength,wearbitrarilyselected20needlesfromeachseedling,countedthenumberof adult female scales on each needle, and measuredneedle length. To estimate percent parasitism andpredation, we arbitrarily selected 50 adult femalescales per tree and examined them under a dissectingmicroscope. In many cases the scale insects them-selves were missing, but the condition of their waxcovers provided evidence of activity by any naturalenemies (characterized in part during studies of pre-dation, described above). A circular exit hole was1306 E
Vol. 29, no. 6
evidence of parasitism; predators with chewingmouthparts tore ragged holes in the cover, whereasthose with sucking mouthparts left only the shriveledbodies of their prey. We excluded adult females thathad been killed by fungi, as well as those from theprevious generation, which were identiÞed by havingeither discolored tests or discarded chorions undertheir tests.
Impact of Parasitoids on Scale Populations Exclu-sive of Predators.
Predators of pine needle scale ap-parently consumed parasitized scales and even para-sitoidpupae,andthereforecouldinßuencetheroleof parasitisminregulationofscalepopulationdensity.Toexamine the impact of parasitoids in scale populationdensity, we excluded predators from potted pines inwooded habitats where potential natural enemies of the scale were most abundant (see below), and mea-suredparasitismrate.Toexcludepredators,weplaceda cylinder of aluminum window screen (1 mm mesh,60 cm tall) around scale-infested potted trees, closedatthetopwithstaplesandsecuredtothepotwithducttape.Preliminaryexperimentsdemonstratedthatwin-dow screen excluded most, if not all, predators whilepermitting parasitoids access to scales. Female scalesonpottedtreeswereinthesecondorthirdinstar.Wepositioned these potted pines next to nine Scotchpinesineachofthreewoodedareas.After5wk,pottedpines were returned to the laboratory to estimatepercent parasitism (as described above).
To evaluate dispersal abilities of natural ene-mies over small spatial distances, we positioned threescale-infestedpottedpines(femalescalesinsecondorthird instar) in each of three impoverished sites so asto contact the canopy of resident trees. In the samesites, we also positioned three scale-infested pines30 cm from the canopy edge (bare ground or gravelseparatedpottedpinesfromresidenttrees).Thestudywas set up on 13 August 1997 and potted pines weresubsequently watered as needed. After 5 wk, pineswere returned to the laboratory and held for 2 wk toallow parasitoids to develop, and then scales wereexaminedtoestimateratesofpredationandparasitism(as described above).
Influence of Plant Community Structure on Abun-dance and Diversity of Arthropods.
To examine theinßuence of habitat on arthropod communities asso-ciated with host plants of pine needle scale, we tookbeatingsamplesofall73studytreeson19Juneand27August 1997. Samples were taken at mid-canopy fromfour branches per tree, one at each cardinal point.Eachbranchwasbeatenfourtimeswitha925-grubbermallet and falling arthropods were captured in anenamel pan partly Þlled with 70% ethanol. Branchsamples from each tree were combined into a singlesample.We categorized all collected arthropods into mor-phospecies and counted the specimens of each. Wefurther categorized these morphospecies into guilds:phytophagous insects, predators, parasitoids, and po-tential predators of pine needle scale (our method of sampling was not effective in capturing aphelinidparasitoids).Weestimatedspeciesdiversityofarthro-pods with the Shannon-Wiener index (
Peet 1974,Hayek and Buzas 1997).
We used analysis of variance (ANOVA;SAS Institute 1988) to test the effect of habitat ondensity of pine needle scale per tree, percent parasit-ism, percent predation, arthropod density, and theeffect of proximity to resident pines on percent par-asitism and predation. We determined whether thedata met the assumptions of ANOVA by conÞrminghomogeneity of variances between treatments (
-test)andnormalitywithintreatments(ShapiroÐWilkstest; Sokal and Rohlf 1995). Data not suitable forANOVA were analyzed by the nonparametricKruskal-Wallis test (Sokal and Rohlf 1995). Differ-ences between individual means were tested by theleastsigncantdifference(LSD)test(Ott1993).Wecompared ShannonÐWiener indices with the Student
-test (Magurran 1988). Data are presented as means
1 SE unless stated otherwise.
ResultsInfluence of Plant Community Structure on Pop-ulation Density of Pine Needle Scale.
Density of pineneedle scale varied signiÞcantly across habitats andwas highest in impoverished habitats (mean of 60scales/tree/3-min search), intermediate in turf habi-tats (17 scales), and lowest in wooded habitats (twoscales; ANOVA
11.7; df 
2, 62;
0.0001). EachmeanwassigniÞcantlydifferentfromtheothers(LSDtest
Influence of Plant Community Structure on Ratesof Predation and Parasitism.
Of four parasitoid mor-phospeciesrearedfromscale-infestedclippings,threewere aphelinids:
sp. poss.
Encarsia aurantii
Howard, and
Coccobius varicornis
(Howard). The fourth parasitoid species was in thegenus
(Hymenoptera: Eulophidae),mostspeciesofwhicharebelievedtobeparasitoidsof leafminers (Schauff et al. 1997). The relatively largebody size of the
sp. (
2Ð3 mm long)also suggests it is probably not a parasitoid of pineneedle scale and that it emerged from other hosts onour pine clippings.Snowy tree crickets, Þeld crickets, and green lace-winglarvaeallreadilyfedonfemalepineneedlescalesin cages. Twicestabbed ladybird beetles and harvest-men did not eat scales on individual needles, but didso on scale-infested Scotch pine trees in an indepen-dent laboratory experiment. Thus, we considered allof these species potential predators of pine needlescale.On scale-infested potted pines placed in the Þeld,parasitism rate was highest in impoverished habitats(wherethescalewasmostabundant),intermediateinturfhabitats,andverylowinwoodedhabitats(wherethe scale was rare; means signcantly different;KruskalÐWallis statistic
16.7, df 
0.0002;Table1).ParasitismratesacrossallthreehabitatsweresigniÞcantlyandpositivelycorrelatedwithhostabun-dance on resident trees (Fig. 1), at least at the spatialDecember 2000 T
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