Materials and Methods
has long been confused with
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
in the Þeld (Shour 1986). The two speciesalso overlap in geographic distribution,
occurring over most of North America while
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)
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),
L. (1.5Ð4m), or
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,wasidentiÞedbyJ.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 [
(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