Arthropods Associated with Above-Ground Portions of the Invasive Tree, Melaleuca Quinquenervia, in South Florida, USA Author(S): Sheryl L

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Arthropods Associated with Above-Ground Portions of the Invasive Tree, Melaleuca quinquenervia, in South Florida, USA Author(s): Sheryl L. Costello, Paul D. Pratt, Min B. Rayamajhi and Ted D. Center Source: The Florida Entomologist, Vol. 86, No. 3 (Sep., 2003), pp. 300-322 Published by: Florida Entomological Society Stable URL: http://www.jstor.org/stable/3496526 . Accessed: 10/09/2014 14:40

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This content downloaded from 158.135.136.72 on Wed, 10 Sep 2014 14:40:21 PM All use subject to JSTOR Terms and Conditions 300 Florida Entomologist 86(3) September 2003

ARTHROPODSASSOCIATED WITH ABOVE-GROUNDPORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA, IN SOUTH FLORIDA,USA

SHERYL L. COSTELLO, PAUL D. PRATT, MIN B. RAYAMAJHI AND TED D. CENTER USDA-ARS, Invasive Plant Research Laboratory, 3205 College Ave., Ft. Lauderdale, FL 33314

ABSTRACT

Melaleuca quinquenervia (Cav.) S. T. Blake, the broad-leaved paperbark tree, has invaded ca. 202,000 ha in Florida, including portions of the Everglades National Park. We performed prerelease surveys in south Florida to determine if native or accidentally introduced arthropods exploit this invasive plant species and assess the potential for higher trophic levels to interfere with the establishment and success of future biological control agents. Herein we quantify the abundance of arthropods present on the above-ground portions of saplings and small M. quinquenervia trees at four sites. Only eight of the 328 arthropods collected were observed feeding on M. quinquenervia. Among the arthropods collected in the plants adventive range, 19 species are agricultural or horticultural pests. The high percentage of rare species (72.0%), presumed to be transient or merely resting on the foliage, and the paucity of species observed feeding on the weed, suggests that future biological control agents will face little if any competition from pre-existing plant-feeding arthropods.

Key Words: Paperbark tree, arthropod abundance, Oxyops vitiosa, weed biological control

RESUMEN

Melaleuca quinquenervia (Cav.) S. T. Blake ha invadido ca. 202,000 ha en la Florida, inclu- yendo unas porciones del Parque Nacional de los Everglades. Nosotros realizamos sondeos preliminares en el sur de la Florida para determinar si los art6podos nativos o accidental- mente introducidos explotan esta especie de planta invasora y evaluar el potencial de los ni- veles tr6ficos superiores para interferir con el establecimento y 6xito de futuros agentes de control biol6gico. En cuatro sitios, nosotros cuantificamos la abundancia de art6podos presentes en las porciones sobre el terreno de los renuevos y pequenios arboles de M. quinquenervia. Solamente ocho de los 328 art6podos recolectados fueron observados alimentandose en la M. quinquenervia. Entre los art6podos colectados en las dreas no nativas de la planta, 19 especies son plagas agricolas 6 de hortalizas. El alto percentaje de especies raras (72.0%), presumidos de ser transeuintes o meramente descansando en el follaje, y la escasez de especies observadas alimentandose de la maleza, sujiere que los futuros agentes de control biol6gico enfrentaran poca o ninguna competencia de los art6podos herbivoras ya presentes en la planta.

Melaleuca quinquenervia (Cav.) S.T. Blake, the Melaleuca infested areas can be restored broad-leaved paperbark tree, was introduced into through removal of existing trees, followed by south Florida during the late 1800s (Thayer & Bo- measures to preempt reinvasion and subsequent dle 1990). Although threatened in its native range recruitment. Conventional control tactics com- along the east coast of Australia and a few nearby bine mechanical and chemical means to eliminate South Pacific islands, life history characteristics seedlings, saplings, entire stands of mature trees, of M. quinquenervia (melaleuca) combine with fa- or isolated plants in sensitive areas (Stocker & vorable ecological characteristics of Everglades Sanders 1981; Bodle et al. 1994). However, biolog- habitats to make this tree an explosive weed in ical attributes of this weed necessitate repeated south Florida (Meskimen 1962; Myers 1983; Bal- mechanical and chemical treatments, which im- ciunas & Center 1991; Hofstetter 1991). Cur- pose an accumulation of negative impacts on non- rently, melaleuca occurs on about 202,000 ha of target organisms, including endangered plants. Florida wetlands (Bodle et al. 1994) and has his- These adverse impacts limit the frequent use of torically spread at a rate of about 2,850 ha/yr such methods. In contrast, classical weed biologi- (Center et al. 2000). The negative impacts of cal control has been described as the most ecolog- melaleuca on native flora and public health prob- ically benign tactic for controlling exotic pests lems have been documented (Di Stefano & Fisher (McEvoy & Coombs 1999) and has been consid- 1983; Myers 1983; Molnar et al. 1991; Bodle et al. ered a desirable addition to conventional methods 1994). Diamond et al. (1991), for instance, deter- (Browder & Schroeder 1981; Bodle et al. 1994). mined that if unchecked, potential losses to the Development of a weed biological control pro- Florida economy as a result of this invasive tree gram typically proceeds in a stepwise fashion, could reach $169 million annually. including: selection of a natural enemy, risk

This content downloaded from 158.135.136.72 on Wed, 10 Sep 2014 14:40:21 PM All use subject to JSTOR Terms and Conditions Costello et al.: Arthropods of Melaleuca quinquenervia 301 analysis, release, monitoring establishment, and To suppress melaleuca growth, land managers finally assessing the effectiveness and ecological mowed trees at ca. 6-month intervals, resulting in impact of the introduced biological control agent coppices 0.5-2 m in height. These coppicing (Harris 1975; McEvoy & Coombs 1999). An often clumps formed a dense, nearly continuous canopy recommendedinitial phase in a classical weed bio- of leaves with 4,406 clumps/ha. In contrast to the logical control program includes surveys of herbi- previous sites, the soil type was primarily sand, vores associated with the invasive weed in the new consistent with an invaded pine flatwoods habitat (adventive) geographic range (Harris 1975; Olck- type (Anonymous 1990). Other than melaleuca, ers & Hulley 1995). Such surveys are intended to the subdominant vegetation included Ludwigia identify herbivores already exploiting the weed sp., Centella asiatica (L.) Urb., Rhynchospora and to ascertain whether niche competition could globularis (Chapm.) Small, Rhynchosporaeximia influence agent establishment and impact (Harris (Nees) Boeck., and Rhynchosporafilifolia Gray. 1971). Although surveys for natural enemies were Site 4 consisted of a 1 ha area within histori- performedin Australia during 1987 to 1991 (Balci- cally mesic flatwoods in the Picayune Forest, unas et al. 1995), surveys of arthropodsassociated Collier Co., FL (N26.10478 and W81.63392) with melaleuca in its adventive range had never (Anonymous 1990). A fire burned much of the been done. Failure to perform such surveys could melaleuca dominated areas during June 1998, re- increase costs due to wasted effort associated with sulting in recruitment of 129,393 trees/ha com- selecting, screening and releasing herbivores that posed of primarily small 1-2 m tall saplings, may already be present, having accompaniedthe interspersed with an occasional large, mature invasive weed upon introduction or thereafter. tree. Pinus elliottii Engelm. and a parasitic (dod- Therefore,specific objectives of this study were: 1) der-like) species growing on the melaleuca were assess the current abundance of arthropods asso- the only other common vegetation. ciated with melaleuca in south Florida, 2) deter- Surveys were conducted monthly at each site mine if native herbivores are exploiting the from November 2000 through June 2001. Sites invasive plant, 3) determine if co-evolvednatural were surveyed between 10 a.m. and 2 p.m. on enemies from the native range inadvertently ac- days without precipitation. To survey arthropods companied melaleuca into south Florida, and 4) associated with melaleuca canopies, we swept fo- inventory those higher trophic levels associated liage, and occasionally trunks, with a 90-cm-di- with the plant that could potentially interfere with ameter sweep net. One sample consisted of 100 the establishment or impact of introducedbiologi- sweeps in a 1800 sweeping motion spaced ca. 1.0 m cal control agents. apart along a randomly selected 100 m transect. Four samples along separate transects were col- MATERIALS AND METHODS lected each month. The contents of the net after 100 sweeps were emptied into a 3.78 liter sealable Arthropodsurveys were performedat four loca- plastic bag and frozen at -19 (?1) ?C until pro- tions in south Florida. Site 1 was located near cessed. Arthropods were then separated from Ft. Lauderdale, Broward Co., FL (N26.05606 and plant material, sorted by morphological types, W80.25168).The site was a 0.5 ha field consisting and stored in 70%ethanol. of 2 to 5 m tall trees occurringat a plant density of One limitation of our sweep sampling method ca. 21,560 trees/ha. In general, melaleuca trees included collecting arthropods that were not were growing in high organic soils typical of re- closely associated with melaleuca, but were tran- claimed 'glades' systems. Although melaleuca was sients, merely resting on the plant foliage or dis- the dominant species, other plants commonly oc- turbed from understory vegetation while curring in the site included Blechnum serrulatum sampling.Additionally, this method was biased to- Rich.,Ampelopsis arborea (L.) Koehne, Vitis aesti- wards those species that are poor fliers or slow to valis Miclhx.,and Ludwigiaperuviana (L.) H. Hara. disperse from a disturbance and, unlike previous Site 2 was located under a power line right-of- Australian surveys, endophages were not in- way near Weston, Broward Co., FL (N26.035483 cluded. Therefore, caution should be used when and W80.43495).Prior to 1997 land managers cut drawing inferences from these data due to the un- melaleucatrees near their bases, resulting in multi- known relationships between some of these ar- stemmed branches re-growing from the stumps. thropods and melaleuca. For this reason, a The survey area was ca. 0.5 ha and trees were 2-5 m minimum of two observers searched for direct her- tall, occurringat a density of 2,517 trees/ha.The site bivory on the above ground portions of melaleuca was swale-like with commonvegetation other than trees for 30 min./month at each site. Arthropods melaleuca including:Sagittaria lancifolia L., Cla- observed feeding on melaleuca are reported inde- dium jamaicensis Crantz, and Andropogonglomer- pendently from those collected in sweep samples. atus (Walt.)B.S.P. (Anonymous 1990). For each species collected, species abundance Site 3 was located near Estero, Collier Co., FL per site was calculated for the entire survey pe- (N26.4255 and W81.81033) and consisted of an 8 riod by first averaging the number of specimens ha area of drained wetland converted to pasture. from the four monthly samples and then averag-

This content downloaded from 158.135.136.72 on Wed, 10 Sep 2014 14:40:21 PM All use subject to JSTOR Terms and Conditions 302 Florida Entomologist 86(3) September 2003 ing among all sample dates. Average species sufficient densities to cause appreciable damage abundance among all sites was determined by to- to trees in south Florida. For instance, of the 18 tal specimens collected throughout the entire sur- orders, 117 families, and 328 species collected in vey (rare = 1-5 specimens, occasional = 6-10 this study, only 54 species were classified as com- specimens, common = >10 specimens). Occasion- mon and 33 species were classified as occasional ally, arthropods were collected by hand to facili- (Tables 1 and 2). Of the most commonly occurring tate identification. Where possible, arthropods species, 33 (66.7%) were predators or detritivores were identified to species. Identifications that (Table 2), and 11 (20.4%) were herbivores (Table could not be confirmed are indicated by "poss." 1). Both adult and immature stages of H. coagu- (possibly) before the scientific name. Some lata, the glassy-winged sharpshooter, were ob- Diptera were not sent for identification because served on melaleuca, suggesting that melaleuca specialists were not available or specimens were may serve as an alternative host for this insect. damaged and lacked key identifying features. However, during the sampling period none of Such specimens were combined into an "unidenti- these arthropods were directly observed feeding fied spp." group and the number of morphological on melaleuca. Furthermore, out of 409 herbivo- types is denoted in parentheses. All morphologi- rous arthropods found attacking melaleuca in cal types, except for immatures that could be as- Australia, none were found on melaleuca in south sociated with their adult forms, were included in Florida indicating that no co-evolved natural ene- the total species count. mies accompanied melaleuca into south Florida All specimens, except formicids, were submit- upon introduction or thereafter (Balciunas et al. ted to and deposited at the Florida State Collec- 1995). The most intuitive explanation for these tion of Arthropods (FSCA, Division of Plant findings is probably due to the fact that all known Industry (DPI), Gainesville, FL) for identification importations of the invasive tree were in the form and incorporated into their taxonomic database. of seed (F. A. Dray, pers. comm.). Most formicids were identified and retained by In contrast, we have observed several arthro- L. Davis at the Fire Ant Unit, Agricultural Re- pod species feeding on melaleuca that were never search Service, USDA, Gainesville, FL. A few recovered in the sweep samples. Both early and formicids were identified by M. Deyrup at the late instars of the polyphagous saddleback cater- Archbold Biological Station, Lake Placid, FL. pillar, Sibile stimulea (Clem.), were observed feed- Several dipteran specimens were identified at the ing on mature melaleuca leaves at Site 3. Larvae Systematic Entomology Laboratory, Agricultural of the caterpillar were concentrated on a single Research Service, USDA, Beltsville, MD. sapling, defoliated much of the tree, and were only present during late winter. After inspection of a RESULTS AND DISCUSSION single damaged sapling (5 cm diam), larvae of the generalist cerambycid Neoclytus cordifer (Klug) Surveys of herbivores associated with an inva- were also collected, allowed to pupate and suc- sive plant in its adventive range are often recom- cessfully emerged as adults (2 males and 1 fe- mended as a prelude to a weed biological control male). Two phytophagous mites, Oligonychus project (Harris 1975). Historically, scientists have coffeae (Nietner) and Brevipalpus obovatus Don- ignored this recommendation, possibly due to the nadieu, were observed feeding and developing supposition that native herbivores are already large (>100 individuals), although isolated popu- suppressing the weed to the greatest level pos- lations. Populations of these generalist mites oc- sible. In contrast, native arthropods can cause curred on mature leaves and were only observed considerable damage to non-indigenous weeds once. The Florida red scale, Chrysomphalus aoni- (Newman et al. 1998). The native weevil, Euhry- dum L., the stellate scale, Vinsonis stellifera, and chiopsis lecontei Deitz, for instance, shows promise an unidentified Coccus sp. often co-occurred on for control of Eurasian watermilfoil, Myriophyllum mature Melaleuca leaves. Although the scale oc- spiacata (L.) (Newman & Beisoer 2000). In addi- curred in surprisingly high densities (>10 per tion to natives, co-evolved herbivores and diseases leaf), no apparent foliar damage was visible. Two may also be accidentally introduced from the polyphagous aphids, Aphis gossypii Glover and plant's native range. The biological control agents Toxoptera aurantii (Boyer de Fonscolombe), were Megastigmus aculeatus (chalcid wasp) and Phyllo- observed feeding on stems of developing branches. coptes fructiphilus (an eriophyoid mite), for exam- Infestations of both polyphagous aphids were ple, were collected in West Virginia during surveys slight (<50 individuals per plant). Although these of arthropods associated with the exotic weed arthropod species were observed feeding on mela- Rosa multifloria (Thunb.). The eriophyoid mite, leuca, no damage was visible. These observational and the virus it transmits, is considered the most findings suggest that, unlike some invasive plants effective agent for the suppression of R. multiflora that can be stressed by native arthropods in the (Amrine 1996). adventive range, the arthropod community cur- In its adventive range, however, it appears that rently associated with melaleuca provides little if melaleuca has not acquired native herbivores at any suppressive effect on the exotic tree. The pau-

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This content downloaded from 158.135.136.72 on Wed, 10 Sep 2014 14:40:21 PM All use subject to JSTOR Terms and Conditions Costello et al.: Arthropods of Melaleuca quinquenervia 319 city of herbivores indicates that direct competi- sient, and considering the dearth of commonly tion between natives and introduced biological collected arthropods,caution should be exercised control agents will be minimal. when making conclusions concerning the func- Habitats dominated by invasive plants are of- tional well being of melaleuca invaded ecosystems. ten assumed to be sterile environments with few The role of invasive species as facilitators of wildlife species utilizing the ecosystem (Bodle et other invasive species has received little attention al. 1994). However, Mazzotti et al. (1981) deter- in the literature (Simberloff & Von Holle 1999). mined that differences exist among invasive One example of this interaction may include the plants in their ability to support native fauna, in- ability of nonindigenousplants to modifythe habi- dicating that habitats invaded and dominated by tat in a way that favors exotics over natives. In this non-indigenous plants are not necessarily biologi- study, 20 exotic species were collectedin the mela- cal deserts. After eight months of surveying ar- leuca habitat (Tables 1 and 2). Among the exotic thropods in melaleuca dominated ecosystems, species, Solenopsis invicta Buren, the red imported rarefaction curves of both herbivorous and non- fire ant, was common (Table 2) and is included as herbivorous arthropods suggests that continued one of the most ecologically destructive invasive surveying efforts would result in the collection of species in the southeastern U.S. These ant colonies additional species (Figs. 1 and 2; Magurran 1988). not only cause human disturbance, but also are The variety of arthropods,both collected (Tables 1 known to cause 70%mortality of freshwater turtle and 2) and predicted (Figs. 1 and 2), reported hatchlings (Pseudemys nelsoni Carr), can nega- herein indicates that melaleuca dominated habi- tively impact the endangered Schaus swallowtail tats do support an arthropod community. How- (Papilio aristodemus porceanus), and can dramati- ever, this does not necessarily imply that cally change arthropodcommunities (Porter et al. melaleuca is a superior habitat for such fauna as 1988;Allen et al. 2001; Forys et al. 2001). Although indicated by the paucity of basal trophic levels native to Florida, the glassy-winged sharpshooter (i.e., herbivores). Without the ability to compare is an invasive species in California,where it vec- arthropod diversity in surrounding native habi- tors Xylella fastidiosa Wells et al., the causal agent tats, the probability that many species are tran- of Pierce's disease in vineyards. Because the

140

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240.

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Fig. 2. Rarefaction curve for cumulative non-herbivorous species collected from M. quinquenervia (Nov.-June). glassy-winged sharpshooter is commonly associ- histories involving insect introductions for weed ated with melaleuca in Florida, it may be predicted control (Goeden & Louda 1976). Parasitoids and that the plant also provides a refuge for the inva- pathogens, for instance, caused 24% larval mor- sive sharpshooter in California. In this manner, tality of the introduced moth, Samea multiplica- melaleuca may serve as a reservoir for these and lis Guenee (Semple & Forno 1987). Herein, we other invasive species in Florida and beyond. collected several generalist predators that may In addition to the facilitation of ecological im- potentially impact current and future biological pacts by exotic species, invasive weeds may also control agents, including Euthyrhynchus florida- harbor agricultural pests. For instance, 1/3 of the nus (Pointer) (Pentatomidae), Podisus mucronatus phytophagous insects associated with Salsola Uhler (Pentatomidae), Podisus saggita (Fabri- kali L. var. tenuifolia Tausch (Russian thistle) cius) (Pentatomidae), Stiretrus anchorago (Fabri- and 1/2 of the insect species on Carduus pycno- cius) (Pentatomidae), and Zelus longipes (L.) cephalus L. (Italian thistle) proved to be pests of (Reduviidae), as well as various ant and spider agricultural importance (Goeden & Ricker 1968). species. Predation on populations of the recently In our study, 18 arthropods collected from mela- released biological control agent Boreioglycaspis leuca canopies are major or minor economic pests melaleucae Moore (melaleuca psyllid, Psyllidae) of agricultural crops. Three species, Aphis spirae- by various pentatomid and coccinellid species has cola Patch (Aphididae), T aurantii (Aphididae), been observed in the field and may be negatively and S. invicta (Formicidae), were commonly asso- affected by generalist predators. During host ciated with melaleuca. Both aphid species are cos- specificity testing and under mass rearing condi- mopolitan, phytophagous pests of Citrus spp. and tions prior to its introduction, B. melaleucae was many other plants. An infestation of these aphid attacked by multiple arachnid species. However, species can result in abortion of Citrus flower the level of predation observed in the field or un- buds and both aphids produce honeydew, thus fa- der laboratory conditions does not appear to im- voring the development of sooty molds. pact colonies in a significant way (P D. Pratt, Native predators, parasitoids, and pathogens pers. obs.; S. A. Wineriter pers. comm.). Studies have interfered with half of the published case on other psyllids, Psylla pyricola Forester (pear

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psyllid) and Diaphorina citri Kuwayama (Asian ACKNOWLEDGMENTS citrus psyllid), have shown that their populations are reduced by generalists predators such as: The authors thank the following specialists and as- sociates for identification of most arthropod species: Chrysopa sp. (Chrysopidae), Anthocoris sp. J. Bramblia, G. B. Edwards, G. A. Evans, S. E. Halbert, (Anthicoridae), and Olla v-nigrum (Mulsant) J. B. Heppner,J. M. Kingsolver,B. F. Mauffray,C. C. Por- (Coccinellidae) (Watson & Wilde 1963; Michaud ter, L. A. Stange, G. J. Steck, M. C. Thomas, W. C. Wel- 2001). Furthermore, Watson & Wilde (1963) and bourn, J. R. Wiley of the FSCA, Division of Plant Santas (1987) demonstrated a reduction in psyl- Industry, Gainesville, FL. We also thank formicid spe- lid populations by generalist predators. Never- cialists L. R. Davis of the Fire Ant Unit, AgriculturalRe- theless, in each study psyllid populations were search Service, USDA, Gainesville, FL, and M. A. suppressed by generalist predators at different Deyrup of the ArchboldBiological Station, Lake Placid, FL.; diptera specialists F. C. Thompsonand N. E. Wood- levels, suggesting that predicting the acquisition ley of the Systematic Entomology Laboratory,Agricul- and impact of these predators on introduced bio- tural Research Service, USDA, Beltsville, Maryland; logical control agents is tenuous. Collembola specialist R. J. Snider at Michigan State During our study, we also collected several par- University; and Psocopoteraspecialist E. L. Mockfordat asitic hymenopteranspecies associated with mela- Illinois State University. leuca in south Florida (Table 2). Hymenopteran species in Australia parasitized ca. 40% of galls REFERENCES CITED formed by the potential biological control agent Fergusonia spp. (gall fly) (Davies et al. 2001). ALLEN,C. R., E. A. FORYS,K. G. RICE,AND D. P. WOJCIK. Davies et al. (2001) suggested the impact by Fer- 2001. Effects of fire ants (Hymenoptera:Formicidae) on hatching turtles and prevalenceof fire ants on sea gusonia spp. as biological control agents of mela- turtle nesting beaches in Florida. Florida Entomol. leuca will likely be reduced due to parasitism from 84(2): 250-253. local hymenopteran species in Florida. However, AMRINE,J. W. 1996. Phyllocoptes fructiphilus and bio- predicting which parasitoids may exploit this or logical control of multiflora rose, pp. 741-749. In other proposed biological control agents is diffi- E. E. Lindquist, M. W. Sabelis and J. Bruin [eds.], cult. Initial steps may include a taxonomic com- Eriophyoid Mites: Their Biology, Natural Enemies parison among the co-evolved parasitoids in the and Control.Elsevier Science, 790 pp. agent's native and adventive ranges. For instance, ANONYMOuS.1990. Guide to the natural communities of Cirrospilus sp. (Eulophidae), Eupelmus sp. (Eu- Florida.Florida Natural Areas Inventory.Florida De- were partment of Natural Resources,Tallahassee, 118 pp. pelmidae) and Eurytoma sp. (Eurytomidae) BALCIUNAS,J. K., AND T. D. CENTER.1991. Biological collected in Australia associated with Fergusonia control of Melaleuca quinquenervia: prospects and spp. and during our survey we also collected para- conficts, pp. 1-22. In T. D. Center, R. F. Doren, R. L. sitoids belonging to these genera in south Florida Hofstetter, R. L. Myers and L. D. Whiteaker [eds] (Goolsbyet al. 2001). Unfortunately,species deter- Proc. Symp. Exotic Pest Plants, Miami, Florida, 2-4 mination was not possible for those reported Nov. 1988, U.S. Dept. Interior, National Park Ser- herein. Due to the diversity of both genera, geo- vice, Denver, CO, 387 pp. graphic separation over evolutionary time, and BALCIUNAS,J. K., M. F. BURROWS,AND M. F. PURCELL. lack of Fergusoninidaein the New World,it is un- 1995. Australian insects for the biological control of the paperbark tree, Melaleuca quinquenervia, a seri- likely that the species occurringin Australia and ous pest of Florida, USA, wetlands, pp. 247-267. In Florida are the same. Other genera found during E. S. Delfosse, and R. R. Scott [eds.], Proc. Symp. on our survey do not correspond to those genera Biological Controlof Weeds. 2-7 February 1992, Lin- known to parasitize current and candidate biolog- coln University, Canterbury,New Zealand, 735 pp. ical control agents in their native range, including BODLE,J. J., A. P. FERRITER,AND D. D. THAYER.1994. Fergusonina spp., B. melaleucae, Poliopaschia li- The biology, distribution, and ecological conse- thochlora (Lower) (tube-dwelling moth) and Lo- quences of Melaleuca quinquenervia in the Ever- phyrotoma zonalis (Rohwer) (melaleuca sawfly) glades, pp. 341-355. In S. M. Davis and J. C. Ogden (Jensen 1957; Riek 1962; Burrows & Balciunas [eds.], Everglades: The Ecosystem and Its Restora- 1997; Davies et al. 2001; J. A. Goolsby,USDA/ARS, tion. St. Lucie Press, Delray Beach, 826 pp. BROWDER,J. A., ANDP. B. SCHROEDER.1981. Melaleuca Aust. Bio. Cont. Lab., pers. comm.). Predictions seed dispersal and perspectives on control, pp. 17-21. based solely on this survey may grossly underesti- In R. K. Geiger [ed.], Proceedings of a Melaleuca mate parasitoid acquisition as additional species Symposium. Florida Dept. of Agriculture and Con- may be recruited to the system after introduction sumer Services, Division of Forestry, Tallahassee, of the biologicalcontrol agent. In the future a more Florida, 140 pp. accurate assessment may be obtained by survey- BURROWS,D. W., AND J. K. BALCIUNAS.1997. 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