19 Recent Adventive Scale Insects (Hemiptera: Coccoidea) and Whiteflies (Hemiptera: Aleyrodidae) in Florida and the Caribbean Region

Ian Stocks Division of Plant Industry, Florida Department of Agriculture & Consumer Services, 1911 SW 34th Street, PO Box 147100, Gainesville, Florida 32614-7100, USA

19.1 Introduction close association with their host plant; thus, they are likely to be introduced already attached The flora and fauna of Florida and the Caribbean to a suitable host. The ecological and economic region are in continual flux. Underlying this flux impact of adventive scales and whiteflies is is the dynamic and complex international trade of enhanced primarily by the fact that the natural plant material involving the USA, the countries of complement of predators and parasitoids is the Caribbean region, South America, Europe and often absent in the new environment, but also Asia. Vast quantities of fruits and vegetables, and by the fact that, given an expanded diversity of propagative and non-propagative plants, are dis- novel potential host plant species, the insects tributed through ports in the Caribbean region become far more polyphagous than they are in and Florida, and inspection of this material their native environment. results in a constant struggle against the threat Complicating still further attempts to con- of adventive species. Among the most prominent trol the flow of adventive scales and whiteflies is of potentially and actually adventive animal the fact that, in many cases, we have very little ­species are hemipterous insects in the family information about the species beyond the name, Aleyro­didae (the whiteflies) and numerous known host(s) and collection locale (which may ­families in the superfamily Coccoidea, such as or not be the original provenance). Also, it is not Diaspididae (armored scales), Pseudococcidae uncommon that undescribed species are discov- (mealybugs) and soft scales (Coccidae). Several ered because they are adventive and have become, factors conspire to make insects in these groups at least initially, prominent members of the some of the most noxious of plant pests. First, plant–pest insect community. they are typically quite small, and are therefore This chapter will focus on recent invasions more likely to pass unnoticed through inspection of scales and whiteflies as a result of international sites. Second, many species are parthenogenic. trade among Florida and the countries and terri- Therefore, it is possible that a new infestation tories that comprise the Caribbean region. The could be started from the introduction on plant data presented in table format are derived prima- material of a single juvenile female, or even a sin- rily from the extensive database maintained by gle egg mass. Third, all of these insects are plant the Florida Department of Agriculture, Division parasites, and are therefore almost always in of Plant Industry (FDACS-DPI), and ScaleNet, an

342 © CAB International 2013. Potential Invasive Pests of Agricultural Crops (ed. J. Peña) Recent Adventive Scale Insects in Florida and the Caribbean Region 343

online database that endeavours to maintain an and surveying that takes place within Florida. exhaustive compendium of literature pertaining Thus, Florida overall has a better-known fauna to scale insects.i ScaleNet offers numerous than the islands in the Caribbean region. The taxa ­database search options that enable the user to considered in this review are presented in Table 19.1. ‘data-mine’ under various parameters, such as To place these recent developments in per- distribution, host and literature citation infor­ spective, of ~7400 described species of Coccoidea mation. Other information presented is derived (Miller et al., 2005), there are c. 1300 species of from the primary literature, especially Miller Coccoidea in the Neotropical (NT) region, and et al. (2005), which is an excellent synopsis of the c. 320 species of this total are recorded from adventive Coccoidea fauna of the USA. at least one territory in the Caribbean region The length of time under consideration, and (75 species of Pseudococcidae; 132 species of therefore the list of relevant taxa, is somewhat Diaspididae; 55 species of Coccidae; 54 species ­arbitrary, since the number of adventive taxa in in assorted families). By comparison, ScaleNet Florida and the Caribbean region has been ­growing retrieves ~1020 species of Coccoidea in the conti- inexorably since record-keeping began. Also, the nental USA, of which ~340 species are recorded list is biased toward a description of those taxa from Florida (67 species of Pseudococcidae; introduced into Florida. This is primarily due to the 165 ­species of Diaspididae, 45 species of Coccidae, fact that FDACS-DPI has a long history of detailed 61 species in assorted families). A comparison of record-keeping, but also to the intensive sampling the ­respective lists of species indicates that while

Table 19.1. Taxa discussed in this chapter. The year of first report in Florida is given, and the known ranges as reported by ScaleNet and Evans (2008).

First Recorded Family Scientific Name in Florida Range

Pseudococcidae Nipaecoccus viridis (Newstead) Pseudococcus dendrobiorum Williams 2009 AF, AU, OR, PA Palmicultor lumpurensis (Takahashi) 2009 AU, OR, PA Palmicultor palmarum (Ehrhorn) 2003 AU, NE, OR, PA Chaetococcus bambusae (Maskell) 1999 AU, NE, NT, OR, PA Paracoccus marginatus Williams 1998 AF, AU, NA, NT, OR, PA and Granara de Willink 1998 AU, AF, NE, NT Maconellicoccus hirsutus (Green) Planococcus minor (Maskell) 2002 AF, AU, OR, PA, NT, NA 2010 AF, AU, OR, NA, NT Kerriidae Paratachardina pseudolobata Kondo and Gullan 1999 NA Tarchardiella mexicana (Comstock) 1985 NA Coccidae Pulvinaria psidii Maskell 1985 AU, AF, NA, NT, OR, PA Protopulvinaria longivalvata Green NR, intercepted AF, AU, NT, OR Phalacrococcus howertoni Hodges & Hodgson 2008 NA, NT Diaspididae Poliaspis cycadis Comstock 2007 NA, OR, PA Unachionaspis tenuis (Maskell) 2007 OR, PA Aulacaspis yasumatsui Takagi 1996 AU, NA, NT, OR, PA Aulacaspis tubercularis Newstead 2002 AF, AU, OR, PA, NT, NA Duplachionaspis divergens (Green) 2000 AU, OR, PA, NA, NT Monophlebidae Crypticerya genistae (Hempel) 2006 NT, NA Aleyrodidae: Aleurodicinae Aleurodicus dugesii Cockerell 1996 NT, AU, OR Aleurodicus rugioperculatus Martin 2009 NT, NA Aleyrodidae: Aleyrodinae Siphoninus phillyreae (Haliday) 2010 NA, NT, AU, AF Dialeurodes schefflerae Hodges and Dooley 1986 NA, NT, AU, OR

OR, Oriental; PA, Palearctic; AU, Australasian; NT, Neotropical; AF, Afrotropical; NR, not recorded in Florida. 344 I. Stocks

there is substantial similarity, there are nonethe- the recently introduced and established species less significant differences. It is these differences A. dugesii and A. rugioperculatus; however, that are of concern, because they are the primary although many of the Caribbean records per- source of species that are available for transport tain to Trinidad, whose fauna has a distinctly between Florida and the Caribbean region. Given South American character (Peck et al., 2002), the diversity in numerous characteristics of the there are almost certainly more species of territories and countries that comprise the coun- Aleurodicus more widely distributed in the tries and territories in the Caribbean region, it is Caribbean region, but that are as yet officially not surprising that species distribution across unrecorded. the Caribbean region is uneven; some countries Several species will be excluded from con­ are relatively well sampled and support a large sideration in this chapter because they will be number of species, while others, for various rea- ­discussed elsewhere in this book: Planococcus sons, are less well sampled or support a lower minor (Maskell) (Pseudococcidae), Praelongorthe­ diversity. Even many common and polyphagous zia prae­longa (Douglas) (Ortheziidae) and species are unrecorded from many Caribbean Singhiella simplex (Singh) (Aleyrodidae). region countries. For instance, the seriously ­pestiferous mealybug Maconellicoccus hirsutus (Green) (Pink hibiscus mealybug), is officially 19.2 Mealybugs: Pseudococcidae recorded from only ten of the 27 territories in the Caribbean region; Coccus viridis (Green) (green Some of the most seriously noxious Coccoidea are scale) from 11 territories; Ceroplastes floridensis mealybugs, attacking not just plants grown and Comstock (Florida wax scale) from 13 territories; sold for decoration and aesthetic purposes, but Coccus hesperidum L. (brown soft scale) from 12 numerous fruit and vegetable crops. In Florida, a territories; Pinnaspis strachani (Cooley) (lesser large proportion of the known species are adven- snow scale) from 15 territories. tive. Two particularly pestiferous species have Compilation of detailed distribution and been introduced in recent years, the papaya species-richness data for Aleyrodidae is compli- mealybug (Paracoccus marginatus) and the pink cated by the lack of a whitefly database similar to hibiscus mealybug (Maconellicoccus hirsutus), but ScaleNet; however, Evans (2008) enumerated each appears to be under significant natural species richness for the biogeographic regions of ­control by parasitoid wasps and predatory lady the world, so rough numbers are available. The beetles. Given the very recent arrival in Florida distributional and host data given for the species of two additional species of mealybugs, the leb- discussed in this review are derived from that beck mealybug (Nipaecoccus viridis) and the pas- source. sionvine mealybug (Planococcus minor), it is too Worldwide, there are 1560 species of white- early to know what effect these two well-known fly in 161 genera (Martin and Mound, 2007). pest species will have on Florida agriculture and In the NT region, there are 15 genera with c. 120 horticulture. species in the subfamily Aleurodicinae, and 42 genera with c. 210 species in the subfamily Aleyrodinae. The FDACS-DPI database contains 72 species recorded from Florida, but, as with 19.2.1 Nipaecoccus viridis (Newstead) the scales, this number increases almost yearly. No data are readily available for the number of Common names: lebbeck mealybug, spherical whitefly species in the Caribbean region, but mealybug, karoo thorn mealybug. Distribution: as the number of monographic systematic Australasian (AU), Afrotropical (AF), Oriental works increases, and our knowledge of the dis- (OR), Palearctic (PA), Nearctic (NE) (USA: Florida). tribution of species improves (Martin, 2008), ScaleNet reports that it is present in Mexico and compiling those data will become easier. For the Bahamas, but these records are ­apparently instance, Martin (2008) lists 14 species from based on misidentifications (Meyerdirk et al., the Caribbean region in the mainly NT genus 1988; Dr G. Evans and Dr D. Miller, pers. comm.). Aleurodicus (Aleurodicinae), and there are three Five other species of Nipaecoccus occur from Florida, including A. dispersus Russell and throughout the Caribbean region: N. annonae Recent Adventive Scale Insects in Florida and the Caribbean Region 345

Willams and Granara de Willink (Guadeloupe and parasitoids of the mealybugs, the predatory flies Martinique), N. filamentosus (Cockerell) (Turks or perhaps both, as both mealybugs and flies are and Caicos, Puerto Rico, Haiti), N. neogaeus listed as known hosts of Pachyneuron wasps. If Willams and Granara de Willink (Trinidad) and they are parasitoids of the mealybugs, they would N. pitkini Willams and Granara de Willink join the other wasp fauna that might exert some (Trinidad and Tobago). Nipaecoccus floridensis measure of biological control, such as Anagyrus Beardsley, which is possibly native to Florida, kamali Moursi (Encyrtidae). After it was reared appears to be restricted to Florida. Only the and released for the control of pink hibiscus potentially pestiferous species N. nipae (Maskell) mealybug (PHM), A. kamali can now be found in is widespread in the Caribbean region, occurring on both Florida and the Caribbean region (Hoy et al., 17 islands. In general, specimens of Nipaecoccus 2006; Division of Plant Industry, 2004). Overall, require slide mounting for positive identifica­ some 65 species of Hymenoptera primary para­ t­ion, but there are some recognizable differences sitoids have been recorded from N. viridis (Noyes, between species in the colour and shape of the 2003), with A. indicus (Subba Rao) having an wax produced. exceptional record of control in Guam and Jordan Lebbeck mealybug was first recorded in (Meyerdirk et al., 1988). Florida in late 2009 from Palm Beach County attacking dodder (Cuscuta exaltata Engelm; Cuscutaceae) growing in a natural area, and its 19.2.2 Pseudococcus discovery prompted the release of a pest alert dendrobiorum Williams (Stocks and Hodges, 2010). A ground survey to determine the extent of the infestation was initi- Common name: dendrobium mealybug. Distri­ ated, which revealed that it had become wide- bution: AU, OR, PA, NE (USA: Florida, Hawai’i). spread in the natural area, feeding on a variety of This species was reported from Florida in host plants, and was possibly responsible for the 2009 from a population of Phalaenopsis orchids demise of the dodder (K. Griffiths and A. Derksen, maintained in a greenhouse on the University of pers. comm.); however, it was of only limited dis- Florida campus. Although it prompted the publi- tribution in the surrounding suburban areas, cation of a pest alert (Hodges and Buss, 2009), it where it was found to be feeding most frequently has not been collected since, and the population on Citrus sp. and Gardenia jasminoides Ellis may have been eradicated. Other than distribu- (Rubiaceae). As of February 2011, the host list tion information and host data, nothing has been had grown to 14 plant species, with Citrus spp., published on the biology, ecology or economic G. jasminoides and dodder seeming to be preferred impact of this species. Hosts are exclusively hosts (FDACS-DPI database). In citrus-growing Orchidaceae, including the genera Ascoglossum, areas of Jordan, where lebbeck mealybug had Cymbidium, Dendrobium, Phalaenopsis, Pholidota been a pest prior to the introduction of biological and Promatocalpum. Records from USDA-ARS controls, it caused such extensive damage that report 16 interceptions in the past 25 years, all orchards were burned in an effort to eradicate it. but one from orchid shipments, and all originat- Through their feeding, developing fruits are ing from Asia (15 interceptions) and Australia aborted and drop prematurely, and toxins in the (one interception). saliva kill the terminal branch tip near feeding sites, causing die-back (Meyerdirk et al., 1988). Almost contemporaneous with the discov- ery of the mealybug in Florida was the discovery 19.2.3 Palmicultor browni (Williams) of predators and parasitoids associated with N. viridis populations. Adult flies reared from No common name. Distribution: AU, NE (USA: masses of mealybug ovisacs were identified as Florida) Leucopis sp. (Chamaemyiidae), known predators Nothing was known about the biology or of mealybugs. Also reared from sequestered ecology of this host prior to its discovery in Florida infested plant material was the parasitoid wasp in 2002, and very little has been learned since. Pachyneuron eros Girault (Pteromalidae). In the literature, it is recorded from coconut, oil Curiously, it is not clear whether these wasps are palm (Elaeis guineensis Jacq.), Howea forsteriana 346 I. Stocks

(F. Muell. ex H. Wendl.) Becc. and Veitchia spp. little has been published about this species, so In Florida, it has a broad palm-host range, with presumably it is not of economic concern in its ­coconut (Cocos nucifera L.), Phoenix roebelenii native range, nor are there reports of its status in O’Brien and Adonidia merrillii (Becc.) Becc. Australia, where it has also been recorded. In (Arecaceae) accounting for 60% of records. The Florida, it has been collected 65 times, primarily majority of records (58%) are from Miami-Dade from Bambusa spp., but also Arundinaria sp., County, but it is also recorded from Broward, Dendrocalamus hamiltonii Nees and Arn. ex Collier, Monroe and Orange counties. Heavy Munro, Gigantochloa atroviolacea Widjaja, Guadua ­infestations of this species may cause dieback angustifolia Kunth and Phyllostachys nigra (Lodd. (Dr G. Hodges, pers. comm.) ex Lindl.) Munro. It appears to have been intro- duced into Florida via the international trade in exotic bamboo, in particular into the landscaping of an amusement park (Hodges, 2004a; Hodges 19.2.4 Palmicultor palmarum (Ehrhorn) and Hodges, 2004). It is now widely distributed across Florida, but does not appear to be having a Common name: Ehrhorn’s palm mealybug. large negative impact. Distribution: AU, PA, OR, NT (Bahamas, Bermuda, Guadeloupe, Jamaica, St Barthélémy, St Martin, St Croix), NE (Mexico, USA: Florida). 19.2.6 Chaetococcus bambusae (Maskell) This pest was described from Hawai’i from specimens feeding on palms. Ali (1987), citing Common name: giant bamboo scale. Distribution: Beardsley (1966), speculated that it may be AF, AU, OR, PA, NE (USA: Florida), NT (Bermuda, indigenous to the Pacific region, and Williams Guadeloupe, Jamaica, Puerto Rico, Trinidad, US and Martin (2003) speculated that the natural Virgin Islands). hosts are Arecaceae and possibly Pandanaceae Giant bamboo mealybug was first intro- (Freycinetia sp.). ScaleNet also records Fabaceae duced into Florida in 1956, but was thought to (Kentia sp.) and Poaceae (Phyllostachys sp.) as have been eradicated (Hodges and Hodges, 2004). additional hosts, but these are such aberrant It was subsequently re-collected in Miami-Dade records that they could be erroneous. Very little is County 1998, and has been infrequently and known about the biology or ecology of this spe- ­sporadically collected around the state. Hosts cies, but it does not seem to be a large threat to are exclusively species of grasses, primarily bam- palms in general. It was first recorded in the New boo, including Bambusa spp., Phyllostachys sp., World in Nassau, Bahamas, in 1980 (Williams, Dendrocalamus latiflorus Munro and D. asper 1981), and has been recorded from Florida Backer ex K. Heyne (FDACS-DPI database), and 26 times since its first collection in 1999. In Gigantochloa spp., Lingnania chungii (McClure) Florida, it has been recorded from the following McClure, Miscanthus sp. and Schizostachyum sp. counties: Broward, Flagler, Miami-Dade, Monroe, (ScaleNet). Hodges and Hodges (2004) reported Palm Beach, Polk and Sarasota, although most that since the species was not causing economic frequently from Miami-Dade. Collection records damage, the initial infestation in 1998 was not indicate that numerous species of palms are suit- discovered until older leaf sheaths were peeled able hosts, but Phoenix roebelenii O’Brien and back from the bamboo for cosmetic reasons. coconut are preferred.

19.2.7 Paracoccus marginatus 19.2.5 Palmicultor lumpurensis Williams and Granara de Willink (Takahashi) (=Trionymus lumpurensis) Common name: papaya mealybug, la cochenille No common name. Distribution: AU, OR, PA, NE du papayer. Distribution: AU, AF; NE (Mexico, (USA: Florida). Florida); NT (Antigua, Cuba, Dominican Republic, This mealybug is presumably endemic to the Guadeloupe, Haiti, St Kitts, Martinique, Puerto Old World, possibly China or Malaysia. Hosts are Rico, St Martin and St Barthélémy, British Virgin exclusively Poaceae, especially Bambusa spp. Very Islands, US Virgin Islands). Recent Adventive Scale Insects in Florida and the Caribbean Region 347

Papaya mealybug is an excellent example of 19.2.8 Maconellicoccus hirsutus (Green) an insect species moving from the relative obscurity of the newly described, to an increas- Common names: pink hibiscus mealybug, grape ingly wide distribution as a pest species as it mealybug, La cochenille de l’Hibiscus, la cochin- moves globally through international trade, and illa rosada del hibisco. Distribution: AU, AF, OR, back to a status as pest of negligible importance. PA, NE (USA: California, Florida, Louisiana, As such, it is a testament to the power of classi- Texas), NT (Cuba, Guadeloupe, Grenada, Haiti, cal biological control. Papaya mealybug is St Kitts and Nevis, Martinique, Puerto Rico, thought to be a native of Mexico or Central St Barthélémy and St Martin, Trinidad and America (Walker et al., 2006), and was described Tobago). relatively recently (Williams and Granara de Pink hibiscus mealybug (PHM) is one of the Willink, 1992; Miller et al., 1999; Miller and most recognizable names in the pantheon of pest Miller, 2002); however, it moved rapidly through insect species, familiar throughout much of the the Caribbean region, and by 2006 was present tropical to sub-tropical world to agricultural in at least 12 Caribbean islands. It was first workers and the ornamental growing public alike. recorded in Florida in 1998 and spread rapidly This pest of worldwide distribution was first throughout the state. ScaleNet records papaya recorded from Florida in 2002 (Hodges, 2002), mealybug from 45 species in 20 families, but in and its invasion into Florida from the Caribbean Florida, hibiscus accounts for 67% of host region, where it had been moving from island to records, with Jatropha integerrima Jacq. and island since the mid-1990s, proved to be inevita- Acalypha wilkesiana Muell. Arg. (Euphorbiaceae), ble despite robust quarantine measures. PHM and Carica papaya L. (Caricaceae) also common is one of the most polyphagous species of all hosts. Twenty-five percent of records are from known Coccoidea, having been recorded from Mimi-Dade County. over 340 plant species in 75 families, with The wide host range, attraction to certain Euphorbiaceae, Fagaceae and Malvaceae appear- hosts (such as hibiscus and papaya) and rapid ing to be the most preferred of host families worldwide spread made papaya mealybug an (ScaleNet). In Florida, it has been recorded from ideal candidate for classical biological control. 175 host ­species, but by far the majority of Thus, parasitoids collected from mealybug popu- records (90%) are from Hibiscus rosa-sinensis. lations in Mexico were artificially reared as part Other common hosts are Viburnum odoratissimum of ­various USDA programs (Meyerdirk et al., Ker-Gawl (Caprifoliaceae), Trema micrantha (L.) 2004; Muniappan et al., 2006; Amarasekare et al., Blume (Ulmaceae), Talipariti tiliaceum (L.) Fryxell 2009). The parasitoids Acerophagus papayae (Malvaceae), Senna polyphylla (Jacq.) Irwin and Noyes and Schauff, Anagyrus loecki Moyes and Barneby (Fabaceae) and Calophyllum spp. Pseudoleptomastix mexicana Noyes and Schauff (Clusiaceae). Of c. 3600 Florida records since (Encyrtidae) were released almost contempora- 2002, 25% are from Miami-Dade County and neously in Florida (2003), Guam (2002) and 15% from Broward County. Palau (2003–2004) from cultures maintained in In Florida, although PHM is particularly Puerto Rico. Amarasekare et al. (2009) reported destructive to hibiscus, various cultivars differ in that, of the three parasitoid species reared and their degree of susceptibility and degree of dam- released in Florida, only A. papayae and A. loecki age incurred through feeding (Vitullo et al., 2009). were recovered from experimental plots, with With hibiscus, PHM feeding typically induces A. papayae accounting for the highest mealybug abnormalities in the length of the internode, mortality. Papaya mealybug population reduc- resulting in a gall-like condition known as tion in Florida, as measured by the number of ‘bunchy-top’ or ‘rosetting’, which negatively samples submitted to FDACS-DPI, has been affects the aesthetic value of the plant (Meyerdirk ­dramatic, with a decline from a peak in 2004 of et al., 2001). Prolonged feeding results in defolia- 230 samples, to 28 samples in 2010. In quantita- tion and eventually death of the plant. tive studies of field effectiveness of parasitoid In anticipation of its eventual arrival into the controls, a reduction in mealybugs of as much NT region, the USDA, in conjunction with state as 97% has been observed (cited in Meyerdirk and university cooperators, and the agricultural et al., 2004). agencies of several countries in the Caribbean 348 I. Stocks

region, initiated biocontrol programs to help Sterculiaceae). However, Fernando and reduce the economic impact of PHM on horticul- Kanagaratnam (1987) reported that in Sri Lanka, ture and agriculture in the Caribbean region and P. lilacinus was found feeding on the peduncle and Florida. Economic losses in the Caribbean region stalk of the inflorescence of coconut, causing it to following introduction were significant, with dry up, and Waite and Martinez Barrera (2002) Grenada reporting losses of US$3.5–10 million reported that it was a minor pest of avocado over the 1996–1997 season, and Trinidad and (Persea americana Mill; Lauraceae) in the Tobago anticipated losses in excess of US$100 Philippines, where it occasionally caused early million per annum (Meyerdirk et al., 2001). PHM fruit drop. In a study to determine which biocon- was found in Louisiana in 2006 (Hodges et al., trol agents would be most suitable for control of 2007), Texas in 2007 (Anonymous, 2011a) and P. lilacinus in various crop systems in India, Mani Georgia in 2008 (Anonymous, 2011b), each infes- (1995) found that several predators and parasi- tation presumably the result of shipments of toids from Java and the Philippines were associ- infested nursery stock from Florida, even though ated with the mealybug, and were possibly FDACS-DPI quarantines shipments of stock posi- suitable for release in India. The species tive for PHM. Tetracnemoidea indica Ayyar and Leptomastix Of the 30 species of parasitoid wasps listed ­dactylopii Howard (Encyrtidae) and Aprostocetus by the Universal Chalcidoid Database as docu- purpureus Cameron (Eulophidae), and the mented primary host for PHM, the two encyrtid ­predators Spalgis epius (Westwood) (Lycaenidae), species Anagyrus kamali Moursi and Gyranusoidea Brumus sp., S. coccivora and Cryptolaemus indica Shafee, Alam and Agarwal were deemed ­montrouzieri Mulsant (Coccinellidae), Triommata most suitable for importation, rearing and coccidivora (Felt) (Cecidomyiidae), and Cacoxenus release. Each parasitoid has a preferred host perspicax (Knab.) (Drosophilidae) were recovered stage for the deposition of the egg and devel- from infested crops, but T. indica, A. purpureus opment of the larva, with Anagyrus kamali and S. epius appeared to be the most effici­­ females pre­ferring 3rd-instar PHM larvae, and ent at ­reducing mealybug populations, with G. indica females preferring late 2nd-instar lar- L. dactylopii having little effect. In a separate vae (Sagarra and Vincent, 1999; Ahmed et al., study, Mani and Krishnamoorthy (1990a) found 2007; Roltsch, 2007). Additionally, two coccinel- that C. montrouzieri eliminated P. lilacinus popula- lid predators (Cryptolaemus montrouzieri Mul­ tions on pomegranate (Punica granatum L.; sant and Scymnus coccivora Ayyar) were released, Lythraceae) fruits in India. In the laboratory, but only C. montrouzieri, the mealybug destroyer, Mani and Krishnamoorthy (1990b) found that appears to be effective in Florida. Through these the predatory larvae of the lacewing Mallada biocontrol efforts, the reduction in PHM pest ­desjardinsi (Navás) (=Mallada boninensis (Okamato)) load has been dramatic, with countries in the (Chrysopidae) was also effective against pas- Caribbean region experiencing persistent reduc- sionvine mealybugs. tions of 90–95% (Osborne et al., 2011). In Florida, PHM is a quarantinable pest, which helps reduce the overall pest load in the environment. However, it remains a common landscaping pest. 19.3 Soft Scales: Coccidae

Numerous pestiferous soft-scale species are found in Florida and the Caribbean region, 19.2.9 Planococcus lilacicus (Cockerell) ­including the recently described croton scale, Phalacrococcus howertoni Hodges and Hodgson, Common names: coffee mealybug, oriental cacao and there are fears that several other pest species mealybug. Distribution: AF, AU, OR, PA, NT could be introduced. The soft wax scale, (Dominican Republic, Haiti). Ceroplastes destructor Newstead, is a major pest of ScaleNet lists 89 recorded host plants in 36 Citrus in parts of its introduced range, but thus families, but this mealybug is most widely known far is not known from the New World. Vinsonia in the NT region as a pest of coffee (Coffea arabica stellifera (Westwood) (=Ceroplastes stellifera), L.; Rubiaceae) and cacao (Theobroma cacao L.; believed in 1954 to have been eradicated from Recent Adventive Scale Insects in Florida and the Caribbean Region 349

Florida (Hodges, 2004b), is being collected 19.3.2 Pulvinaria psidii Maskell more frequently, predominantly in Miami-Dade (=Chloropulvinaria psidii Borchsenius) County, and occurring most commonly on Schefflera spp. and Ixora sp. Also, in 2007, it was Common names: green shield scale, guava mealy newly reported from New Providence, Bahamas, scale, guava scale. Distribution: AF, AU, NE on Melaleuca quinquenervia (Cav.) S.F. Blake (USA: Alabama, DC, Florida, Georgia, Missouri, (Myrtaceae), a previously unreported host Mississippi, New York, Pennsylvania); NT (South (Blackwood and Pratt, 2007). America, Central America, Antigua, Bahamas, Bermuda, Cuba, Dominican Republic, Guadeloupe, Haiti, Jamaica, St Kitts, Montserrat, Martinique, Puerto Rico, Trinidad and Tobago, St Vincent and 19.3.1 Protopulvinaria longivalvata Green the Grenadines, US Virgin Islands). Green shield scale was first recorded in No common name. Distribution: AF, AU, OR, NT Florida in 1985 (FDACS-DPI database), but it has (South America, Guadeloupe, Haiti, Martinique, been moving slowly through the Caribbean islands Puerto Rico, US Virgin Islands), NE (USA: Florida, since the late 1890s (Jamaica 1895, Cuba 1926, interceptions only). Guadeloupe 1957), being present in 17 Caribbean This presumably Old World species began countries by 1993 (ScaleNet, 2011). ScaleNet lists appearing in the Caribbean region in the 1950s 207 host species in 62 families, with Moraceae, (Guadeloupe). It is frequently intercepted by the Myrtaceae, Rubiaceae and Rutaceae the most United States Department of Agriculture, with commonly reported families. The FDACS-DPI 200 quarantine interceptions from 1985 to 2010 database lists 118 host species, with a distinct from shipments or passengers originating in preference for Schinus terebinthifolius Raddi South and Central America and the Caribbean (Anacardiaceae), Ficus spp. (Moraceae), Ixora sp. region, including Antigua and Barbuda, Dominica, and Gardenia spp. (Rubiaceae), and agreeing in Jamaica, St Vincent and the Grenadines, and general with that reported by ScaleNet. Nada et al. Trinidad and Tobago. The most frequent intercep- (1990) reported that P. psidii (as Cribropulvinaria tions are from the Dominican Republic (23%), psidii) was one of the three most important pests Puerto Rico (18%) and Jamaica (12%) (USDA-ARS of mango in Egypt. El-Minshawy and Moursi database). This scale has been intercepted in quar- (1976) reported that P. psidii was also a serious antine in Florida by FDACS-DPI inspectors seven pest of guava in Egypt, but was also found times in the past 2 years, all from Schefflera throughout the year attacking S. terebinthifolius, ­arboricola (Hayata) Merr. (Araliaceae) plants origi- Meyerta sinclairii Seem. (Araliaceae) and Jasminum nating in Costa Rica. ScaleNet lists 21 host plant humile L. (Oleaceae). In Bangalore, India, Putta­ species in 12 families, including Mangifera indica rudriah and ChannaBasavanna (1957) reported L. (Anacardiaceae), Psidium guajava L. (Myrtaceae), that P. psidii was attacked by Cryptolaemus Gardenia spp. and Citrus spp. The most frequent ­montrouzieri, Mulsant (Coleoptera: Coccinellidae) host interceptions by the USDA were on Citrus (mealybug destroyer). In Bermuda, Microterys spp. (29%), Schefflera sp. (15%) and M. indica kotinskyi (Fullaway) and Coccophagus (= Aniseristus) (12%). Protopulvinaria longivalvata is very similar ceroplastae (Howard) were released as biocontrol to P. pyriformis Cockerell (pyriform scale), and agents, but M. kotinskyi was primarily responsible needs to be slide mounted for accurate identifica- for control (Bennett et al., 1976). The predators tion. There is relatively little information on C. montrouzieri and Azya luteipes (Mulsant) P. longivalvata, but thus far it does not appear to be (Coccinellidae) were also released and became as pestiferous as pyriform scale, which has a host established. In combination with the parasitoid list of 68 species in 34 families (but does not wasps, green shield scale was reduced to noneco- include S. arboricola as a known host). No data are nomic levels in Bermuda. Similarly, in South available regarding natural enemies, but observa- Africa, Annecke and Moran (1982) found that tions on submitted samples with parasitoid coccinellid predators were common and that ­emergence holes indicate that there may be at ­populations of green shield scale were parasitized least one species of parasitoid wasp attacking it so heavily that the scale is no longer of economic (Anonymous, 2009). importance (cited in de Villiers, 2001). 350 I. Stocks

19.3.3 Phalacrococcus howertoni Bursera ­simaruba L. (Burseraceae; gumbo-limbo) Hodges and Hodgson is the next most common host, with 7% of records, but Ficus spp. (Moraceae) and P. guajava Common name: croton scale. Distribution: NE L. are also common hosts. Thus far, naturally (USA: Florida); US Virgin Islands (St Thomas, occurring infestations are limited to the south- St Croix; FDACS-DPI database); Figs 19.1 ern-most regions of Florida, with 56% and 10% and 19.2. of records from Miami-Dade County and Broward Croton scale was first detected in Florida in County, respectively; however, due to the trans- 2008 and soon thereafter was determined to be port of infested nursery stock, FDACS-DPI has an undescribed species (Hodges, 2008). The records of croton scale from 27 counties. description followed in 2010, by which time At this time, relatively little is known regard- ­croton scale was recorded from 72 host plant ing the biology of this pest species, but some life ­species in 34 families. As of February 2011, the history details have been discovered (Hodges and host list had grown to 90 species in 36 families, Hodgson, 2010; C. Mannion and S. Brown, with 60% of the host records from Codiaeum unpublished data). In primary areas of infesta- ­variegatum (L.) A. Juss. (Euphorbiaceae; croton). tion in south Florida, year-round warm weather allows for overlapping generations throughout the year, with females taking roughly 30 days to complete development. Female fecundity is high, with egg production of up to 400 eggs per female. Natural predators have been associated with cro- ton scale infestations, including C. montrouzieri Mulsant, Azya orbigera Mulsant (Coccinellidae) (H. Liere, pers. comm.) and Laetilia coccidivora Comstock (as Laelilla in Hodges and Hodgson, 2010). The commercially available parasitoid wasp Metaphycus flavus (Howard) (Hymenoptera: Encyrtidae) has also been collected in the field, 1 but is apparently at such low levels that it has ­little effect on population size. Infestations can build rapidly and can cover Fig. 19.1 Croton scale Phalacrococcus howertoni Hodges and Hodgson (Coccidae) adult female on almost all above-ground parts of the plant (espe- croton (Codiaeum variegatum). Photograph, Ian cially croton). Females prefer the stems and Stocks. ­petioles, while males prefer the adaxial leaf sur- face (personal observation). A heavily infested croton becomes highly stressed through loss of nutrients and the production of honeydew, and can eventually succumb. At Sanibel Island (Lee County, Florida), conservation staff have observed this scale on both native and non-native plants in a wildlife refuge. They report that gumbo limbo and strangler fig (Ficus aurea Nutt.) are showing signs of stress, with branch and twig die-back attributed to heavy scale infestations (J. Evans and S. Brown, pers. comm.). They report that, in addition to gumbo-limbo and strangler fig, firebush (Hamelia patens Jacq.), wild coffee 2 (Psychotria nervosa Sw.), and Bahama wild ­coffee (P. ligustrifolia (Northrop) Millsp.) Fig. 19.2 Croton scale immature male puparia (Rubiaceae), paradisetree (Simarouba glauca DC; (left), adult male under puparium (center), Simaroubiaceae), and marlberry (Ardisia escallon­ immature female (right). Photograph, Ian Stocks. oides Schiede and Deppe ex Schltdl. and Cham.) Recent Adventive Scale Insects in Florida and the Caribbean Region 351

and myrsine (Myrsine cubana DC) (Myrsinaceae) lobate lac scale (as P. pseudolobata) has an are also heavily affected. However, horticultural unknown provenance, which hinders the task of experience with this scale is still so limited that it finding parasitoids for biocontrol considerably. is too early to tell what effect it will have on the Lobate lac scale was first discovered in natural landscape in the long term. Florida in 1999, but subsequent examination of slides in the Florida State Collection of Arthro­ pods (FSCA) revealed that the scale has been in 19.4 Lac Scales: Kerriidae several Bahamian islands since at least 1992. In Florida, by 2004 the scale had spread to six ­counties and was by that time considered to be Two lac scales are known from Florida, both of a serious pest (Anonymous, 2004; Mannion et al., which are adventive, and either or both of which 2005), prompting the publication of several fact- could easily become established widely in the sheets (Hamon and Hodges, 2004; Mannion Caribbean region. Paratachardina pseudolobata et al., 2005; Howard et al., 2009). The host list for Kondo and Gullan is seriously pestiferous, this species is extensive, with up to 307 plant whereas the pest status of Tachardiella mexicana species enumerated in a report by Howard et al. (Comstock) is still under investigation. Although (2006). While many plants from the list of an additional 17 species of Kerriidae are found in known host plants are variable in their suita­ the NT region, only Austrotachardiella gemmifera bility as hosts or susceptibility once infested, (Cockerell) (=Tachardia gemmifera), which is there are a number of species that are clearly endemic to Jamaica on Chrysobalanus spp. ­preferred hosts. Howard and Pemberton (2003) (Chrysobalanaceae), and P. pseudolobata, which is list wax myrtle (Morella cerifera (L.) Small = adventive, are currently found in the Caribbean Myrica ­cerifera; Myricaceae), Chrysobalanus region (ScaleNet, 2011). Other than the report icaco L. (Chrysobalanaceae), Conocarpus erectus by Cockerell (cited in ScaleNet, 2011) that L. (Combretaceae), Myrsine guianensis (Aublet) A. ­gemmifera was destroying Chrysobalanus spp. Kuntze, Psychotria spp., Annona spp. (Anno­ trees in Kingston, no life-history data are pub- naceae), Averrhoa carambola L. (Oxali­daceae), lished for this species. Litchi chinensis Sonn. (Sapindaceae), Ficus spp., H. rosa-sinensis and M. indica as preferred hosts. Wax myrtle is so susceptible that several plants 19.4.1 Paratachardina pseudolobata in one survey plot in south Florida died within Kondo and Gullan 1 year after becoming heavily infested (Howard et al., 2006), and another experimental host had Common name: lobate lac scale, escama lobada to be found because the wax myrtles died soon de laca. Distribution: AU, NE (USA: Florida), after experimental infestation. NT (Bahamas: Andros, Grand Bahamas, New Controlling this pest scale has proved chal- Providence). lenging. The hard resinous test is extremely Confusion as to the identity of this scale was ­effective at preventing topically applied insecti- finally clarified by Kondo and Gullan (2007) in cides from reaching the scale. Soil drenches of their revision of the genus Paratachardina. The lac imidacloprid were effective at suppressing scale insect christened lobate lac scale was originally populations (Howard and Pemberton, 2003), but identified as P. l. lobata (Chamberlin), but was long-term suppression will require finding, found by Kondo and Gullan (2007) upon a more ­rearing and releasing parasitoid biocontrol detailed analysis to be a distinct and undescribed agents. Initial reports of effective biocontrol species. Thus, the literature on the lobate lac scale agents are now known to refer to P. lobata lobata, is confused, with some publications referring to and are therefore unsuitable candidates; however, what is actually P. l. lobata, and others referring to field populations of lobate lac scale monitored the new species P. pseudolobata; this should be in south Florida yielded Metaphycus sp. and borne in mind when consulting the literature. For Ammonoencyrtus sp. (Encyrtidae), but the num- instance, it was originally claimed that the lobate bers of these parasitoids relative to the number lac scale (as P. l. lobata) was native to India and Sri of potential hosts was very low (Howard and Lanka (Howard and Pemberton, 2003). In fact, Pemberton, 2003). 352 I. Stocks

19.4.2 Tachardiella mexicana that they feed directly from plant cells as opposed (Comstock) (also including T. texana to xylem or phloem (which reduces their expo- as a junior synonym) sure to ­systemic insecticides), they are notori- ously hard to control once established. Five No common name. Distribution: NE (Mexico, adventive species have recently been detected USA: Texas, Florida). in Florida: Unachionaspis tenuis (Maskell), Native to the south-west of the USA and to Aulacaspis yasumatsui Takagi, Poliaspis cycadis Mexico, this lac scale was first discovered in (Comstock), Aulacaspis tubercularis Newstead and Florida in 1985 infesting Acacia sp. at an amuse- Duplachionaspis divergens (Green). Unfortunately, ment park in Orange County, and a second dis- other than host and distribution information, no junct population was discovered at a nursery in information is available about the biology of Seminole County in 1987 infesting Texas ebony either P. cycadis or U. tenuis. (Ebenopsis ebano (Berl.) Barneby and Grimes = Pithecellobium flexicaule). Since then, this scale has been found in other locations and on other 19.5.1 Unachionaspis tenuis (Maskell) hosts, including Lysiloma latisiliquum L. (Benth.), L. sabicu Benth., Acacia pinetorum F.J. Herm. and No common name. Distribution: PA, OR, NE A. cornigera (L.) Willd. Overall, this lac scale shows (USA: Florida, South Carolina). a marked preference for Fabaceae, including Hosts are grasses (Poaceae), including: Mimosa sp. in its native range. Thus, it is discon- Bambusa sp., Phyllostachys spp., Pleioblastus spp., certing that another population of this scale was Sasa sp. and Shibataea kumasaca (Zoll. ex Steud.) discovered in 2010 infesting wax myrtle at the Makino. In Nassau County, Florida, it has been same amusement park. taken exclusively from Bambusa sp. There is one Nothing is known about the biology of this additional unpublished record of this species scale in its native environment. In Florida, it from one collection in South Carolina in a botani- can have profound consequences for the tree or cal garden, also on Bambusa sp. shrub, leading to defoliation and death of the plant (S. Brown, unpublished data). In Lee County, massive populations killed several wild 19.5.2 Aulacaspis yasumatsui Takagi tamarind (L. latisiliquum) trees, and numerous small wax myrtleshrubs were dead or dying in Common names: cycad aulacaspis scale, cycad Orange County (pers. obs.). Massive quantities scale, sago palm scale. Distribution: AU, OR, PA, of honeydew are secreted, leading to a prolifera- NE (USA: Florida, Texas, Louisiana), NT (Cayman tion of sooty mold on the plant, surrounding Islands, Puerto Rico, US Virgin Islands: St John, plants and ground. Although initially each scale Barbados); Figs 19.3–19.4. starts growth in isolation, in heavy infestations Thought to be native to Thailand, cycad larger females will coalesce into a hard resinous scale has spread widely across the globe and mass that completely envelops the branch. Thus far, no parasitoids are known, and control efforts using horticultural oils are only mini- mally effective, even at low infestation levels.

19.5 Armored Scales: Diaspididae

Armored scales are notorious tramp species, pri- 3 marily due to their small size and the attendant difficulty in finding them during inspections. Also, because of the protective cover that they Fig. 19.3 Cycad scale, Aulacaspis yasumatsui manufacture (which substantially protects Takagi (Diaspididae) adult female scale covers on them from topical insecticides), and the fact adaxial leaflet surface. Photograph, Ian Stocks. Recent Adventive Scale Insects in Florida and the Caribbean Region 353

and may in fact be the primary native host for A. yasumatsui, it has also been collected from six other commercially available species, Microcycas calocoma (Miq.) DC (native to Cuba and endan- gered; Vovides et al., 1997), the Zamiaceae hosts Dioon spp., Encephalartos spp. and Macrozamia moorei F.J. Muell. and Stangeria eriopus (Kunze) (Stangeriaceae). A heavily infested plant will have scales on all parts of the plant, including the roots (Howard et al., 1999). 4 In its native range, naturally occurring para- sitoids and predators help maintain populations Fig. 19.4 Cycad scale, primarily male puparia on below levels that cause significant damage to the adaxial leaflet surface. Photograph, Ian Stocks. plants (Hodgson and Martin, 2001). However, outside its native range, pest load can cause from 70% to 100% mortality of infested C. revoluta can now be found wherever Cycas spp. (Cycadales: (Hodgson and Martin, 2001). Howard et al. Cycadaceae) and Zamia spp. (Cycadales: Zamia­ (1999) reported that leaves of recently infested ceae) grow naturally or are cultivated (Hodgson experimental C. revoluta plants were necrotic and Martin, 2001; Germain and Hodges, 2007). within 112 days of exposure, with the death of Cycad scale was first collected in Florida in 1996, the plant occurring in 1 year post exposure. In presumably having been introduced via the inter- control experiments, levels of pesticide applica- national trade in ornamental cycads (Weissling tion (imidacloprid) exceeding that practicable by and Howard, 1999). Other regions of the south- home gardeners were only marginally successful, ern US that are suitable for the outdoor cultiva- perhaps because the plants were rapidly rein- tion of cycads (Louisiana and Texas) also have the fested from the root populations untouched by scale (Germain and Hodges, 2007), but thus far the pesticide application (Howard et al., 1999). To in California it remains unrecorded outside of date, the only pesticide products with known effi- nursery settings (G. Watson, pers. comm.). ScaleNet cacy are certain formulations of horticultural oils, reports that the Caribbean region distribution the primary benefit being a reduction in the includes Cayman Islands, Puerto Rico and the US crawler stage (Howard and Weissling, 1999). Virgin Islands (St John). The Puerto Rico records Natural enemies, which often provide the are derived from the FDACS-DPI sample submis- best and most efficient long-term control, have sion database, but the Cayman Islands and US been discovered and have provided partial con- Virgin Islands reports were unverified at the time trol. In Florida, the scale predator Cybocepahlus of their publication (Howard and Weissling, ­nipponicus Endroudy-Younga (Nitidulidae, originally 1999). As reported by the Barbados Ministry of identified as C. binotatus) and parasitoid Coccobius Agriculture and Rural Development website, it fulvus (Compere and Annecke) (Encyrtidae) were has been present there since 2003 (Lavine, 2010). bred for a release program. These are now estab- Reports of the scale’s presence in Martinique lished throughout the cycad scale infestation since 2005 were cited by Germain and Hodges range, and can eliminate a scale population (2007), and it is now considered established; (Howard and Weissling, 1999). Recently, other however, given the rapidity of the worldwide predators and parasitoids have been recovered spread of this pest, and the size of the popula- from cycad scale-infested areas. In south Texas, tions that can build up, it is very unlikely that the where ornamental cycads are grown commercially, Caribbean region in general will escape the estab- the predator Rhizobius lophanthae Blaisdell lishment of this scale. For a general discussion of (Coccinellidae) and Aphytis sp. lingnanensis group the biology of cycad scale, the reader is referred (Aphelinidae) appear to be keeping the cycad scale to Howard et al. (1999), and the synopsis availa- at relatively low levels (Flores and Carlson, 2009). ble at Ben-Dov et al. (2011). Field recognition of this species is rela- While the commercially popular Cycas revo­ tively straightforward, although slide-mounting luta is by far the most common host in Florida, adult female specimens is necessary for positive 354 I. Stocks

­identification. The adult female scale is white to not very significant, pest of mango (FDACS-DPI off-white, tear-drop to roughly circular in outline database). ScaleNet lists 44 host species records and 1–2 mm along the longest axis (Plate 2). in 17 families, with a heavy bias to members of Beneath the scale cover, the insect is orange in the Lauraceae; however, thus far, Lauraceae of colour. Females preferentially colonize the adax- Florida appear to have not become common host ial surface of the leaflet, with relatively lower den­ plants, with one record each from Cinnamomum sity on the rachis and abaxial surface; ­however, zeylanicum Garcin ex Blume and Persea palustris the male scale covers, which are <1 mm long and (Raf.) Sarg. Seventy-five percent of records are parallel sided, preferentially colonize the abaxial from Miami-Dade County, but we have records leaflet surface and both sides of the rachis. In from five other primarily southern Florida heavily infested plants, almost the entire surface counties. of the leaf will be covered. Care should be taken in identifying cycad scale in the field, because the white magnolia scale (= false oleander scale), 19.5.5 Duplachionaspis divergens Pseudaulacaspis cockerelli (Cooley) has a similar (Green) scale cover with a similarly shaped body; how- ever, white magnolia scale adult females and their No common name. Distribution: AU, OR, PA, NT, eggs are yellow, and they typically do not build up NE (USA: Florida, intercepted from Alabama and large populations on the leaflets. Texas, possibly established in Texas). This species was first discovered in 2002 on Miscanthus sp., but an examination of slides of 19.5.3 Poliaspis cycadis Comstock unidentified scales from Miscanthus revealed that it has been in Florida since at least 2000 No common name. Distribution: OR, PA, NE (Hodges, 2004c). Evans and Hodges (2007) pub- (USA: California, Florida, DC). lished a brief article on this species in which Hosts include Cycas spp. (Cycadaceae), Dioon they provide a synopsis of the known ecology, edule (Zamiaceae), Gaultheria spp. (Ericaceae) and economic impact and biological control issues. Microsemia sp. (Brassicaceae). UK and Scottish This species is now widely distributed around records are presumably from hothouse plants, Florida, known from 35 counties. In 2010, and in fact the type series is from Washington DC FDACS-DPI received a ­sample from St Croix, on Cycas revoluta Thunb., Dioon edule Lindl. and US Virgin Islands, on Bothriochloa pertusa (L.) Microsemia sp. plants grown in a conservatory. In A. Camus. It has been collected exclusively from Florida, it has been collected in Miami-Dade Poaceae, including Andropogon spp., Cimbopogon County from Dioon sp., but appears to be either citratus (DC ex Nees) Stapf, Eustachys sp., not established or at very low levels in the envi- Miscanthus sinensis Andersson, Pennisetum sp., ronment (FDACS-DPI database; Hodges and Saccharum spp., Stenotaphrum secundatum Dixon, 2007). (Walt.) O. Kuntze, Tripsacum dactyloides (L.) L. and T. floridanum ­Por­ter ex Vasey. In regions where sugarcane is grown, it has caused minor economic impact, but parasitoids appear to sup- 19.5.4 Aulacaspis tubercularis Newstead press the scale to below economic levels. In Florida, Aphytis sp. lingnanensis group and Common names. cinnamon scale, mango scale, Encarsia citrine (Craw) (Aphelinidae) have been white mango scale, escama del mango, escama reared from populations of D. divergens. blanca del mango. Distribution: PA, OR, AF, AU, NE (USA: Florida), NT (Antigua, Aruba, Barbados, Bermuda, British Virgin Islands, Dominican Republic, Grenada, Guadeloupe, 19.6 Cushiony Scales: Jamaica, Martinique, Puerto Rico, Saint Croix, Monophlebidae St Lucia, Trinidad and Tobago, US Virgin Islands). This scale was first observed in Florida in Nine species of monophlebid scales are known 2002, and has since become a common, though from the Caribbean. Perhaps the most seriously Recent Adventive Scale Insects in Florida and the Caribbean Region 355

pestiferous is Icerya p. purchasi Maskell (citrus 19.7 Whiteflies: Aleyrodidae fluted scale, cottony cushion scale), which is present on 14 Caribbean islands, but Crypticerya This family contains many well-travelled species, genistae (Hempel), known from Guadeloupe, also several of which have had profound and long-last- causes economic injury. Icerya s. seychellarum ing impacts on agriculture, namely Dialeurodes (Westwood) has recently been recorded from citri (Ashmead) (citrus whitefly), Aleurocanthus Guadeloupe on M. indica and Citrus sp., and from woglumi Ashby (citrus blackfly) and Aleurodicus Martinique on P. guajava (Matile-Ferrero and dispersus Russell (spiraling whitefly). In recent Étienne, 2006), but nothing is known yet of its years, several species of whiteflies have become biology on these islands. Williams and Butcher established either in Florida, the Caribbean region (1987) reported that this species is capable of or both. Four species are of interest for this killing Citrus trees, and Beardsley (1966) reported review, at least three of which either could be, or that it was a severe pest of breadfruit (Artocarpus are currently, found in the Caribbean region. altilis (Parkinson) Fosberg).

19.7.1 Dialeurodes schefflerae Hodges 19.6.1 Crypticerya genistae (Hempel) and Dooley (Aleyrodinae) (=Icerya genistae) Common name: schefflera whitefly. Distribution: Distribution: NT (Barbados, Guadeloupe), NE Indonesia, USA (Florida, Hawai’i); NT (Puerto Rico). (USA: Florida). Dialeurodes schefflerae was described in There is no accepted common name for 2007, but a systematic review of unidentified this species, but images of it are returned after slide specimens collected from Schefflera species an internet search using the name ‘white par- in preparation for the published description indi- tridge pea bug’. Little has changed regarding cated that it has been in Hawai’i since c. 1960, our knowledge of this species since a Pest Alert California since 1988–1990 and Florida since was created in 2006 (Hodges, 2006) and short 1986 (Hodges and Dooley, 2007). Various lines of technical note written in 2008 (Hodges et al., evidence, such as host-plant affiliation and affin- 2008). This species was first collected in Florida ity with other Dialeurodes species, suggest that it in Broward County in 2006, and has since is native to Asia. Collection data indicate that it is spread throughout Broward County and into polyphagous within the genus Schefflera, but has Miami-Dade and Palm Beach Counties. The host a marked preference for the popular landscape list, which is quite broad at over 50 ­species, con- and potted-plant species S. arboricola (Hayata) tinues to grow. There is little evidence of host Merr. Hodges and Dooley (2007) noted that no ­preference pattern other than a slight bias to serious damage occurs to the plant, even though Asteraceae, Fabaceae and Euphorbiaceae (espe- the whiteflies can reach quite high densities on cially Chamaesyce spp.), an observation in the leaf (pers. obs.). The current distribution in Florida that was also made for this species in Florida includes 16 counties, with greatest abun- Guadeloupe (Étienne and Matile-Ferrero, 2008). dance in two of the southernmost counties, Hodges et al. (2008) reported that it is a pest of Broward and Miami-Dade. The records for San several vegetable crops in Barbados, where the Juan, Puerto Rico are based on pre-departure Ministry of Agriculture is currently surveying quarantine interceptions of infested propagative for this pest and searching for biocontrol agents. Schefflera sp. plants (Hodges and Dooley, 2007). Étienne and Matile-Ferrero (2008) also reported that the coccinellid beetle Rodolia cardinalis (Mulsant), the vedalia beetle, was found associ- 19.7.2 Siphoninus phillyreae (Haliday) ated with the mealybugs. In Guadeloupe, Gagné (Aleyrodinae) and Étienne (2009) found the presumably pred- atory midge Pectinodiplosis erratica (Felt) Common names: ash whitefly, pomegranate (Cecido­myiidae) along with remains of C. genis­ whitefly, la mosca blanca del Fresno, mosca blanca tae feeding on Mimosa sp. (Fabaceae). del granado. Distribution: Western PA; AU; NT 356 I. Stocks

(Mexico, Central America, USA: California, Florida, Giant whitefly was first discovered in the USA Georgia, Hawai’i, North Carolina, Virginia; also in Texas in 1991, and by 1996 was established in reported from Arizona, New Mexico and Nevada). California, Louisiana and Florida. This whitefly is a Throughout much of its range, ash whitefly highly polyphagous species that can build to severe has historically been a severe pest on a wide variety infestation levels, producing such large quantities of commercial and non-commercial plant species, of long stringy wax that the plant itself becomes especially pomegranate and Citrus spp. (Abd-Rabou, obscured (Hodges, 2004d; Smith and Fox, 2004; 2006). Ash whitefly was discovered in California in Martin, 2008). The FDACS-DPI database lists 51 1988, where it quickly became a severe pest that host plant species, but 66% of the samples received resulted in millions of dollars of economic loss. were from H. rosa-sinensis; however, there appears A similar situation ensued when ash white­fly app­ in recent years to be a substantial decrease in the eared on ornamental pear trees (Pyrus ­calleryana populations, reflected in a decrease in samples Decne. ‘Bradford’; Rosaceae) in North Carolina in submitted to FDACS-DPI. The reasons for this are 1993. Taking advantage of the results of ­biological not entirely clear, but the decrease may be due to control programs developed in the Middle East, suppression by parasitoids. Shortly after the giant whitefly parasitoid E. inaron Walker (=E. partenopea whitefly’s discovery in Texas, Entedononecremnus Masi) (Aphelinidae) was imported into California krauteri Zolnerowich and Rose (Eulophidae) was and reared for eventual release (Pickett et al., 1996). collected from puparia, and subsequently reared Two years after release, the parasitoid had effected and released in California. The other parasitoids a nearly complete suppression of ash whitefly on associated with giant whitefly are: Encarsia both major hosts, ash (Fraxinus sp.) and pomegran- ­guadeloupe Viggiani, E. brasiliensis Hempel, ate. Additionally, several other ­parasitoids and E. meritoria Gahan, E. noyesi Hayat (=Encarsiella the beetle predator Clitostethus arcuatus (Risso) noyesi) (Eulophidae) and Idioporus affinis LaSalle (Coccinellidae) have been used in biological control and Polaszek (Pteromalidae) (Noyes, 2003). The programs in other parts of the ash whitefly’s beetle Delphastus catalinae Horn (Coccinellidae), a range. In Mexico, Myartseva (2006) discovered generalist whitefly predator, has also been recov- Eretmocerus sp. (Aphelinidae) and the hyperpara- ered from giant whitefly infestations (Evans, site Signiphora aleyrodis Ashmead (Signiphoridae) 2008). This beetle is present in Florida (FSCA attacking ash whitefly, and she concluded that the museum records), but several related Delphastus parasitoids had been ­introduced along with the species are found throughout the Caribbean whitefly. Overall, 23 Hymenoptera parasitoids or region, including D. chapini Gordon (Trinidad), associates of ash whitefly are reported in the D. guiniculus Gordon (Dominican Republic), Universal Chalcidoid Database. D. nebulosus Chapin (Puerto Rico), D. pallidus In Florida in 2010, a small population of ash (LeConte) (Bahamas, Cuba, Dominican Republic, whitefly was discovered on containerized pome- Virgin Islands), D. pusillus (LeConte) (Cuba, granate trees in the nursery grounds of an amuse- Jamaica) (Gordon, 1970; FSCA museum data). ment park in Orange County, the discovery of In Florida, release of E. krauteri was initiated which prompted the release of a FDACS-DPI Pest in 1997 in Seminole, Indian River, St Lucie and Alert (Stocks and Hodges, 2010b). Later in 2010, a Volusia Counties. In 1998, E. noyesi and Encarsia second population was discovered on Citrus trees sp. were added to the biocontrol program. Survey on private property in Bay County, and is presumed data from 2001 found that E. krauteri and E. noyesi to be the result of a separate introduction. The had become established and appeared to be parasitoid E. inaron was reared from the whiteflies ­having a significant effect on giant whitefly from Orange County and an unidentified parasi- populations. toid was reared from the whiteflies in Bay County.

19.7.3 Aleurodicus dugesii Cockerell 19.7.4 Aleurodicus rugioperculatus (Aleurodicinae) Martin

Common name: giant whitefly. Distribution: Proposed common name: rugose spiraling white- NE (USA: Arizona, California, Florida, Hawai’i, fly. Distribution: USA (Florida); Belize, Guatemala, Louisiana, Texas; Mexico) NT; Pakistan; Java. Mexico (Martin, 2008) (Figs 19.5–19.6). Recent Adventive Scale Insects in Florida and the Caribbean Region 357

Miami-Dade County, and by December 2010, FDACS-DPI had accumulated 87 records. In ­res­ponse, two factsheets were released to inform home­owners, landscapers and plant regulatory officials (Mannion, 2010; Stocks and Hodges, 2010c). The majority of records are from Miami- Dade County, but through the spread of infested nursery stock, it is now recorded from two addi- tional counties. In Miami-Dade County, rugose spiraling whitefly has spread rapidly through the environment on its own to infest an ever- increasing area. Furthermore, the host list, which is currently at 37 species, keeps expand- ing as more collections are made. Bursera sima­ ruba (L.) Sarg. (Burseraceae) is the most common, 5 with 23% of the host records, followed by coco- nut with 10%, Bucida buceras L. (Combretaceae) with 9% and P. americana with 7% of records. Fig. 19.5 Puparia of rugose spiraling whitefly, While coconut is a preferred palm host, at least Aleurodicus rugioperculatus Martin. Photograph, five other palm species­ are suitable hosts. Lyle Buss. Rugose spiraling whitefly is, at present, con- sidered to be seriously pestiferous, causing exten- sive aesthetic damage to the host plants and physical damage by the production of prodigious quantities of honeydew. The time since infesta- tions were noted has been too brief to determine if it causes significant health problems to the tree, but it seems likely that it will have severe negative consequences. In many cases, the leaves of Bursera simaruba and palms have been entirely covered with puparia and the sticky flocculent wax they produce. The honeydew promotes the growth of sooty mold, further compromising the aesthetic quality of the tree and hindering nor- 6 mal tree physiology. Whether rugose spiraling whitefly continues to increase its range and effect Fig. 19.6 Rugose spiraling whitefly, adult female. in the landscape will almost certainly depend on Photograph, Lyle Buss. the role of parasitoids and predators. Martin (pers. comm.) noted parasitoid emergence holes in puparia from Belize, and at least one species of Rugose spiraling whitefly was described parasitoid wasp (Aleuroctonus vittatus Dozier; from Belize in 2004 from Cocos nucifera L. (Martin, Eulophidae) has been associated with infesta- 2004), with subsequent records from Mexico and tions in Florida, often at high levels, and samples Guatemala on Caryocar amygdaliferum Mutis received by FDACS-DPI frequently show signs of (Caryocaraceae), P. guajava, P. americana Mill., parasitism (I. Stocks, pers. obs.). Musa sp. (Musaceae) and Melia sp. (Meliaceae). No additional information about the biology or ­ecology of this whitefly was published, so all Acknowledgements that is currently known about rugose spiraling whitefly is derived from limited experience with The author would like to thank several people for it in Florida. The first specimens received by their assistance during the preparation of this FDACS-DPI were collected in March 2009, from chapter. Beverly Pope (FDACS-DPI) helped to 358 I. Stocks

locate several obscure articles; Dr Greg Evans Anonymous (2011a) Pink Hibiscus Mealybug in (USDA-APHIS) extracted interception data from Texas. Available at http://etipm.tamu.edu/ USDA databases; Dr Catharine Mannion insect_alerts/ph_mealybug.cfm. (UF-IFAS), Dr Greg Hodges (FDACS-DPI), Jenny Anonymous (2011b) Pink Hibiscus Mealybug Found in North Georgia. Available at www.plantman- Evans (Sanibel-Captiva Conservation Foundation) agementnetwork.org/pub/php/news/2008/ and Stephen Brown (UF-IFAS); Gillian Watson PHMealybug, accessed 9 August 2012. (CDFA) provided unpublished information on Beardsley, J.W. (1966) Insects of Micronesia, several species of scales and whiteflies discussed Homoptera: Coccoidea. Insects of Micronesia in this article. Lyle Buss, UF, kindly allowed repro- 6, 377–562. duction of photographs. Ben-Dov, Y., Miller, D.R. and Gibson, G.A.P. (2011) Remarks Concerning the biology of a scale query results. www.sel.barc.usda.gov/scalecgi/ Note bionote.exe?Family=Diaspididae&genus=aula caspis&species=yasumatsui&subspecies=&r mktype=biology, accessed 7 August 2012. i www.sel.barc.usda.gov/ScaleNet/ScaleNet.htm, Bennett, F.D., Rosen, D., Cochreau, P. and Wood, B.J. accessed 8 August 2010. (1976) Biological control of pests of tropical fruits and nuts. In: Huffaker, C.B and References Messenger, P.S. (eds) Theory and Practice of Biological Control. Academic Press, New York, New York, pp. 359–395. Abd-Rabou, S. (2006) Biological control of the Blackwood, J.S. and Pratt, J.D. (2007) New host pomegranate whitefly, Siphoninus phillyreae and expanded geographic range of stellate (Homoptera: Sternorrhyncha: Aleyrodidae) by scale (Hemiptera: Coccidae: Ceroplastinae). using the bioagent, Clitostethus arcuatus Florida Entomologist 90, 413–414. (Coleoptera: Coccinellidae). Journal of de Villiers, E.A. (2001) Pulvinaria psidii (Cockerell). Entomology (Faisalabad) 3, 331–335. Guava scale. In: van den Berg, M.A., de Ahmed, S.A., El-Saedi, A.A, Ahmed, H.M and Villiers, E.A. and Joubert, P.H. (eds) Pests and Sayed, A.M. (2007) Host-parasitoid relation- Beneficial Arthropods of Tropical and Non- ship between the parasitoid, Anagyrus citrus Subtropical Crops in South Africa. ARC- kamali Mourse (Hymenoptera: Encyrtidae) Institute for Tropical and Subtropical Crops, and the pink hibiscus mealybug, Maco­ Nelspruit, South Africa, pp. 174. nellicoccus hirsutus (Green), (Homoptera: Division of Plant Industry (2004) Pink hibiscus Pseudococcidae). Egyptian Journal of mealybug biological control. Available at www. Biological Pest Control 17, 107–113. freshfromflorida.com/pi/methods/phm.html. Ali, M. (1987) Palm mealybug, Palmicultor pal- El-Minshawy, A.M. and Moursi, K. (1976) Biological marum (Homoptera: Pseudococcidae), new to studies on some soft scale-insects (Hom., the Indian subcontinent. Annals of the Coccidae) attacking guava trees in Egypt. Entomological Society of America 80, 501. Zeitschrift fur Angewandte Entomologie 81, Amarasekare, K.G., Mannion, C.M. and Epsky, N.D. 363–371. (2009) Efficiency and establishment of three Étienne, J. and Matile-Ferrero, D. (2008). Crypticerya introduced parasitoids of the mealybug genistae (Hempel), nouveau danger en Paracoccus marginatus (Hemiptera: Guadeloupe (Hemiptera, Coccoidea, Pseudococcidae). Biological Control 51, 91–95. Monophlebidae). Entomologique d’Egypte Annecke, D.P. and Moran, V.C. (1982) Insects 113, 517–520. and Mites of Cultivated Plants in South Evans, G.A. (2008) The whiteflies (Hemiptera: Africa. Butterworths, Durban, South Africa, Aleyrodidae) of the and their host plants and pp. 383. natural enemies. Available at www.sel.barc.usda. Anonymous (2004) Lobate lac scale and gov:8080/1WF/World-Whitefly-Catalog.pdf. Melaleuca: a devastating insect aided by an Evans, G.A and Hodges, G.S. (2007) invasive tree. Available at http://tame.ifas.ufl. Duplachionaspis divergens (Hemiptera: edu/pdfs/publications/Lobate.pdf. Diaspididae), a new exotic pest of sugarcane Anonymous (2009) Protopulvinaria longivalvata. and other grasses in Florida. Florida Available at www.freshfromflorida.com/pi/enpp/ Entomologist 90, 392–393. triology/4802/triology_4802_entomology.html, Fernando, L.C.P. and Kanagaratnam, P. (1987) accessed 9 August 2012. New records of some pests of the coconut Recent Adventive Scale Insects in Florida and the Caribbean Region 359

inflorescence and developing fruit and their Diaspididae). Available at www.freshfromflorida. natural enemies in Sri Lanka. Cocos 5, 39–42. com/pi/enpp/ento/d.divergens.html. Flores, D. and Carlson, J. (2009) Fortuitous estab- Hodges, G.S. (2004d) Giant whitefly, Aleuro­ lishment of Rhizobius lophanthae (Coleoptera: dicus dugesii Cockerell, in Florida. Available Coccinellidae) and Aphytis lingnanensis at www.freshfromflorida.com/pi/pest-alerts/ (Hymenoptera: Encyrtidae) in south Texas on aleurodicus-dugesii.html, accessed 8 the cycad aulacaspis scale, Aulacaspis yasu- August 2012. matsui (Hemiptera: Diaspididae). Southwestern Hodges, G.S. (2006) Crypticerya genistae Naturalist 34, 489–491. (Hemiptera: Margarodidae). A new exotic Gagné, R.J. and Étienne, J. (2009) Note on the scale insect for Florida. Available at www. Cecidomyiidae from Guadeloupe (West Indies) freshfromflorida.com/pi/enpp/ento/c. with description of a new species of genistae.html Paracalmonia (Diptera). Bulletin de la Société Hodges, G.S. (2008) A new exotic soft scale insect Entomologique de France 114, 337–350. on croton in south Florida (Hemiptera: Germain, J.F. and Hodges, G.S. (2007) First Coccoidea: Coccidae). Available at www.fresh- report of Aulacaspis yasumatsui (Hemiptera: fromflorida.com/pi/enpp/ento/coccoidea_­ Diaspididae) in Africa (Ivory Coast), and coccidae.html. update on distribution. Florida Entomologist Hodges, G.S. and Buss, L.J. (2009) An orchid 90, 755–756. mealybug, Pseudococcus dendrobiorum Gordh, G. and Headrick, D.H. (2001) A Dictio­ Williams (Hemiptera: Pseudococcidae. nary of Entomology. CABI Publishing, Available at www.freshfromflorida.com/pi/pest- Wallingford, UK. alerts/pseudococcus-dendrobiorum.html, Gordon, R.D. (1970) A review of the genus accessed 8 August 2012. Delphastus Caset (Coleoptera: Coccinellidae). Hodges, G.S. and Dixon, W.N. (2007) The Proceedings of the Entomological Society of ­poliaspis cycad scale Poliaspis cycadis Washington 72, 356–369. Comstock (Hemiptera: Diaspididae): A new Hamon, A.B. and Hodges, G.S. (2004) Lobate lac exotic scale insect for Florida. Available at scale, Paratachardina pseudolobata Kondo & www.freshfromflorida.com/pi/enpp/ento/ Gullan (Hemiptera: Kerriidae). Available at poliapsis_cycadis.html. www.freshfromflorida.com/pi/enpp/ento/para- Hodges, G.S. and Dooley, J.W. (2007) A new spe- tachardina.html. cies of Dialeurodes Cockerell (Hemiptera: Hodges, A., Roda, A. and Hodges, G.S. (2007) Aleyrodidae) on Schefflera Forst and Forst in Pink hibiscus mealybug. Available at www. Florida. Insecta Mundi 16, 1–5. plantmanagementnetwork.org/proceedings/ Hodges, G.S. and Hodges, A.C. (2004) New inva- npdn/2007/posters/6.asp, accessed 8 Au­­ sive species of mealybugs, Palmicultor gust 2012. lumpurensis and Chaetococcus bambusae Hodges, G. and Hodgson, C.J. (2010) Phala­ (Hemiptera: Coccoidea: Pseudococcidae), crococcus howertoni, a new genus and on bamboo in Florida. Insecta Mundi 87, ­species of soft scale (Hemiptera: Coccidae) 396–397. from Florida. Florida Entomologist 93, 8–23. Hodgson, C.J. and Martin, J.H. (2001) Three note- Hodges, G.S. (2002) Pink hibiscus mealybug, worthy scale insects (Hemiptera: Coccoidea) Maconellicoccus hirsutus (Green). Available at from Hong Kong and Singapore, including www.freshfromflorida.com/pi/pest-alerts/ Cribropulvinaria tailungensis, new genus and maconellicoccus-hirsutus.html, accessed 8 species (Coccidae), and the status of the August 2012. cycad-feeding Aulacaspis yasumatsui Hodges, G.S. (2004a) A bamboo mealybug (Diaspididae). Raffles Bulletin of Zoology 49, Palmicultor lumpurensis (Takahashi) 227–250. Coccoidea: Pseudococcidae. Available at Hodges, G.S., Hodges, A.C. and Unruh, C.M. (2008) www.freshfromflorida.com/pi/enpp/ento/ A new exotic pest from Florida’s natural t-lumpurensis.html. areas: Crypticerya genistae (Hemiptera: Hodges, G.S. (2004b) Stellate scale, Vinsonia stel- Monophlebidae). Florida Entomologist 91, lifera (Westwood) Coccoidea: Coccidae. 335–337. Available at www.freshfromflorida.com/pi/ Howard, F.W. and Pemberton, R.W. (2003) The enpp/ento/v.stellifera.html. lobate lac scale, a new pest of trees and Hodges, G.S. (2004c) Duplachionaspis diver- shrubs in Florida: implications for the gens (Green) - A new pest of sugarcane Caribbean region. Proceedings of the and other grasses in Florida (Hemiptera: Caribbean Food Crops Society 39, 91–94. 360 I. Stocks

Howard, F.W. and Weissling, T.J. (1999) Questions pest alert. Available at www.ncipmc.org/alerts/­ and answers about cycad aulacaspis scale lobatelacscale/lobate_lac_scale.pdf. insect. Proceedings of the Florida State Martin, J.H. (2004) Whiteflies of Belize (Hemi­ Horticultural Society 112, 243–245. ptera: Aleyrodidae). Part 1- introduction and Howard, F.W., Hamon, A., McLaughlin, M., account of the subfamily Aleurodicinae Weissling, T. and Yang, S. (1999) Aulacaspis Quaintance & Baker. Zootaxa 681, 1–119. yasumatsui (Hemiptera: Sternorrhyncha: Martin, J.H. (2008) A revision of Aleurodicus Diaspididae), a scale insect pest of cycads Douglas (Sternorrhyncha, Aleyrodidae), with recently introduced into Florida. Florida two new genera proposed for palaeotropical Entomologist 82, 14–27. natives and an identification guide to Howard, F.W., Pemberton, R.W., Hodges, G.S., world genera of Aleurodicinae. Zootaxa Steinberg, B., McLean, D. and Liu, H. (2006) Host 1835, pp. 100. plant range of lobate lac scale, Paratachardina Martin, J.H. and Mound, L. (2007) An annotated lobata, in Florida. Proceedings of the Florida check list of the world’s whiteflies (Insecta: State Horticultural Society 119, 398–408. Hemiptera: Aleyrodidae). Zootaxa 1492, 1–84. Howard, F.W, Pemberton, R., Hamon, A., Matile-Ferrero, D. and Étienne, J. (2006) Cochenilles Hodges, G.S., Steinberg, B., Mannion, C.M., des Antilles françaises et de quelques McLean, D. et al. (2009) Lobate lac scale, autres îles Caraïbes (Hemiptera, Coccoidea). Paratachardina lobata lobata (Chamberlin) Revue Française d’Entomologie 28, 161–190. (Hemiptera: Sternorrhyncha: Coccoidea: Meyerdirk, D.E., Khashimuddin, S. and Bashir, M. Kerriidae). Available at http://edis.ifas.ufl.edu/ (1988) Importation, colonization and establi­ in471, accessed 8 August 2012. sh­ment of Anagyrus indicus (Hym: Encyrtidae) Hoy, M., Hamon, A. and Nguyen, R. (2006) Pink on Nipaecoccus viridis (Hom: Pseudo­­coccidae) hibiscus mealybug, Maconellicoccus hirsutus in Jordan. Entomophaga 33, 229–237. (Green) (Insecta: Hemiptera: Pseudococcidae) Meyerdirk, D.E., Warkentin, R., Attavian, B., Available at http://entnemdept.ufl.edu/crea- Gersabeck, E., Francis, A., Adams, M. and tures/orn/mealybug/mealybug.htm, accessed Francis, G. (2001) Biological control of 8 August 2012. pink hibiscus mealybug project manual. Kondo, T. and Gullan, P.J. (2007) Taxonomic review of United States Department of Agriculture, the lac insect genus Paratachardina Balachow­ Animal and Plant Health Inspection Service, sky (Hemiptera: Coccoidea: Kerriidae), with a Plant Protection and Quarantine, Available at revised key to genera of Kerriidae and descrip- www.aphis.usda.gov/import_export/ tion of two new species. Zootaxa 1617, 1–41. plants/­manuals/domestic/downloads/phm.pdf, Lavine, G. (2010) Identifying and controlling the pp. 194. Sago Palm Scale (Aulacaspis yasumatsui). Meyerdirk, D.E., Warkentin, R., Attavian, B., Available at www.agriculture.gov.bb. Gersabeck, A., Francis, A., Adams, M. and Mani, M. (1995) Studies on the natural enemies of Francis, G. (2002) Biological control of oriental mealybug, Planococcus lilacinus (Ckll.) pink hibiscus mealybug project manual. (Homoptera: Pseudococcidae) in India. Journal Available at www.aphis.usda.gov/import_ of Entomological Research 19, 61–70. export/plants/manuals/domestic/downloads/ Mani, M. and Krishnamoorthy, A. (1990a) Outbreak phm.pdf. of mealybugs and record of their natural ene- Meyerdirk, D.E., Muniappan, R., Warkentin, R., mies on pomegranate. Journal of Biological Maba, J. and Reddy, G.V.P. (2004) Biological Control 4, 61–62. control of papaya mealybug, Paracoccus Mani, M. and Krishnamoorthy, A. (1990b) Predation ­marginatus (Hemiptera: Pseudococcidae) in of Mallada boninensis on Ferrisia virgata, Guam. Plant Protection Quarterly 19, 110–114. Planococcus citri and P. lilacinus. Journal of Miller, D.R. and Miller, G.L. (2002) Redescription Biological Control 4, 122–123. of Paracoccus marginatus Williams and Mannion, C. (2010) Gumbo limbo spiraling whitefly, Granara de Willink (Hemiptera: Coccoidea: a new whitefly in south Florida. Available at Pseudococcidae), including descriptions of http://miami-dade.ifas.ufl.edu/documents/ the immature stages and adult male. GumboLimboSpiralingWhitefly.pdf, accessed Proceedings of the Entomological Society of 8 August 2012. Washington 104, 1–23. Mannion, C., Howard, F., Hodges, G. and Hodges, A. Miller, D.R., Williams, D.J. and Hamon, A.B. (1999) (2005) Lobate lac scale, Paratachardina Notes on a new mealybug pest in Florida lobata (Chamberlin). Regional IPM center and the Caribbean: the papaya mealybug, Recent Adventive Scale Insects in Florida and the Caribbean Region 361

Paracoccus marginatus Williams and Granara Roltsch, W. (2007) Insectary rearing of the pink de Willink. Insecta Mundi 13, 179–181. hibiscus mealy bug and its parasitoids in a Miller, D.R., Miller, G.L. and Watson, G.W. (2005a) desert environment. Available at www.cdfa. Invasive species of mealybugs (Hemiptera: ca.gov/phpps/ipc/biocontrol/pdf/insects/ Pseudococcidae) and their threat to U.S. phm_parasite-production.pdf, accessed 8 ­agriculture. Proceedings of the Entomological August 2012. Society of Washington 104, 825–836. Sagarra, L. and Vincent, C. (1999) Influence of Miller, D.R., Miller, G.L, Hodges, G.S and host stage on oviposition, development, sex Davidson, J.A. (2005b) Introduced scale ratio, and survival of Anagyrus kamali Moursi insects (Hemiptera: Coccoidea) of the United (Hymenoptera: Encyrtidae), a parasitoid of Sates and their impact on U.S. agriculture. the hibiscus mealybug, Maconellicoccus hir- Proceedings of the Entomological Society of sutus Green (Homoptera: Pseudococcidae). Washington 107, 123–158. Biological Control 15, 51–56. Muniappan, R., Meyerdirk, D.E., Sengebau, F.M., Smith, T. and Fox, A.J. (2004) Biological control of Berringer, D.D. and Reddy, G.V.P. (2006) giant whitefly, Aleurodicus dugesii Cockerell, Classical biological control of the papaya in Florida. Available at www.freshfromflorida. mealybug, Paracoccus marginatus (Hemiptera: com/pi/methods/giant-whitefly-bc.html, Pseudococcidae) in the republic of Palau. accessed 8 August 2012. Florida Entomologist 89, 212–217. Stocks, I.C. and Hodges, G.S. (2010a) Nipaecoccus Myartseva, S.N. (2006) Siphoninus phillyreae viridis (Newstead), a new exotic mealybug in (Haliday) (Hemiptera: Sternorrhyncha: South Florida (Coccoidea: Pseudococcidae). Aleyrodidae) and its parasitoid, Encarsia Available at www.freshfromflorida.com/pi/ inaron (Walker) (Hymenoptera: Encyrtidae): pest_alerts/pdf/nipaecoccus-viridis-pest- two new records of insects for Mexico. alert.pdf. Entomological News 117, 451–453. Stocks, I.C. and Hodges, G.S. (2010b) Ash Nada, S., Abd Rabo, S. and El Deen Hussein, G. ­whitefly, Siphoninus phillyreae (Haliday), (1990) Scale insects infesting mango trees in a new exotic whitefly (Hemiptera: Aley­ Egypt (Homoptera: Coccoidea). In: Koteja, J. rodidae) in central Florida, and Encar­ (ed.) Proceedings of the Sixth International sia inaron, its parasitoid (Hymenoptera: Symposium of Scale Insects Studies, Part II. Aphelinidae). Available at www.freshfromflorida. Cracow, pp. 133–134. com/pi/pest_alerts/pdf/ash-whitefly-pest- Noyes, J.S. (2003) Universal Chalcidoidea alert.pdf. Database. Available at: www.nhm.ac.uk/ Stocks, I.C. and Hodges, G.S. (2010c) Aleurodicus entomology/chalcidoids/index.html, accessed rugioperculatus Martin, a new exotic whitefly 8 August 2012. in South Florida (Hemiptera: Aleyrodidae). Osborne, L., Mannion, C., Griffiths, K., Hornby, P., Available at www.freshfromflorida.com/pi/ Roda, A. and Meyerdirk, D. (2011) Pink hibiscus pest_alerts/pdf/aleurodicus-rugioperculatus- mealybug (PHM) Maconellicoccus hirsutus pest-alert.pdf. (Green). Available at www.aphis.usda.gov/plant_ Vitullo, J., Zhang, A., Mannion, C. and Bergh, J.C. health/plant_pest_info/phmb/downloads/ (2009) Expression of feeding symptoms project-information.pdf, accessed 8 August 2012. from pink hibiscus mealybug (Hemiptera: Peck, S.B., Cook, J. and Hardy Jr, J.D., (2002) Pseudococcidae) by commercially important Beetle fauna of the island of Tobago, Trinidad cultivars of hibiscus. Florida Entomologist 92, and Tobago, West Indies. Insecta Mundi 16, 248–254. 9–23. Vovides, A.P., Ogata, N., Sosa, V. and Pickett, C.H., Ball, J.C., Casanave, K.C., Peña-García, E. (1997) Pollination of Klonsky, K.M., Jetter, K.M., Bezark, L.G. and ­endangered Cuban cycad Microcycas Schoenig, S.E. (1996) Establishment of the ­calocoma (Miq.) A.DC. Botanical Journal of ash whitefly parasitoid Encarsia inaron the Linnean Society 125, 201–210. (Walker) and its economic benefit to ornamen- Waite, G.K. and Martinez Barrera, R. (2002) Insect tal street trees in California. Biological Control and mite pests. In: Whiley, A.W., Schaffer, B. 6, 260–272. and Wolstenholme, B.N. (eds) The Avocado. Puttarudriah, M. and ChannaBasavanna, G.P. Botany, Production and Uses. CAB (1957) Notes on some predators of mealybugs International, Wallingford, UK. (Coccidae: Hemiptera). Mysore Agricultural Walker, A., Hoy, M. and Meyerdirk, D. Journal 32, 4–19. (2006) Paracoccus marginatus. Available at 362 I. Stocks

http://entnemdept.ufl.edu/creatures/fruit/ Williams, D.J. and Butcher, C.F. (1987) Scale insects mealybugs/papaya_mealybug.htm, accessed (Hemiptera: Coccoidea) of Vanuatu. New 8 August 2012. Zealand Entomologist 9, 88–99. Weissling, T.J. and Howard, F.W. (1999) Aulacaspis Williams, D.J. and Granara de Willink, M.C. (1992) yasumatsui. Available at http://entnemdept. Mealybugs of Central and South America. ufl.edu/creatures/orn/palms/cycad_scale. CAB International, Wallingford, UK, pp. 635. htm. Williams, D.J. and Martin, J.H. (2003) A palm Williams, D.J. (1981) New records of some impor- ­mealybug, Palmicultor palmarum (Ehrhorn) tant mealybugs (Hemiptera: Pseudococcidae). (Hem. Pseudococcidae), now found in the Bulletin of Entomological Research 71, Canary Islands. Entomologist’s Monthly 243–245. Magazine 139, 178.