18 Potential Invasive Species

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1 18 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin Gregory A. Evans 1 and John W. Dooley 2 1 USDA/APHIS/PPQ/National Identification Service, Baltimore Avenue, BARC-West, Bldg. 005, Rm 09A, Beltsville, Maryland 20705, USA; 2 USDA/ APHIS/PPQ 389 Oyster Point Blvd, Suite 2A, South San Francisco, California 94080, USA 18.1 Introduction History has shown that when an exotic pest enters and establishes in a country outside its native range, it often takes only a little time for the to spread to other countries in the region. Therefore, it is mutually beneficial for those working in quarantine and crop protection in all regions of the world to work together to stop, or at least to slow down, the movement of pests. This chapter deals with potential invasive scale insect pests for the USA and the Caribbean Basin. There are approximately 7500 known of scale insects (Coccoidea) belonging to 45 families (extant and fossil); however, the most common, and the ones that are usually intercepted at US ports of entry on plant material, belong to one of the following eight families: Asterolecaniidae, Coccidae, Dactylopiidae, Diaspididae, Monophlebidae (formerly part of Margardodidae), Ortheziidae, Pseudococcidae and Stictococcidae. The number of in these families that are known worldwide is shown in Table In addition, the number of known to occur in the USA and in the following seven zoogeographic regions: Nearctic (NA), Neotropical (NT), Afrotropical (AF), Palearctic (PA), Oriental (OR), Australasian (AU) and Antarctica (AN) is provided, with the percentage of the total number of represented in the USA and the respective zoogeographic regions indicated beside the number of (Table 18.1). These data were extracted from ScaleNet (2011), an online database that includes all published information on scale insect. A few sub might be included as part of the total number of ; in addition, the creators of ScaleNet place the northern part of Mexico in the NA, whereas southern Mexico is in the NT region. Northern mainland China is included in the PA, whereas southern China is in the OR region. This differs from the interception data presented herein, in that all of Mexico is included in the NT region; all of mainland China, Japan and Korea are placed in the Eastern PA region; and the records from the AU region are separated into those from Australia and those from the PI. All scale insects are obligate plant feeders, and therefore are almost always found closely associated with plants, commonly being intercepted on imported plant material. In its native area, an ecological balance usually exists between the population of the scale insect and its natural enemies, which maintains the population of the scale insect below economic injury levels. However, when a is separated from its natural enemies, primarily due to human mediation 320 CAB International Potential Invasive Pests of Agricultural Crops (ed. J. Peña)

2 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 321 Table Number of scale insect of selected families by zoogeographic region, and the percentage of the total number of known (ScaleNet, 2011). Asterolecaniidae Coccidae Diaspididae Pseudococcidae Total % total Total % total Total % total Total % total USA NA NT PA AF OR AU AN < WW Dactylopiidae Monophlebidae Ortheziidae Stictococcidae Total % total Total % total Total % total Total % total USA NA NT PA AF OR AU AN WW NA, Nearctic; NT, Neotropical; AF, Afrotropical; PA, Palearctic; OR, Oriental; AU, Australasian; AN, Antarctica; WW, Worldwide (transported to a new area or by the non-selective use of chemical pesticides), then high infestations of the scale often occur, resulting in economic damage. Therefore, any scale insect that has been separated from its natural enemies and survives the environmental conditions in its new environment, has the potential to become a pest in its non-native habitat. Virtually all of the major scale insect pests in the USA and Caribbean basin are non-native. The introduction of an exotic has often been followed by large population outbreaks and subsequent economic plant damage. The cycad aulacaspis scale, Aulacaspis yasumatsui, was an obscure in South-East Asia until it arrived in Florida in 1996; thereafter, its population exploded and has now spread to many other countries. However, not all invasive become major pest. In fact, Miller and Miller (1990) reported that only about 8% of the known of armored scales had been reported as pests somewhere in the world. Nevertheless, since it is difficult to predict the impact an invasive might have, all invading scale are of interest to plant-protection workers. Regardless of the efforts by plant-protection officers, discovery of an invasive scale often happens after the scale population has become established and population outbreaks have occurred; however, some have been discovered shortly after they entered the country, and measures were taken to eradicate the before it had a chance to become established. Examples of invasive of armored scales that have recently been introduced and have been established in a region include: Aulcaspis yasumatsui Takagi and Poliaspis cycadis Comstock on cycads

3 322 G.A. Evans and J.W. Dooley in Florida, and Parlagena bennetti Williams, a pest of palms that is now known to occur in Trinidad, St Andres Island (Colombia) and the Bay Islands (Honduras). In Table 18.1, the number of potential invasive for a country or region was determined by subtracting the number of known to occur in the country or region from the total number of known worldwide. This number does not take into account the number of cryptic and other that have not been described. For example, currently 329 of armored scales are known to occur in the USA, representing 13.3% of the total number of (2465) known worldwide. Therefore, 2136 (86.7%) are not known to occur in the USA but could potentially invade and establish there and perhaps become economic pests. The potential of an exotic to enter a region and become an economic pest is a function of the characteristics of the, and the probability that it will arrive, gain entrance and establish in the country and/or region. Some of the important characteristics of a scale-insect are: (i) ability to adapt to a new habitat; (ii) reproductive rate; (iii) life cycle (number of generations per annum); (iv) natural enemies (native and exotics); (v) host plants (economic crops and alternate hosts); and (vi) the part(s) of the plant on which it feeds (fruit, leaves, roots). Information on its known plant hosts, distribution and other information can be gathered quickly from ScaleNet (2011). Knowing which scale insect are associated with each imported plant, and their distribution, may provide important clues to plant protection and quarantine workers with respect to possible presence of a scale on the imported hosts and its identity, and information to determine the threat that the poses and the appropriate action that must be taken. Other factors that influence the ability of a scale insect to arrive and gain entrance into the region include: (i) its pest status in the exporting country; (ii) its plant hosts, including imported and alternate hosts; (iii) the frequency and volume of the host that is imported; and (iv) pest frequency on the imported plant material at the ports of entry. The increase in the diversity and volume of agricultural products being shipped throughout the world and their rapid movement on the world market has also increased the movement of these pests and the likelihood of their introduction and establishment in the region. Plant material is being imported into the USA from nearly every country in the world. In addition, there is much greater diversity and a much higher volume of plant being imported than ever before. Exporting countries are bringing new lands into production, which sometimes results in native moving from their usual hosts onto the crop (an alternate host). In addition, the management of these is sometimes complicated by the lack of taxonomic expertise to identity the, and insufficient knowledge about the fauna of many regions of the world. The Animal and Plant Health Inspection Service (APHIS/USDA) and the Department of National Homeland Security, US Customs and Border Protection (NHS/CBP) are responsible for preventing the introduction of exotic into the USA. Currently, there are several US inspection stations distributed throughout the USA and its territories at airports, seaports and land border ports. APHIS has for many years maintained the AQAS (Agricultural Quarantine Activity Systems) database, which is an internal database of records of insects, mites, mollusks and other pests intercepted at US ports of entry (Table 18.2). These data include the pest, host plant, the part of the host plant on which it was found and the country from which the shipment originated. The data are limited, however, because some pests are not identified to the level, usually because the identifiable stage of the was not intercepted, or the condition of the specimen was unsatisfactory for identification. The true origin of the interception is sometimes questionable, especially when a specimen is intercepted in a passenger s baggage, and to a lesser extent in cargo, even though certification of origin is usually provided. When were intercepted on imported plant material, this indicates that they are known to feed on at least one plant, and because they were intercepted at a port of entry this increases the probability that they may gain entrance into the country and region. The number of interceptions of scale insects has varied by zoogeographic region (Table 18.2). In this table, the NA region includes only Canada and the USA; all records from Mexico are included

4 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 323 Table Interceptions of soft scales (Coccidae), armored scales (Diaspididae) and mealybugs (Pseudococcidae) at US ports of entry ( ) by zoogeographic region (USDA/APHIS, 2012). Coccidae Diaspididae Pseudococcidae Total interceptions % Total Total interceptions % Total Total interceptions % Total NA NT 1, , , WPA , , AF EPA OR , , Australian PI , , UNK , , Total 3, , , NA, Nearctic; NT, Neotropical; AF, Afrotropical; OR, Oriental; AU, Australasian; WPA, Western Palearctic; EPA, Eastern Palearctic; PI, Pacific Islands; UNK, Unknown in the NT region. China (entire), Japan and South Korea are in the Eastern PA region, and the records from the AU region are separated into those from Australia and those from the PI. For scales, armored scales and mealybugs, there were many more interceptions made from the NT countries, which accounted for 49.2%, 74.6% and 65.9% of the interceptions of soft scales, armored scales and mealybugs, respectively. Many of the that were intercepted from this region were already present in the Caribbean basin region. The next highest number of interceptions of soft scales was from the PI region which had 28.7% of the total interceptions, with the majority of those from Hawai i. The Western PA had the next highest number of interceptions of armored scales (9.3%); and the OR region had the next highest number of interceptions of mealybugs (14.0%). The ability of an exotic scale insect to gain entry and establish in the USA and Caribbean basin region is also influenced by its host and the part of the host that is imported (Table 18.3). Scale insects, especially armored scales, that have been found on imported fresh fruit are believed to be less likely to survive and establish in the new habitat than those imported on leaves or material for propagation. Possible exceptions must be made for soft scales and mealybugs, since most in these families are mobile in their nymphal and adult stages. The number of interceptions of soft scales, armored scales and mealybugs from the top 12 host plants of each of the three families is shown in Table For soft scales, Gardenia (19.6%) was the most common host, followed by Schlefflera (11.2%). Both of these hosts are ornamental that are imported as leaves and/or propagated ma terial. For armored scales, Citrus (50.4%), Mangifera indica (18.0%), Persea americana (15.9%) and Psidium guajava (1.5%) were the most common hosts. Together they accounted for 85.8% of all interceptions of armored scales. Material of all of these fruit is imported as fruit and the probability of a being found on these hosts gaining entry and establishing is less than for pests on hosts that are imported as leaves. Only 9.2% of the armored scales were found on hosts other than those listed in Table For mealybugs, Musa (22.6%), Ananas comosus (5.9%), Annona spp. (5.7%) and Nephelium (4.6%) were the most common hosts. All of these hosts are imported as fruit. The numbers of different scale insect intercepted at US ports of entry between 1 January 2007 and 31 December 2010 are categorized by their relative distribution in the Caribbean basin region as being either: (i) widespread; (ii) limited; or (iii) not known to occur in the Caribbean basin region (Table 18.4). If the is known to occur in the USA, Mexico and at least one of the countries of Central America and/or a Caribbean island, it is designated as being widespread; if the

5 324 G.A. Evans and J.W. Dooley Table Interceptions (Total int.) of soft scales (Coccidae), armored scales (Diaspididae) and mealybugs (Pseudococcidae) at US ports of entry ( ) by their top 12 imported hosts (USDA/APHIS, 2012). Coccidae Diaspididae Pseudococcidae Host genus Total int. % Total Host genus Total int. % Total Host genus Total int. % Total Gardenia (L) Citrus (F) 20, Musa (F) 5, Schlefflera (L) Mangifera (F) 7, Ananas (F) 1, Citrus (F) Persea (F) 6, Annona (F) 1, Nephelium (F) Psidium (F) Nephelium (F) 1, Mangifera (F) Chamaedorea (L) Ocimum (L) Alyxia (L) Musa (F) Garcinia (F) Laurus (L) Areca (L) Citrus (F) Psidium (F) Annona (F) Alpinia (L) Annona (F) Cocos (F) Psidium (F) Alpinia (L) Draecaena (L) Salvia (L) Carica (F) Dimocarpus (F) Carica (F) Musa (F) Murraya (L) Sechium(F) Other 1, Other 3, Other 10, Total 3, , , F, imported fresh fruit; L, leaves has not been known to occur in at least one of the countries of these four areas, then it is designated as being limited in distribution. For armored scales (Diaspididae), 71 (50.7%) of the 140 that have been intercepted are widespread in the Caribbean Basin. Since they are already present in all or many of the countries in the region, they are not considered as potential invasive, and therefore are not included in the discussion of each family. Of the remaining that have been intercepted, 41 (29.3%) are limited in distribution and 28 (20%) are not known to occur in the region. These are considered to be potential invasive for the region or for other countries within the region. The number of interceptions of different scale insect at US ports of entry between 2007 and 2010 has been categorized by that are widespread, limited or not known to occur in the Caribbean basin region (Table 18.5). For example, for armored scales (Diaspididae), 20,063 (50.2%) of all interceptions (40,000) were of that are widely distributed in the Caribbean basin region, 16,048 (40.1%) of the interceptions were of that occur in the region but are limited in their distribution, and 190 (0.5%) were interceptions of that are not known to occur in the region. Specimens identified at the family or genus level represented 3699 (9.2%) of the interceptions. Although the 28 intercepted that are not known to occur in the region only represented 190 (0.5%) of all interceptions, their potential impact should not be underestimated. The entry and establishment of any of these in the region could have major repercussions (e.g., the introduction and establishment of Aulacaspis yasumatsui). The number of interception records of scaleinsect that are not known to occur in the Caribbean basin or have limited distribution in the region, their region of origin, their distribution in the Caribbean basin region, if the is known to occur in the USA, the country and host plant on which the was intercepted and other notes are shown in Table Since the focus of this chapter is on the potential invasive scale-insect to the Caribbean basin region, we have excluded those that are widely distributed in the region. Species of the eight families of scale insects are discussed separately by family. The of scale insects that have been intercepted at US ports between 2007 and 2010 are discussed below. An asterisk (*) indicates those that are of particular interest as potential pest.

6 Table Number of (# sp.) and percentage of total number of scale insect (% sp.) intercepted at US ports of entry ( ) by family and their distribution in the Caribbean basin region (USDA/APHIS, 2012). Distribution in Caribbean basin Widespread in region Limited in region Not known in region Total # intercepted Asterolecanidaeae Coccidae Diaspididae Pseudococcidae Dactylopidae Monophlebidae Ortheziidae Strictococcidae # sp./(%) sp. # sp./(%) sp. # sp./(%) sp. # sp./(%) sp. # sp./(%) sp. # sp./(%) sp. # sp./(%) sp. # sp./(%) sp. 4 50% 2 25% 2 25% % % % % % % % % % % % 0 0% % % % % % % 4 0 0% 0 0% 3 3 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 325

7 Table Interceptions of different scale insect (# int.) at US ports of entry ( ), and the percentage of total number of (% sp.) intercepted by distribution in the Caribbean basin region (USDA/APHIS, 2012). Distribution in Caribbean basin Widespread in region Limited in region Not known in region Unidentified to genus or Total # intercepted Asterolecanidae Coccidae Diaspididae Pseudococcidae Dactylopidae Monophlebidae Ortheziidae Strictococcidae # int./ (% int.) # int./ (% int.) # int./ (% int.) # int./ (% int.) # int./ (% int.) # int./ (% int.) # int./ (% int.) # int./ (% int.) 4 4% 52 52% 6 6% 38 38% 100 1, % % % 1, % 3,273 20, % 16, % % 3, % 40,000 6, % 1, % % 15, % 24, % 0 0% 0 0% % % % 5 1.3% % % % 1 0.5% % % 0 0% % % G.A. Evans and J.W. Dooley

8 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 327 Table Interceptions of scale insects of limited distribution or not known to occur in Caribbean basin (USDA/APHIS, 2012). Species Total int. of orig. of dist. In USA Interceptions and notes Asterolecaniidae (pit scales) Asterolecanium 2 OR N no Thailand on Durio ungulatum Palmaspis inlabefactum 50 NT L no Mexico on Chamaedorea *Palmaspis phoenicis 4 WPA N no India, Jordan, Pakistan, Turkey on Phoenix Total = Coccidae (soft scales) *Ceroplastes japonicus 7 EPA N no Armenia, France, Georgia, Germany, Russia on Laurus, Prunus; pest of kiwi and persimmon Coccus moestus 4 AF L no Ecuador, Honduras, Puerto Rico on Nephelium, Prunus and palms Drepanococcus chiton 16 OR N no Asia on Dimocarpus, Garcinia, Lansium, Nephelium Kilifia deltoides 1 AF N no India on Mangifera *Lichtensia viburni 1 WPA N no Ireland on Hedera *Philephedra broadwayi 1 NT L no Grenada on Annona; also in South America and Caribbean on Mangifera, Spondias, Theobroma Protopulvinaria longivalvata *Taiwansaissetia formicarii 7 OR L no Costa Rica, Hawai i, Jamaica, Puerto Rico on Cinnamomum, Morinda, Piper, Plumeria, Schefflera 8 OR N no India, Thailand, Vietnam on Areca, Durio, Lansium, Mangifera; also on Garcinia, Lagerstroemia, Persea Udinia catori 8 AF N no Cameroon, Gambia, Nigeria, Senegal on Cola, Mangifera; also on Citrus, Ficus, Persea, Psidium Udinia farquharsoni 1 AF N no Kenya on Mangifera; also on Citrus, Coffea, Durio, Ficus, Gardenia, Theobroma Total = 10 3,273 Diaspididae (armored scales) Abgrallaspis aguacatae 4,691 NT L no Mexico on Persea Abgrallaspis mendax 10 NT L no Guatemala, Mexico on orchids Acutaspis scutiformis 15 NT L yes Costa Rica on Aglaonema Acutaspis umbonifera 15 NT L yes Mexico, Central and South America; various plants Aonidiella comperei 16 OR L no Brazil, India, Korea, Sri Lanka; in Central America and Caribbean on Carica, Citrus Aspidiella hartii 102 NT L no Africa, Barbados, Haiti, Thailand, Vietnam on roots of Dioscorea and Zingiber; pest of yams Aspidiotus capensis 1 AF N no South Africa on Encephalartos, cycads *Aulacaspis crawii 2 EPA L no China on Citrus, Cymbidium; also on Hibiscus, Murraya, Poncirus, orchids; Asia, El Salvador Continued

9 328 G.A. Evans and J.W. Dooley Table Continued. Species Total int. of orig. of dist. In USA Interceptions and notes Aulacaspis ima 1 EPA N no India on Cinnamomum Aulacaspis rosarum 1 EPA L no Mexico on Rosa; also in EPA, OR and PI region Aulacaspis tubercularis 6,245 OR L yes Widespread; many hosts but most common on Mangifera (mango); USA (Florida) *Chrysomphalus pinnulifer 13 OR L no Australia, Asia, Italy, Colombia, Dominican Republic on Citrus, Garcinia, Mangifera, Persea, pest of tea 5 OR N no Philippines, Vietnam on Annona, Cocos, Garcinia 3 AF L no Peru, Virgin Islands on Cymbopogon; also in Brazil Chrysomphalus propsimus Duplachionaspis natalensis Fiorinia coronata 2 PI N no Hawai i, Vietnam on Chrysalidocarpus, Cocos, palms *Fiorinia proboscidaria 7 OR N no China, Hawai i, Indonesia, Vietnam on Citrus, roses, palms Greenaspis sp. 1 Asia L no Dominican Republic, Grenada, Puerto Rico on Cymbopogon Leonardianna sp. 58 NT L no Dominican Republic,Trinidad on Canella, Guatteria Lepidosaphes cornuta 3 OR N no Singapore, Thailand on Michelia, Piper *Lepidosaphes laterochitinosa *Lepidosaphes malicola *Lepidosaphes pistaciae Lepidosaphes rubrovittata 679 WPA L no Asia, Australia, Costa Rica on many hosts including Citrus, Mangifera and ornamentals 4 WPA L no Dominican Republic, Italy on Malus, Prunus, Pyrus 68 WPA N no India, Egypt, Iran, Jordan, Syria, Tunisia, Turkey on pistacios; also on Malus, Prunus, Pyrus and others 7 OR L no Asia, Jamaica, Puerto Rico, South Africa on cycads, Psidium, Murraya, Nepenthes and others Lepidosaphes similis 15 PI N no Asia, PI, Brazil on Areca, Cocos, Dracaena, Musa, palms *Lepidosaphes 1 OR N no Asia, Guam; ornamental pest tubulorum *Leucapis riccae 1 WPA N no Europe on Olea Lindingaspis misrae 6 OR N no India on Cycas, Mangifera, Murraya, Tamarindus Pseudoparlatoria mammata 30 NT L no Mexico on Chamaedorea, palms; also in Panama Melanaspis calura 5 NT L no Costa Rica, Ecuador, Guatemala, Honduras, Mexico on Spondias; also on Crataegus, Mammea Melanaspis inopinata 26 WPA N no Middle East on pistacios, Dioscorea; Malus, Prunus Parlatoreopsis chinensis 1 EPA N no Taiwan on Acer; also Hibicus, Malus, Prunus, Pyrus Parlatoria citri 6 OR N no China, Indonesia, Vietnam on Citrus, Pyrus *Parlatoria crypta 17 OR N no India, Pakistan on Mangifera, Murraya Continued

10 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 329 Table Continued. Species Total int. of orig. of dist. In USA Interceptions and notes Parlatoria fluggeae 1 AF N no Germany on Adenia; also on Ficus, Hibiscus; Hawai i Parlatoria multipora 1 EPA N no South Korea on Cornus *Parlatoria pseudaspidiotus 101 OR L no India, Mexico on Mangifera; Caribbean on orchids *Parlatoria ziziphi 9,864 WPA L no widespread on Citrus; eradicated from Florida, USA *Pinnaspis theae 1 OR L no Brazil on Thea; Colombia, Guadeloupe, Martinique Pseudaonidia corbetti 1 OR N no China on Garcinia, ornamentals Pseudaonidia curculiginis Pseudaulacapis brimblecombei Pseudaulacaspis eugeniae 1 OR N no Philippines on Pholidota; also on Gardenia, palms; Malaysia, Thailand, Vietnam 2 AUS N no Australia, New Zealand on Macademia, Proteaceae 8 AUS N no Australia on Proteaceae (ornamentals); also reported on Prunus and ornamentals from Asia Pseudaulcaspis barberi 5 OR N no India on Mangifera Rolaspis leucadendri 2 AF N no South Africa on Leucadendron, Proteaceae *Salicicola archangelskyae 1 WPA N no Central Asia on Malus, Pyrus; pest in Central Asia Selenaspidopsis browni 2 NT L no Mexico on Chamaedorea (palms) Silvestraspis uberifera 3 OR N no Vietnam on Cinnamomum, Machilus, Pimenta Thysanofiorinia leei 1 OR N no Hawai i, Korea, India, Taiwan on Dimocarpus, Nephelium, Litchi *Unaspis yanonensis 192 EPA L no Asia, Australia, China, Greece, Hawai i, Japan; citrus pest, Araceae, Alyxis, Citrullus, Fortunella Total = 46 40,000 Dactylopiidae Dactylopius coccus 4 NT L no on Opuntia from Mexico Total = Monophlebidae (giant scales) *Crypticerya genistae 14 NT L yes Grenada, Puerto Rico, Trinidad, Virgin Is. on Cajanus, Caesalpina, Origanum, Spondias; in Florida, USA *Drosicha sp. 3 OR N no India, Pakistan, Philippines; on Annona, Psidium *Icerya seychellarum 9 OR L no Cook Islands, Philippines, Taiwan on Alyxia, Lycium, Piper, Rosa; reported from Guadeloupe Total = Ortheziidae (ensign scales) Arctorthezia cataphracta 1? N yes UK, on Cantharellus; in Alaska, Canada Insignorthezia pseudoinsignis 4 NA W yes El Salvador, Guatemala, Mexico; on Citrus, Tillandsia, Thymus; in Louisiana and Texas, USA Continued

11 330 G.A. Evans and J.W. Dooley Table Continued. Species Total int. of orig. of dist. In USA Interceptions and notes *Praelongorthezia praelonga 13 NT L no Dominica, Jamaica, Puerto Rico, Virgin Islands; on Cajanus, Croton, Kalanchoe, Mentha, Rosmarinus Total = Pseudococcidae (mealybugs) *Crisicoccus 3 EPA N no Japan, South Korea on Pyrus matsumotoi Delottococcus proteae 3 AF N no South Africa on Brunia, Leucadendron, Protea Dysmicoccus 1 AU N no Australia on Leucospermum hypogaeus *Dysmicoccus lepelleyi 26 OR N no China, Indonesia, Taiwan, Thailand, Vietnam on Garcinia, Lansium, Musa, Nephelium, Punica Dysmicoccus 1 AU N no Australia on Allocasuarina queenslandianus Exallomochlus camur 2 OR N no Philippines on Annona, Lansium *Exallomochlus hispidus 57 OR N no Asia, on Dimocarpus, Garcinia, Lansium, Murraya, Nephelium Exallomochlus philippinensis 5 OR N no Philippines on Dimocarpus, Durio, Lansium Ferrisia terani 3 NT L no Guatemala, Mexico on Rosmarinus, Citrus Formicococcus robustus 7 OR N no India on Mangifera; also on Citrus, Ficus, Vitis Hordeolicoccus 4 OR N no Thailand, Vietnam on Nephelium heterotrichus Hordeolicoccus nephelii 1 OR N no Vietnam on Garcinia; also on Artocarpus, Nephelium Laminicoccus pandani 1 PI N no Hawai i on Pandanus *Maconellicoccus multipori 11 OR N no Philippines, Singapore, Vietnam on Garcinia, Lansium, Nephelium Nipaecoccus annonae 1 NT L no Puerto Rico on Annona; also Cocos, Persea, Psidium Nipaecoccus jonmartini 3 NT L no Costa Rica, Puerto Rico on Calathea, Musa, Nephelium *Nipaecoccus viridis 11 OR L yes widespread; in Costa Rica; on many hosts; recently found in Florida, USA Palmicultor browni 7 PI L no Australia, Bahamas, Colombia, Puerto Rico on Cocos Palmicultor palmarum 23 PI L no Widespread on palms; in Mexico and Bahamas *Paracoccus burnerae 5 AF N no China, India, South Africa on Citrus, Nephelium, Pyrus Paracoccus ferrisi 12 NT L no El Salvador, Mexico on Citrus, Eriobotrya, Fernaldia, Ocimum, Origanum, Rosmarinus Paracoccus herreni 2 NT L no El Salvador, Mexico on Annona, Fernaldia *Paracoccus interceptus 81 OR N no widespread in Asia on Dimocarpus, Garcinia, Lansium, Litchi, Nephelium Paracoccus mexicanus 9 NT L no Mexico on Ocimum, Rosmarinus Continued

12 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 331 Table Continued. Species Total int. of orig. of dist. In USA Interceptions and notes Paracoccus solani 19 NT L no Mexico on Agave, Ananas, Leucaena, Mentha, Salvia Paraputo odontomachi 4 OR N no Cambodia, Vietnam on Garcinia, Nephelium *Phenacoccus franseriae 254 NT L no Mexico, common on Salvia; also intercepted on Ocimum, Origanum, Rosmarinus, Thymus (herbs) 1 WPA L yes New Zealand on Feijoa; in South Africa and California, USA Phenacoccus graminicola Phenacoccus hakeae 8 AU N no Australia on Banksia, Leptospermum, Persoonia Phenacoccus stelli 6 AF N no South Africa on Brunia, Leucadendron, Symphoricarpos Planococcus halli 75 NT L no AF common on Dioscorea; widespread in NT *Planococcus kenyae 1 AF N no Kenya on Musa; pest of coffee *Planococcus lilacinus 88 OR L no Asia on Annona, Dimocarpus, Garcinia, Lansium, Litchi, Nephelium; also intercepted Costa Rica, Peru Planococcus litchi 5 OR N no China, Philippines, Thailand on Garcinia, Litchi, Nephelium Pseudococcus 11 OR N no Asia on Garcinia, Nephelium aurantiacus *Pseudococcus baliteus 165 OR N no Asia on Garcinia, Lansium, Nephelium, Litchi *Pseudococcus cryptus 47 NT L no widespread on Garcinia, Lansium, Litchi, Nephelium Pseudococcus 3 AU N no Taiwan, Thailand on orchids dendrobiorum Pseudococcus elisae 1,219 NT L no NT, common on Musa Pseudococcus 1 AU N no Australia on Chamelaucium eucalypticus Pseudococcus landoi 21 NT L no Costa Rica, Haiti, Honduras, Mexico, South America on Citrus, Musa, Nephelium, Sechium Pseudococcus zamiae 1 AU N no Australia on Macrozamia Puto barberi 22 NT L no NT on Cajanus, Hibiscus, Origanum Rastrococcus 1 OR N no Vietnam on Dimocarpus expeditionis Rastrococcus iceryoides 6 OR N no India, Thailand on Ficus, Murraya, Psidium *Rastrococcus 1 OR N no Philippines on Psidium invadens Rastrococcus spinosus 1 OR N no Vietnam on Artocarpus Rastrococcus tropicasiaticus 3 OR N no India, Philippines, Vietnam on Murraya, Nephelium Rhizoecus amorphophalli 4 OR L no Thailand on roots of Curcuma, Zingiber, Typhonium; in Guadeloupe Spinococcus convolvuli 1 WPA N no Israel on Salvia Vryburgia distincta 1 AF N no AF on Echeveria Vryburgia viator 1 AF N no AF on Leucospermum Total interceptions = 52 24,933 Continued

13 332 G.A. Evans and J.W. Dooley Table Continued. Species Total int. of orig. of dist. In USA Interceptions and notes Stictococcidae Parastictococcus multispinosus Stictococcus intermedius 2 AF N no Nigeria on Dennettia 6 AF N no Cameroon, Nigeria on Annonaceae, Cola, Persea *Stictococcus sjostedti 17 Cameroon, Nigeria on Annonaceae, Cola, Persea, Piper, Uvaria Stictococcus sp. 7 AF N no Nigeria on Annonaceae, Bactris, Dennetiia, Phaseolus, Solanum Total interceptions = 3 32 Total int., total intercepted; of orig., region of origination; of dist., region of distribution; NA, Nearctic; NT, Neotropical; AF, Afrotropical; OR, Oriental; AU, Australasian; WPA, Western Palearctic; EPA, Eastern Palearctic; PI, Pacific Islands; W, widespread; L, limited; N, not known to occur in region; *Species of particular interest as potential invasive based on their pest status, known hosts and distribution, and the number of times they have been intercepted Family Asterolecaniidae (Pit Scales) Asterolecaniids are a relatively small family of scale insects with 240 known worldwide; of these, 19 (7.9%) and 51 (21.3%) are known to occur in the USA and the NT region, respectively. Russell (1941) reported that in this family are destructive and some are potential enemies of economic plants, but their impact is probably underestimated because of their inconspicuousness. Asterolecanium coffeae Newstead was reported to cause heavy damage to coffee in Kenya. Of the eight of asterolecaniids that have been intercepted, four are widespread in the region and represent 61.5% of the interceptions. Two of limited distribution in the region have been intercepted; Palmaspis inlabefactum (Russell) was intercepted on Cham aedorea palms from Mexico and is only known to occur in Mexico, and Planchonia stentae (Brain) was intercepted on Ocimum from Antigua and is known to occur in the USA (California and Florida), Puerto Rico and Martinique, and on a wide variety of hosts. Two are not known to occur in the region. A. ungulatum Russell was intercepted on durian Durio zibethanus from Thailand, and Palmaspis phoenicis (Ramachandra Rao) on Phoenix sp. palms from Pakistan and the Middle East. All of these are tropical or subtropical that could become established in the southern USA, Mexico, Central America and Caribbean islands. Palmaspis phoenicis is reported to be a pest of date palms (Phoenix dactylifera) in Israel, Iraq, Egypt and elsewhere in North Africa (Howard, 2001) Family Coccidae (Soft Scales) Soft scales are the third largest family of scale insects, with 1131 known worldwide, of which 106 (9.4%) are known to occur in the USA. Many are known to attack agricultural, ornamental and greenhouse crops throughout the world. They damage plants by their direct feeding that removes the plant sap and also by the secretion of large amounts of honeydew. This serves as a medium for sooty mold fungus, which not only inhibits phytosynthesis but also greatly reduces the aesthetic qualities of the plants. Most of the soft scales of economic importance in the USA are Ceroplastes, Coccus and Saissetia (Hamon and Williams, 1984). Of the 29 of soft scales that have been intercepted, 24 (61.5%), representing 60.6% of all interceptions of soft scales, are widely distributed in the region. Seven (17.9%) are limited in distribution and represent 0.6% of the interceptions. Of these, Coccus capparidis, Eulecanium tiliae, Kilifia americana and

14 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin 333 Parthenolecanium corni are known to occur in the USA. The following three are limited in distribution in the region, and are not known to occur in the USA or to be pest. Coccus moestus, an AF that is present in Central America and the Caribbean region, was intercepted on Nephelium spp., Prunus spp. and palms; *Philephedra broadwayi, an NT, was intercepted from Grenada on an Annona sp. and occurs in other Caribbean countries on such economic hosts as Mangifera indica, Spondias spp. and Theobroma cacao. Protopulvinaria longivalvata, an OR that is known to occur in Central America and on some of the Caribbean Islands, was intercepted on Cinnamomum, Morinda, Piper, Plumeria and Schefflera. The following seven (20.5%) are not known to occur in the region and represent 20.5% of the interceptions: Ceroplastes japonicus, Drepanococcus chiton, Kilifia deltoides, Lichtensia viburni, Taiwansaissetia formicarii, Udinia catori and Udinia farquharsoni. Of these, Miller and Miller (2003) listed Ceroplastes japonicus as being a major pest or threat to the USA. This was described from Japan but occurs throughout much of Europe on a wide range of ornamental and fruit and is reported to be a pest of kiwi fruit (Actinidia deliciosa) and persimmons (Diospyros kaki). It is Holarctic (i.e. Nearctic and Paleactic) in distribution, and may not be well adapted to the Caribbean basin region, but poses a threat to the central and northern parts of the USA. Miller and Miller (2003) list Drepanococcus chiton, Lichtensia viburni and Taiwansaissetia formicarii as potential threats to the USA. The first is an Asian known to attack several tropical fruit of economic importance that are grown in the Caribbean basin region; Lichtensia viburni is a European that is a pest of ornamentals but also attacks Prunus; and Taiwansaissetia formicarii is an Asian that attacks several ornamental and fruit that could become an important pest in the Caribbean basin region. Udinia catori and U. farquharsoni are AF that attack several ornamental and fruit and could become important pest in the Caribbean basin region Family Diaspididae (Armored Scales) Armored scales (Hemiptera: Diaspididae) comprise the largest and most diverse group of scale insects and are found on terrestrial plants on every continent (including AN). There are 2465 known worldwide; of these, 329 (13.3%) are known to occur in the USA and 365 (14.8%) in the NT region. They include many economically important pest of forest, fruit and ornamental crops throughout the world, and are among the most common found on imported plant products. A review of the armored intercepted at US ports of entry between 2007 and 2010 (Table 18.3) showed that 71 (50.7%) of the identified were widely distributed in the Caribbean basin region and comprised 50.2% of all interceptions of armored scales; 41 (29.3%) were limited in distribution and comprised 40.1% of the interceptions; 28 (20.0%) were not known to occur in the region and comprised 0.5% of the interceptions; unidentified genera and accounted for 9.2% of the interceptions. Three, Parlatoria ziziphi, Aulcaspis tubercularis and Abgrallaspis aguacatae were exceptionally common, accounting for 20,800 (52%) of the total interceptions. The are discussed by their zoogeographic region of origin Neotropical (8) 1. Abgrallaspis aguacatae Evans, Watson and Miller was recently discovered on avocados imported from Mexico (Evans et al., 2009). Currently, it is only known to occur on avocados in Mexico. The impact that this scale would have on agriculture in the USA and elsewhere in the region is unknown, but it does not appear to cause significant damage to avocados in Mexico. 2. Abgrallaspis mendax (McKenzie) was described on orchids from Guatemala and was intercepted on orchids from Mexico. 3. Acutaspis scutiformis (Cockerell) was described on avocados from Mexico. In the Caribbean basin region it is known to occur in Texas (USA), Mexico and Guatemala, but has not been reported from the Caribbean islands. It was intercepted on plants of Aglaonema exported from Costa Rica for propagation. 4. Acutaspis umbonifera (Newstead) was described on Lecythis from Guyana and is reported from other South American countries and New York (USA) on Anthurium, Attalea, Pereskia and

15 334 G.A. Evans and J.W. Dooley Heliconia. It has been intercepted on various plants for propagation from Mexico and Central and South America. 5. Aspidiella hartii (Cockerell), the yam scale, was described from Trinidad and occurs throughout much of the Caribbean basin region. It is reported as a pest of yams (Dioscorea), especially in storage, and also occurs on Curcuma (tumeric) and Zingiber (ginger) roots. It is a phloem feeder; when large populations are present, yellowing, defoliation and reduction in fruit set and loss in plant vigour may occur. Stored yams encrusted with scales become dry and fibrous. It was intercepted on Zingiber and Curcuma from Asia, on Dioscorea from Africa and Haiti and on Zingiber from Barbados. 6. Hemiberlesia gliwicensis (Komosinka) was described on Billbergia from Poland. It was intercepted on Tillandsia from Brazil. 7. Pseudoparlatoria mammata (Ferris) was intercepted on Chamaedorea palms from Mexico. It is not reported as a pest and its potential impact on agricultural crops in other countries in the region is unknown. 8. Selenaspidopsis browni Nakahara was intercepted on Chamaedorea palms from Mexico. It is not reported as a pest and its potential impact on agricultural crops in other countries in the region is unknown Afrotropical (4) 1. Aspidiotus capensis Newstead was described from South Africa on cycads and is only known to attack cycads. It was intercepted Encephalartos sp. from South Africa in 2006 and Although it has not been reported as a pest, the absence of known natural enemies indicates that it has the potential to become a pest of cycads outside its native region. 2. Duplachionaspis natalensis (Maskell) is an AF that was intercepted on lemongrass (Cymbopogon) from the Caribbean basin region, and may pose a threat to other grass in the region. 3. Parlatoria fluggeae Hall was described on Fluggea from Zimbabwe. It occurs in Algeria, Hawai i, Morocco and Taiwan. Its reported hosts include Ficus and Hibiscus, and it was intercepted on Adenia from Germany. 4. Rolaspis leucadendri (Brain) was described from South Africa and is only known to attack Proteaceae. It was intercepted on Berzelia and Leucadendron (Proteaceae) from South Africa Western Palearctic (5) 1. Lepidosaphes malicola Borchsenius was described from Armenia, and is also reported in other European and Middle Eastern countries, China and India. It was intercepted on apples from Denmark and Italy and on Prunus from the Dominican Republic. It should be considered as a potential economic pest since it attacks apples, pears, roses and other hosts, and Miller and Davidson (1990) list it as a pest. 2. Lepidosaphes pistaciae Archangelskaia was described from Turkmenistan on pistachios (Pistacia vera), and is present in other Central Asian countries, the Middle East and Pakistan. It was intercepted on pistachios from India, Egypt, Iran, Jordan, Kuwait, Lebanon, Syria, Tunisia and Turkey. It is not reported as a pest, but hosts include important plant such as Malus, Prunus, Pyrus, pistachios and azaleas. 3. Leucaspis riccae (Targioni Tozzetti) was described from Italy and reported in other European and Central Asian countries and in Argentina. Its hosts include ornamentals (Nerium, Erica, Ephedra), figs (Ficus) and olives (Olea). It was intercepted on olives from Italy. Miller and Davidson (1990) list it as a pest. 4. Melanaspis inopinata (Leonardi) was described from Italy on pears and mandorlo and is also reported from other European countries and Pakistan. Its hosts include Malus, Pistacia, Prunus and ornamentals. It was intercepted on pistachios from Iran, Jordan, Lebanon and Syria, and on Dioscorea from Nigeria. It is a pest of fruit trees, mainly of Rosaceae, in southern Europe, the Middle East and Armenia, and therefore should be considered a potential threat to the region. 5. Salicicola archangelskyae (Lindinger) was described on pears (Pyrus communis) from Uzbekistan. Its host also include Cydonia, Malus, Punica, Olea and ornamental. It was intercepted on a plant, and The Netherlands was listed as the origin.

16 Potential Invasive Species of Scale Insects for the USA and Caribbean Basin Eastern Palearctic (6) 1. Aulacaspis crawii (Cockerell) is an Asian, known to attack citrus, mango, orchids and other ornamentals. It was intercepted on mango fruit in baggage from El Salvador in 2001; and on Citrus fruit, and leaves of Aglaia, Buxus, Cymbidium, Pelea and Syringa from Asia and Hawai i. It is not reported as a pest, but should be considered as a potential threat since it attacks several economic hosts. 2. Aulacaspis ima Scott was described from China on Lindera and intercepted on Cinnamomum from India in Aulacaspis rosarum Borchsenius is an Asian also known to occur in the PI. Its hosts include Rosa, Rubus and Ficus. It was intercepted on roses from Mexico in 1986 and Parlatoreopsis chinensis (Marlatt) was described on crab apple (Malus) from China. It is also known to occur in the USA (California, Florida, Missouri), Iran and Japan; its hosts also include Hibiscus, Prunus and Pyrus. It was intercepted on Acer from Taiwan. 5. Parlatoria multipora McKenzie was described from China, and intercepted on Cornus from South Korea. 6. Unaspis yanonensis (Kuwana), the arrowhead scale, was described from Japan where it is a major of pest of Citrus. It is of economic importance in China. Heavy infestations of the scale cause dieback of terminal branches, and if severe, may kill the tree. Old World interceptions of the scale include Citrus (fruit) from Asia, Australia, Greece, Hawai i; Araceae (leaves) from Australia; Alyxia (leaves) from the Cook Islands; Citrullus (fruit) from Japan; and Fortunella (fruit) from China and Japan. New World interceptions include: Citrus (fruit) from Panama in 2001; Mangifera indica (fruit) from Nicaragua in 1992; and Pouteria (fruit) from Mexico in Since all of the New World interceptions were taken from baggage, it is uncertain whether U. yanonensis is established in the New World, and must be confirmed by collections made on plants in these countries or that are known to have originated there Oriental (23) 1. Aonidiella comperei McKenzie is an Asian, known to occur in the Caribbean basin region and Central America on Carica papaya, Citrus, Cocos nucifera and other economic crop, but is not reported to be a pest. 2. Aulcaspis tubercularis Newstead, the white mango scale, was described from Java on Cinnamomum. It is intercepted primarily on mango and occurs throughout the region. A mode rate infestation of this was discovered in Miami, Florida, in 2002, but it has not been reported in Mexico or Central America. It injures leaves and fruits and is reported to cause significant damage to mangoes in South Africa (Colyn and Schaffer, 1993; Watson, 2002). 3. Chrysomphalus pinnulifer (Maskell) is an Asian reported as a pest of tea in Kenya and India, and a potential invasive pest of Citrus. Among its economic hosts are Annona spp., Citrus, Cocos nucifera, Cycas spp., Garcinia mangostana, Mangifera indica, Musa, Prunus spp. and Vitis vinifera. It is widespread in the Old World and was intercepted on Citrus from the Dominican Republic in 2007 and on Persea americana from Guatemala in It should be considered as a potential threat since it is reported as reaching pest status and has many economic hosts. 4. Chrysomphalus propsimus Banks is an Asian. It is also reported from Hawai i and usually is found on leaves of palms and Pandanus. It was intercepted on Cocos nucifera and Garcinia mangostana from Vietnam and on Annona spp. from the Philippines. 5. Fiorinia coronata Williams and Watson was described from palms from the Solomon Islands, and intercepted on Cocos nucifera from Vietnam and on palms from Hawai i. It is not reported as a pest. 6. Fiorinia proboscidaria Green was described from Sri Lanka, and is also reported in Hawai i and other PI and Asia. It was intercepted on Citrus from Indonesia and Vietnam and on Areca palms from Hawai i. Its hosts include Mangifera indica, Citrus, roses, palms and other. It is not reported as a pest, but its hosts include several economically important. 7. Greenaspis sp. This is in an Asian genus. It was intercepted on lemongrass (Cymbopogon) from the Caribbean basin region, and may pose a threat to other grass in the region. 8. Lepidosaphes cornuta Ramakrishna Ayyar was described from India on Piper nigrum. It was intercepted on Michelia from Thailand, and is not reported as a pest.

17 336 G.A. Evans and J.W. Dooley 9. Lepidosaphes laterochitinosa was described from a garden in the UK, but is probably native to Asia. It has been intercepted from Asia, Australia and Costa Rica on many hosts including Citrus, Mangifera indica, orchids, palms and various ornamental. It has not been reported as an economic pest, but since its hosts include many economic plants and very little is known of its natural enemies, it could pose a threat to agriculture in the region. 10. Lepidosaphes rubrovittata Cockerell is an Asian ; it is known to attack Anthurium, palms, cacti, cycads, Ficus, Heliconia and other ornamental. It is known to occur in Puerto Rico, and was intercepted on Alpinia from Costa Rica. 11. Lepidosaphes similis Beardsley was described from Micronesia, and is present in other PI. Its hosts include Areca, Coco nucifera and other palms and Musa. It was intercepted on palms from Asia and the PI, and on Musa from the Philippines. 12. Lepidosaphes tubulorum Ferris was described on Sapium from Taiwan. Its hosts include Malus, Prunus spp., Vitis vinifera and various ornamental. Miller and Davidson (1990) list it as a pest. 13. Lindingaspis misrae (Laing) was described on Tamarindus from India. Its reported hosts also include Capparis and cycads. It was intercepted on Azadirachta, Mangifera indica and Murraya from India and is not reported to be a pest. 14. Parlatoria citri McKenzie was described from Indonesia and is also reported in Nigeria, India and Thailand. It was intercepted on Citrus from Indonesia and Vietnam, and on Pyrus from China. 15. Parlatoria crypta McKenzie was described from India, and is also reported in the Comoros, Iran, Iraq, Saudi Arabia and Pakistan; it is a pest. It was intercepted on Mangifera indica from India and Pakistan, and on Murraya from India. 16. Parlatoria pseudaspidiotus Lindinger is an Asian that is most often associated with orchids but is also known to occur on Mangifera indica and other hosts. Miller and Davidson (1990) list it as a pest. It was intercepted on M. indica fruit from India and Mexico and on orchids from the Caribbean islands. 17. Parlatoria ziziphi (Lucas), the black parlatoria scale, is by far the most frequently intercepted. It is almost always intercepted on Citrus fruit, and on shipments originating in all the regions where Citrus is grown. An infestation was discovered in 1995 on Citrus trees in Miami, Florida, but apparently the was eradicated before it could become established. It is a pest and considered to be a threat to agriculture in the USA. 18. Pinnaspis theae (Maskell) is an Asian known to occur in Guadeloupe and Martinique. It was intercepted on Thea from Brazil, but its hosts also include Camellia, Cordyline and Punica. Miller and Davidson (1990) list it as a pest. 19. Pseudaonidia corbetti Hall and Williams was described from Malaysia. It primarily attacks ornamentals, and was intercepted on Garcinia mangostana from China. 20. Pseudaonidia (=Stringaspidiotus) curculiginis (Green) was described from Indonesia, and is also known from Malaysia, Philippines, Vietnam and Thailand. Its hosts include Gardenia and palms; it was intercepted on Pholidota from the Philippines. 21. Pseudaulacaspis barberi (Green) was described from Sri Lanka. Its hosts include Mangifera indica and ornamentals. It was intercepted on mango fruit from India. 22. Silvestraspis uberifera (Lindinger) was described from the Philippines on Mallotus; it is also known from China, Taiwan and Hong Kong. Its reported hosts also include Artocarpus, Cinnamomum, Machilus and Syzygium and it was intercepted on Pimenta from Vietnam. 23. Thysanofiorinia leei Williams was described from Hong Kong on Nephelium, and is also reported in Hawai i, India, Taiwan on Litchi chinensis and Nephelium lappaceum. It was intercepted on Dimocarpus longan from South Korea Australian (2) 1. Pseudaulacaspis brimblecombei Williams was described from Australia on Macadamia integrifolia. It has only been reported to occur in Australia, and was intercepted on Protea and Telopea from New Zealand. 2. Pseudaulacaspis eugeniae (Maskell) was described from Australia. Its reported hosts include Eucalyptus, Melaleuca, Michelia, palms and other ornamentals. It was intercepted on Leucospermum, Protea (Proteaceae) and Scaevola from Australia.