Research Note. Merdelyn T. Caasi-Litl*, Ireneo L. Lit, Jr.2' 3 and Ariel R. Larona3

Transcription

1 Philippine Journal of Crop Science (PJCS) April 2012, 37 (1):47-56 Copyright 2012, Crop Science Society of the Philippines Research Note Expansion of Local Geographic and Host Ranges of Nipaecoccus nipae (Maskell) (Pseudococcidae, Hemiptera) in the Philippines with New Records of Predators and Attending Ants Merdelyn T. Caasi-Litl*, Ireneo L. Lit, Jr.2' 3 and Ariel R. Larona3 'Institute of Plant Breeding, Crop Science Cluster, College of Agriculture, University of the Philippines Los Banos (UPLB), College, Laguna 4031, Philippines; 2 Institute of Biological Sciences, College of Arts and Sciences, UPLB, College, Laguna 4031, Philippines; 3 Museum of Natural History, OVCRE, UPLB, College, Laguna 4031, Philippines; * Corresponding Author, mtcaasi_lit@yahoo.corn The local geographical distribution of the buff mealybug (BCM), Nipaecoccus nipae (Maskell) (Pseudococcidae, Hemiptera) has widened since its initial invasion of Luzon Island in the early 2000s, being now known to occur in other islands and provinces of the Philippines. Surveys were conducted in several regions by initially inspecting known hosts of BCM and other plants. Samples of BCM and of host plants were collected, the latter identified to species. Seventeen (17) plant species belonging to 12 families are recorded as new hosts of BCM, bringing its total known host plants in the Philippines to 71 species in 20 families. An updated host list was prepared to include all local plants that serve as hosts of this invasive species. New associated organisms are also reported for the first time. These include larvae of the green lacewing, Mallada basalis (Walker) (Chrysopidae, Neuroptera) and the brown lacewing, Micromus igorotus Banks (Hemerobiidae, Neuroptera) that have been observed to prey on nymphs and adults, the first known locally available natural enemies of BCM. The black ants, Dolichoderus thoracicus (Smith), fire ants, Solenopsis gem inata (Jerdon), and crazy ants, Anoploiepis gracilipes (Fabricius) (Formicidae, Hymenoptera), are also now occasionally observed to attend to BCM colonies. Keywords: Anoplotepis gracilipes, Buff mealybug, Dolichoderus thoracicus, Mal lada basalis, Micromus igorotus, Nipaecoccus nipae INTRODUCTION The buff mealybug (BCM, Figure 1), Nipaecoccus nipae (Maskell), is a relatively recent introduction (ca. 2000) into the Philippines and was formally reported as an invasive pest by Lit et al. (2005, 2006a, 2006b). Locally, concern about its effects on the industry was first expressed by farmers and gatherers of tuba, the sweet liquid that is processed into lambanog', a locally produced alcoholic beverage similar to gin or vodka. Already an export product, lambanog producers were worried they may not be able to sustain and fulfill orders from abroad if the decline in sap yield continues due to the sucking insects. Other farmers like those raising sweetsop (atis), Annona squamosa L., in Lobo, Batangas also reported declining fruit yield, dying trees and abruptly diminished or lost potential income (Caasi-Lit et al. 2008; Caasi-Lit et al. 2009a). Since its introduction, the buff mealybug has gradually spread to other localities and colonized other host plants (Caasi-Lit et al. 2008; 2009). The lack of basic biological studies especially under local tropical conditions and even in other countries where the BCM is known to occur has consequently prevented entomologists from prescribing effective, practical and long-term solutions to affected local government units and concerned farmers. Realizing such need, the authors are continuing studies, albeit limited, despite the lack of funds or other support for a sustained series of biological and ecological studies. This paper aims to: i) update local distribution data and reflect therein the spread of its geographic range in the Philippines; ii) report new results gathered in the continuing documentation of host plants infested by BCM from 2000 to June 2011 by updating the list of host plants of this polyphagous insect; and iii) record its new associated ants and local natural enemies in the Philippines. MATERIALS AND METHODS Survey of BCM and Documentation of Infested Host Plants Surveys were conducted in different areas of the country whenever possible, from January 2006 to September 2011 and involved convenience or opportunity sampling, without structured sampling design as long as the criteria for host-herbivore association are satisfied. Such surveys generally started among plants identified as hosts of BCM after the initial survey in 2002 (Lit et al. 2006) and onwards. New locality records were plotted on a map as soon as occurrences of BCM therein were confirmed. Some areas were visited more regularly than the

2 Figure 1. The buff mealybug, Nipaecoccus nipae (Maskell) on Freycinetia multiflora (A), Annona squamosa (B) and Carludovica palmata (C,D), showing the different stages: female (F); males, in pupal wax cocoons (MP); and nymphs (N), the first instar of which is called crawler (Cr). others. These include various places in the CALABARZON provinces (except Rizal) and Metro Manila, Davao City and Davao del Norte (at least twice a year), Polillo Island (1-2 times a year since 2002); Isabela, Nueva Ecija, Nueva Vizcaya, Pangasinan, Saranggani and South Cotabato (6-8 times a year from ), Marinduque (1-2 times a year since 2008) and Bulacan,!locos provinces (once a year since 2009). In June,, a survey sponsored by the Bureau of Plant Industry was conducted in Regions 11 and 12 in Mindanao to document the extent of BCM infestation in several agricultural areas in 6 provinces, namely: Davao del Norte, Davao del Sur (including Davao City), North Cotabato, Saranggani, South Cotabato and Sultan Kudarat. The rest of the provinces listed among the new locality records and the provinces of Benguet, Mountain Province, and Ifugao were only visited once or twice within Updating the List of Host Plants of BCM To qualify as a host, a plant had to be infested not just by a single individual but by colonies which would include nymphs, adult females and adult males, usually in their waxy 'cocoons'. There should be clear signs of feeding such as yellowing of portions of plants where the BCM individuals were found, honeydew deposits, and/or growth of sooty molds on surfaces where honeydew dropped or gathered. Photographs of host plants and also the 48 Expansion of Host Ranges of Nipaecoccus nipae

3 corresponding herbarium specimens, when necessary, were taken. Specimens of mealybugs were collected and preserved at least in 95% ethanol for proper documentation and verification. To determine the extent of BCM infestation, the list from Lit et al. (2006) was updated. Botanical names were checked and verified with nomenclatural databases and/or confirmed with the assistance of botanist curators of the University of the Philippines Los Banos (UPLB) Museum of Natural History. Search for natural enemies and associated ants Alongside the continuing survey of BCM and its host plants, natural enemies and associated ants were noted, collected and preserved together with the mealybugs. Predators were reared to adult stage whereas attending ants were preserved in vials of 95% ethanol. All predators and ants were also identified to species level whenever possible. RESULTS AND DISCUSSION Local Geographical Distribution The geographical range of N. nipae in the Philippines continued to widen since its introduction in the early part of this millennium. Lit et al. (2006) reported its initial occurrence in Laguna province among ornamental palms and then in the neighboring provinces of Quezon, Batangas and Cavite (Figure 2). They also noted the introduction to Davao City in Mindanao probably also via ornamental palms. During that time, no BCM individuals or colonies were found in other places visited in the Southern Tagalog region, as well as in the Davao and Cotabato provinces. Since 2006, the local distribution expanded and BCM is now known to be present in nearly all major islands and provinces (Table 1, Figure 2). The presence of BCM in Rizal (part of the CALABARZON subregion) and in Metro Manila is now confirmed. It is also present in the island of Polillo which is politically a part of Quezon. South of the CALABARZON areas, the BCM is now also known in Albay, Camarines Sur and Sorsogon, as well as in three islands/provinces of MIMAROPA subregion (Marinduque, Oriental Mindoro and Palawan. Towards the northern part of Luzon, the occurrence of BCM is now confirmed in the provinces of Bataan, Benguet, Isabela, Nueva Ecija, Nueva Vizcaya, Pangasinan and Zambales. All occurrences of BCM in the Visayan islands of Leyte (Leyte Province), Bohol, Cebu, Negros Occidental Province, Panay (Iloilo Province) and Guimaras are new local geographical records. In Mindanao, from a small area in Davao City, BCM is now confirmed to be present in the provinces of Bukidnon, Davao del Norte, Davao del Sur, Misamis Oriental, North Cotabato, Saranggani, South Cotabato and Sultan Kudarat. The specific localities visited in these new areas are enumerated in Table 1. MT Caasi-Lit et at Figure 2. Geographical distribution of the buff mealybug, Nipaecoccus nipae (Maskell) in the Philippines. Legend: black circles - distribution as of 2005 (based on Lit et al. 2006); 0- new locality records determined through surveys from 2006 to To emphasize the expansion of geographic range of BCM, it is worth mentioning the survey that was conducted in Regions 11 and 12 in Mindanao in June. Initially, the data may be said to only confirm the presence of BCM in the surveyed areas. However, it is noted that there were no traces of BCM in around Tagum and Panabo cities in Davao del Norte, which was just about km from Davao City and yet the incidence of BCM was observed to be abundant among s and bananas in The infestation in Tagum City has been observed on various species of palms planted around the city since Moreover, considering the general biology and ecology of scale insects, as well as the broad view of dispersal as an attribute of insect populations, the great difference between the previous distribution data (Lit et al. 2006) and the new data presented herein all suggest that there has been an expansion in geographical range, and that the set of available data is an indication of the spread from Calabarzon and Davao City to other areas. Similarly, the presence of BCM was only observed in 2008 in Polillo island, just 49

4 Table 1. New Philippine locality records for the buff mealybug, Nipaecoccus nipae (Maskell), based on data gathered from Island Luzon Po lillo Marinduque Mindoro Palawan Leyte Bohol Cebu Negros Panay Guimaras Mindanao Province Albay Bataan Benguet Camarines Sur Isabela Metro Manila Nueva Ecija Nueva Vizcaya Pangasinan Rizal Sorsogon Zambales Quezon Marinduque Mindoro Oriental Palawan Leyte Bohol Cebu Negros Occidental Iloilo Guim aras Bukidnon Davao del Norte Davao del Sur Misamis Oriental North Cotabato Saranggani South Cotabato Sultan Kudarat Places Surveyed Guinobatan, Legazpi City, Ligao Limao, Mariveles Baguio City, La Trinidad City Naga City, Pi li Cauayan City, Ilagan, Gamu, Santiago City Makati City, Manila, Quezon City, Taguig City Cabanatuan City, San Jose City Bambang, Bayombong, Solana, Sta. Fe Bolinao, Dagupan City, San Carlos City, Sta Maria Pililla Gubat, Sorsogon Subic Burdeos, Polillo Boac, Sta. Cruz Calapan City, Naujan Puerto Princesa City Baybay Panglao, Tagbilaran City Cebu City, Mandaue City Bacolod City, La Carlota, La Granja, San Carlos City Iloilo City Port Area Malaybalay City, Musuan Panabo City, Tagum City Digos City, Sta. Cruz Cagayan de Oro City Kabacan City, Makilala Kiamba, Maitum Gen.Santos City, Marbel, Polomolok, Tampakan, Tupi Tacurong * refers to the year when the presence of BCM was confirmed, especially for places visited more than once. Host Plants Observed, guava palmera, royal palm, atis, guyabano, palmera, guava, avocado Year* Royal palm,, palmera, atis, guava, Indian 2009 lanutan, guava 2009, guava, caimito 2009, guava, atis, guava, bun, nipa, mistletoe, palmera, Manila palm, atis, palmera, banana, royal palm, palmera,, banana, palmera, phoenix, bottle palm, guava palmera, banana, atis, nipa h by boat ride from Real in mainland Quezon Province, eventhough visits to the island have been conducted 1-2 times a year since The same is true for Marinduque where BCM was only observed to occur in 2011 although yearly visits were done since Nevertheless, the occurrence of BCM in Baguio City and La Trinidad City in Benguet Province seems to be limited to the commercial areas there. The BCM has not been observed yet in Data, Mountain Province and Lagawe, lfugao during the visits in, and in the!locos Region during yearly visits there since Dispersal refers to the spreading of an organism's propagules or movement of individuals into or out of a population or area (Spellerburg and Sawyer 1999; Odum and Barrett 2005). Spread or movement by itself, may be difficult to directly observe or monitor particularly for minute animals. The study of spread among small arthropods is probably one of the areas "around which ecological generalizations have yet to be developed" (Walter 2003). The spread can be 50 active as in the case of animals which are highly mobile and/or capable of long-distance dispersal, or passive like many sessile and slow-moving arthropods. Among scale insects, including the mealybugs, dispersal is mainly through wind-carried first instar nymphs (Figure 1) called crawlers (Gullan and Kosztarab 1997). Dispersal aided by ants is generally restricted to species with obligate mutualistic relationships with attendant ants as observed by Klein et al. (1992). The spread or expansion of range (or 'ranging' according to Walter 2003) may not be always to escape adverse conditions but could be part of positive response to favorable environmental conditions. Thus, dispersal through wind probably occurs more regularly than previously thought (den Boer. 1990). As mentioned, mealybugs, like other scale insects, disperse through crawlers having a free ride with wind currents. However, there was no chronological order of occurrences on islands and provinces that would Expansion of Host Ranges of Nipaecoccus nipae

5 coincide with known occurrence of major wind directions in the country. New locality records of BCM as listed in Table 1 were dominated by urban and urbanized areas, and the major trading centers and ports of entry. The occurrence of BCM in those new locality records as well as the initial local outbreaks on ornamental plants and other commercially valuable crops indicate that the spread of this invasive species is mainly through human activities. An example of such human activity is the flourishing trade on ornamental plants which has largely capitalized on consumer fancy for trendy exotics rather than on native ornamentals but without the necessary strict implementation of phytosanitary regulations. Obviously, the polyphagous nature of BCM is a significant factor in its spread. As far as inter-island and other relatively long dispersal routes are concerned, however, wind and humans could be the major agents of BCM dispersal and spread. This is supported by the trend of first occurrences in urban and trade centers and ports of entry as well as the lack of pattern attributable or coincident to distribution of host plants, including the apparently more preferred hosts (e.g. s and other palms). The Philippine Department of Agriculture - Bureau of Plant Industry issued strict quarantine measures for and other palms to halt or arrest the spread of pests, targeting specifically the beetle Brontispa longissima Gestro, another invasive species. The implementation of the ban on transport and import of produce, seedlings and other products has slowed down the spread of Brontispa and by default also of the BCM. The main problem however, is that is not the only host of BCM. The BCM can spread to other places through the movement of agricultural crops and products. Unlike Brontispa however, BCM apparently easily succumbs to heavy rains brought about by seasonal monsoon rains and strong typhoons but spreads probably faster during prolonged periods of drought. This is mainly the reason why BCM is easier to control and manage than Brontispa. Extent of BCM Infestation on Several Plant Species An updated list of plants that are now confirmed hosts of BCM is presented in Table 2. The list includes 17 species (in boldface letters) which are new host records for the Philippines. They are the: Mexican sword-plant (Echinodorus palaefolius (Nees & Mart.) J.F. Macbr.); Indian Lanutan (Polyalthia longifolia Benth. & Hook. f.); Lukmoy or Tabatib (Rhaphidophora merrillii Engl.); Queensland palm (Archantophoenix alexandriae H. Wendl. & Drude); Buri (Cotypha utan Lam.); Spindle palm (Hyophorbe verschaffeltii H. Wendl.); Anahau (Livistona rotundifolia (Lam.) Mart.); Abiki (Pinanga copelandii Becc.); Stepladder plant (Costus malortianus H. Wend land); Spiral ginger (Costus speciosus (J. Koenig) Sm.); Dapong Kahoy or Mistletoe (Loranthus philippinensis Cham. & Schlecht.); Ornamental banana (Musa ornata Roxburgh); River red gum (Eucalyptus camaldulensis Dehnh); Climbing pandanus (Freycinetia multiflora Merr.); Iron bamboo (Guadua angustifolia Kunth); Satinleaf or caimitillo (Chrysophyllum oliviforme Linn.); Traveller's palm (Ravenala madagascariensis Sonn.); and Torch ginger (Alpinia purpurata (Vieill.) K. Schum.). The new host records bring the total host range to 20 families, 47 genera and 71 species. Of the 17 new host records, 6 species are shown in Figure 3, each representing the families of plants (Alismataceae, Costaceae, Loranthaceae, Pandanaceae, Poaceae and Zingiberaceae) now reported as new hosts of BCM in the Philippines. Based on the report of Lit et al. (2006), the number of species being infested rose from four during the first survey in 2001 to 12 in 2004 (Figure 4). In 2005, the number increased to 25 plant species. This went up drastically in 2006 and with 53 and 59 host plants recorded, respectively. The present survey conducted to include areas in Mindanao confirmed that BCM infestation had reached this part of the country (Table 1). The long dry spell during could be the reason for such increase. Mealybugs like most sternorrhynchans are regarded as dry season pests as their reproduction is generally favored by the warm temperature and other environmental conditions during dry seasons. In addition, the lower relative humidity and less precipitation during dry months that favor scale insects are not favorable for their pathogens. Although mealybugs are covered with waxy secretions, these secretions are not enough to protect them against attack particularly by fungi like Neozygites fumosa (Speare) Remaudiere & Keller as reported by Villacarlos (2000). Since then, 2-3 species per year were added to the list, and a total of 71 host plants were finally recorded in this paper. Again, this indicates the rapid spread of the pest to other areas. In other countries where BCM has been previously known to occur, this species has also been known to be polyphagous. Comparison of the Philippine Records with World Record of Host Plants Worldwide, the host plants of BCM consist of 96 species that belong to 85 genera and 42 families (Ben -Dov 1994; Ben-Dov and Miller 2009). Combining those reported by Lit et al. (2006) and the new host records in this paper, the Philippine host plant record includes 71 species in 47 genera and 20 families. Potentially, therefore, there are still 25 species in 38 genera and 22 families that may be discovered as host plants in the Philippines. It is also worth noting that the Alismataceae, Loranthaceae and Poaceae (=Gramineae), here reported as among the six new MT Caasi-Lit et al 51

6 Table 2. Updated host list of the buff mealybug, Nipaecoccus nipae (Maskell), in the Philippines. New host records are written in boldface letters. Family ALISMATACEAE ANNONACEAE AR ACEAE ARECACEAE CANNACEAE CO NVALLAR IACEAE COSTACEAE CYCLANTHACEAE HELICONIACEAE LAURACEAE LORANTHACEAE MAR ANTACE AE MORACEAE MUSACEAE MYRTACEAE English and Other Names Mexican sword-plant Atemoya Soursop, Guyabano Anonas Sweetsop, Sugar apple, Atis Indian Lanutan Biriba Dumb cane, Bakya Dumb cane Lukmoy, Tabatib Gabing San Fernando Manila palm, Bunga de Jo lo Queensland palm Betel nut palm Fishtail palm, Pugahan Large fishtail palm, Takipan Bamboo palm Coconut, Niyog, Lubi Buri Red palm Blue palm, Triangle palm Golden Palm, Palmera Madagascar palm Normandy palm African oil palm Saguisi Marighoi Champagne/ Bottle palm Spindle palm Licuala, Balatbat-bilog Spiny licuala, Balatbat Anahau Nipa, Sasa Date palm Date palm, Voiavoi Date palm Abiki Kuhl abiki Gasigan Sarauag Tigbisa Banisan MacArthur palm Rhapis, Lady palm Royal palm Foxtail palm Bandera Espanola Ti plant Baston de San Jose Stepladder plant Spiral ginger Panama hat palm Yellow plantain False bird of paradise Alligator pear, Avocado Dapong Kahoy, Mistletoe Arrow root, Uraro Himbabao Ornamental Banana Banana, saba, lakatan River red gum Bagras Guava, Bayabas Scientific Name Echinodorus palaefolius (Nees & Mart.) J.F. Macbr. Annona atemoya Hort. Annona muricata L. Annona reticulata L. Annona squamosa L. Polyalthia longifolia Benth. & Hook. f. Rollinia deliciosa Saff. Dieffenbachia amoena Bull. Dieffenbachia maculata (Lodd.) G. Don Rhaphidophora merrillii Engl. Xanthosoma sagittifolia (L.) Schott. Adonidia merrillii (Becc.) Becc. Archantophoenix alexandriae H. Wendl. & Drude Areca catechu L. Caryota cumingii Lodd. ex Mart Caryota rumphiana Mart var. philippinensis Becc. Chamaedorea seifrizii Burret Cocos nucifera L. Corypha utan Lam. Cyrtostachys renda Blume Dypsis decaryi (H. Jumelle) Beentje & J. Dransf. Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. Dypsis madagascariensis (Becc.) Beentje & J. Dransf. Dypsis sp. Elaeis guineensis Jacq. Heterospathe elata Scheff. Heterospathe intermedia (Becc.) Fernando Hyophorbe lagenicaulis (L.1-I. Bailey) H.E. Moore Hyophorbe verschaffeltii H. Wendl. Licuala grandis H.Wendl. Licuala spinosa Wurmb. Livistona rotundifolia (Lam.) Mart. Nypa fruticans Wurmb. Phoenix dactylifera L. Phoenix loureiroi Kunth Phoenix roebelinii O'Brien Pinanga copelandii Becc. Pinanga coronata (Blume ex. Mart) Blume Pinanga heterophylla Becc. Pinanga insignis Becc. Pinanga maculata Porte ex. Lam. Pinanga speciosa Becc. Ptychosperma macarthurii Wendl. Rhapis excelsa (Thunb.) Henry Roystonea regia (HBK) O.F. Cook Wodyetia bifurcata A.K. Irvine Canna indica L. Cordyline fruticosa (L.) A. Chev. Cordyline terminalis (L.) Kunth. var. ferrea (L.) Endl. Costus malortianus H. Wend land Costus speciosus (J. Koenig) Sm. Carludovica palmata Ruiz & Pay. Heliconia bihai (L.) L. Heliconia platystachys Baker Persea americana Mill. Loranthus philippinensis Cham. & Schlecht. Maranta arundinacea 0. Petersen Broussonetia luzonica (Blanco) Bur. Musa ornata Roxburgh Musa x paradisiaca L. Eucalyptus camaldulensis Dehnh Eucalyptus deglupta L. Psidium.quaiava L. 52 Expansion of Host Ranges of Nipaecoccus nipae

7 Table 2. Continued... Family ANDS A EAE POACEAE SAPOTACEAE STRELITZIACEAE ZINGIBERACEAE English and Other Name s hmbing pan anus Iron bamboo Star apple, Caimito Satinleaf, Caimitillo Sapodilla, Chico Traveller's palm True bird of paradise Torch ginger Scientific Name reycinetia multi lora Merr. Guadua angustifolia Kunth Chtysophyllum cainito L. Chrysophyllum oliviforme Linn. Manilkara zapota (L.) P.van Royen Ravenala madagascariensis Sonn. Strelitzia reginae L. Alpinia purpurata (Vieill.) K. Schum. Figure 3. Six of 17 new host plant records of Nipaecoccus nipae, representing the six new host plant families in the Philippines. A. Alpinia purpurata (Zingiberaceae). B. Freycinetia multiflora (Pandanaceae). C. Costus malortianus (Costaceae). D. Echinodorus palaefolius (Alismataceae). E. Guadua angustifolia (Poaceae). F. Loranthus philippinensis (Loranthaceae). MT Caasi-Lit et at 53

8 No. of Host 40 Plants \ ;1' Year Figure 4. The increasing number of host plants infested with the buff mealybug from 2001 to (Source of data from 2001 to 2006: Lit et al. 2006). host plant families in the Philippines are also not among the families listed as hosts by Ben-Dov (1994) in his world catalogue. In addition, the hosts reported in this paper and in the previous one (Lit et al. 2006) are backed up by specimen documentations, with a clear criterion for classifying a plant as host. Definitely, there are host plant families, genera and species reported in other countries that are not present in the Philippines. It is also possible that locally available varieties of known host plants might be resistant to BCM infestation. Based on the known host range in the Philippines, BCM is primarily associated with palms in the family Arecaceae comprising about 35 species or 49.3% of all known host plants (Table 3). According to Maskell (1893), this pest was originally described from Nypa fruticans, a palm species abundantly growing in mangroves, in Demerara, Guyana in South America. Several other palms were recorded including the economically important, Cocos nucifera and almost all kinds of palms. These numerical values include the single species (Panama hat palm, Carludovica palmata Ruiz & Pay.; Figure 1) that belongs to the Cyclanthaceae, a close ally of the Palmae (Arecaceae). Associated Ants and Local Natural Enemies New organisms associated with the BCM are also reported for the first time. These include larvae of the green lacewing, Mallada basalis (Walker) (Chrysopidae, Neuroptera), and the brown lacewing, Micromus igorotus Banks (Hemerobiidae, Neuroptera), that were observed to prey on nymphs and adults, the first known locally available natural enemies of BCM. The black ants, Dolichoderus thoracicus (Smith), fire ants, Solenopsis geminata (Jerdon), and crazy ants, Anoplolepis gracilipes (Fabricius) (Formicidae, Hymenoptera), were occasionally observed to attend to BCM colonies. Lit et al. (2006) noted that only the weaver ants, Table 3. Number of plant species collected from each family as host plants of the buff mealybug, Nipaecoccus nipae (Maskell), in the Philippines. Family Number of Species Percent Alismataceae Annonaceae Araceae Arecaceae Cannaceae Convallariaceae Costaceae Cyclanthaceae Hel iconiaceae Lauraceae Loranthaceae Mara ntacea e Moraceae Musaceae Myrtaceae Pandanaceae Poaceae Sapotaceae Strelitziaceae Zingiberaceae TOTAL Oecophylla smaragdina Fabricius, and an undetermined species of Dolichoderus attend to colonies of BCM. The undetermined Dolichoderus sp. mentioned by Lit et al. (2006) is confirmed in the present study as D. thoracicus. These ant species are common in most agroecosystems and their attendance to BCM colonies is most probably facultative, like the relationships of these ants to other mealybugs. As BCM is relatively new locally, the significance of the presence of these ants to BCM occurrence, distribution and spread needs further study. SUMMARY AND CONCLUSION This paper is a sequel to the study by Lit et al. (2006) where host plants infested with BCM were recorded. In this paper, local geographical distribution and host plant range data are updated by reporting new results gathered in the continuing documentation of BCM spread and infestation from to June The results in this study confirm that the geographical range of N. nipae in the Philippines has widened since the last documentation of its local distribution in During that time, Philippine populations of BCM were known only from the provinces of Batangas, Cavite, Laguna and Quezon, and in Davao City. The new areas of infestation are: i) the provinces of Albay, Bataan, Benguet, Camarines Sur, Isabela, Metro Manila, Nueva Ecija, Nueva Vizcaya, Pangasinan, Rizal, Sorsogon and Zambales in Luzon; ii) the provinces of Bukidnon, Davao del Norte, Davao del Sur, Misamis Oriental, North Cotabato, Saranggani, 54 Expansion of Host Ranges of Nipaecoccus nipae

9 South Cotabato and Sultan Kudarat in Mindanao; and iii) in the islands of Polillo, Marinduque, Mindoro (Mindoro Oriental Province), Palawan, Leyte (Leyte Province), Bohol, Cebu, Negros (Negros Occidental Province), Panay (Iloilo Province) and Guimaras. Among the places visited within , BCM is not known or confirmed yet to occur in Mountain Province, Ifugao and the provinces of the!locos Region. There was also 20% increase in number of host plant species infested by BCM. As of 2011, the list includes 20 families, 47 genera and 71 species with 17 species as new host records for the Philippines. Based on the known host range in the Philippines and elsewhere, BCM is primarily associated with palms (family Arecaceae). New associated organisms are also reported for the first time in the Philippines. These include lacewing larvae that have been observed to prey on nymphs and adults, the first known locally available natural enemies of BCM. Five species of ants are also now known to occasionally attend to BCM colonies. ACKNOWLEDGEMENT The authors would like to thank Ms. Wilma Cuaterno for inviting Dr. M.C. Lit to travel to Region 11 and Region 12 and monitor the infestation level of BCM in the area; Ms. Lilian V. Punzalan and Ms. Regina U. Herrera for their assistance in the on-going life history studies, rearing and insect preservation; Dr. Ma. Dulce J. Mostoles, Dr. Joan C. Orense and Mabel A. Capricho for sharing information on BCM occurrences in the Bicol Region and Cebu; and Mr. Russel C. Larona for his help in drawing the map shown in Figure 2. LITERATURE CITED Ben-Dov Y A Systematic Catalogue of the Mealybugs of the World (Insecta: Homoptera: Coccoidea: Pseudococcidae and Putoidae) with Data on Geographical Distribution, Host Plants, Biology and Economic Importance. Andover, UK: Intercept Limited. 686 pp. Ben-Dov Y, Miller DR Scale Net. barc. usda. gov/scalenet/scalenet. htm. Last updated 15 August Caasi-Lit MT, Lit ILJr, Larona AR, Manzanilla C, Plata R, Egea RD Invasive mealybug: the culprit in the decline of the atis industry in Lobo, Batangas. In Abstracts of papers presented during the 30th NAST Annual Scientific Meeting, July 9-10, Manila Hotel, Metro Manila. Trans NAST Philipp 30(1): 104. Caasi-Lit MT, Lit ILJr, Larona AR, Plata R, & Egea RD. 2009a. Need for rehabilitating the mealybuginfested atis plantations in Lobo, Batangas: lessons to learn. Philipp Ent 23(2): 180. Caasi-Lit MT, Lit ILJr, Larona AR, Dizon TO. 2009b. Invasion of the buff mealybug (Nipaecoccus nipae) in UP Los Banos campus, elucidation of the confounded "Mealybug burn" damage and practical control. In Abstracts of papers presented during 30th NAST Annual Scientific Meeting, July 9-10, Manila Hotel, Metro Manila. Trans NAST Philipp 31(1): Caasi-Lit MT, Lit ILJr, Larona AR Boom and bust and other updates among local populations of the buff mealybug, Nipaecoccus nipae (Maskell) (Hemiptera: Pseudococcidae). In Abstracts of papers presented during 30th NAST Annual Scientific Meeting, July 9-10, Manila Hotel, Metro Manila. Trans NAST Philipp 30(1): Den Boer PJ The survival value of dispersal in terrestrial arthropods. Biol Cons 54: Gullan PJ, Kosztarab M Adaptations in scale insects. Annu Rev Entomol 42: Klein RW, Kovac D, Schellerich A, Maschwitz U Mealybug-carrying by swarming queens of a Southeast Asian bamboo-inhabiting ant. Naturwissenschaften 79: Lit ILJr, Caasi-Lit MT, Larona AR. 2006a. Nipaecoccus nipae (Maskell), an invasive mealybug attacking agricultural crops and forest trees. In Abstracts of papers presented during the Annual Scientific Conference of the Philippine Association of Entomologists, Inc., held during the 37th Annual Convention of the Pest Management Council of the Philippines, May 2-5, Grand Regal Hotel, Davao City, Philippines. Philipp Ent 20(2): 179. Lit ILJr, Caasi-Lit MT, Larona AR. 2006b. The buff mealybug, Nipaecoccus nipae (Ma ske II), a new invasive pest in the Philippines with a synopsis of mealybugs (Hemiptera: Coccoidea: Pseudococcidae). Philipp Agric Scientist 89 (1): Lit ILJr, Caasi-Lit MT Island hopping and burning without fire: The case of the buff mealybug, a new invasive pest. UPLB Museum of Natural History Seminar Series, UP Los Banos, College, Laguna. July 25. Lit ILJr, Caasi-Lit MT Unprecedented multispeci es outbreaks of scale insects (Hemiptera: Coccoidea) in Mindanao Island, Philippines: effects of climate change or poor quarantine practices. In Abstracts of papers presented during the 10th International Congress of Ecology (INTECOL 2009), August MT Caasi-Lit et at 55

10 Brisbane Convention & Exhibition Centre, Brisbane, Queensland, Australia (CD Rom). Maskell WM Further coccid notes: with descriptions of new species from Australia, India, Sandwich Islands, Demerara, and South Pacific. Trans N Z lnst 25: Odum EP, Barrett GW Fundamentals of Ecology. 5th Ed. Belmont, CA, USA: Thomson Brooks/Cole. xviii p. Spellerberg IF, Sawyer JWD An Introduction to Applied Biogeography. Cambridge, UK: Cambridge University Press. xiv p. Villacarlos LT Two Neozygites species (Zygomycetes: Entomophthorales) infecting aphids and mealybugs on Leyte Island. Philipp. Ent. 14(1): Walter GH Insect Pest Management and Ecological Research. New York, USA: Cambridge University Press. xii p. 56 Expansion of Host Ranges of Nipaecoccus nipae