Professional Documents
Culture Documents
Bioinvasion
Threat to Food, Health, Livelihood,
Cultural and Environment Security in the Pacific Islands
R. R. Thaman
R. R. Thaman
2018
Coffee Brown Scale (Saisettia coffeae)
Dedication
1 Introduction
This paper is about the recent biological Most IAS that have historically invaded
invasion of Fiji’s cities, villages, homes, oceanic island groups, such as Fiji, Samoa,
garden lands and islands by ants and Tahiti and Hawaii, have come from the con-
mealybugs, scale insects and a number tinents of Asia, Australia, Africa and South
of other sap-sucking insects and diseases America or from continental islands, such
that form alliances with, and are tended by as the islands of Indonesia and Papua New
this highly invasive black ant. This multi- Guinea, where they evolved in competition
pronged invasion, hereafter referred to as with their own predators, parasites and
the 2016 Fiji Ant-Mealybug bioinvasion, is diseases which had controlled their num-
perhaps the latest historical example of the bers. When introduced into new, less-com-
devastation of indigenous peoples, plants, petitive oceanic island ecosystems, where
animals, islands and the ecosystem goods there are few indigenous predators, para-
and services they depend on by invasive sites and diseases, these more competitive
alien species (IAS) in the Pacific Islands. continental organisms, which are common-
ly introduced without their original preda-
Since Elton (1958) reported the serious- tors, parasites and diseases, have a distinct
ness of the impacts of bioinvasions on competitive advantage and outcompete
native biological communities, invasive and displace the naïve indigenous island
alien species (IAS) have been recognised organisms that have evolved in isolation in
as a major driver of the loss of biodiversity less-competitive oceanic island ecosystems.
and ecosystem services (BES)2, particularly Moreover, most IAS reach sexual maturity
on islands (MacArthur and Wilson 1967; early, reproduce in large numbers and are
Williamson 1981; Simberloff 1995; Jourdan well-adapted to disturbed habitats, which
1997; UNEP 2014), with over 75% of all gives them further competitive advantages
extinctions since 1600 having occurred on with clearly increasing disturbance and
islands, particularly oceanic islands (Goom- habitat degradation due to human actions
bridge 1992). There are innumerable cases and extreme events, such as devastating
within the Pacific Islands where IAS (e.g., tropical cyclones (Simberloff 1995; CGAPS
alien mammals, birds, reptiles, amphibians, 1996; MA 2005; Thaman 2011; Simberloff
fishes, molluscs, insects, echinoderms, ter- et al. 2013; Thaman 2013; Butler and Bax
restrial and aquatic weeds, fungi and a vast 2014). Table 1 highlights some of the more
array of infectious diseases and micro-or- serious IAS and the impacts they have had
ganisms) have led to extinctions of indige- on islands.
nous mammals, birds, amphibians, reptiles,
land crabs and snails and insects; the loss
of traditional plant cultivars; and negative
economic, social and environmental costs
and threat to human health associated
with of the use of toxic pesticides to control
IAS (Thaman 1984; Thaman 2011; Thaman
2013).
2
Ecosystems, the plants, animals and organisms they contain and the goods and services (tangible and intan-
gible) they provide to humans (Diaz et al. 2013.)
2 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Table 1. Examples of some of the more serious IAS and the historical impacts they have
had on the islands of the APR.
• Extinction of endemic birds, fruit bats and geckos and the reduction of bird-dispersal
and reproduction of new trees by as much as 60—90% in Guam due to the introduc-
tion of the brown tree snake shortly after World War II (Rodda et al. 1992; CGAPS
1996; Baskin 2003; McCrae 2017; Rogers et al. 2017).
• Widespread extinction and drastic population declines of birds, land snails and land
crabs in Hawaii, French Polynesia, Rotuma and many other Pacific Islands due to
combinations of avian malaria, rats, mongooses, cats, pigs, goats, ants, predatory
land snails, flatworms and habitat degradation (Howarth 1985; van Ripper et al. 1986;
CGAPS c. 1996; Thaman 2013; Brodie et al. 2014; PIAT 2016).
• Widespread loss of biodiversity and human discomfort and negative social impacts
due to alien ants, such as the little fire ant (Wasmannia auropunctatus), which has
been introduced into New Caledonia, Solomon Islands, Vanuatu and Wallis and
Futuna, where it has made subsistence food gardening difficult and become a major
pest of coconuts, cocoa and citrus fruits and other plants and forms negative symbi-
otic relationships with scale insects, mealybugs and other crop pests (Jourdan 1997;
Le Briton et al. 2003; Fasi et al. 2013); and the yellow crazy ant (YCA) (Anoplolepis
gracilis), which also forms alliances with scale insects, has had devastating impacts on
land crabs, seabirds, indigenous insects and spiders, pigs, chickens, tree regeneration
and crops on Christmas Island in the Indian Ocean (O’Dowd et al. 2003) and in Samoa,
Tuvalu, Tokelau and elsewhere in the Pacific (Auina et al. 2011; Gruber et al. 2012;
Vaqalo et al 2014; PIAT 2016). In Tuvalu the YCA has displaced or supressed other ants
on even some of the smallest uninhabited islets (Vaqalo et al. 2014), and in Australia,
the YCA has invaded over 200 hectares of rainforest in and adjacent to the Wet Trop-
ics World Heritage Area near Cairns and over 500 hectares of adjacent residential land
and cane farms, A draft 2012 cost-benefit analysis by the Queensland government,
considering only limited impacts on agriculture and dwellings, found that costs would
range from $115 million to over $3 billion if the ants were not controlled (Lasch and
Hoskin 2015).
• Invasion of cropland and secondary forest in Fiji by the invasive African tulip tree
(Spathodea campanulata), which takes up to one-third of the cropland in some areas
and outcompetes and impedes the natural regeneration of a wide range of cultural-
ly valuable pioneer tree species that were formerly part of sustainable multispecies
fallow cropping systems (Thaman 2011, 2013, 2014);
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 3
• Devastation of taro production in Samoa by the taro leaf blight (Phytophthora coloca-
siae); in Fiji, Solomon Islands and PNG by the taro beetle (Papuana spp.) and/or along
with Alomae and Bubone viral complex in Solomon Islands and PNG (Alolii et al. 1993;
Hunter et a. 1998; Thaman 2014; Tsatsia and Jackson 2016).
• Discovery and recent devastation of the papaya export industry and other fruit trees
in the Cook Islands in 2013 by the oriental fruit fly (Bactrocera dorsalis), which report-
edly attacks over 150 species of fruits and vegetables and native fruits (Allwood et al.
1999; Vargas et al. 2007; Weems et al. 2012; Wilson 2013).
• The virtual disappearance of dadap or coral tree (Erythrina variegata var. orientalis)
an important nitrogen-fixing, live fencing and cultural tree in Hawaii, Fiji, Samoa and
other countries due to the highly invasive African Erythrina gall wasp (Quadrastichus
erythrinae) (Campbell 2010; Thaman 2011); and in 2012, on Nanumea Atoll, Tuva-
lu, the devastation of beach cordia (Cordia subcordata), the main woodcarving and
coastal protection species, by infestations of kou leafworm (Ethmia nigroapicella)
(Thaman 2011; Thaman and O’Brien 2012).
• Asian subterranean termites (Coptotermes gestroi) have, in Fiji, since the mid-2000s
caused millions of dollars losses in housing and destroyed livelihoods, in addition to
over a millions dollars in control costs and the need for the use of very toxic chemicals
(BAF 2014)
• Marine invasive species, such as algae, plankton and echinoderms, such as the crown-
of-thorns starfish (Acanthaster planci) are also an increasing concern (Coles et al.
1999; Butler and Bax 2014; N’Yeurt and Iese 2015).
As suggested above, the 2016 Ant-Mealybug Bio-invasion of Fiji’s cities, villages, homes,
garden lands and islands by alien ants in symbiotic relationships with a range of mealy-
bugs, scale insects and other associated insects and diseases constitutes, perhaps, the
most recent and serious IAS invasion in the Pacific. With little or no assistance from desig-
nated authorities, this seemingly unstoppable biological juggernaut invades and reinvades
our properties and destroys our plants, leaving homeowners, gardeners, horticulturalists
and conservationists helpless. For many, the 2016 Fiji Ant-Mealybug Bio-invasion seems
as serious and certainly more bothersome, more immediate and less understandable than
climate change, natural disasters and environmental degradation. Like climate change, but
unlike natural disasters and oil spills, these IAS constitute a living pollution that seemingly
won’t go away and will probably only get worse over time (Thaman 2013b).
4 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
The paper discusses 1) the origin, timing and nature of the 2016 Fiji Ant-Mealybug Bioinvasion;
2) the diversity of invasive organisms and host plants that have been affected; 3) emerging
environmental, economic and social impacts of the invasion; 4) possible reasons for the
seriousness of the invasion; 5) a preliminary assessment of what can be done to manage
the invasion without further damaging our health, cultures, economies and environment;
and 6) suggestions of areas of investigation for future studies. The study is based on; 1) a
review of information on the ants, mealybugs and other insects that are part of the inva-
sion, infested host plants, information on other ant-mealybug bio-invasions, and control
measures that have or could be employed to control the invasion; 2) a four-month-long
field study of the presence and diversity and seriousness of impacts of this invasion at
over 40 sites in Fiji, during which voucher photographs were taken of ants, mealy bugs
and other associated insects and pathogens on infested host plants, and a tabular data-
base prepared of infested species, IAS responsible and levels and locations of infestations;
3) discussions with gardeners, horticulturists, agriculturalists, conservationists, homeown-
ers, tourism operators and hospitality providers in Suva and other parts of Fiji’s main is-
land Viti Levu about the invasion, its impacts; and 4) discussions and correspondence with
entomologists, plant scientists and other authorities on biological invasions. The main
motivation behind the study is to better understand and raise awareness of the serious-
ness of the invasion, which had seriously impacted the author’s own garden and property
and the gardens and properties of many friends and other Fiji residents, and to explore
what has been, and might be, done to arrest the invasion. All photograph are those of the
author, unless otherwise stated.
Acknowledgements
During the study, many people have shared their knowledge and allowed me to examine
the plants in their gardens. Among those who I would like to give thanks (in no special
order) are Richard Titoko, Dick Watling, Bob Gillett, Petero Manoa, Mosese Uluiciciya, Ken
MacDonald, Kiran Lal, Linikoni Vakauta, Konai Helu Thaman, Seini Tuiteci, Semi and Doro-
thy Duaibe, Josua Wainiqolo, Maclean Vaqalo, Monica Gruber, Gilliane Watson, Gilianne
Brodie, Mani Mua, Peter Maddison and Grahame Jackson). Special thanks are also given
to Professor Elisabeth Holland, Director, and Dr. Morgan Wairiu, Deputy Director, of the
Pacfic Centre for Environment and Sustainble Development (PaCE-SD) for strongly sup-
porting the publication and distribution of this report; Dr. Antoine N’Yeurt who proof-read
the manuscript; and Chris Ward who was responsible for the layout and production of the
final edition. To those of you who helped and I have failed to acknowledge, please accept
my sincere thanks and apologies.
More recently (for most of us early 2017), many of us began to see our citrus, breadfruit,
jackfruit, guava, avocado, soursop, other fruit trees and our hibiscus, ixoras, gardenias,
island musk (uci) and other ornamentals and culturally important plants invaded by these
same ants and encrusted with white, downy, cotton-like accumulations or yellow-white
scale-like infestations that seemed to be accompanied, tended or farmed and protected
by the ants. The leaves of many affected plants were also infested on the upper surfac-
es by black sooty mould that inhibits photosynthesis by the leaves (Schmaedick 2007).
Together these seemingly inseparable allies have become an extremely irritating, hard-
to-control and costly invasion of our buildings, properties and gardens, killing and dam-
aging many of our most important food, multipurpose and ornamental plants. With little
or no assistance or guidance from designated authorities, this seemingly unstoppable
juggernaut has invaded and reinvaded our properties, destroyed our plants and left most
homeowners, gardeners, horticulturalists and conservationists helpless, save for those
who have discovered the “silver bullet” (a simple solution to a complicated problem) or
have resorted to the use of large amounts of Mortein, Raid, Orthene or other insecticides,
the use of which constitutes a serious, but poorly understood, health and environmental
hazard.
Fig. 1. Super-abundant white-footed ants (WFA) (Technomyrmex sp.), which have spearheaded the Fiji 2016
Ant-Mealybug Bioinvasion of Fiji, on a kitchen sink, Suva September 2017 (left) and WFS tending Seychelles
Scale (SS) on the underside of an avocado leaf, Suva, June 2017 (right).
As the survey progressed, it became clear that, in addition to these more common spe-
cies, there were other mealybugs, scale insects, aphids and other sap-sucking insects and
plant pathogens that that were also associated with WFA and part of the invasion, often
infesting the same plant hosts along with SS and/or PHM (Table 2).
Table 2. Polyphagous mealybugs, scale insects, other sap-sucking insects and plant patho-
gens observed and identified (some tentatively) as being present on host plants in appar-
ent symbiotic infestations with WFA, based on field studies in Suva and other selected
locations on Viti Levu Island, Fiji by R. Thaman from May to September 2017 (Notes: 1)
the main host plants include those plants seen to be most commonly infested in Fiji and/
or reported to be important hosts overseas; 2) the identification of some organisms is
tentative pending microscopic analysis of specimens).
Among the more common of these other insects or pathogens are coconut mealy bug
(CMB) (Nipaecoccus nipae); citrus mealybug (CTM) (Planococcus citri); pineapple mealy-
bug PMB) (Dysmiscoccus brevipes); possibly breadfruit mealybug (BMB) (Icerya aegyp-
tiaca); coffee brown scale (CBS) (Saisettia coffeae) and/or nigra scale (NS) (Parasaisettia
nigra); Asian cycad scale (ACS) (Aulacaspis yasumatsui); soft bamboo scale (SBS) (Bambu-
saspis bambusae); spiralling white fly (SWF) (Aleurodiscus dispersus); aphids (APH) (Aph-
ididae); bele or Abelmoschus shoot borer (ASB) (Earias vittella); brown and yellow aphids
(Aphididae); black sooty moulds (BSM) (Capnodium and Tripospermum spp. plus others)
and plumeria rust (PR) (Coleosporium plumeriae) (Table 2)(See figures below related to
individual plant taxa).
Apparently native to Madagascar, where it was first reported present in 1892, the difficult
WFA began spreading through Southeast Asia and Oceania more than 60 years ago and
was first reported in the New World in 1986 where, until 2007, it was misidentified as
Technomyrmex albipes (Forel 1892, 1911; Wetterer 2013), with Bolton (2007), believing
that all recently published records of T. albipes from Australia and some of the published
records from Pacific Islands (Wilson and Taylor 1967) were probably also T. difficilis,
although, as Bolton (2007) stressed, in the absence of specimens, the actual identity of
many published T. albipes records “must remain equivocal” (Wetterer 2013).
The relatively recent arrival of WFA (T. difficilis) in most of the Pacific is supported by
Wetterer (2013) who reported that it was neither among the 15 tramp ant species spread
by human commerce reported by Forel in 1911 to have achieved almost cosmopolitan
distributions (eight of which had had already become major ecological, agricultural, and/
or household pests), nor was it among several other serious invasive ant species to subse-
quently become cosmopolitan during 20th century.
10 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
In the Caribbean, WFA was first recorded at a nursery in Florida in 1986 and was subse-
quently found present in Puerto Rico in 1996 and on at least 7 other islands, most recent-
ly in the Bahamas and Jamaica in 2007. Although first recorded in the western Pacific in
Guam and the Federated States of Micronesia as early as 1946 and 1953 (possibly intro-
duced during or just after WW II) and in Australia in 1972, it was first reported from in Ha-
wai’i in 1994 and Papua New Guinea in 2007 (Wetter 2008; Hodgson and Lagowska 2011;
Wetterer 2013); and it was not reported present by Sarnaat and Economos’ in Ants of Fiji
in 2012, although the related, apparently endemic or indigenous, species, Technomyrmex
vitiensis Mann, which is difficult to differentiate from the WFA, was reported present near
Nadarivatu in central Viti Levu in 1921; and T. albipes (F. Smith), which is considered to be
synonym for T. vitiensis, a species reportedly indigenous to New Guinea and Melanesia,
was reported present on all of the larger islands in Fiji by Ward and Wetterer in 2009.
It was only in 2013, however, that Wetterer said: “One emerging cosmopolitan pest ant
species, now spreading rapidly through Florida and the West Indies is Technomyrmex
difficilis FOREL, 1892”, which, as stressed above, until 2007, was commonly misidentified
as Technomyrmex albipes (Wetterer 2013: 93). Given, its rapid recent spread in the Carib-
bean, it is probably safe to suggest that WFA has recently spread from Australia, Papua
New Guinea, Micronesia or Hawai’i into Fiji and other parts of the Western and Central
Pacific.
Fig. 2. Seychelles scale (SS) (Icerya seychellarum), also known as the Seychelles mealybug or breadfruit scale
or mealybug seen on leaves of avocado (Persea americana)(left) and chenille plant (Acalypha hispida), Suva,
Fiji, 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 11
Like the WFA, the SS is possibly native to Africa or the Indian Ocean Islands, such as the
Seychelles (after which it was named in 1855), and was one of the earliest reported
mealybugs from the Mascarene Islands, where it was reported present in Mauritius and
Reunion as Coccus saccharin in 1867 (Pestnet 2014), It subsequently spread to South
Asia, with many of the earliest documented reports coming from India (Rao 1951), and
the type specimen, which is lodged at the Smithsonian Institution National Museum of
Natural History, collected in 1905 in Manila, Philippines (ALA 2017). Interestingly, in 1896,
W.W. Markell, the eminent New Zealand entomologist and acknowledged world expert on
coccid insects, identified a specimen with no recorded place of origin which is lodged in
the DSIR Entomology Division in New Zealand; and, given the fact that SS was not report-
ed from Australia, NZ or the Pacific Islands until recently, it almost certainly came from
Asia, from where he was reportedly sent coccid specimens for study and identification
(Morales 1993). The SS has also been reported from other Indian Ocean islands, including
Zanzibar, Comoros, Mauritius, Reunion, Rodrigues, Andaman and Nicobar Islands and the
British Indian Ocean Territory (CABI/EPPO2008; EPPO 2014; CABI 2016). Seychelles scale
was also first discovered in 1968 on Aldabra atoll in the far north of the Seychelles, the
world’s second largest atoll and one of the most isolated uninhabited islands on Earth,
where the SS seriously impacted the native vegetation, threatening the survival of some
woody species (SIF 2017). It had not been reported present by early expeditions to the
atoll (Hill and Newbery 1980). There are also studies of interactions between SS and ants
on Aldabra, where SS were reportedly attended by large numbers of ants that seemed to
stimulate the SS to produce more honeydew (Hill and Blackmore 1980).
The presence of SS has also been documented from most of Africa and throughout South
and Southeast Asia, including Nepal, Pakistan, Sri Lanka, China, Hong Kong, Thailand,
Malaysia, Brunei, Indonesia, Irian Jaya, Japan (including the Ryukyu Islands) and Taiwan;
(CABI/EPPO2008; EPPO 2014; CABI 2016). It has also been reported present in parts of
southern Europe and was first collected in Australia in 1990 (CABI/EPPO2008). In the
Pacific, it is now widespread and was first collected and identified present in the Pacific
Islands in the Cook Islands and Niue in 1933 (ALA 2016); on Tutuila in American Samoa at
least by the 1950s (Dumbleton 1954; Schmaedick 2007); in Fiji by the mid-1940s (Lever
1946, 1947); in the Federated States of Micronesia in 1985; in Australian Northern Territo-
ry, Papua New Guinea, Solomon Islands, Vanuatu, New Caledonia, Kiribati, Tuvalu, Samoa
by 1990; and in Fiji, Tonga, Palau, Nauru, and French Polynesia and New Zealand by 2008
(Williams 1985; Williams and Watson 1990; CABI/EPPO2008; EPPO 2014; CABI 2016).
Most recently, it has been reported present as a serous pest in the Caribbean islands of
Guadeloupe, Martinique and Dominica (DNO 2015; CABI 2016; ALA 2016). These data
indicate that SS was probably present in Fiji by at least 2008.
12 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Figure 3. Pink hibiscus mealybugs (Maconellicoccus hirsutus) on beach hibiscus, vau (Hibiscus tiliaceus) leaves
(left) and on the growing tip of a hibiscus hybrid (Hibiscus rosa-sinensis) (right), Suva, Fiji, May 2017.
The PHM is one of about nine species in the genus Maconellicoccus, which is, like DWF
and SS, probably of East Asian and/or possibly tropical Australian origin, where five of
the nine species are found. Like the WFA, the PHM is the only species of its genus that
has now spread worldwide throughout the tropics and subtropics (Meyerdirk et al. 2001;
OEPP/EPPO 2005), and is now considered a serious pest in tropical and subtropical regions
of Africa, Southeast Asia and Northern Australia and more recently in the Caribbean,
where it was first reported present in Granada 1993 and subsequently spread to most
islands in the region (Serrano et al. 2001), as well as to Florida, California and Louisiana,
and was reported from Jamaica for the first time in 2007 (Warner and Scheffrhan 2002).
Although it was first reported present in the Pacific in Hawai’i, Guam, Palau, Federated
States of Micronesia, Solomon Islands, Vanuatu, Tonga, Samoa and Tuvalu by 2004 (CABI/
EPPO 2004; OEPP/EPPO 2005), it was first reported from Fiji by Hodgson and Lagowska
in 2011. Given their close association and the similar times of reported presence in the
Caribbean and parts of the Pacific, it is not implausible that the WFA and the PHM were
introduced together into some of these locations.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 13
WFA are strongly attracted to sugar and sweet foods and feed voraciously on protein, es-
pecially tinned fish, pieces or blood from meat that are left on kitchen counters or tables
and dead insects, such as cockroaches and moths. They are most commonly found forag-
ing along branches and trunks of trees and shrubs and on leaves and flowers, in symbiosis
with sap-sucking and honeydew producing insects, such as mealy bugs, scales and aphids.
Large groups of foragers also leave nests to search for new food sources, after which they
lay trails of pheromones between the food sources and nests to lead new recruits to the
food sources, with some trails being observed for months at a time. Outside homes and
other structures, foragers tend to follow lines, such as along edges of wall panelling, doors
or window frames or other paths that eventually lead to small openings to the interior,
where ants that enter become more noticeable to occupants, frequently finding their
way inside walls or ceilings where they follow electrical cables and emerge into rooms,
especially kitchens and bathrooms, where liquid, sugars, proteins and other foods can be
found (Warner and Scheffrhan 2002).
14 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
As such, the WFA is considered a serious hard-to-control nuisance and a health concern
by home owners, businesses, tourism facilities and other institutions; and a particularly
devastating agricultural and horticultural pest because they feed in large numbers on
plant nectars and “farm”, protect, and feed on the honeydew produced by sap-sucking
insects, such as mealybugs, scale insects and aphids, which also help promote other
plant diseases, such as black sooty moulds (BSM) that feed on honeydew residues on
the leaves. As the ants move from one plant to another, the mealy bugs are transported
with the ants to other plants (Warner and Scheffrhan 2002; JIS 2007; ISSG 2009; Wetterer
2013). Ants also interfere with natural and introduced biological control agents by attack-
ing parasites and predators, thus further protecting mealybugs and other sap-sucking
insects (Bugs for Bugs 2015a).
In Florida. where by 2005 WFA had spread to all southern and central countries, it was
predicted that within the “next few years” the ant “will likely saturate urban and suburban
habitats in central and south Florida . . . and possibly spread throughout the state”, where
one of the most important means of spread appeared to be via transportation of infested
residential landscaping plants and materials (Warner and Scheffrhan 2002; Warner et al.
2016). This is borne out by our experience in Fiji. Unlike some other alien tramp ant spe-
cies, WFA also seems to be able to colonise intact native forest, as of 2013, it had become
the dominant arboreal ant in numerous areas of Florida and the West Indies where it can
impact native plant and animal species (Wetterer 2013), something that must be further
investigated as a grim prospect in Fiji.
Seychelles scale is found on leaves, stems, twigs, flower buds, fruit and roots, most com-
monly or on the underside of leaves near the lower midribs and on terminal stems. Heavy
infestations lead to yellowing of leaves, abnormal leaf fall, death of young shoots, stunting
and dwarfing and, in some cases, the death of entire plants.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 15
Honeydew is produced by SS when feeding on the phloem, which is in turn fed upon by
ants and fungi, the later which forms the black sooty moulds that infest the topsides of
leaves and the fruit of many host plants, thus limiting photosynthesis and making the
plants and the fruit unattractive. Studies also show that the serious damage caused by
mealy bugs and scale insects, which can lead to the death of plants, may be due more to
the toxic substances in their saliva that are injected into the plant tissue during feeding
(ALA 2016; CABI 2016 Jackson 2016).
Between 1995 and 2012, SS was intercepted 30 times on a variety of hosts at U. S. ports-
of-entry; some of the countries of origin along with genera of infested host plants that
were intercepted include: American Samoa (Alyxia), Cambodia (Mangifera), Cook Islands
(Alyxia), Fiji (Murraya), India (Citrus, Murraya), Indonesia (Citrus), Malaysia (Nephelium),
Mexico (Leucaena), Philippines (Arachnis, Artemisia, Citrus, Codiaeum, Lagerstroemia,
Lansium, Musa, Nephelium, Psidium), Singapore (Euphorbia) and Tahiti (Gardenia, Mangif-
era, Musa, Psidium) (ALA 2016; CABI 2016). Most of these are genera also infested in Fiji.
The most relevant long-term study of the impacts of SS is from the isolated Indian Ocean
atoll of Aldabra, where SS was first discovered in 1968 and where, by 1975, Renvoiz’s
(1975) study of all the main vegetation types showed that of 65 infested woody species,
27 were heavily infested and 38 lightly infested. He predicted a subsequent loss of one-
third of the species in mixed scrub and up to two-thirds reduction in the entire vegetation
cover. Later studies of 55 woody plant taxa showed that by 1975, when SS reached peak
densities, 21 species were ‘heavily’ infested and 21 ‘lightly’/’moderately’ infested (76%
of all species). Interestingly, by the end of 1978 no species were considered to be heav-
ily infested but 37 remained ‘lightly’/’moderately’ infested, with the state of vegetation
showing negligible change between the 1976/77 and 1978 surveys (Hill and Newbery
1980). Studies in the late 1970s, did show, however, that in association with ants, SS had
seriously threatened some woody species including the small indigenous tree, Euphorbia
pyrifolia, the widespread coastal shrub, Scaevola taccada, and the mangrove, Avicennia
marina (Hill and Blackmore 1980, Newberry 1980a, b, c). The studies showed that infested
E. pyrifolia plants produced about one-third of the leaves and had a growth rates less than
50% of non-affected plants (Newberry 1980a); that SS was the major cause of reduced
leaf production by about 60% and the main cause of the death of up to 50% of the stud-
ied Scaevola plants. The studies also showed that SS densities were greatest on senescing
leaves with infestations increasing with distance from the sea; and that SS infestation
were most serious on older Avicennia mangroves, many which were subjected to poor
drainage, which may have increased their susceptibility to attack and possibly also be-
cause salt secretion on the leaves of young trees served as a barrier to the establishment
of SS (Newberry 1980abc). Most recently, 1983 surveys showed that, although the overall
abundance of SS had changed little between 1978 and 1983, its spatial distribution over
the atoll had markedly changed with the level of infestation having risen in the SE, where
tree mortality was largely density independent, but had remained low in the NW, where
tree mortality was more density dependent; and several susceptible host tree species had
showed ten-fold or higher median infestation levels in the SE of the atoll (Newbery and
Hill 1985).
16 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
In American Samoa, since the 1990s, repeated severe outbreaks of SS reportedly heavily
impacted breadfruit and other plants on Ta’u in American Samoa (Schmaedick 2007).
There were no available reports of SS infestations and resultant damage from other Pacific
Island countries.
If you think fire ants and armadillos are terrible exotic, invasive pests, just wait until
you have a houseplant infested with pink hibiscus mealybug (PHM). The damage
this insect does to plants makes them look like weed killer was sprayed on them
(Reeves n.d.).
The PHM is considered a serious pest in both tropical and subtropical regions including
Africa, Southeast Asia, northern Australia and most recently the Caribbean, the south-
eastern USA and the Pacific Islands where it attacks food plants, tree crops, vineyards,
ornamentals, weeds and forest trees, as well as attacking indoor and glasshouse plants.
In 1995, in Grenada and Trinidad and Tobago, it reportedly caused over US$150 million
direct damage in addition to export losses and property and environmental damage
(Meyerdirk et al. 2001; RADA 2015; Bugs for Bugs 2015); and it was estimated that PHM
could cause losses of $750 million if it were to spread across the southern USA (OEPP/
EPPO 2005). The destruction has not been as serious in Hawaii, where it seems that its
natural enemies from South Asia or Australia were fortuitously introduced along with it
(Meyerdirk et al. 2001).
As stressed above, like SS, PHM is dispersed, tended and protected from its predators by
ants, by WFA in Fiji, which feed on the sap or honeydew produced by the PHM. It is also
spread on plants, fruits and other plant parts by humans and by other animals and wind.
PHM breeds rapidly and, even without a male, one mature female can lay over 600 eggs
which hatch in in 3—8 days. Both the eggs within the white egg sacs, which are initially
orange but turn pink on maturity, as well as the immature and adult females are pink in
colour. The newly hatched mealybug nymphs, which are known as crawlers, settle on host
plants and start their development that takes 10 to 25 days. Under optimum laborato-
ry conditions PHM can produce 15 generations per year and 10 generations per year in
the subtropics, where, if there is a cold or winter season, all growth phases hibernate or
remain quiescent, remaining hidden in cracks in the bark, stems, in fruit bunches or in the
soil, until it is warmer again and host food plants return or bear flowers and fruit (APIS
1996; Meyerdirk et al. 2001; JIS 2007, RADA 2015).
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 17
Although adult female PHM are not particularly mobile, the nymphs, crawlers and males
can migrate with air currents; and the females, nymphs and crawlers are mobile enough
to move from host to host within infested areas. It is often hard to see, unless you check
closely because it is most commonly found on the tender undersides of leaves, commonly
concentrated along the lower part of the leaf midrib near the petiole and may be hidden
in cracks/crevices, such as in the bark, on spathes (e.g., on palms), within folded petioles
or stems of leaves or within the shrivelled growing tips or flowers of plants such as hibis-
cus. Although they prefer the growing tips, flowers buds and tender parts of host plants,
the crawlers also infest other plant parts where they develop into winged male and
wingless female adults that migrate to the lower parts of host plants as the apical portions
wither away. Although adult females normally lay their eggs on the apical or terminal
parts of plants, when the weather get colder eggs are laid in more sheltered places on the
plant. Because of these cryptic characteristics and because ants protect it from natural
enemies, effective chemical control of PHM is very difficult given the diversity of its hosts,
its rapid reproduction and ease and rapidity of spread (Meyerdirk et al. 2001; JIS 2007,
RADA 2015).
Like SS, during feeding, PHM injects toxic poison into plants which causes malformation,
distortion, shrivelling and stunting of leaves and shoots, and in many cases ultimately
death of the plant. On heavily infested plants, such as hibiscus (after which the PHM is
named), the infested leaves becoming twisted or crinkled leading to “bunchytop”, growth
is stunted and flowers either do not form or dry up, remain in a stunted or shrivelled state
or, if directly infested, flowers may be small or drop off. Similar but less distinctive damage
may occur on the trunks, stems, leaves, flowers, fruits and even roots systems of other
culturally and ecological plants. On some plants, such as guava and gardenia, a heavy,
black sooty mould which feeds on the PHM’s honeydew secretions develop on the leaves
(APHIS 1996; Meyerdirk et al. 2001; JIS 2007; NISIC 2012), which inhibits photosynthesis
and makes the affected plants very unattractive which serves as a disincentive to planting
vulnerable ornamental species in home gardens, botanical gardens, tourism facilities and
other areas.
18 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
As stressed above, a range of other ant-associated mealybugs, scale insects and other
sap-sucking insects and pathogens, some quite common, are found on the same host
plants as ants and SS and PHM (Table 2). Most common are coconut or spiked mealybug
(CMB) (Nipaecoccus nipae)(Fig. 4), first reported from Fiji in 2009 (Pestnet 2006); and
probably citrus mealybug (CTM) (Planococcus citri)(Fig. 5), which was first reported pres-
ent in Australia in the 1980s and from most Pacific Islands in 1999 (Watson 2016). Like SS
and PHM, both are very polyphagous sap-sucking insects that are found on many of the
same host plants as SS and PHM (e.g., citrus, guava, banana, papaya, soursop, bullock’s
heart, red ginger, heliconias, ginger, gardenia, Indian mulberry, coconut palm and a range
of other palms, including the endemic, Fiji fan palm (Pritchardia pacifica).
Also common are a number of scale insects, including coffee brown scale (Saisettia coffei))
(Fig. 6), the adult which has a shell- or coffee-bean shape and shiny brown colour, which is
found on soursop, breadfruit, guava, citrus, coffee, and cycad (Tsatsia and Jackson 2016);
nigra scale (Parasaisettia nigra), another a polyphagous scale with an elongate slightly
convex oval body, ranging in colour depending on the host from translucent yellow to
brown or black with no obvious wax covering, the distribution of which is now almost
cosmopolitan and has been recorded on hosts from over 94 plant families (Mau and
Kessing 2007; Idtools 2014); brown soft scale (BSS) (Coccus hesperidum); pyriform scale
(PS) (Protopulvinaria pyriformis), a light brown scab-like scale surrounded by a thin white
waxy fringe, another polyphagous species that is particularly common on Cordyline (Fig.
6), which was first reported from Western Australia in mid-2015, but which was intercept-
ed at US ports of entry between 1995 and 2012 on host plants originating in all regions
except Australasia, but including Hawaii, Micronesia, Asia and most of South America and
the Caribbean (IDtools 2013; NSW DPI 2015); and circular scale (Chrysomphalus aonidum),
which can be mistaken for juveniles of other scales species, which especially affects citrus
trees, commonly leading to death of the trees (NISIC c. 2015). Like the other mealybugs
and scale insects, these damage plants by direct feeding and injecting toxins that cause
leaves to be spotted, deformed, wilting or dying early and the fruits to be smaller and
produce honeydew that encourages the development of black sooty moulds (Table 2).
Fig. 6. Tree cassava or Ceara rubber tree (Manihot carthagenensis ssp. glaziovii) leaves, Muanikau, Suva, Sep-
tember 2017 (left) and white ixora, sinu ni vavalagi (Ixora finlaysoniana) trunk, Governors Museum Themed
Restaurant, Suva, May 2007 (right) both infested with coffee brown scale (Saessetia coffeae) tended by
white-footed ants (Technomyrmex sp.), September 2017.
20 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 7. Ti-plant, vasili (Cordyline fruticosa) leaf infested by pyriform scale (Protopulvinaria pyriformis), Mac-
Gregor Rd., Suva (left) and fish-poison tree, vutu rakaraka (Barringtonia asiatica) leaf base infested by both
circular scale (Chrysomphalus aonidum)(below) and coffee brown scale (Saisettia coffeae), Queen Elizabeth
Dr., Suva (right), both tended by white-footed ants, August 2017.
The breadfruit or Egyptian mealybug or Egyptian fluted scale (Icerya aegytiaca) has also
been recorded from Fiji, Samoa and is reportedly widespread in Micronesia and a major
pest in Kiribati and atolls of the Federated States of Micronesia. It is possibly still present,
but now rare. Found mostly on breadfruit, it is also found on avocado, banana, citrus,
jackfruit, mango, soursop, taro, ornamentals and occasionally on giant taro, pandanus,
and young coconuts. On breadfruit, the mealybugs, like the current infestation of SS, are
found along the midribs and larger veins on the undersides of the leaves and also on the
fruit. They suck the sap from the leaves, and heavy infestations cause the leaves to dry up
and die (Jackson 2015). In Fiji, they seem to have been replaced by the super-dominant
SS.
Other ant-associated invasive insects include spiralling white fly (Aleurodiscus dispersus)
(Fig. 8) and aphids, possibly the brown citrus aphid (Toxoptera citrida), one of the world’s
most serious citrus pests (Halbert and Brown 2014) and some other aphids, including an
unidentified yellow aphid, all of which are tended my WFA. A serious problem, in addition
to sucking sap and injecting toxins into host plants, is that aphids and some of the other
sap-sucking insects spread viruses and other diseases, such as the aphid transmitted cit-
rus tristeza closterovirus (CTV) (Halbert and Brown 2014), with one of the most devastat-
ing citrus crop losses ever reported following the introduction of brown citrus aphid into
Brazil and Argentina, where 16 million citrus trees on sour orange rootstock were killed by
CTV (Carver 1978). Although the root cause has not been identified in this survey, there
are numerous citrus trees, currently infested by WFA and SS and other associated pests
that have been seriously defoliated or have almost died, possibly due to such disorders
that clearly started to affect the trees before or during the onset of the current ant-mealy-
bug bio-invasion.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 21
Fig. 8. Spiralling white fly (Aleurodiscus dispersus) infestation on the underside of a kassod tree (Senna siamea)
leaf, USP Upper Campus, Suva, September 2017 (left); and on the underside of a guava (Psidium guajava) leaf,
Beachhouse Resort, Nadroga, May 2017 (right), the latter of which is also infested by Seychelles scale tended
by white-footed ants.
Other widespread pathogens associated with the WFA-led invasion included a number of
fungal diseases, including black sooty moulds (BSM)(Fig. 9) and plumeria rust (PR). BSM,
which are caused by Capnodium and Tripospermum spp. and a number other species,
grow on the honeydew produced by the sap-sucking insects and cover the upper leaf
surfaces of many plants, such as guava, citrus, mango, soursop, frangipani, gardenias
and palms, with a dense, black, soot-like deposits that inhibit photosynthesis and cause
leaf yellowing and loss and stunted plant growth (Kohler et al., 1997; Tsatsia and Jackson
2016d).
Fig. 9. MacArthur palm (Ptychosperma macarthuri) with the flowers spathes infested with coconut mealybugs
(Nipaecoccus nipae) tended by white-footed ants (Technomyrmex sp.) and the leaves infested with black sooty
mould (BSM), Service St., Suva, June 2017 (left); and leaves of beach cordia, nawanawa (Cordia subcordata)
infested with BSM, USP Lower Campus, Suva, May 2017 (right).
22 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 10. White frangipani, bua ni vavalagi (Plumera obtusa) infested by frangipani or plumeria rust (Coleosper-
mum plumeriae) and Seychelles scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.),
Beachhouse Resort, Nadroga (left); and red frangipani (Plumeria rubra) infested by frangipani or plumeria
rust, USP Lower Campus, Suva (right), both May 2017.
Table 3. Total numbers of families, genera and species of plants observed to be affected
by DFA and associated mealybug, scales and other insects and pathogen infestations in Fiji
based on field studies by R. Thaman from May—September 2017.
Table 4. Number of host plant species from specified genera and families observed to be
affected in Fiji by WFA in symbiotic infestations along with SS, PHM, other mealybugs or
scales, other sap-sucking or damaging insects and pathogens, based on field studies by
the author from May—August 2017. Ferns and fern allies and gymnosperms are listed first
followed by dicotyledons and monocotyledons (greater detail is provided in Appendix I,
which is a listing of the number of species within different genera and families, in the ap-
proximate order of severity of infestation and number of species infested, along with the
scientific, common and local Fijian names and notes on selected species; and Appendix II
is a listing by families in alphabetical order, including information on severity of observed
infestations, infesting organisms, parts infested and the study locations at which infesta-
tions were observed) (Note: * indicates that the number of species within a given genus,
e.g. Hibiscus, includes a number of distinct hybrid cultivars and species).
DOCOTYLEDONS (217 species from 137 genera and 47 families) (Note: numbers
after family name are the number of genera in the family with the total species in paren-
theses, e.g. 20 genera and 25 species for Fabaceae)
Fabaceae 21 (26): Flemingia (2), Bauhinia (2), Cassia (2), Vigna (2), Calliandra (2), Intsia
(1), Phaseolus (1), Derris (1), Tamarindus (1), Arachis (1), Psophocarpus (1), Centrosoma
(1), Senna (1), Samanea (1), Inocarpus (1), Cynometra (1), Peltophorum (1), Brugmansia
(1), Adenanthera (1), Pterocarpus (1), Millettia (1), Gliricidia (1)
Malvaceae 7 (17) Hibiscus (9), Abelmoschus (3), Thespesia (1), Abutilon (1), Malvaviscus
(1), Urena (1), Gossypium (1)
Rubiaceae 5 (14): Gardenia (3), Ixora (7), Coffea (1), Mussaenda (2), Pentas (1)
24 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Rutaceae 4 (12): Citrus (8), Euodia (1), Murraya (2), Micromelum (1)
Euphorbiaceae 7 (11): Acalypha (3), Euphorbia (2), Excoecaria (1), Aleurites (1), Codiaeum
(1)
Solanaceae 5 (11): Solanum (4), Capsicum (4), Brunfelsia (1), Brugmansia (1)
Endospermum (1), Macaranga (1), Manihot (1), Jatropha (1)
Moraceae 4 (10): Ficus (6), Artocarpus (2), Broussonetia (1), Morus (1)
Myrtaceae 3 (9): Syzygium (6), Psidium (2), Eugenia (1)
Acanthaceae 9 (10): Pachystachys (2), Hemigraphis (1), Sanchezia (1), Blechum (1), Eran-
themum (1), Asystasia (1), Odontonema (1), Pseuderanthemum (1), Thunbergia (1)
Asteraceae 7 (7): Eleutheranthera (1), Synedrella (1), Mikania (1), Ageratum (1); Cyanthil-
lium (1), Crassocephalum (1), Youngia (1)
Annonaceae 3 (7): Annona (4), Polyalthia (2), Cananga (1)
Verbenaceae 3 (6): Clerodendrum (4), Duranta (1), Vitex (1)
Cucurbitaceae 5 (5): Luffa (1), Cucurbita (1), Citrullus (1), Coccinia (1), Cucumis (1)
Lamiaceae 3 (4): Ocimum (2), Tectona (1), Congea (1)
Convolulaceae 3 (5): Ipomoea (3), Merremia (2)
Lythraceae 3 (4): Lagerstroemia (2), Cuphea (1), Punica (1)
Oxalidaceae 2 (4): Averrhoa (2), Oxalis (2)
Piperaceae 1 (3): Piper (3)
Anacardiacee 3 (3): Mangifera (1), Dracontomelon (1), Spondias (1)
Lecythidaceae 2 (3): Barringtonia (2), Couroupita (1)
Sapindaceae 3 (3): Acer (1), Filicium (1), Pometia (1)
Phyllanthaceae 3 (3): Bischofia (1), Flueggea (1), Phyllanthus (1)
Combretaceae 2 (2): Quisqualis (1), Terminalia (1)
Apocynaceae 2 (3): Plumeria (2), Alyxia (1)
Araliaceae 2 (4): Polyscias (3), Schefflera actinophylla (1)
Casuarinaceae 2 (2): Gymnostoma (1), Casuarina (1)
Boraginaceae 1 (2): Cordia (2)
Clusiaceae 2 (2): Calophyllum (1), Garcinia (1)
Urticaceae 2 (2): Boehmeria (1), Pilea (1)
Passifloraceae (2): Passiflora (1), Turnera (1)
Lauraceae (1): Persea (1)
Goodeniaceae 1 (1): Scaevola (1)
Sapotaceae 1 (1): Manilkara (1)
Thymelaeaceae 1 (1): Phaleria (1)
Amaranthaceae 1 (1): Amaranthus (1)
Santalaceae 1 (1): Santalum (1)
Rosaceae 1 (1): Rosa (1)
Basellaceae 1 (1): Basella (1)
Apiaceae 1 (1): Eryngium (1)
Moringaceae 1 (1): Moringa (1)
Bignoniaceae 1 (1): Mansoa (1)
Chrysobalanaceae (1): Atuna (1)
Gesneriaceae (1): Chrysothemis (1)
Polygonaceae (1): Antigonon
Caricaceae (1): Carica (1)
Vitaceae (1): Vitis (1)
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 25
The severity of impacts of the ant-mealybug juggernaut is clearly variable over space
and time, with some localities, individual gardens and different plant species, genera or
families more seriously affected than others. Whereas some plants are infested with thou-
sands of ants, mealybugs and other sap-sucking insects, others show only low, limited
or incidental presence or no evidence at all of infestation or damage. Other plants have
clearly been invaded by ants, but with little clear evidence yet of other insect or patho-
gens. On infested plants, ants can be seen around the bases of the plants and in a steady
streams moving up and down the trunks, branches or stems, something that can be quite
dramatic when seen against the lighter coloured bark of trees such as vesi, Fiji’s national
tree (Intsia bijuga)(Fig. 11).
Fig. 11. Borneo teak, vesi (Intsia bijuga), Fiji’s national tree, infested by white-footed ants (WFA)(Technomyr-
mex sp.) on the bark (left) and tending Seychelles scale (Icerya seychellarum) on the leaves (right), both USP
Lower Campus, Suva, May 2017.
Also seen occasionally on infested plants are other insects that as known predators or
parasites of mealybugs, such as the mealybug ladybird, also known mealybug destroyer
(Cryptolaemus montrouzieri), an endemic species from Australia that was introduced into
the US mainland, Hawai’i and New Zealand over 100 years ago and into Fiji, Vanuatu,
Solomon Islands, the Cook Islands and other Pacific Islands prior to WW II, in all cases to
control mealybugs and scale insect infestations (Bugs for Bugs 2015b; Martin 2016).
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 27
1. Plant form or types, with ferns being virtually unaffected, none of the common
gymnosperms, apart from cycads, being infested, and trees, shrubs, lianas and
woody plants being preferred hosts over smaller herbaceous species, with virtual-
ly no grasses (Poaceae) or sedges (Cyperaceae) being affected;
2. Plant origin and cultivar diversity within a given species, with species and culti-
vars from, or developed in, Africa or Asia, such as the “Malaysia Dwarf” coconut,
showing more resistance than species or cultivars from other regions, such as
tropical America, or island endemic plant or traditional Pacific Island food plant
cultivars that evolved or were selected in the absence of these IAS of mainly Asian
or Australasian origin;
3. Whether a plant is deciduous, drops its leaves and regrows them again after the
infestation subsides, e.g. Terminalia catappa and Erythrina variegata;
The degree of infestation may also vary seasonally and with the weather, with infestation
being more virulent during warmer months or during dry spells, something that is sup-
ported by the tendency of the PHM to hibernate or hide during the cold season in colder
climates, such as in Florida, and remerging when plants have new growth and are in flow-
er or with fruit (Meyerdirk et al. 2001). This has been supported by studies in the early
2000s showing positive correlation between temperature and humidity and dew point
and the population density of SS on the host plant Dodonaea viscosa in Egypt (Mesbah et
al. 2012).
28 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
4.1 Ferns
As stressed above, checking most of the common ferns found in the study sites (e.g.
Microsorum, Davallia, Pteris spp. and the tree fern, balabala (Cyathea lunalata), only one
fern, a black-stiped edible fern, ota loa (Tectaria latifolia) was seen with a low or inciden-
tal level of infestation.
4.2 Gymnosperms
Although, none of the common larger indigenous gymnosperms (e.g. Agathis, Dacrycar-
pus, Dacrydium, Retrophyllym and Podocarpus spp.) and introduced gymnosperms (e.g.
Araucaria, Cupressus, Pinus and Thuja, spp.) showed any signs of infestation in any of the
study sites, among the most heavily infested of all taxa were cycads, including the indig-
enous queen sago palm or cycad, logologo (Cycas seemannii) and king or Japanese sago
palm (C. revoluta)(Fig.13), with fronds becoming very chlorotic or dying (Fig. 12). All were
infested with varying combinations of Seychelles scale (SS), coconut mealybug (CMB),
coffee brown scale (CBS) and Asian cycad scale (ACS) (Table 2, Appendix II).
Fig. 12. Queen cycad, logologo (Cycas seemannii) plant with dead fronds (left) and underside of a frond infest-
ed with Seychelles scale (SS) (Icerya seychellarum) and with coffee brown scale (Saessetia coffeae) tended by
white-footed ants (Technomyrmex sp.)(right), both Nailuva Rd., Suva, June 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 29
Fig. 13. Japanese or king cycad (Cycas revoluta) with dead fronds, USP Upper Campus (left); and undersides
of fronds infested with both San Jose scale (SJS) (Quadraspidiotus perniciosus)(light with nuclear dark-brown
central spot) and coffee brown scale (Saessetia coffeae), all tended by white-footed ants (Technomyrmex sp.)
(right), both Nailuva Rd., Suva, June 2017.
The vulnerability of cycads to scale insects is strongly supported by evidence from both
Florida and Guam, where the rampant spread of the recently introduced cycad scale
insect (CSI) (Aulacaspis yasumatsui), which is possibly native from Thailand to S. China,
devastated cycad populations. In Florida, where CSI was first found in 1996, both C.
revoluta and C. seemannii were heavily infested, as were extensive ornamental plantings
of other cycads, including irreplaceable collections in Florida’s Fairchild Tropical Garden
and Montgomery Botanical Centre, both of which have important world collections of
cycads, including rare and endangered species that were threatened by the scale, which
also posed an additional threat to the large concentration of nurseries in southern Florida
that grow and ship cycads throughout the US and internationally (Weissling and Howard
2017). More recently, in 2003, CSI was discovered and spread rapidly throughout Guam,
infesting both horticultural cycads and the endemic cycad, Cycas micronesica - a culturally
iconic dominant forest and savanna plant. Like many island endemics, C. micronesica was
particularly susceptible to cycad scale insect, with 100% mortality in infested areas, with
the rapid spread threatening the entire population of 1.5 million trees (Campbell 2005;
Moore et al. 2005).
30 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
4.3 Dicotyledons
Of the 47 infested dicotyledons families, along with the genera and species affected,
shown in Tables 3 and 4 and Appendices I and II, the 22 most seriously affected families,
in approximate order based of severity of infestation of individual species, numbers of
species affected, levels of infestation, damage to plants and the cultural, economic and
environmental costs represented by their loss, are:
Rutaceae (Citrus or Rue Family): All of Fiji’s important citrus trees (Citrus spp.)(Fig. 14)
and island musk (Euodia hortensis) (Fig. 15) are seriously infested, almost exclusively
by SS, with many also heavily infested with black sooty mould (BSM), which may be an
indication that the more competitive SS has replaced citrus mealybug (CTM) and other
species known to infest Citrus spp.
Fig. 14ab. Pommelo, moli kana (Citrus maxima), an important threatened multipurpose Fijian cultural plant
infested with Seychelles scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.), Borron
Rd., Suva, August 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 31
Fig. 15. Island musk, uci (Euodia hortensis), a native sacred Fijian plant, the leaves and flowers of which are
used medicinally, in garlands (salusalu) and to perfume coconut oil, the underside of the leaves and stems of
which are infested with Seychelles scale (Icerya seychellarum) tended by white-footed ants
(Technomyrmex sp.), National Trust of Fiji, May 2017.
Moraceae (Mulberry or Fig Family): Important food and multipurpose plants that are
infected include, breadfruit, uto (Artocarpus altilis)(Fig. 16), an important seasonal staple
food and multipurpose plant; jackfruit, uto ni Idia or katthar (Artocarpus heterophyllus),
an important Indian staple food and; paper mulberry (Broussonetia papyrifera)(Fig; 17),
an important women’s crop that is used for making tapa (bark) cloth. Also infested, mainly
by SS, are a wide range of culturally important indigenous and introduced figs and ban-
yan trees (Ficus spp.), with the universally most highly infested species being the Fijian
endemic figs, Barclay’s fig, losilosi (F. barclayana)(Fig. 18), which is also seen infested with
citrus mealy bug (CTM), and Fiji fig, lolō (F. vitiensis) (Fig 19).
32 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 16. Breadfruit, uto (Artocarpus altilis), an important staple food crop and multipurpose tree, the under-
sides of the leaves which are infested by Seychelles scale (Icerya seychellarum) tended by white-footed ants
(Technomyrmex sp.), Laucala Bay Rd. Suva, June 2017.
Fig. 17. Paper mulberry, masi (Brousonnetia papyrifera), an important handicraft plant, the inner bark or bast
fibre of which is used to make tapa cloth (masi), with the leaves (left) and stems (right) infested by Seychelles
scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.), Laucala Beach Estate, Suva, June
2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 33
Fig. 18. Barclay’s fig, losilosi (Ficus barclayana), an important endemic Fijian medicinal plant, infested by
Seychelles scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.), Nailuva Rd. (left) and
Knolly St., Suva (right), May 2017.
Fig. 19. Fiji fig, lolō (Ficus vitiensis), an important endemic Fijian medicinal plant, emergency food plant and
fruit bat and bird food, infested by Seychelles scale (Icerya seychellarum) tended by white-footed ants (Tech-
nomyrmex sp.), Nailuva Rd., Suva, August 2017.
34 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 20. Common hibiscus, senitoa (Hibiscus rosa-sinensis) with the terminal buds and immature shrivelled
flowers infested with pink hibiscus mealybugs (Maconellicoccus hirsutus) tended by pink hibiscus mealybugs
(Maconellicoccus hirsutus), Governors Museum Themed Restaurant, Suva, May, 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 35
Fig. 21. Hybrid hibiscus with the Fiji endemic, Storke’s hibiscus (Hibiscus x storkei), the dying stunted flowers
and growing tips of which are infested with pink hibiscus mealybugs (Maconellicoccus hirsutus) tended USP
Lower Campus, Suva, May, 2017.
Figure 22. Beach hibiscus, vau (Hibiscus tiliaceus) leaves infested with pink hibiscus mealybugs (Maconellicoc-
cus hirsutus), Oceania Centre for Arts and Culture, USP, Suva, August 2017 (left above and below); and a young
beach hibiscus trunk infested by Seychelles scale (Icerya seychellarum) tended by white-footed ants (Techno-
myrmex sp), Nailuva Rd., Suva, Fiji, May 2017.
36 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Also affected, but mainly by SS, are okra, roselle and hibiscus spinach or bele (Abelmo-
schus spp.); and the important cultural medicinal and woodcarving plant, Thespian’s
tree, mulomulo (Thespesia populnea) is also affected by SS, citrus mealybug (CTM), and
possibly by PHM. The weedy species, Pink hibiscus bur (Urena lobata), was infested with
SS, what appears to be nigra scale (NS) and white fly (SWF). Bele (Ablelmoschus manihot)
is also seriously affected by the Abelmoschus shoot borer (ASB) (Earias vittella);
Fabaceae (Legume, Bean or Pea Family): Also infected are also important leguminous
food plants (Vigna, Phaseolus, Psophocarpus and Inocarpus spp.)(Fig. 23), nitrogen-fixing
legumes (Flemingia, Centrosoma, Vigna and Calliandra spp.), ornamental species (Albizia,
Bauhinia, Cassia, Samanea and Senna spp.)(Fig. 24) and Fiji’s culturally important national
tree, vesi (Intsia bijuga), are all infested at variable levels by SS, with Bauhinia and Cassia
spp., also infested with black sooty mould (BSM).
Fig. 23. French beans (Phaseolus vulgaris) leaves (left), Laucala Beach Estate (left) and longbeans (Vigna ses-
quidpedalis) stems and fruit, Flagstaff (right), both infested by Seychelles scale (Icerya seychellarum) tended
by white-footed ants (WFA), Suva, July 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 37
Fig. 24. Golden shower tree, vaivai (Cassia fistula) leaves infested by Seychelles scale (Icerya seychellarum)
tended by white-footed ants (WFA) (Technomyrmex sp.), USP Upper Campus, Suva, August 2017 (upper and
lower left), showing flowers of unaffected tree (right).
Myrtaceae (Myrtle Family): A wide range of culturally useful food and multipurpose
plants (Syzygium spp.) and guavas (Psidium spp.), particularly common guava (P. guajava)
(Fig. 25), are seriously infested by SS, coconut mealy bug (CMB), spiralling whitefly (SWF)
and black sooty mould (BSM), with some cherry guava (P. cattleyanum), also infested with
SS and SWF. Also heavily infested with SS were pomegranate (Punica granatum) and leba
(Syzygium neurocalyx)(Fig. 26), a threatened Fijian medicinal and garland plant.
38 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 25. Guava, quwawa (Psidium guava) leaves infested with coconut mealybugs (Nipaecoccus nipae), Nas-
ese, Suva, June, 2017 (left); and infested with both citrus mealybug (Planococcus citri) and spiralling whitefly
(Aleurodiscus dispersus), Beachhouse Resort, Nadroga May 2017 (right), both tended by white-footed ants
(Technomyrmex sp.)
Fig. 26. Leba (Syzygium neurocalyx), a rare endemic sacred Fiji plant, the fruit of which is used to make
garlands (salusalu)(left) and the underside (right) of a leaf infested by Seychelles scale (Icerya seychellarum)
tended by white-footed ants (Technomyrmex sp.), Laucala Beach Estate, Suva, Fiji, May 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 39
Annonaceae (Custard Apple Family): Soursop (Fig. 27) sweetsop and bullock’s heart (Fig.4
, above) (Annona spp.), ylangylang or perfume tree (Cananga odorata) (Fig. 28), and trees
of cultural importance to both Fijian and Indian people, mocelolo (Polyalthia laddiana)
and the ashok tree, ashoka (Polyalthia longifolia) are infested by SS, with soursop and
other Annona spp. also infested with coconut mealybug (CMB), black sooty mould (BSM)
and the fruit of soursop with citrus mealybug (CTM).
Fig. 27. Soursop, seremaia (Annona muricata) fruits infested with citrus mealybug (Planococcus citri) tended
by white-footed ants (WFA), on Drew St., Suva, July 2017 (left) and at Legalega Research Station, Nadi, Sep-
tember 2017 (right, closup).
Fig. 28. Ylangylang or perfume tree, makosoi (Cananga odorata) flowers and leaves, Laucala Beach Estate,
Suva, July 2017 (left) and upper stems very heavily infested with Seychelles scale (Icerya seychellarum) tended
by white-footed ants (WFA), Nailuva Rd., Suva, December 2017 (right).
40 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Rubiaceae (Coffee or Madder Family): Important ornamental plants, the flowers of which
are used in garlands, such as gardenias (Gardenia spp.) and a range of ixoras (Ixora spp.),
mussaenda (Mussaenda spp.), Indian mulberry or noni (Morinda citrifolia)(Fig. 5 above)
and coffee (Coffea sp.) are all infested with a range of IAS including SS, PHM, CTM and CBS
and NS, with the citrus mealybug (CTM) and the coffee brown scale (CBM) and/or nigra
scale (NS), always tended by WFA. Black sooty moulds (BSM) also heavily infest most Rubi-
aceae (Fig. 29). On the positive side, also found on gardenias, ixoras and Indian mulberry
were the larvae of the mealybug destroyer (MBD), which seems to be more common in
association with citrus mealy bugs (CTM) and scale insets (CBS and NS), which it may have
been reduced in numbers before the current invasion.
Fig. 29. Cape jasmine (gardenia), jiale ni vavalagi (Gardenia jasminoides) infested with citrus mealybug (Plano-
coccus citri) tended by white-footed ants, USP Upper Campus, Suva, Sept. 2017 (left) and Tahitian gardenia or
tiare Tahiti leaves infested with black sooty mould and white-footed ants (Nailuva R., Dec. 2017) (right).
Lauraceae (Laurel Family): Avocado (Persea americana) is one of the most heavily and
widely infested species, by both SS, CMB and BSM, with many gardeners’ mature trees
dying due to high levels of infestation (Fig. 30 and Fig. 2 above).
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 41
Fig. 30. Avocado, pea (Persea americana) leaf infested by Seychelles scale (Icerya seychellarum), Desvoux Rd.,
Suva, May 2017 (left) and by coconut mealybug (Nipaecoccus nipae), USP Upper Campus, Sept. 2017 (right),
both tended by white-footed ants (Technomyrmex sp.),
Euphorbiaceae (Spurge Family): A range of ornamental and culturally useful plants (Aca-
lypha, Codiaeum, Euphorbia and Macaranga spp.) are infested, with the common orna-
mental chenille plant or red-hot poker (A. hispida) seriously affected by SS (Fig. 3 above);
the common ornamental, A. wilkesiana is affected by both SS and SWF; croton (Codiaeum
variegatum, is affected by both citrus mealybug (CTM)(Fig. 31) and nigra scale (NS); the
endemic Fiji spurge (Euphorbia fidgiana) is infested with SS; cassava (Manihot spp.) (Fig.
6 above) infested with both BCS and SWF; and the native timber tree, whitewood or
kauvula (Endospermum macrophyllum) is the only species seen affected by the longtailed
mealybug (LTM) (Pseudococcus longispinnus).
Fig. 31. Croton, sacasaca (Codiaeum variegatum) infested with citrus mealybugs (Planococcus citri) tended by
white-footed ants (Technomyrmex sp.), USP Laucala Campus, Suva, September 2017.
42 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Goodeniaceae (Goodenia Family): The culturally important multipurpose and coastal pro-
tection plant (Scaevola taccada) showed very high levels of infestation by SS in Suva (Fig.
32), which is supported by the fact that it was one of the most seriously affected native
species on Aldabra Atoll in the Indian Ocean in the 1970-80s (Newbery, 1980; Newbery
and Hill 1985).
Fig. 32. Beach half-flower or scaevola, vevedu (Scaevola taccada), an important indigenous multipurpose
cultural plant and coastal protection species, infested with Seychelles scale (Icerya seychellarum) tended by
white-footed ants (Technomyrmex sp.), Oceania Centre for Arts and Culture, USP (left) and USP Lower Campus
(right), August 2017.
Casuarinaceae (Casuarina Family): Both the endemic Fiji casuarina (Gymnostoma vi-
tiense) and common casuarina (Casuarina equisetifolia) are infested, the former at high
levels.
Fig. 33. Fiji casuarina, velau (Gymnostoma vitiensis) leaves (left) and stems (right) infested by Seychelles scale
(Icerya seychellarum) tended by white-footed ants (WFA), USP Lower Campus, Suva, August 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 43
Piperaceae (Pepper Family): Both betel pepper, paan (Piper betle) (Fig. 34), an important
Indian cultural plant the invasive spiked pepperbush (Piper aduncum), a tropical American
species, is also often heavily infested with SS, tended by WFA.
Fig. 34. Betel pepper, paan (Piper betle) the leaves of which are chewed with betelnut by South Indian and
Solomon Island residents, infested by Seychelles scale (Icerya seychellarum) and tended by white-footed ants
(Technomyrmex sp.), Nailuva Rd., August 2017.
Solanaceae (Nightshade Family): Eggplant, tomato and other edible and weedy plants,
such as chili peppers and capsicums (Capsicum spp.) are infested with SS and SWF, the
weedy shrub, prickly solanum (Solanum torvum) with SS, with eggplant being among the
most heavily infested by SS (Fig. 35).
Fig. 35. Eggplant, baigan (Solanum melongena) leaves (left) and stem (right) infested by Seychelles scale
(Icerya seychellarum)(left) and tended by white-footed ants (WFA), Fiji National Food and Nutrition Centre,
May 2017.
44 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Anacardiaceae (Cashew Family): Mangoes, often depending on the cultivar, can be heavi-
ly infested with SS, coconut mealy bug (CMB) and black sooty mould (BSM).
Acanthaceae (Acanthus Family): A range of ornamental plants are infested, mainly at low
levels by SS, the most heavily infested species being Pachystachys spp., natives to tropical
America, and Hemigraphis alternata, a common groundcover native to Malaysia that has
become a minor invasive in lawns and pot plants.
Convolvulaceae (Morning-glory Family): Sweet Potato and a number of weedy vines (Ipo-
moea and Merremia spp) and omemetal plants are infested by SS, with only the woodrose
(M. tuberosa), native to Mexico, showing high levels of infestation (Fig. 36).
Fig. 36. Hawaiian woodrose or yellow morning-glory (Merremia tuberosa) flower (left) and underside of leaf
infested with Seychelles scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.)(right),
USP Botanical Garden, Suva, August 2017.
Fig. 37. Healthy beach cordia, nawanawa (Cordia subcordata)(left); fruit stems infested with coffee brown
scale (Saisettia coffaea) tended by tended by white-footed ants (Technomyrmex sp.), USP Lower Campus,
August 2017 (right).
Fig. 38. Philippine glorybower (Clerodendrum quadriloculare) leaf (left) infested with coconut mealybugs
(Nipaecoccus nipae); and flowers (right) infested with citrus mealybugs (Planococcus citri), both tended by
white-footed ants (Technomyrmex sp.), USP Laucala Campus, Suva, September 2017.
46 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Lamiaceae (Mint Family): Basils (Ocimum spp,), particularly holy or sacred basil (O. te-
nuiflorum), and teak (Tectona grandis), an important plantation timber tree, the common
ornamental climber, and woolly congea or shower orchid (Congea tomentosa)(Fig. 39), are
all infested with SS, with basils sometimes also infested with spiralling white fly (SWF).
Fig. 39. Wooly congea or shower orchid (Congea tomentosa) flower (left) and leaf infested with Seychelles
scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.)(right), Beachhouse Resort, Nad-
roga, Fiji, May 2017.
Fig. 40. Maile, vono (Alyxia stellata) stem and leaves (left) and close-up of stem (right), heavily infested with
Seychelles scale (Icerya seychellarum) tended by white-footed ants (Technomyrmex sp.), Laucala Beach Estate,
Suva, July 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 47
Cucurbitaceae (Gourd or Cucurbit Family): Infested species include ridged gourd, taroi
(Luffa acutangula), pumpkin (Cucurbita moschatus) and cucumber (Cucumis sativus),
which are mainly infested with low levels of SS, but also widely affected by downy mildew
fungal diseases and viral infections, some of which do not seem to be ant-related.
Sapindaceae (Soapberry Family): fernleaf tree (Filicium decipiens), Japanese maple (Acer
palmatum) and oceanic lichi, dawa (Pometia pinnata) are infested with SS.
Other dicotyledons families with species infested at mostly low levels are shown in Table
4 and Appendices I and II. Of these, species most commonly infested include: the Tongan
national tree or flower, heilala (Garcinia sessilis)(Clusiaceae), a tree native to Fiji and ap-
parently an aboriginal introduction into Tonga: carambola (Averrhoa carambola (Oxalida-
ceae); Mascarene Islands leaflower (Phyllanthus tenellus) (Phyllanthaceae); damask rose
(Rosa x damascena); and sinu ni baravi or mataivi (Phaleria disperma) (Thymeliaceae), an
important Fijian cultural plant.
4.4 Monocotyledons
Of the 14 infested dicotyledons families, along with the genera and species listed in Table
4 and Appendices I and II, the 5 most seriously affected families in approximate order
degree of infestation based on a combination of numbers of species affected, levels of
infestation, damage to plants and the cultural, economic and environmental costs repre-
sented by their loss, along with mention of some of the important species most seriously
affected are:
Arecaceae (Palm Family): At least 21 palm species were affected, the most widely infest-
ed including the coconut palm (Fig. 41), betelnut palms (Areca catechu), Bismarck palm
(Bismarckia nobilis) (Fig. 42), fishtail palm (Caryota urens), golden cane palm (Chrysali-
docarpus lutescens), ruffled fan palm (Licuala grandis), Macarthur palm (Ptychosperma
macarthuri), bush palmetto (Sabal minor) and the indigenous or aboriginally introduced
Fiji or Pacific fan palm (Pritchardia pacifica) and a wide range of introduced ornamental
palms, most of which are infested with coconut mealybug (CMB) and/or Seychelles scale
(SS), with many species also infested with black sooty mould (BSM).
48 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 41. Coconut palm, niu (Cocos nucifera) with flower spathes and fruit infested by coconut mealy bug (Nipa-
ecoccus nipae) tended by white-footed ants (Technomyrmex sp.), Muanikau, Suva, Sept. 2017.
Fig. 42. Young Bismarck palm (Bismarckia nobilis)(left) and underside of leaf infested by coconut mealy bug
(Nipaecoccus nipae) tended by white-footed ants (WFA), USP Botanical Garden, July 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 49
Fig. 43. Golden cane palm (Chrysalidocarpus lutescens) flower spathes infested by Seychelles scale (Icerya sey-
chellarum), Tamavua, Suva, August 2017 (left) and Chinese fan palm (Livistona chinensis) frond infested with
coconut mealybug (Nipaecoccus nipae), USP Botanical Upper Campus, Suva, September 2017 (right), in both
cases tended by white-footed ants (WFA)(Technomyrmex sp.).
Araceae (Arum of Taro Family): The 14 infested Araceae species included the indigenous
forest vine, yalu (Epipremnum pinnatum); a number of introduced ornamental species,
including anthuriums (Fig. 44), philodendrons (Philodendron spp., and cultivars), arrow-
head vine (Syngonium podophyllum) and Aglaonema spp. which are infested with SS and/
or CMB; and important food plants taro, dalo (Colocasia esculenta), and American taro or
tannia (dalo ni tana) and belembe or tanier spinach (Xanthosoma spp.), which are infest-
ed with SS and/or aphids.
Fig. 44. Anthurium or flamingo flower (Anthurium andraeanum) (left) and underside of leaves (right) infested
by Seychelles scale (Icerya seychellarum) tended by white-footed ants (WFA), Vatuwaqa, Suva, July 2017.
50 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Fig. 45. Parakeet or golden torch heliconia cultivar (Heliconia psittacorum) flower (left) and underside of leaf
infested with coconut mealybug (Nipaecoccus nipae) tended by white-footed ants (Technomyrmex sp.), Nas-
ese, Suva, July 2017 (right).
Fig. 46. Lobster claw heliconia (Heliconia rostrata) flower infested with coconut mealybug (Nipaecoccus nipae)
(left) and underside of leaf infested with Seychelles scale (Icerya seychellarum), both tended by white-footed
ants (Technomyrmex sp.), Nailuva Rd., Suva, December 2017.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 51
Zingiberaceae (Ginger Family): Some 6 species are infested, the most commonly infested
of which are red ginger and variegated gingers (Alpinia spp.) and white and pink gingers
(Hedychium spp.), which are infested with SS and/or CMB, with some species also infested
with aphids.
Fig. 47. Red ginger, cevuga damu (Alpinia purpurata) infested with Seychelles scale (Icerya seychellarum) tend-
ed by white-footed ants (Technomyrmex sp.), Rifle Range, Suva, July 2017 (left) and variegated ginger (Alpinia
vittata) infested with white-footed ants, Nailuva Rd., Suva, August 2017 (right).
Poaceae (Grass Family): Although members of the grass family were uncommonly infest-
ed, bamboo was seen to be infested by the soft bamboo scale (Bambusapis bambusae)
tended by WFA and sugarcane was seen infested WFA (Fig. 48) and, at times, with citrus
mealybug (CTM) (Planococcus citri)
Fig. 48. Common giant bamboo, bitu ni vavalagi (Bambusa vulgaris) infested with soft bamboo scale (Bam-
busaspis bambusae) tended by white-footed ants (Technomyrmex sp.), USP Upper Campus, September 2017
(left) and sugarcane, dovu (Sacchraum officinarum) was seen infested by WFA tending their eggs (USP Upper
Campus, Sept. 2017).
52 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Marantaceae (Arrowroot Family): Ornamental species infested with CMB and/or SS in-
clude rattlesnake plant (Calathea crotalifera) and other Calathea spp.
Other infested monocotyledons families that have a number of species infested at mostly
low levels are listed in Table 2 and Appendices I and II).
1. Native to Africa, the Indian Ocean or South Asia, the apparent source areas of WFA,
SSS, HMB, CMB and many other sap-sucking insects, where they have possibly co-
evolved with and developed resistance to these invasive species;
2. Toxic or medicinal plants, the sap of which is milky or latex-like and unattractive or
toxic to, or that do not produce honeydew targeted by, mealybugs, scales and other
sap-sucking insects;
3. Widespread Indo-Pacific coastal species that have in-built resistance to the infesta-
tions;
Table 5. Families and species of native and introduced, mainly woody plant species that
are unaffected or almost never affected by the WFA-mealybug and sap-sucking insect
bio-invasion in Fiji, based on field studies by R. Thaman from May—September 2017
(Notes: 1) * = common species showing very limited or incidental infestation in only one
or few locations, but uninfected in almost all study sites; 2) see Appendix IV for details of
common and Fijian names, origin, characteristics of plants that may help to explain their
resistance to infestation, and main sources of information.)
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 53
Araliaceae (5): *Polyscias filicifolia, P. fruticosa, *P. guilfoylei, *P. scuttelaria, Schefflera
actinophylla, S. arboricola
Some of the more common un-infested or rarely-infested species (Appendix III) include:
• Plants native to India, South Asia or Australasia: neem tree (Azadirachta indica), an
important medicinal plant renowned for its use as an organic insecticide and insect
repellent; crape jasmine, false gardenia or chameli (Tabernaemontana divaricata), a
medicinal plant with milky sap sacred to Hindus; bush thunbergia (Thunbergia erecta);
crown flower (Calotropis gigantea), a plant with poisonous milky sap; two banyan or
fig trees, weeping banyan and the Indian rubber tree (Ficus benjamina and F. elastica);
and Queensland umbrella tree (Schefflera actinophylla), an invasive species in Fiji,
which is native from Indonesia to New Guinea and N. Australia, and dwarf Schefflera
(S. arboricola), which is native to Taiwan.
• Uninfested highly aromatic or toxic beverage plants or plants used to protect gardens
from evil spirits include kava yaqona (Piper methysticum), the important Pacific mild
narcotic social beverages and medicinal plant; Indian borage and variegated coleus,
(Plectranthus amboinicus and P. scutellarioides) and bayrum tree, sinamoni (Pimenta
racemosa), an important medicinal plant
1. Possible recent introduction of more virulent genetic strains of SS and/or the recent
introduction, spread and population explosion of the WFA, which has replaced the
less competitive and productive Technomyrmex albipes and other previously common
ants such as the longhorn crazy ant (Paratrechina longicornis), a slightly larger, fast-
er-moving ant than the WFA that emits a slight fruity odour when crushed and which
seemed to be the dominant ant in our households and kitchens two or three years
ago (Wetterer 2008; Warner and Scheffrhan 2016). This which is supported by a listing
all of the ants in association with pineapple mealy bugs throughout the world (includ-
ing South Africa, South and Central America, the Caribbean, Malaysia, Philippines,
Australia Hawaii, and Fiji), which included most of the well-known tramps ants (e.g.,
Pheidole, Camponotus, Crematogaster, Linepithema, Tetromorium Tapinoma, Paratre-
china, Solenopsis, Wasmannia spp.), but did not list WFA from any location in associa-
tion with mealybugs, except South Africa (El Wanis (John 1990; E Wanis c.2010).
2. Possible introduction or reintroduction of the ants and/or SS along with relief supplies
in the aftermath of Tropical Cyclone Winston (TCW), the strongest cyclone ever affect-
ing Fiji on 20 February 2016.
3. The highly invasive SS has been a relatively recent re-introduction into Fiji where
there are no natural enemies or where introduced predators and parasites were in
insufficient numbers to deal with the invasion, possibly due to the indiscriminate use
of insecticides that have led to a pest backlash in the absence of predators (Thaman
1984). This is supported by reports that recent outbreaks of tea scale (Fiorinia theae)
that threaten the tea industry in Assam, India may be due to over-dependence on
synthetic pyrethroid insecticides that may have been responsible for reducing pred-
ator populations (Staff reporter 2017), something supported by mealybug and scale
expert, Gillian Watson (2017 pers. Com.)
4. The possible, eradication or reduction to extremely low levels of the natural preda-
tors, parasites and diseases that suppress mealybugs after TCW, which according to
Watson (2017 pers. Com.), a renowned authority on mealybugs and scale insects, is
known to happen periodically when violent storms hit small islands.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 57
Long-term control, not to mention “eradication” of the species involved in the 2016 Fiji
Ant-Mealybug Bioinvasion will, however, be extremely difficult and require integrated pest
management and the combination of a range of cultural and chemical controls and, in the
long-term, probably biological control agents. This is due to: 1) the multi-species nature of
the invasion and the strong symbiotic relationships that WFA have with mealybugs, scales
and other infesting organisms; 2) the extent of the invasion, the large size of ant colonies,
and the high reproductive rates, mobility and dispersal ability of these invasive organisms;
3) the high diversity of suitable habitats and host plants and plant parts that are infested;
4) their ability to hide in cracks, crevices, tree bark, etc. which protects them from both
natural enemies and humans control, including the protection from insecticides and other
control agents afforded to mealybugs and scales by their natural coatings of waxy egg sacs
and hard scale-like, structures (which are reportedly almost impossible to penetrate with
many pesticides) with the last resort and possibly most sustainable long-term option be-
ing the use of biological control agents (McKenzie 1967; Meyerdirk et al. 2001; Bugwood
2003; Mesbah et al. 2009; Mani 2016).
As stressed in Jamaica, because of the close relationship between WFA and SS mealybugs
and scale insects, the control of WFA is key to the successful containment of the insects
that the ants tend and protect from natural and introduced predators, parasites and
diseases (JIS 2007), which without the ants, both natural and introduced predators and
pathogens can help control the mealybugs and other insects (Jackson 2015). This is borne
out in Fiji by the fact that some of the only areas found largely free of both pests are
those, such as tourist resorts, hotel, restaurants and other institutions, that routinely, in
some cases almost daily, spray insecticides to rid their properties, including their gardens
and landscaping, from bothersome and off-putting (to tourists and diners) ants, cock-
roaches, flies, mosquitos and bedbugs, thus inadvertently also controlling WFA, mealy
bugs and other associated insects and resultant fungi or sooty moulds.
58 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
6.1.1 Removal or destruction of ant nests and nesting sites: The removal or destruc-
tion of outdoor and indoor ant nests and potential nesting sites, such as woodpiles,
rubbish or waste accumulations, or by burning or the use of boiling water to destroy ant
nests, and continual cleaning of cupboards and ant nesting areas in garages, washrooms
and waste areas around buildings and homes and in gardens is the first step to controlling
WFAs.
6.1.2 Use of adhesives, sticky tapes and banding: By banding the lower trunks or
stems of plants or placing or painting adhesives pastes, strong-smelling substances such
as turmeric paste or other products (some of which are sold to intercept and/or kill rats,
cockroaches and other pests) on cardboard or small pieces of wood, to catch or impede
the movement of ants, is a technique that has been used successfully by some people in
Suva to stem the flow and reduce populations of WFA on their properties.
6.1.3 Physical removal: Hand removal or wiping or spraying jets of water or misting oils
(e.g. white oil™) to physically remove mealybugs, scales and other insects from plants has,
when combined with other interventions, has also helped control the level of infestation.
6.1.4 Weed control and removal, destruction and control of movement of infest-
ed plant materials: One of the most sustainable, long-term ways of controlling the
WFA-mealybug invasion is the control of weeds and removal, disposal and destruction or
burning of infested plants and plant materials (e.g., cuttings, flowers and fruits) to en-
sure the removal and destruction of the white egg sacks and pink mealybugs. This would
include pruning of infested lower branches to stop ants from reaching and infesting other
parts of the plant. Because immature mealybugs move on wind currents, it is important,
where feasible to burn infested material or to place suspected infested materials into
sealed plastic bags for transport because the movement of materials in open vehicles is
particularly risky, because of the virulence of mealybug infestations, under a Pink Hibiscus
Mealy Bug Order of 2007, Jamaica mounted an awareness campaign and strictly prohib-
ited the movement of any plants, seedlings, cuttings or plant products or material into or
out of affected areas (RJR News 2007); and the Jamaica Ministry of Agriculture and Lands
established a “Mealy Bug Hotline” that people were to phone when they experienced an
infestation so that authorities could visit and assess the infestation; and because of the
virulence of the infestation, the Rural Agricultural Development Agency (RADA) did not
want farmers to take samples which could further contribute to the spread of this pest. It
was stressed that when farmers pruned trees and plants where they suspected the pres-
ence of the pest on their property, they should burn all plant parts or place the remnants
in garbage bags and bake them in the sun in order to kill the pests (JIS 2007).
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 59
6.1.5 Horticultural practices and plant selection: Horticultural practices such as plant
spacing, intercropping and species and cultivar selection, including the planting of resis-
tant, poisonous or aromatic plants, such as those discussed above, that seem to be unaf-
fected by, repel or do not produce the type of sap favoured by mealybugs, scales, aphids
and other sap-dependent species, are also important cultural control options.
Use of ant baits seems to be quite effective. Among the less toxic options, some people
have had success using a combination of boric acid or borax, mixed with sugar or honey
left in saturated gauze, cotton wool or cloth in saucers in areas of ant traffic, from which
small quantities can be carried by armies of foraging workers to ant nests where it is fed
to queens and larvae to kill ant colonies. It takes some time, but eventually this action
can reduce in size or kill the colony. The important thing about an ant bait is that it should
not be so toxic or concentrated so as to kill individual ants quickly before they have had a
chance to return to the nest to transfer the toxicant to the colony at large (Bugs for Bugs
2017). More toxic ant baits include the use of the widely available commercial ant baits
that contain Fipronil™, a fast-acting ingredient that is also taken back to nests by foraging
workers to kill the colony.
6.2.2 Banding, painting or spraying the lower trunk: The banding, painting or spraying
the lower trunk or stems of infested plants with insecticide such as Dieldrin™ or Mortein™
has also been used to stop or reduce plant infestation by ants and mealybugs
6.2.4 High toxicity chemicals: Although a range of highly toxic and dangerous contact
and systemic insecticides have been successfully used to control ants, adult mealybugs,
scales, etc., they have significant negative human health and environmental impacts, and
must be used with utmost care wearing appropriate protective clothing and stored in safe
places out of the reach of children and pets, something that is often not done in Fiji. The
use and impacts of some of these toxic pesticides are discussed below.
In recent years the use of Rogor and other organophosphate insecticides has been
debated because of their suspected links with cancer and respiratory and cardiac disor-
ders (Morello-Frosch et al. 2011), although some gardeners still used them because of
their effectiveness. And most retailers have now stopped selling dimethoate products in
Australia, where in 2011, based a dietary risk and assessment report, the Australian Pes-
ticides and Veterinary Medicines Authority determined that it was illegal to use Rogor on
any edible crop because dietary exposures exceeded the Australian health standard (the
acute reference dose) and prohibited the use of dimethoate on all food producing plants
in home gardens, on certain horticultural crops, a finding that is highly relevant to the
situation in Fiji (Campbell 2012; APVMA 2017).
In summary, all of these toxic insecticides that are currently available and used in Fiji have
widespread and increasingly documented long-term negative cascading impacts on a wide
range of other non-target organisms, ecosystems services and human health. Their use re-
quires storage in safe, preferably locked places, and the use of protective clothing, gloves,
hats, and googles or face protection, preferably a half-face respirator, by people handling
or applying the insecticides, all which should be washed afterwards and kept separately
from other clothing and equipment. They should never be applied to indoor surfaces; pets
and humans should not be allowed in the area during application; they should normally
not be applied to edible plants; and should not be used if heavy rains are expected to
occur within 48 hours of application or when soil is wet.
This is supported by studies of HMB and other polyphagous scale insects, including the
Columbian fluted scale (Crypticerya multicicatrices), that were unintentionally introduced
with plants from the mainland to Columbia’s offshore Caribbean islands, but only con-
trolled when the a native predator of the scale insect was unintentionally introduced from
mainland Columbia (Kondao et al. 2014). In Jamaica, where the mealy bugs had caused
an estimated $3 million crop loss per year in the early 2000s, the only way to control the
mealybug in the long-term was believed to be the introduction of obligate predators and
parasites, such as parasitic wasps, natural enemies of the pest, which when released lay
their eggs on the immature stages of the mealy bugs and after hatching the larvae feed
internally on the mealybugs. Where these wasps had been released in the Caribbean and
the United States, there was a 90 to 95 per cent drop in the mealybug populations (JIS
2007); studies 5 months after releasing Anagyrus kamali, a parasitic wasp from China,
there was reportedly an 80—90 % reduction in population density of the PHM at release
sites (Mani and Gul 2016); and effective control of SS was reportedly achieved in Mauri-
tius by introducing the parasitic fly (Chryptochetum monophlebi) (HUJI 2016).
Closer to home, predatory ladybird beetles from Australia, such as Rodolia limbata were
successfully introduced to control breadfruit mealybug (Icerya aegyptiaca) in FSM, Kiri-
bati, the Marshall Islands and Palau in Micronesia where it was reducing fruit yields by up
to 50% (Mani and Gul 2016); Rodolia cardinalis was reportedly introduced into the Cook
Islands in the late 1930s to control SS, which had become established on Aitutaki and
Rarotonga; and R. pumila was released on Rarotonga in 1974 (Walker and Deitz 1979).
Although the SS is recorded present in the Cook Islands Biodiversity Database, there is no
photo and no associated data, indicating that it may no longer be a problem there and
possibly controlled by these previously introduced predators (McCormick 2007). Finally,
the SPC has reportedly successfully controlled mealybugs around its Suva headquarters in
Fiji using an imported Australian ladybird, the mealybug destroyer (Cryptolaemus mon-
trouzieri) (MBD), which is recognised worldwide as a voracious predator of mealybugs, ca-
pable of eating 3,000-5,000 mealybugs in various life stages during its lifetime (Mani and
Gul 2016; Martin 2016). This is supported by the discovery, during the current study, of
remnant populations of MBD larvae and adults on mealybug and scale infested gardenia
and ixora plants. One complicating factor is that because of their heavy wax coating, the
larvae of MBD, have apparently evolved to look very similar to and are often mistaken for
large mealybugs, and unsuspectingly inadvertently killed by the use of insecticides, which
are very toxic to ladybirds and other insects. It is consequently recommended that insec-
ticides use be avoided where possible, and, if used, that four weeks should elapse before
biocontrol insects such as Cryptolaemus ladybirds are released (Bugs for Bugs 2015b).
However, as stressed above, the most serious obstacle to the successful introduction and
effectiveness of biocontrol agents is that the toxic insecticides discussed above kill and
threaten the long-term effectiveness of biological control.
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 63
7 FURTHER STUDIES
This is only a first exploratory study focused on trying to understand the extent and nature
of the 2016 Fiji Ant-Mealybug Bio-invasion and opportunities for, and obstacles to its
control and management, and there is obviously need for further, more-in-depth studies.
Some suggested areas for further study include:
1. More-in-depth studies of the geographical extent, habitat preferences and the range
of invasive and host species involved in the bio-invasion,
3. Studies of natural, mainly forest, ecosystems on both main inhabited and uninhabited
offshore islets, such as the studies carried out on Aldabra Atoll in the Seychelles in the
1970s—80s to asses impacts on less-humanised ecosystems.
4. Studies in Fijian village gardens, rural household gardens and other urban gardens,
landscaping and botanical gardens and nurseries.
5. Studies in rural agricultural areas, such as the Fiji sugarcane belt, ginger and banana
growing areas, the Sigatoka Valley, Taveuni taro farming areas and areas of Fijian shift-
ing cultivation.
6. Studies of the presence or absence of predators and parasites, both native and intro-
duced, and the impact of ants and the use of insecticides on them.
8. Studies of the history of the use and effectiveness of biological control of ants, mealy-
bugs scale insects and other sap-sucking insects, both locally and internationally and
the identification of promising biocontrol agents and strategies for biocontrol imple-
mentation in Fiji and elsewhere in the Pacific Islands.
9. Economic evaluation of the economic, social and environmental costs of the 2016 Fiji
Ant-Mealybug Bio-invasion.
64 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
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82 T H E 2 0 1 6 F I J I A N T- M E A LY B U G B I O I N VA S I O N
Appendices
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 83
Appendix I
Families, genera and species of cultivated and wild plants affected by symbiotic infes-
tations of WFA along with SS and/or CMB, ACS, PHM, scales and other insects such as
aphids based on field studies in Suva and other selected locations on Viti Levu Island,
Fiji by R. Thaman from May to August 2017. Common and Fijian names (including Hindi
names) are, when available, provided in bold for some local indigenous and culturally
important plants. Ferns and then Gymnosperms are listed first, followed by Dicotyledons
and Monocotyledons (Notes: Families, genera and species are listed in the approximate
order of severity of infestation and number of genera and species infested; * indicates
that a given genus or family, e.g. Hibiscus may be represented by a number of distinct
cultivars)(Appendix II is a detailed species-by-species listing including approximate order
of levels and incidences of infestation along with full scientific, common and vernacular
Fijian (Fiji and Indo-Fijian) names, levels of observed infestations, parts infested and num-
ber of locations at which infestations were observed).
FERNS AND FERN ALLIES (PTERIDOPHYTES) (2 sp. from 2 genera and 2 families)
Aspidiaceae (Rounder Sori Fern Family) = 1 genus, 1 spp.
• Tectaria latifolia, black-stiped fern, otaloa
Nephrolepidiaceae (Sword Fern Family) – 1 genus, 1 sp.
• Nephrolepis biserrata, sword fern, digi
• Cassia: golden shower tree (C. fistula), pink-and-white shower tree (C. javanica) (2)
• Tamarandus indicus, tamarind, imli (1)
• Arachis hypogaea, peanut, groundnut (1)
• Phaseolus vulgaris, French bean, haricot bean (1)
• Psophocarpus tetragonolobus, winged bean (1)
• Arachis hypogaea, peanut, groundnut, vinati (1)
• Samanea saman, raintree, monkeypod, vaivai ni vavalagi, vaivai moce, sirsa (1)
• Inocarpus fagifer, Tahitian chestnut, ivi (1)
• Intsia bijuga, Pacific teak, ipil-ipil, vesi (1)
• Centrosoma pubescens, centro (1)
• Derris trifoliata, fish-poison vine (1)
• Senna siamea, Siamese cassia, kassod tree (1)
• Peltophorum pterocarpum, yellow poinciana, yellow flamboyant, copperpod (1)
• Cynometra insularis, cibicibi, moivi (1)
• Pterocarpus indicus, Burmese rosewood, Amboyna wood, red sandalwood (1)
• Milletia pinnata, beach walnut, pongam, vesiwai (1)
• Gliricidia sepium, Mother-of-cocoa, Mexican lilac, quickstick, bainicagi (1)
• Adenanthera pavonina, red-bead tree, lera, diridamu (1)
Appendix II
Plant families, genera and species affected by DFA, SS, PHM, CMB and/or other mealy-
bugs, scale insects, other sapsucking insects or other pathogens infestations in Fiji based
on field studies by R. Thaman from May—September 2017. Ferns and fern allies and
gymnosperms are listed first followed by dicotyledons and monocotyledons (Note: the
deliberately introduced mealybug destroyer or mealybug ladybird (Cryptolaemus montro-
zieri), a notorious predator of mealybugs and scale insects, is also listed with hostplants
on which it was observed and photographed)
Key.
1. Under species: 1) * indicates species (e.g., Hibiscus rosa-sinensis) that are affected by
PHM, whereas all other species are affected by SS, the main active invasive mealybug/
scale and or other sap-sucking insects tended by WFA; 2) E = endemic Fiji species, I =
indigenous species; A = aboriginal introduction, with all other species being considered to
be non-indigenous post-European-contact introductions into Fiji.
2. Under common names: 1)The common English or other widely recognized names listed
followed by local indigenous Fijian or Indian names for a given plant.
3. Under Inf. Lev. (Infestation Level), which has been assessed in terms of both the level
of infestation of individual plants and the frequency it is seen in study areas): H = high, M
= medium, L = low or incidental, A = ants only.Under Pts Inf (parts infested): Bd = buds,
Br = branches, Fl = flowers, Fr = fruit; Fs = flower spathe/ stalk/cluster; Lm = leaf mid-vein,
Ln = leaf node, Ls = leaf sheath; Lv = leaves,; Ps = pseudostem (bananas); Pt – petiole, Sc –
found along with another scale insect; Sp = spathe; St = stem or stalk;, Tb = terminal buds,
Tr = trunk, u = underside of leaves, t = topside of leaves, y = young leaves.
4. Under IAS (Invasive Alien Species responsible for infestation along with WFA): ACS -
Asian cycad scale (Aulacaspis yasumatsui); APH = aphids (Aphididae); ASB = Ablemoschus
(bele) shoot borer (Earias vittella); BMB = breadfruit mealy bug (Icerya aegyptiaca); BSM =
black sooty mould (Capnodium and Tripospermum spp. plus others); CBS = coffee brown
scale (Saisettia coffaea); CMB - Coconut mealybug (Nipaecoccus nipae.); CS = circular scale
(Chrysomphalus aonidum); CTM – citrus mealybug (Planococcus citri); LYM = longtailed
mealybug (Pseudococcus longispinus); MBD = mealybug destroyer or mealybug ladybird
(Cryptolaemus montrouzieri); NS = nigra scale (Parasaisettia nigra); PHM = Pink hibiscus
mealybug (Maconellicoccus hirsutus); PMB = papaya mealybug (Dysmicoccus brevipes);
PR = Plumeria or frangipani rust (Coleospermum plumeriae); SBS = Soft bamboo scale
(Bambusaspis bambusae); SS = Seychelles scale (Icerya seychellarum); STS = stellate scale
(Ceroplastes stellifera); SWF = Spiralling whitefly (Aleurodicus disperses); UKS = unknown
scale; YA = yellow aphid (ID unknown)
T H R E AT T O F O O D , H E A LT H , L I V E L I H O O D , C U LT U R A L A N D E N V I R O N M E N TA L S E C U R I T Y I N T H E PA C I F I C I S L A N D S 93
Appendix III
Common native and introduced mainly woody plant species that are un-affected or
almost never affected by the WFA-mealybug and sap-sucking insect bio-invasion in Fiji
based on field studies by R. Thaman from May—September 2017 (Notes: 1) * = com-
mon species showing very limited incidental infestation in only one or few locations, but
uninfested in almost all sites; 2) main sources (for origen/comments): Quisumbing 1951;
Whistler 1980, 1994, 2000; Hammond 1995; Guevara et al. 1996; Argueta 1994; Thaman
1999, 2011; Quattrocchi, U. 2012; Thaman et al. 2012; Thaman et al. 2013; Brown 2013;
Gilman and Watson 2013; Moghadamtousi et al. 2013; Dinesh et al. 2014).