HroceedingsP
istory of Wof
asmannia
the Hawaiian
auropunctata
Entomologicalin HawaiiSociety (2015) 48:39–50

39

The History of Little Fire Ant Wasmannia auropunctata Rogerin the Hawaiian Islands:
Spread, Control, and Local Eradication
Casper Vanderwoude1, Michelle Montgomery, Heather Forester,
Ersel Hensley, and Michael K. Adachi

Hawaii Ant Lab, Pacific Cooperative Studies Unit, University of Hawaii at Manoa, 16 E.
Lanikaula St Hilo, Hawaii. 1Corresponding author: casperv@hawaii.edu

Abstract. The islands of Hawaii have been the battleground for successive “invasion waves” by exotic ants for over a century. The arrival of Pheidole megacephala(Fabricius) (the big headed Argentine ant) in the late nineteenth century, was followed in 1939 by Linepithema humile (Mayr) (the Argentine Ant) and Anoplolepis
gracilipes (fr. Smith), (the longlegged Ant) in 1953. The most recent arrival is thelittle fire ant (Wasmannia auropunctata Roger) which was first recorded in 1999.
This paper chronicles the subsequent spread of W. auropunctata through the
Hawaiian archipelago. Initially introduced and spread via the import and sale ofnursery plants, W. auropunctata is now well-established on the island of Hawaii.
Ubiquitous on the windward side of Hawaii island, W. auropunctata are now being transported not only via nursery plants but also via non-agricultural products.
The prevention, detection and response to W. auropunctata introductions is addressed by informal and ad hoc partnerships between a number of agencies, eachcontributing to preventing and reducing spread of this species. The draft HawaiiInter-Agency Biosecurity Plan recognizes and strengthens these partnerships andwill contribute positively to Hawaii’s biosecurity system.
Key words: invasive ants, Hawaii, Wasmannia auropunctata, biosecurity, biological invasions, Pacific, little fire ant
Introduction
Native ants are thought to be naturally
absent from the islands of the eastern
Pacific, including those of the Hawaiianarchipelago (Wilson and Taylor 1967). All
ant species currently recorded in Hawaii
are widespread cosmopolitan tramp species that have been introduced by humantravel and commerce (Krushelnycky et al.
2005). The biota of Hawaii has evolved in
the complete or nearly complete absenceof ants, which most likely resulted in an
ecological predisposition to invasions byexotic ant species along with increased
impacts such invasions may cause (Reimer

et al. 1990). The number of new ant species
has accumulated steadily over time to 47(Krushelnycky et al. 2005), with the current number of species a little higher due
mostly to taxonomic revisions.
Of these, four ant species are especially
noteworthy due to their ecological andeconomic impacts worldwide, featuring prominently in the IUCN list of theworld’s worst invasive species (Lowe etal. 2000). The bigheaded ant (Pheidole
megacephala (Fabricius)) was first recorded in Hawaii as early as 1879 (Smith
1879), at which time it was already wellestablished. In the years that followed,

40
entomologists lamented the dearth of native Coleoptera wherever P. megacephala
had become established (Perkins 1913).
Their association with mealybugs andother common plant pests caused crop
losses, especially in pineapple (Beardsley
et al. 1982, Jahn and Beardsley 1994). In
the 1939, the Argentine ant (Linepithema
humile (Mayr)) was detected on the island of Oahu (Zimmerman 1940, Reimer1994). Primarily considered a nuisancespecies, Argentine ants spread quickly
to the neighboring islands. The ensuing
battle for territory between L. humile andP. megacephala saw the new invader restricted to higher elevation habitats where
it caused considerable impacts to native
ecosystems (Medeiros et al. 1986, Cole
1992, Krushelnycky and Gillespie 2008).
In 1953, a new invader, the longlegged
ant (Fr. Smith) (Anoplolepis gracilipes,
also known as the yellow crazy ant) arrived at the US Naval base, Pearl Harbor(Clagg 1953). A shade-tolerant species,
A. gracilipes thrived in shaded lowland
environments, preying on birds and invertebrates (Gillespie and Reimer 1993).
Capable of episodic population explosions,A. gracilipes forms dense super-colonies
that drive out other fauna and at somelocations, can cause the collapse of plantcommunities (O’Dowd et al. 2003).
In 1999, the little fire ant (Wasmannia auropunctata Roger) was detectedon the island of Hawaii (Conant andHirayama 2000). This ant species has anative range that includes South America
and the Caribbean (Wetterer and Porter2003), but has invaded many Pacificislands, West Africa, Australia, Florida,
and Israel (Wetterer 2013). Genetic comparisons with material from native andintroduced locations suggest Florida is theputative source of the Hawaii introduction(Mikheyev and Mueller 2007, Foucaud etal. 2010). Here, we describe the spread of
this species through the Hawaiian islands

Vanderwoude et al.
between 1999 and 2016 and discuss likely
introduction pathways.
Methods and Materials
We used published and unpublishedliterature as well as personal communications and observations from others
involved with the response to this introduction to document the spread of W.
auropunctata from the date of the initialdetection to the present (2016).
History of Introductionand Spread
The state of Hawaii is located in thecentral Pacific Ocean, approximately
between longitudes 154–160° west, andlatitudes 19–22° north. It is made up ofeight separate islands, of which, six areaccessible by the general public: Hawaii,
Oahu, Maui, Kauai, Molokai, and Lanai.
Since the initial discovery in 1999, W.
auropunctata have become establishedon the four most populous islands (Oahu,
Hawaii, Maui, and Kauai). The spread, toand within, each island, is detailed below.
Hawaii island. In 1999, Conant andHirayama (2000) reported the presenceof W auropunctata at 13 locations in theSouth Hilo district on the island of Hawaii(the Big Island). Initially, W. auropunctata
were observed on three infested properties in Hawaiian Paradise Park south of
Hilo. Soon thereafter, additional infested
locations were discovered at Kapoho
and Paipaikou. Most infested locations
were commercial nurseries or agricultural properties that had recently planted
windbreaks of Caryota sp. (fish-tailpalm) (P. Conant pers. com). Subsequent
public outreach, e.g. Gruner (2000), andsurveys revealed that W. auropunctata
infestations were more widespread thanfirst estimated, likely spread through thesale and movements of infested potted
plants. Despite this challenge and a lackof resources, the Hawaii Department ofHistory of Wasmannia auropunctata in Hawaii

41

Number of infested proper0es

60
50
40
30
20
10
0
1999

2000

2001

2002

2003

2004

2005

2006

2007

year
1

Figure 1. Number of known locations infested with Wasmannia auropunctata on Hawaii island between 1999 and 2007. Data sourced from Conant and Hirayama (2000);
Motoki et al. (Motoki et al. 2013), P. Conant (pers. com.) and informal reports from
Hawaii Department of Agriculture.
Agriculture (HDOA) responded by treating all known infested properties withbaits. Between 1999 and 2007, the numberof separate known infestations increased
from an initial 3 properties to 56 by 2007
(Figure 1). These properties were scattered between Kalapana (30 miles SE of
Hilo) and Laupahoehoe (25 miles NW of
Hilo) (Figure 2) spanning some 55 miles
to an elevation of 1,500 ft a.s.l.. However,
the actual number of infested properties
within these boundaries was probably
much higher (P. Conant, pers. com) as thenumber of known sites was a reflection ofsurvey effort, increasing levels of publicawareness and actual spread.
The widespread and mostly unknown
distribution of W. auropunctata, alongwith an inability to treat colonies established in the tree canopy (Souza et al.
2008), resulted in the continued spread of
this species. By early 2010, W. auropunctata had spread to several locations on thewest coast of Hawaii island (Vanderwoude

et al. 2010). New infestations continued to
be detected beyond the original KalapanaLaupahoehoe area and now include mostof the west side of Hawaii island, Waipio
Valley, Hawi, Kapaau, Holualoa, Naalehu,
Captain Cook, and Waimea. In districts
with lower rainfall, W. auropunctata arelimited to favorable microclimates nearhomes and other structures that feature
artificial landscaping and irrigation (C.V.
pers. obs.). This concurs with the observations of Vonshak in Israel (Vonshak et al.
2010). By end 2010, the estimated numberof infested properties island-wide had
exceeded 4,500, growing to an estimated
6,400 by end 2012 (Lee et al. 2015). Figure
3 shows areas on Hawaii island currently
infested with W. auropunctata.
Kauai. At about the same time as theinitial detection (October 1999), plantsfrom an infested nursery on Hawaii had
been shipped to the island of Kauai. These
plants were infested with W. auropunctata colonies. The plants and adjacent142

Vanderwoude et al.

Figure 2. Location of properties infested with Wasmannia auropunctata in January
2007 prepared by Hawaii Department of Agriculture.
areas were immediately treated withbaits to prevent further spread within
Kauai (Conant and Hirayama 2000). This
infestation was assumed eradicated. However, W. auropunctata were recorded in afollow-up survey at the site four years laterin September 2003 (Null and Gundersen2007). The infestation now covered fiveacres and had encroached onto an adjoining
property (see Figure 4). The site was treatedwith granular baits followed by ad hoc
retreatment and periodic surveys throughto 2012. During these years, the infestation
spread mostly westwards eventually spanning 12 acres and extending down a steep
escarpment to Kalihiwai beach.

History of Wasmannia auropunctata in Hawaii

43

Figure 3. Areas of Hawaii island currently infested with Wasmannia auropunctata(2016). (Not all properties in the larger shaded section are infested).

Figure 4. Map of Kauai showing location infested by Wasmannia auropuntata (2012).
Currently this site is putatively ant free.

1

44

Vanderwoude et al.

Figure 5. Locations of all known sites on Maui infested with Wasmannia auropunctata.

In late 2012, a second eradication attempt was implemented. At this time, thecritical issues of bait efficacy (Hara 2013,
Hara et al. 2014, Montgomery et al. 2015)
and arboreal treatment (Vanderwoude andNadeau 2009) had been largely resolved.
Due to the complexity of the site andregulatory issues, this attempt was divided
into two phases: initially focusing on thereadily accessible areas and later addressing the escarpment and taller vegetation.
To date (late 2016), results are encouraging. The entire site is putatively free of W.
auropunctata with only a single known
active colony detected beneath a taller
tree. Monitoring of this site and treatmentof the known small colony continues.
Maui. Wasmannia auropunctata havebeen detected multiple times on the island of Maui (Figure 5). The first LFA
infestation detected on Maui was locatedin Waihee, immediately west of the maincity of Kahului, in September 2009. A

resident reported receiving painful stingsfrom small ants on her property. Samplesof these ants were submitted to the HDOA
entomologist who confirmed it was Wasmannia auropunctata. An inter-agency
taskforce was established, consisting ofstaff from the County of Maui, Maui
Invasive Species Committee (MISC),HDOA, US Geological Survey, University of Hawaii, and the Hawaii Ant Lab(Hawaii Department of Agriculture 2009,
Vanderwoude et al. 2010). Together they
formulated an eradication plan which included treatment, outreach and delimiting
elements (Vanderwoude et al. 2010). The
ants were restricted to a single property
and an island-wide delimiting survey ofprobable high-risk sites did not find additional infestations. The Waihee infestation
was officially eradicated in April 2014.
In December 2013, a Maui resident,
alerted by various outreach programs implemented by MISC, found W. auropunc-

History of Wasmannia auropunctata in Hawaii
tata on a hapuu log (Cibotium sp., a treefern) purchased from a local landscaping
supplier. The discovery prompted a larger
investigation by HDOA who discoveredthat several shipments of hapuu logs, originating from the Big Island, were infested(Hawaii Department of Agriculture 2013).
These shipments and subsequent distribution to retailers were located and inspectedby quarantine officers. A number ofthese also had W. auropunctata. These
were either destroyed or treated in situ.
Two additional nascent infestations were
found in south Maui (Wailea area) during
the first half of 2014 and these have been
eradicated by HDOA and MISC.
In September 2014, MISC field workers
were stung by small ants while conducting other activities in Nahiku (near Hana,
Maui). These ants were later identified as
LFA and subsequent surveys found highdensity LFA in challenging rainforest terrain on both sides of the Hana Highway,
extending 1½ miles along a drainage tothe ocean. Four properties were involved.
The infestation appeared to have spread
downstream from an initial upstream
establishment point to the ocean. The
speed at which W. auropunctata spread
downstream was substantially faster thannormal lateral spread, most likely facilitated by the movement of infested debris
during periodic flooding events. Due tothe challenging nature of this infestation,agencies collaborating on the response(HAL, HDOA, Maui County and MISC)
formulated a containment and aggressive
control plan, first removing LFA fromlocations from which it would be likely to
spread, then to later assess the possibilities
for a more comprehensive approach. This
plan is ongoing.
Another LFA discovery was madein Huelo in January 2015. An eradication plan has been developed and partly
implemented. Activities at this site were
hampered by the refusal of one resident

45

to allow treatment staff access. This resulted in the HDOA taking the unusual
step of obtaining a court order (Hawaii
Department of Agriculture 2016), and later
declaring a quarantine on the property
in order allow the eradication program
to continue at this site. The delays totreatment activities have allowed W. auropunctata to recover and spread further
into this property, necessitating additional
treatment effort.
The site at Waihee, which had been
ant-free since 2010, was surveyed repeatedly between 2010 and 2014. In 2016
another survey was conducted at this site.
W. auropunctata were again detected in anarea immediately adjacent to the original
treatment area. It is possible that some
infested plant trimmings may have been
moved there before the original detection
in 2009. Only spanning an acre or so, thissite is now being treated again to ensure
no live ants remain.
Oahu. The detection of infested shipments of hapuu in Maui prompted HDOA
to investigate other shipments from thesame supplier destined for Oahu andLanai. Some of these were also infested,
and as a result, HDOA staff systematicallysurveyed the retail stores that received
these items. Several of these retail storesalso had become infested, and these weresystematically treated by HDOA staff(Hawaii Department of Agriculture 2013).
The increased publicity surrounding
the infested hapuu led to the discoveryof two well-established infestations on
Oahu, each covering approximately fiveacres (Figure 6). One of these was located in abandoned agricultural land in
Waimanalo and another in a suburban
area of Mililani. Eradication plans weredeveloped for each site and baits wereapplied repeatedly to both sites over thecourse of one year. One year after the lasttreatment was applied (2016), both sites
are putatively free of LFA.

46

Vanderwoude et al.

Figure 6. Locations of known sites on Oahu infested with Wasmannia auropunctata.
(currently the infestation in Mililani and the original infestation in Waimanalo areputatively ant-free)

The movement of W. auropunctata to
Maui and Oahu identified critical gaps in
the biosecurity system. On Oahu, thesegaps were addressed by implementing an
ongoing island-wide survey of high-risk
entry sites that began in January 2015 andcontinues to the present. This program
was designed to complement existing
regulation and inspection systems, witha goal to detect and eradicate infestations
while small. During the past two years,
this program has detected 16 nascentinfestations at Oahu nurseries which weresystematically treated. In late 2016 a large,
4-acre infestation was also discovered in
Waimanalo (not linked to the original
detection in the same district). Without
this early detection, such infestations willgrow too large to manage and become asource-point for jump-dispersal to new
locations (Suarez et al. 2001).

Discussion
The worldwide spread of invasive ants
began at least as early as the 16th century(Gotzek et al. 2015). By the beginning of
the 20th century, the ecological impacts
caused by these invasions were becoming apparent as entomologists lamented
the paucity of other invertebrate fauna
in locations invaded by ant species suchas Pheidole megacephala (Tryon 1912,
Perkins 1913). These invasions are widely
regarded as a consequence of human commerce (Wilson and Taylor 1967, Passera
1994, McGlynn 1999, Holway et al. 2002),
and in this regard, the recent introduction
and spread of W. auropunctata is no exception.
Queens and males in invasive W. auropunctata populations are mostly produced
through thelytokous parthenogenesis(Fournier et al. 2005). Clonal reproduc-History of Wasmannia auropunctata in Hawaiition allows global invasion pathways ofthis species to be accurately reconstructed(Foucaud et al. 2010). Thus, the origin ofW. auropunctata in Hawaii can be attributed to W. auropunctata from Florida, as
one population is a clonal subset of theother (Foucaud et al. 2010). Further, there
is an unambiguous connection with thenursery trade as the original vector, bothfor the initial introduction and subsequent
early spread within Hawaii island.
Potted plants are an ideal vehicle forthe movement of this species. The spaces
between the potting medium, plant roots
and the wall of plant containers areconvenient nesting sites, and forms amoisture gradient that optimizes brooddevelopment (Holldobler and Wilson 1990p374). W. auropunctata colonies are small,interconnected and typically possess aworker:queen ratio between 250 and 500(Ulloa-Chacon and Cherix 1990). Thisvirtually assures every plant within an
infested nursery houses a viable W. auropunctata colony which can remain largely
undetected. Further, by their nature, plant
nurseries are effective distribution points.
Together, these factors contributed to therapid spread of this species within Hawaii
Island, mirroring the historical spread of
this species through southern Florida via
the movement of potted plants and balled
citrus seedlings (Spencer 1941).
The pathways for movement of W.
auropunctata between the Hawaiian
islands have become more diverse as thisspecies became increasingly ubiquitous.
After the initial discovery in 1999, HDOA
further regulated the movement of plants
and propagative plant materials between
islands. Regulatory intervention included
a requirement for exporting nurseries tobe certified by HDOA, or for each shipment to neighbor islands to be inspected
before shipment. Without this increased
watchfulness, the inter-island movementof W. auropunctata would undoubtedly

47

have been much more rapid. However, atleast some of the multiple infestations detected on Maui and Oahu are not linked to
the nursery trade in any way. For example,
no links between the purchase of potted
plants and infestations in Nahiku, Huelo,
Waihee and Mililani could be found.
The majority of ant-infested agricultural commodities shipped between Hawaii Island and other islands is detected
and prevented from arriving by meansof a thorough and careful system ofregulation and inspection implemented
by HDOA. Inspection systems are basedon a risk-management approach thatutilize available resources to optimize
risk reduction. However, not all infested
commodities are (or can be) detected at
the border. As W. auropunctata become
increasingly ubiquitous on Hawaii island,
the variety and proportion of infested
cargoes increases beyond simply “nursery
plants” to include non-agricultural items
such as general cargo, household items
and vehicles. A percentage of infested
plants and other non-regulated material
will continue to arrive as a result of slippage (Whyte 2006)—infested goods thatbypass regulated pathways, escape detection or are in commodity categories thatare not inspected.
By its very nature, slippage is difficult
to quantify, and occurs in four commodity
classes: those that bypass the biosecurity
system, false negatives (infested material
inspected and cleared), commodities excluded from inspection and commodities
that do not fall within the HDOA mandate (Government of Hawaii 1973). Not
all pathways are adequately regulated.
Air passengers carrying plants and otherpropagative material between islands are
not inspected due to a lack of resources.
The rate of false negatives is likely to bevery low, but remains largely unknown.
Hawaii Administrative Rules (Hawaii Administrative Rules 2012) limit commodity

48
inspections to “plants and propagative
material”. The rules also acknowledge thatHDOA has legislative authority to inspect
a wider range of commodities such as foliage, cut flowers and produce, but self-limits
activities to “periodic random inspections”.
Finally, there are no systematic inspectionsof other commodity classes (used vehicles,machinery, household effects etc.) because
HDOA does not have legislative authority
to do so.
Detection and response to these introductions demonstrates the complementaryroles of prevention through regulationand inspection; early detection throughincreased awareness and surveillance, andrapid response through multi-agency collaboration. These elements of the Hawaii
biosecurity framework are performed bydifferent and sometimes multiple agencies(Kraus and Duffy 2010) often throughsemi-formal or ad hoc collaborations. Regardless of the multitude of funding partners, agency governance issues, obstacles
to data sharing, complex legal considerations, and the often difficult operational
impediments, these collaborations can bestartlingly effective, as evidenced by the
rapid detection, response, and treatmentof multiple W. auropunctata infestations
throughout Hawaii. Of the eight infestations on the neighbor islands of Oahu,
Kauai and Maui, five sites are putatively
free of W. auropunctata and the remaining three are contained and continue tobe treated. A biosecurity plan that brings
these agencies closer and recognizes thesecollaborations, is currently being draftedby the State of Hawaii (Anon 2016), andwill serve as a blueprint for biosecurity
activities in the next decade.
As Wasmannia auropunctata spreadthrough the islands of Hawaii, the economic and ecological impacts are likely to
be catastrophic. The predicted economic
costs to the island of Hawaii alone arelikely to exceed $100 million annually

Vanderwoude et al.
(Lee et al. 2015). Continued prevention,
early detection and response to new incursions on islands other than Hawaii island
is an invaluable investment in the futureof the unique and fragile ecosystems thatHawaii has to offer.
Acknowledgments
The Hawaii Department of Agricultureand Hawaii Invasive Species Council provide ongoing funding to the Hawaii Ant
Lab. The authors sincerely acknowledge
the contributions of HISC, HDOA, Forest and Kim Starr the various Invasive
Species Committees, University of Hawaii, the Counties of Hawaii, Maui, andKauai, City and County of Honolulu, theUS Geological Survey, Department ofLands and Natural Resources, the PacificCooperative Studies Unit and all otheragencies and individuals that contributeto the management of Wasmannia auropunctata in Hawaii. We thank P. Conant,
N. Reimer, K. Onuma, D. Arakaki, C.
Kaneshige and others who provided vital
anecdotal information.
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