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History of Wasma nnia auropunctata in Hawaii 39
The History of Little Fire Ant Wasmannia auropunctata Roger
in the Hawaiian Islands:
Spread, Control, and Local Eradication
Casper Vanderwoude1, Michelle Montgomery, Heather Forester,
Ersel Hensley, and Michael K. Adachi
Hawaii Ant Lab, Pacic 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 “inva-
sion waves” by exotic ants for over a century. The arrival of Pheidole megacephala
(Fabricius) (the big headed 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 the little re ant
(Wasmannia auropunctata Roger) which was rst recorded in 1999. This paper
chronicles the subsequent spread of W. auropunctata through the Hawaiian archi-
pelago. Initially introduced and spread via the import and sale of nursery 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 infor-
mal and ad hoc partnerships between a number of agencies, each contributing to
preventing and reducing spread of this species. The draft Hawaii Inter-Agency
Biosecurity Plan recognizes and strengthens these partnerships and will contribute
positively to Hawaii’s biosecurity system.
Key words: invasive ants, Hawaii, Wasmannia auropunctata, biosecurity, biologi-
cal invasions, Pacic, little re ant
Introduction
Native ants are thought to be naturally
absent from the islands of the eastern
Pacic, including those of the Hawaiian
archipelago (Wilson and Taylor 1967). All
ant species currently recorded in Hawaii
are widespread cosmopolitan tramp spe-
cies that have been introduced by human
travel and commerce (Krushelnycky et al.
2005). The biota of Hawaii has evolved in
the complete or nearly complete absence
of ants, which most likely resulted in an
ecological predisposition to invasions by
exotic 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 cur-
rent number of species a little higher due
mostly to taxonomic revisions.
Of these, four ant species are especially
noteworthy due to their ecological and
economic impacts worldwide, featur-
ing prominently in the IUCN list of the
world’s worst invasive species (Lowe et
al. 2000). The bigheaded ant (Pheidole
megacephala (Fabricius)) was rst re-
corded in Hawaii as early as 1879 (Smith
1879), at which time it was already well
established. In the years that followed,
proceedings of th e haWaiian entom ological so ciety (2016) 48:39–50
40 Vanderwoude et al.
entomologists lamented the dearth of na-
tive Coleoptera wherever P. megacephala
had become established (Perkins 1913).
Thei r association with mealybugs and
other 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 is-
land of Oahu (Zimmerman 1940, Reimer
1994). Primarily considered a nuisance
species, Argentine ants spread quickly
to the neighboring islands. The ensuing
battle for territory between L. humile
and P. 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, Anoplolepis
gracilipes (fr. Smith) (the longlegged ant,
also known as the yellow crazy ant) ar-
rived at the US Naval base, Pearl Harbor
(Clagg 1953). A shade-tolerant species,
A. gracilipes thrived in shaded lowland
environments, preying on birds and in-
vertebrates (Gillespie and Reimer 1993).
Capable of episodic population explosions,
A. gracilipes forms dense super-colonies
that drive out other fauna and at some
locations, can cause the collapse of plant
communities (O’Dowd et al. 2003).
In 1999, the little re ant (Wasman-
nia auropunctata Roger) was detected
on the isla nd of Hawai i (Conant and
Hirayama 2000). This ant species has a
native range that includes South America
and the Caribbean (Wetterer and Porter
2003), but has invade d many Pacific
islands, West Africa, Australia, Florida,
and Israel (Wetterer 2013). Genetic com-
parisons with material from native and
introduced locations suggest Florida is the
putative source of the Hawaii introduction
(Mikheyev and Mueller 2007, Foucaud et
al. 2010). Here, we describe the spread of
this species through the Hawaiian islands
between 1999 and 2016 and discuss likely
introduction pathways.
Methods and Materials
We used published and unpublished
literature as well as personal commu-
nications and observations from others
involved with the response to this intro-
duction to document the spread of W.
auropunctata from the date of the initial
detection to the present (2016).
Histor y of Introduction
and Spread
The state of Hawaii is located in the
central Pacific Ocean, approxi mately
between longitudes 154–160° west, and
latitudes 19–22° north. It is made up of
eight separate islands, of which, six are
accessible by the general public: Hawaii,
Oahu, Maui, Kauai, Molokai, and Lanai.
Since the initial discovery in 1999, W.
auropunctata has become established on
the four most populous islands (Oahu,
Hawaii, Maui, and Kauai). The spread, to
and within, each island, is detailed below.
Hawaii island. In 1999, Conant and
Hirayama (2000) reported the presence
of W. auropunctata at 13 locations in the
South Hilo district on the island of Hawaii
(the Big Island). Initially, W. auropunctata
was observed on three infested proper-
ties 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 agricul-
tural properties that had recently planted
windbrea ks of Car yota sp. (fish-tail
palm) (P. Conant pers. com). Subsequent
public outreach, e.g. Gruner (2000), and
surveys revealed that W. auropunctata
infestations were more widespread than
rst estimated, likely spread through the
sale and movements of infested potted
plants. Despite this challenge and a lack
of resources, the Hawaii Department of
History of Wasma nnia auropunctata in Hawaii 41
Agriculture (HDOA) responded by treat-
ing all known infested properties with
baits. Between 1999 and 2007, the number
of separate known infestations increased
from an initial 3 properties to 56 by 2007
(Figure 1). These properties were scat-
tered 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 the
number of known sites was a reection of
survey effort, increasing levels of public
awareness and actual spread.
The widespread and mostly unknown
distribution of W. auropunctata, along
with an inability to treat colonies estab-
lished in the tree canopy (Souza et al.
2008), resulted in the continued spread of
this species. By early 2010, W. auropunc-
tata had spread to several locations on the
west coast of Hawaii island (Vanderwoude
et al. 2010). New infestations continued to
be detected beyond the original Kalapana-
Laupahoehoe area and now include most
of the west side of Hawaii island, Waipio
Valley, Hawi, Kapaau, Holualoa, Naalehu,
Captain Cook, and Waimea. In districts
with lower rainfall, W. auropunctata are
limited to favorable microclimates near
homes and other structures that feature
articial landscaping and irrigation (C.V.
pers. obs.). This concurs with the observa-
tions of Vonshak in Israel (Vonshak et al.
2010). By end 2010, the estimated number
of 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 the
initial detection (October 1999), plants
from an infested nursery on Hawaii had
been shipped to the island of Kauai. These
plants were infested with W. auropunc-
tata colonies. The plants and adjacent
1
0
10
20
30
40
50
60
1999 2000 2001 2002 2003 2004 2005 2006 2007
Numberofinfestedproper0es
year
Figure 1. Number of known locations infested with Wasmannia auropunctata on Ha-
waii 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.
42 Vanderwoude et al.
1
Figure 2. Location of properties infested with Wasmannia auropunctata in January
2007 prepared by Hawaii Department of Agriculture.
areas were immediat ely treated with
baits to prevent further spread within
Kauai (Conant and Hirayama 2000). This
infestation was assumed eradicated. How-
ever, W. auropunctata were recorded in a
follow-up survey at the site four years later
in September 2003 (Null and Gundersen
2007). The infestation now covered ve
acres and had encroached onto an adjoining
property (see Figure 4). The site was treated
with granular baits followed by ad hoc
retreatment and periodic surveys through
to 2012. During these years, the infestation
spread mostly westwards eventually span-
ning 12 acres and extending down a steep
escarpment to Kalihiwai beach.
History of Wasma nnia auropunctata in Hawaii 43
1
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.
44 Vanderwoude et al.
Figure 5. Locations of all known sites on Maui infested with Wasmannia auropunctata.
In late 2012, a second eradication at-
tempt was implemented. At this time, the
critical issues of bait efcacy (Hara 2013,
Hara et al. 2014, Montgomery et al. 2015)
and arboreal treatment (Vanderwoude and
Nadeau 2009) had been largely resolved.
Due to the complexity of the site and
regulatory issues, this attempt was divided
into two phases: initially focusing on the
readily accessible areas and later address-
ing the escarpment and taller vegetation.
To date (late 2016), results are encourag-
ing. 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 treatment
of the known small colony continues.
Maui. Wasmannia auropunctata have
been detected multiple times on the is-
land of Maui (Figure 5). The rst LFA
infestation detected on Maui was located
in Waihee, immediately west of the main
city of Kahului, in September 2009. A
resident reported receiving painful stings
from small ants on her property. Samples
of these ants were submitted to the HDOA
entomologist who conrmed it was Was-
mannia auropunctata. An inter-agency
taskforce was established, consisting of
staff from the County of Maui, Maui
Invasive Species Committee (MISC),
HDOA, US Geological Survey, Univer-
sity of Hawaii, and the Hawaii Ant Lab
(Hawaii Department of Agriculture 2009,
Vanderwoude et al. 2010). Together they
formulated an eradication plan which in-
cluded treatment, outreach and delimiting
elements (Vanderwoude et al. 2010). The
ants were restricted to a single property
and an island-wide delimiting survey of
probable high-risk sites did not nd addi-
tional infestations. The Waihee infestation
was ofcially eradicated in April 2014.
In December 2013, a Maui resident,
alerted by various outreach programs im-
plemented by MISC, found W. auropunc-
History of Wasma nnia auropunctata in Hawaii 45
tata on a hapuu log (Cibotium sp., a tree
fern) purchased from a local landscaping
supplier. The discovery prompted a larger
investigation by HDOA who discovered
that several shipments of hapuu logs, origi-
nating from the Big Island, were infested
(Hawaii Department of Agriculture 2013).
These shipments and subsequent distribu-
tion to retailers were located and inspected
by qua rant ine officers. A number of
these 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 rst half of 2014 and these have been
eradicated by HDOA and MISC.
In September 2014, MISC eld workers
were stung by small ants while conduct-
ing other activities in Nahiku (near Hana,
Maui). These ants were later identied as
LFA and subsequent surveys found high
density LFA in challenging rainforest ter-
rain on both sides of the Hana Highway,
extending 1½ miles along a drainage to
the 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 than
normal lateral spread, most likely facili-
tated by the movement of infested debris
during periodic ooding events. Due to
the challenging nature of this infestation,
agencies collaborating on the response
(HAL, HDOA, Maui County and MISC)
formulated a containment and aggressive
control plan, rst removing LFA from
locations 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 made
in Huelo in January 2015. An eradica-
tion plan has been developed and partly
implemented. Activities at this site were
hampered by the refusal of one resident
to allow treatment staff access. This re-
sulted 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 to
treatment activities have allowed W. au-
ropunctata 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 repeat-
edly between 2010 and 2014. In 2016
another survey was conducted at this site.
W. auropunctata were again detected in an
area 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, this
site is now being treated again to ensure
no live ants remain.
Oahu. The detection of infested ship-
ments of hapuu in Maui prompted HDOA
to investigate other shipments from the
same suppl ier destined for Oahu and
Lanai. Some of these were also infested,
and as a result, HDOA staff systematically
surveyed the retail stores that received
these items. Several of these retail stores
also had become infested, and these were
systematically treated by HDOA staff
(Hawaii Department of Agriculture 2013).
The increased publicity surrounding
the infested hapuu led to the discovery
of two well-established infestations on
Oahu, each covering approximately ve
acres (Figure 6). One of these was lo-
cated in abandoned agricultural land in
Waimanalo and another in a suburban
area of Mililani. Eradication plans were
developed for each site and baits were
applied repeatedly to both sites over the
course of one year. One year after the last
treatment was applied (2016), both sites
are putatively free of LFA.
46 Vanderwoude et al.
The movement of W. auropunctata to
Maui and Oahu identied critical gaps in
the biosecurity system. On Oahu, these
gaps were addressed by implementing an
ongoing island-wide survey of high-risk
entry sites that began in January 2015 and
continues to the present. This program
was designed to complement existing
regulation and inspection systems, with
a goal to detect and eradicate infestations
while small. During the past two years,
this program has detected 16 nascent
infestations at Oahu nurseries which were
systematically 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 will
grow too large to manage and become a
source-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 becom-
ing apparent as entomologists lamented
the paucity of other invertebrate fauna
in locations invaded by ant species such
as Pheidole megacephala (Tryon 1912,
Perkins 1913). These invasions are widely
regarded as a consequence of human com-
merce (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 ex-
ception.
Queens and males in invasive W. auro-
punctata populations are mostly produced
th rough thelytokous par thenogenesis
(Fournier et al. 2005). Clonal reproduc-
Figure 6. Locations of known sites on Oahu infested with Wasmannia auropunctata.
(currently the infestation in Mililani and the original infestation in Waimanalo are
putatively ant-free)
History of Wasma nnia auropunctata in Hawaii 47
tion allows global invasion pathways of
this species to be accurately reconstructed
(Foucaud et al. 2010). Thus, the origin of
W. auropunctata in Hawaii can be attrib-
uted to W. auropunctata from Florida, as
one population is a clonal subset of the
other (Foucaud et al. 2010). Further, there
is an unambiguous connection with the
nursery trade as the original vector, both
for the initial introduction and subsequent
early spread within Hawaii island.
Potted plants are an ideal vehicle for
the movement of this species. The spaces
between the potting medium, plant roots
and the wall of plant conta iners are
convenient nesting sites, and forms a
moisture gradient that optimizes brood
development (Holldobler and Wilson 1990
p374). W. auropunctata colonies are small,
interconnected and typically possess a
worker:queen ratio between 250 and 500
(Ulloa-Chacon and Cherix 1990). This
virtually assures every plant within an
infested nursery houses a viable W. auro-
punctata colony which can remain largely
undetected. Further, by their nature, plant
nurseries are effective distribution points.
Together, these factors contributed to the
rapid 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.
au ropu nctat a betwe en the Hawai ian
islands have become more diverse as this
species 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 to
be certied by HDOA, or for each ship-
ment to neighbor islands to be inspected
before shipment. Without this increased
watchfulness, the inter-island movement
of W. auropunctata would undoubtedly
have been much more rapid. However, at
least some of the multiple infestations de-
tected 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 agricul-
tural commodities shipped between Ha-
waii Island and other islands is detected
and prevented from arriving by means
of a thorough and caref ul system of
regulation and inspection implemented
by HDOA. Inspection systems are based
on a risk-ma nagement approach that
utilize 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 slip-
page (Whyte 2006)—infested goods that
bypass regulated pathways, escape detec-
tion or are in commodity categories that
are not inspected.
By its very nature, slippage is difcult
to quantify, and occurs in four commodity
classes: those that bypass the biosecurity
system, false negatives (infested material
inspected and cleared), commodities ex-
cluded from inspection and commodities
that do not fall within the HDOA man-
date (Government of Hawaii 1973). Not
all pathways are adequately regulated.
Air passengers carrying plants and other
propagative material between islands are
not inspected due to a lack of resources.
The rate of false negatives is likely to be
very low, but remains largely unknown.
Hawaii Administrative Rules (Hawaii Ad-
ministrative Rules 2012) limit commodity
48 Vanderwoude et al.
inspections to “plants and propagative
material.” The rules also acknowledge that
HDOA has legislative authority to inspect
a wider range of commodities such as foli-
age, cut owers and produce, but self-limits
activities to “periodic random inspec-
tions.” Finally, there are no systematic
inspections of other commodity classes
(used vehicles, machinery, household ef-
fects etc.) because HDOA does not have
legislative authority to do so.
Detection and response to these intro-
ductions demonstrates the complementary
roles of prevention through regulation
and inspection; early detection through
increased awareness and surveillance, and
rapid response through multi-agency col-
laboration. These elements of the Hawaii
biosecurity framework are performed by
different and sometimes multiple agencies
(Kraus and Duffy 2010) often through
semi-formal or ad hoc collaborations. Re-
gardless of the multitude of funding part-
ners, agency governance issues, obstacles
to data sharing, complex legal consider-
ations, and the often difcult operational
impediments, these collaborations can be
startlingly effective, as evidenced by the
rapid detection, response, and treatment
of multiple W. auropunctata infestations
throughout Hawaii. Of the eight infesta-
tions on the neighbor islands of Oahu,
Kauai and Maui, ve sites are putatively
free of W. auropunctata and the remain-
ing three are contained and continue to
be treated. A biosecurity plan that brings
these agencies closer and recognizes these
collaborations, is currently being drafted
by the State of Hawaii (Anon 2016), and
will serve as a blueprint for biosecurity
activities in the next decade.
As Wasmannia auropunctata spread
through the islands of Hawaii, the eco-
nomic and ecological impacts are likely to
be catastrophic. The predicted economic
costs to the island of Hawaii alone are
likely to exceed $100 million annually
(Lee et al. 2015). Continued prevention,
early detection and response to new incur-
sions on islands other than Hawaii island
is an invaluable investment in the future
of the unique and fragile ecosystems that
Hawaii has to offer.
Acknowledgments
The Hawaii Department of Agriculture
and Hawaii Invasive Species Council pro-
vide ongoing funding to the Hawaii Ant
Lab. The authors sincerely acknowledge
the contributions of HISC, HDOA, For-
est and Kim Starr the various Invasive
Species Committees, University of Ha-
waii, the Counties of Hawaii, Maui, and
Kauai, City and County of Honolulu, the
US Geological Survey, Department of
Lands and Natural Resources, the Pacic
Cooperative Studies Unit and all other
agencies and individuals that contribute
to the management of Wasmannia auro-
punctata 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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