Biodivers Conserv (2009) 18:36773683

DOI 10.1007/s10531-009-9680-9
REVIEW PAPER

Ecological impacts of feral pigs in the Hawaiian Islands

Sergio L. G. Nogueira-Filho Selene S. C. Nogueira Jose M. V. Fragoso

Received: 13 October 2008 / Accepted: 18 June 2009 / Published online: 2 July 2009
Springer Science+Business Media B.V. 2009

Abstract The foraging habits of exotic ungulate species can directly and indirectly affect
native plant and animal distribution and abundance patterns. Most of the studies on feral pig
interactions with other biota in the Hawaiian Islands have been published as difficult to
access reports to governmental and nongovernmental organizations, graduate student theses, and a few in peer reviewed journals. In this paper we discuss the origins of pig
introductions to Hawaii, their feralization process, population expansion, and interactions
with native and non-native biota. We also consider the environmental effects triggered by
pigs on local ecosystems and biotic communities. Feral pig activities can reduce the
abundance of native plant species, enhance conditions for the establishment of invasive nonindigenous plants, and perhaps indirectly negatively impact native forest bird species. Pig
foraging and traveling patterns may also lead to physical alteration of ecosystems by
increasing soil erosion that may lead to watershed degradation. However, much remains to
be learned about the strength and significance of aforementioned interactions and their longterm effects on Hawaiian biota and ecosystems due to some confounding events. Elucidating the dynamics and long-term ecological effects generated by pigs is a crucial step
towards increasing our understanding of and more effectively managing biotic interactions.
Keywords

Habitat disturbance
Invasive species
Plantanimal interactions

Introduction
Humans were the first large-terrestrial mammals to populate Hawaii. This occurred when
the Polynesians colonized the islands during prehistoric times (Kirch 1982). These early
S. L. G. Nogueira-Filho (&)
S. S. C. Nogueira
Applied Ethology Laboratory, Universidade Estadual de Santa Cruz, Rod. Ilheus Itabuna km 16,
Ilheus, BA 45662-000, Brazil
e-mail: slgnogue@uesc.br
S. L. G. Nogueira-Filho
S. S. C. Nogueira
J. M. V. Fragoso
Department of Botany, University of Hawaii at Manoa, 3190 Maile Way, St. John Lab 101, Honolulu,
HI 96822, USA

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Polynesian settlers (indigenous Hawaiians) introduced the Polynesian breed of pig, probably derived from Sus scrofa vittatus, to the Hawaiian Islands as an important source of
food (Kirch 1979; Diong 1982; Tomich 1986). The direct effects of people and their
introduced animals led to major changes in the ecology of islands (Kirch 1982).
The rate and extent of environmental change probably increased dramatically after the
first European explorers arrived on the Hawaiian Islands. The Europeans released more
pigs (Sus scrofa scrofa) and additional ungulate species, such as goats (Capra hircus), for
the purpose of establishing a readily available food source to support subsequent voyages
(Tomich 1986). Later introductions included more pigs as well as domestic cattle (Bos
taurus) and sheep (Ovis aries) (Stone and Pratt 1994). More recently, game animals,
such as mouflon sheep (Ovis musimon) and axis deer (Axis axis) were introduced to the
islands to establish populations for sports hunting (Giffin 1978b; Gagne 1988). The freeroaming populations of these exotic ungulate species have affected the landscape of
Hawaii (Nichols 1962; Tomich 1986).
Most information on potential mechanisms of pig-induced changes in community
composition and ecosystem processes have been published as reports to governmental and
nongovernmental organizations, with a few published as graduate student theses, and a few
in peer reviewed journals, limiting their accessibility to the research and management
community. We review this literature and provide additional commentary on pig interactions with other biota and ecosystems pointing out the weaknesses and the strengths of
the available studies on the ecological consequences of feral pig habitation of the Hawaiian
Islands.

Pig introductions to Hawaii and their feralization process

At the time of European contact pigs were probably present throughout most of the islands
(McEldowney 1979). Polynesian pigs were maintained around early Hawaiian houses in
the lowlands and were usually kept close by the household (Tomich 1986). These animals
were smaller in size than European pigs (Diong 1982; Tomich 1986). Barret and Stone
(1983) proposed that the reportedly small size of Polynesian pigs might have been caused
by a low protein diet since a diet rich in protein is required for maximum growth of pigs.
Diong (1982), Barret and Stone (1983), and Stone (1985) also posit that feral pig populations in Hawaiian rainforests before European contact occurred at low densities. They
report that these were frequently hunted (indicating the presence of feral populations) and
therefore hypothesized that they did not cause great modification to the interior rainforest
vegetation.
Captain Cook brought European pigs on his first voyage to Hawaii and gave these to
indigenous Hawaiian leaders (Diong 1982; Tomich 1986). Many other importations followed and some point more animals became feral and dispersed throughout the islands; the
general consensus has been that afterward the Asiatic form of pig was absorbed or replaced
by European breeds (e.g., Tomich 1986). However, Diong (1982) considered the Hawaiian
feral pigs to be closely related to the original Polynesian pigs.
In a recent study on feral pig phylogeography using mitochondrial DNA, Larson et al.
(2005) found that modern Hawaiian feral pigs form a monophyletic group with pigs from
New Guinea, Vanuatu, and Halmahera (the Pacific clade) within the large Eastern
Eurasian cluster which is well separated from the other groups, confirming Diongs (1982)
proposition that feral populations already existed before European contact. Land use
changes associated with the post-contact era probably contributed to and accelerated the

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feralization process. This included forest clearing, introduction of western agricultural

practices, ranching, and the uphill recession of the forest line (Diong 1982, p. 74).
Barret and Stone (1983) suggested that the factor that most contributed to the posited
low density of feral pig populations before European contact, was the absence of an
abundant animal protein source such as the earthworms that were introduced after European colonization. Some studies support the hypothesis that earthworms are the most
important source of animal protein for Hawaii feral pig populations (Giffin 1978a; Diong
1982; Anderson 1994). Additionally, Diong (1982) proposed that the introduction of
agriculture along with earthworms contributed to the expansion of feral pig ranges.
However, in conflict with this hypothesis is the observation that pre-European times pigs
would have had accesses to protein sources such as sea turtle eggs and sea birds in breeding
colonies. The establishment of beef cattle industry may also have contributed to feral pig
range expansion in historical times, through the clearing of native forest to establish
pastures (Diong 1982), and the introduction of another protein source, dung beetles which
were introduced to aid in the decomposition of cattle feces (Howarth 1985). Giffin (1978a)
detected dung beetle adults and larvae in pig stomachs. Dung beetles are known to enhance
soil nutrient cycling (Howarth 1985), therefore we suggest that such increment would
increase earthworm population and positively affecting feral pig population.

Interactions with native and non-native biota

Some hypothesize that no Hawaiian endemic plant is evolutionary adapted for coexistence
with herbivorous mammals, making them very vulnerable to damage from pig foraging
habits (e.g., Spatz and Mueller-Dombois 1975). Evidence presented in support of this
hypothesis is that some plant species have lost ancestral traits such as thorns and secondary
leaf compounds that may have served to reduce or prevent herbivory. Plants belonging to
this group, such as the Haleakala greensword (Argyroxiphium virescens), of the bogs on
Maui can be severely damaged by pig foraging (Loope 1983). Evaluating pig impacts on
native plants is further complicated by the reduction in the range and abundance of plants
generated by direct human activity. Teasing apart the impact of these multiple factors can
be difficult.
When pigs forage on rare plant species they represent an additional threat towards a
group that may be on the brink of extinction due to human activities. They may also
threaten the structure and dynamics of native rainforests by altering the dominance pattern
of plant species (Tomich 1986). In one rainforest they foraged on at least 40 plant species,
75% of which were native plants (e.g., tree ferns Cibotium chamissoi and C. splendens;
Barret and Stone 1983). The introductions of fruit-producing non-indigenous plant species,
such as strawberry guava (Psidium cattleianum), may also have contributed to pig population expansion (Barret and Stone 1983).
Pigs are generalist feeders and will eat what is available. Diong (1982) observed that
feral pigs in the Kipahulu Valley, Maui, shifted from a diet composed primarily of hapuu
tree ferns (Citobium spp.) to one comprised mostly of strawberry guava fruits according
to their availability. A similar shift was observed by Giffin (1978a) who observed that
Citobium spp. was the main food item in diets but this shifted almost exclusively to banana
poka (Passiflora mollissima) fruit when these became available.
Pigs in Hawaii compound and intensify the problem of non-indigenous plant invasions
by facilitating their introduction by carrying seeds in their guts (e.g., strawberry guava) and
on their coats (e.g., Carex alligata, Paspalum conjugatum) (Diong 1982). As early as 1936,

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Judd reported that native Hawaiian forests where being replaced by stands of non-indigenous guava established by feral pigs. Diong (1982) posits that feral pigs are the primary
modifiers of remaining native rainforests because of their removal of native tree ferns
(eaten) and their dispersal of guava seeds. Their feeding decreases the populations of native
tree ferns and sub-canopy cover and increases the abundance of guava. It is through this
interactive process that feral pigs facilitate the displacement of native species with a nonindigenous invader (Diong 1982). Pigs can also accelerate damage to native plant populations by trampling individuals of native species, and by increasing soil fertility (Stone
1985). Apparently many native plants are adapted to poor soils and increasing soil fertility
after pig rooting works against their reestablishment and favor the establishment of some
non-indigenous plants (Stone 1985).
Aplet et al. (1991) also reported pigs dispersing Myrica faya seeds into rainforests. This
non-indigenous species is a nitrogen-fixing tree that can generate several effects through
communities. For example, earthworms increased in abundance in areas inhabited by M.
faya, probably due to a response to the increased soil nitrogen generated by M. faya (Aplet
1990). We suggest that an increased abundance of earthworms could lead to an increase in
pig foraging near such trees and, consequently, dispersing more Myrica faya seeds.
Exclosure studies have been used on the Hawaiian Islands to evaluate and demonstrate
impacts of feral pigs upon native vegetation and to assess vegetation recovery from feral
pig damage. To obtain these data, the researchers usually build a pig-proof exclosure,
sampling a proportion of the exclosure interior as well as a corresponding area on the
outside. Loope and Scowcroft (1985) reviewed the exclosure studies and found more than
50 studies using small exclosures established to demonstrate and evaluate vegetation
response after ungulate removal in Hawaii. Only a few studies, however, documented plant
succession through several years and using large fenced areas (e.g., Loope et al. 1991;
Medeiros et al. 1991; Pratt et al. 1999).
In exclosure studies, a variety of methods and sampling intervals have been used for
assessing vegetation change. For example, Jacobi (1981), Katahira (1980), and MuellerDombois (1981) used a point frequency sampling method for estimating percentage ground
cover and species composition. Scowcroft and Giffin (1983) used the line-intercept method
to estimate cover in exclosures located in sparse sub alpine forest on Mauna Kea. Additionally, the percentage of litter, exposed soil, the potential area rooted by feral pigs, the
percentage of the area actually dug, and the degree of root exposure were also examined
(e.g., Higashino and Stone 1982; Stone and Taylor 1984). Sampling has typically been
done annually up to 5 years after the exclosure was built (Loope and Scowcroft 1985).
However, statistical analyses of cover, abundance, and other exclosure data have been
barely used in these studies (Loope and Scowcroft 1985).
Another potential problem in many of these studies is that control areas were actually
located close to or very near the fence. This type of placement may generate a confounding
problem. The fence impedes regular movements and dispersal events and this would tend
to concentrate activities and animals in areas along and adjacent to the fence. This outcome
would magnify pig impacts to some distance beyond the fence because the fence restrains
the pigs usual movement pattern leading to overus around the control areas. Proper
controls should consider animal behavior and spatial components such as pig home range
size, movements, and plant distribution patterns.
The exclosure studies showed that removal of feral pigs can result in the recovery of
native vegetation (e.g., Jacobi 1976; Katahira 1980; Higashino and Stone 1982; Stone et al.
1992). This activity, however, has been a negligible impact on non-indigenous plant
populations within fences (Aplet et al. 1991). Therefore, the feral pigs affected native but

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not non-indigenous plant species. Thus, removing pigs does not always alter plant population and community dynamics, which suggests that in these areas other factors may be
driving plant population dynamics.
Stone (1985) published a review on other consequences of pig introduction on the biota
of Hawaii, and highlighted their indirect negative effects on native forest birds. For
example, the foraging behavior of pigs may reduce the amount of nectar produced by
understory plants, such as Rubus hawaiiensis (Stone 1985). Less nectar has been posited to
negatively impact native nectar feeding birds, such as Oreomystis mana, Loxops coccineus,
Chasiempis sandwichensis ibidis, Myadestes myadestinus, and Moho braccatus (van Riper
and Scott 2001). Pigs may also increase breeding habitat for exotic mosquitoes (Culex
quinquefasciatus), and these are the vector of avian poxvirus (Poxvirus avium) and avian
malaria (Plasmodium relictum). They do this by increasing water pool availability when
they eat the inner core of tree fern trunks (Baker 1979). Diong (1982, pp. 166167) does
not support this hypothesis, arguing that there is an unlimited number of naturally
occurring breeding sites for mosquitoes, such as in tree cavities, trunk axils, forest floor and
at bases of fallen trees. However, the decrease of Culex sp. adults captured inside pig
exclosure areas when compared to places where feral pig activity was high provides some
evidence for a relationship between pig activities and the abundance and distribution of the
disease vector (Lease et al. 1996). More recently, LaPointe (2006) corroborated the
proposition that feral pigs may be pivotally important to the avian diseases system, at least
in remote Hawaiian rain forests. The author observed an increment of rain-filled cavities in
tree fern trunks eaten by pigs and verified that such cavities are the most abundant and
productive habitat for larval mosquitoes.

Conclusions
Direct consumption of native plant species by feral pigs is the most common impact
reported in the literature. Foraging and trampling by pigs can also cause severe erosion and
lead to the degradation of watersheds (Cuddihy and Stone 1993). However, perhaps the
most dramatic ecological consequence of feral pig introduction to the Hawaiian Islands
may be the environmental effects they trigger in biotic communities. They do this, along
with many other animal species, by influencing the population dynamics of native species
and promoting the dispersal and establishment of non-indigenous species, thus altering the
composition and structure of communities (Diong 1982; Cuddihy and Stone 1993). The
feeding habits of pigs and consequent habitat alteration may also indirectly negatively
impact native forest birds (van Riper and Scott 2001; LaPointe 2006). Despite the many
potential negative impacts to native biota and ecosystems generated by pig activities,
eliminating the pig from Hawaiian Islands remains difficult if not impossible, mostly
because many Hawaiians further value it for its cultural, and religious significance (Stone
1985).
There are needed improvements in assessing the adequacy of sample sizes, experimental study designs, issues of scale, spatial perspectives and statistical analyses so that we
can better assess the role of pigs in Hawaii ecosystems. In other words, methods have to be
appropriate to the question asked. Elucidating the dynamics and long-term ecological
effects generated by pigs is a crucial step towards increasing our understanding of and
more effectively managing biotic interactions. An example of this problem is found with
studies that seek to obtain pig home range area and movement pattern data. Many if not all
studies of pig movements in Hawaii occurred during pig eradication efforts, when the

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animals were being chased and killed (e.g., Diong 1982). Data obtained in this manner are
useful in developing methods to eradicate feral pigs more efficiently. However, this
approach is unlikely to produce representative measures of pig home range area and
movement patterns without hunting pressure, which are also crucial information to
establish appropriate control measures. Therefore, much remains to be learned about the
strength of the aforementioned interactions and their long-term effects on Hawaiian biota
and ecosystems. Elucidating the dynamics and long-term ecological effects generated by
pigs is a crucial step towards increasing our understanding of and more effectively managing biotic interactions.
Acknowledgments We thank Edwin D. Johnson, Statewide Hunting Coordinator, for giving us access to
DLNR/DOFAW files, David Duffy, for giving us access to PCSU files, and Biodiversity and Conservation
anonymous reviewers. SSCN and SLGNF were supported by CNPq (Proc. 200335/2005-7) and CAPES
(Proc. 0597-05-8) Brazilian Educational Agencies provided funding through their grants while in Hawaii.
We thank Carter Miller, Janisete Gomes da Silva Miller, Lynn Schnurr and Sean Giery for providing helpful
comments to improve the final manuscript. This research was supported by the Koolau Mountains
Watershed Partnership.

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