Applied Animal Behaviour Science 108 (2007) 111

www.elsevier.com/locate/applanim

Review

Feral pigs in Hawaii: Using behavior and ecology

to refine control techniques
Selene Siqueira da Cunha Nogueira a,c,*,
Sergio Luiz Gama Nogueira-Filho b,c, Michael Bassford c,
Kirsten Silvius c, Jose Manuel Vieira Fragoso c
a

Department of Biology Sciences, Universidade Estadual de Santa Cruz, Rod. Ilheus,

Itabuna km 16, Ilheus, BA 45662000, Brazil
b
Department of Agrarian and Environmental Sciences, Universidade Estadual de Santa Cruz,
Rod. Ilheus, Itabuna km 16, Ilheus, BA 45662000, Brazil
c
Department of Botany, University of Hawaii at Manoa, 3190 Maile Way, St. John Lab 101,
Honolulu, HI 96822, USA
Accepted 15 March 2007
Available online 23 April 2007

Abstract
Early Polynesians settlers were the first to introduce pigs to the Hawaiian Islands. Later Captain Cook
brought European pigs during his first voyage to Hawaii. Many other importations have followed. Animals
from these introductions became feral and dispersed throughout the islands. Free-ranging pigs are now
considered pests with negative impacts on some native biota. Several methods to control the ecological
damage attributed to pigs have been adopted, such as fencing, hunting, live trapping and poisoning.
However, the absence of behavioral knowledge in current control programs has resulted in inefficient
management of this species. Therefore, the feral pig problem continues, and what before was almost strictly
an agricultural and conservation concern has now become an urban problem as well. The aim of this study is
to describe the state of knowledge on feral pig behavior in the Hawaiian Islands, introducing potential
management approaches derived from the principles of behavioral ecology. Considering behavioral aspects
of feral pig ecology, such as cognition and communication could help improve capture techniques, keep
feral pigs away from urban areas and begin to resolve humanwildlife conflicts.
# 2007 Elsevier B.V. All rights reserved.
Keywords: Sus scrofa; Invasive species; Wildlife management; Pest management; Vertebrate pest control

* Corresponding author at: Department of Biology, Universidade Estadual de Santa Cruz, Rod. Ilheus, Itabuna km 16,
Ilheus, BA 45662000, Brazil. Tel.: +55 7336805262.
E-mail address: selene@uesc.br (S. Nogueira).
0168-1591/$ see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.applanim.2007.03.011

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

Contents
1.
2.
3.

4.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
2.1. Description of the study areas . . . . . . . . . . .
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
3.1. Food and feeding habits . . . . . . . . . . . . . . .
3.2. Activity, home range and movement patterns .
3.3. Social aggregation and reproductive behavior
3.4. Sensory behavior . . . . . . . . . . . . . . . . . . . .
3.5. Population estimation from pig-sign . . . . . . .
3.6. Feral pig control techniques in Hawaii. . . . .
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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2
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1. Introduction
The early Polynesians settlers originally introduced pigs to the Hawaiian Islands as an
important source of food (Tomich, 1986). Presumably all eight main islands of Hawaii that
supported Polynesian settlements supported populations of Polynesian pigs, probably animals
descended from the Asiatic form of Sus scrofa (Larson et al., 2005). These animals were smaller
in size than European pigs and some researchers report that they may have remained in
domestication (e.g., Tomich, 1986). Captain Cook brought European pigs on his first voyage to
Hawaii (Cook, 1785). Many imports have followed and some 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 breeds of European origin breeds (Tomich, 1986). However,
in a more recent study on feral pig mitochondrial DNA analyses, Larson et al. (2005) found that
modern Hawaiian feral pigs fall strongly within the Indonesian/Polynesian haplotype and not
within the European one. Hawaiian pigs may therefore be closely related to the original
Polynesian pigs, and not to the introduced European pigs as formerly believed.
Feral pigs are found throughout the Hawaiian archipelago, with the exception of Lanai and
Kahoolawe (Giffin, 1978; Tomich, 1986). However, there once were pigs on Lanai, but they
were eradicated (Edwin Johnson; Hawaii State hunting coordinator, personal communication).
On Kauai, Oahu and Maui they are restricted to dense rainforests and to a few private ranch
areas (Tomich, 1986). On the Island of Hawaii, free-ranging pigs are found from coastal dry
lands through the dense fern forests to the arid upper slopes of Mauna Loa and Mauna Kea
(Giffin, 1978).
Currently, feral pigs are considered pests with negative impacts on native Hawaiian biota.
Several authors have documented the detrimental effects of this animal in forest communities
(e.g., Spatz and Mueller-Dumbois, 1975; Jacobi, 1976; Cooray and Mueller-Dumbois, 1981).
Diong (1982) considered feral pigs the primary modifiers of remaining Hawaiian rainforests.
They consume native tree ferns and effectively disperse strawberry guava (Psidium cattleianum)
seeds, an alien tree that is invading the native forests (Smith and Diong, 1977).
Due these detrimental effects several management plans were devised that attempted to
control the ecological damage caused by pigs (Tomich, 1986; Jenkins et al., 1994). An extensive

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

fence network and intensive hunting program were able to reduce, but not eradicate, the feral pig
population from the Hawaii Volcanoes National Park (Giffin, 1978; Pratt et al., 1999). This
program exemplifies the difficulty of managing this species.
The feral pig problem continues and what before was almost strictly a rural and ecological
concern has now become an urban problem as well (pers. obs. authors). Records of feral pigs
activity in urban areas have recently increased in the local press, causing concern to homeowners
and public health authorities.
Incorporating behavioral approaches, which has not been tried in Hawaii, could help in the
development of more effective control techniques and management plans for feral pigs. Singh and
Kaumanns (2005) demonstrated how information on behavior and life history could improve
strategies for wildlife management. Long-term studies of behavior can lead to the development of
non-lethal control methods such as the development of repellents (Baker et al., 2005; Launchbaugh
and Howery, 2005; Littin and Mellor, 2005; Sato, 2001; Singh and Kaumanns, 2005).
There are no studies published in the scientific literature concerning pig behavior in Hawaii,
although there is some information in technical reports from the Hawaii Division of Forestry and
Wildlife (DOFAW) and the US National Park Service (NPS). However, these behavioral data
were not related to management concerns. Not incorporating this information in control
programs has resulted in inefficient management of this species. Our goal is to review what is
known about Hawaiian feral pig behavior and relate this information to control methods. In
addition to promote discussion on management techniques that will improve feral pig control in
the Hawaiian islands.

2. Methods
This review is a compilation of feral pig behavioral data available through local governmental agencies.
The data were available in technical reports from DOFAW, the United States Geological Survey (USGS),
and the NPS, two theses produced by University of Hawaii students and material published in books
describing invasive species in Hawaii. From these sources we collate information on feral pig food and
feeding habits, activity, home range and movement patterns, social aggregations and reproductive behavior,
sensory behavior and methods adopted to control feral pigs in Hawaii.
2.1. Description of the study areas
These studies comprised areas of the Hawaii Vulcanoes National Park on the island of Hawaii, and
Kipahulu Valley on Maui. The majority of suitable pig habitat lies between 2500 and 3100 m elevation of the
Hawaii Vulcanoes National Park with predominant mountain-pasture vegetation. Temperatures range from
bellow freezing in the winter to 29.4 8C, in the summer (Giffin, 1978). The annual rainfall in this semi-arid
region is approximately 58.9 mm, and the majority of the precipitation occurs during the winter and spring
months. Droughts are common and seriously affect pig reproduction and survival (Giffin, 1978). The lower
portion of this study area is heavily wooded with mamane (Sophora chrysophylla) and naio (Myoporum
sandwicense), while the understory is composed of numerous introduced pasture grasses. These include
orchard grass (Dactylis glomerata), sweet vernal (Anthoxanthum odoratum) and mesquite (Holcus lanatus).
Pukeawe (Styphelia tameiameiao), a native shrub, is also common at higher elevations.
While the rain forest habitats of the island of Hawaii studied included the Koala Watershed,
Laupahoehoe and Piha segments of the Hilo Forest Reserve, Hilo Watershed, Upper Waiakea and Olaa
Forest Reserves. In this habitat, precipitation occurs during all months of the year but is more frequent
during the winter and spring and the average annual rainfall is up to 832 mm. Free water is abundant all year
in the form of small streams and potholes (Giffin, 1978). Vegetation consists mainly of Hawaiian koa

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

(Acacia koa), banana poka (Passiflora mollissima), akala (Rubus hawaiiensis), ferns (Sadleria cyatheoides,
Athirium sandwichianum), waiawi (P. cattleianum) and sphagnum moss bogs are found in some of these
areas (Giffin, 1978).
The Kipahulu Valley is located on the eastern slope of Haleakala Volcano on the island of Maui and is
managed as a closed-entry scientific reserve within Haleakala National Park. The studys areas comprised a
bend in the Valley at 720-m to the Kalapawili grassland at 2380 m elevation. The minimal annual rainfall has
been estimated at about 3080 mm, and temperature ranges from 8 to 25 8C. The coastal lowland ecosystem,
from 460-m to sea level is comprised of grasslands, woodland, streams, ocean sprayzone and the coastal
waters (Anderson, 1994). Patches of Psidium guajava, P. cattleianum, Mangifera indica and Aleurites
moluccana are interspessed in the gulches and exotic grasslandan agricultural area. The dominant species
are Acacia koa, A. moluccana and Cheirodendron trygynum forming the upper stratum and the lower story
formed by P. guajava, P. cattleianum and Cibotium sp. (Diong, 1982).

3. Results
3.1. Food and feeding habits
Nichols (1963), Baker (1975), Giffin (1978), Diong (1982) and Anderson (1994) investigated
feral pig food habits in Hawaii. They describe this species as omnivorous but concluded that
most of its diet consisted of plant matter. In a mountain-pasture habitat in Hawaii Volcanoes
National Park, grasses accounted for 4145% of pigs stomach contents, followed by other plant
parts (flower, bark, fern fronds, rhizomes, etc.) at 29.550%, carrion at 20.5% and unidentified
species of earthworms at around 1% (Baker, 1975; Giffin, 1978). Fruits represented only 1.5% of
the volume of stomach contents. In Hawaiian tropical forests with native Citobium tree ferns and
sweet-fruited alien plant species, trunks and fronds of Citobium represent from 25 to 56% of the
stomach contents, grass and sedge 0 to 45%, fruits from strawberry guava (P. cattleianum) and
passion fruit (Passiflora edulis) 23 to 45%, banana poka (Passiflora mollissima) about 32%,
grasses 2.1 to 45% and earthworms 5 to 6.2% (Giffin, 1978; Diong, 1982). In a habitat with
Citobium tree ferns but without exotic fruits, the trunks and fronds of Citobium represents from
47 to 91% of the volume of stomach contents, sedges and grasses 2.1 to 4.1%, grass and sedge 2
to 10%, earthworms from 1 to 3% and fruits only 0.3% (Anderson, 1994).
In rain forests lacking tree ferns but plentiful in sweet-fruited alien species, guava and papaya
(Carica papaya) represented 70% of the stomach contents of feral pigs, whereas passion fruit
fruits, noni (Morinda citrofolia), kukui nuts (A. moluccana) and grasses totaled 15% and land
snails, land slugs, earthworms, insects and caterpillars represented a further 15% (Diong, 1982).
At least two studies report that earthworms are the most important source of animal protein for
Hawaii feral pig populations (Diong, 1982; Anderson, 1994). Diong (1982) related rooting sign
to searching for earthworms that pigs swallow whole. Carrion, another source of protein, was
important in pig diets during the intensive feral goat and pig control activities of the 1970s in
Hawaii Volcanoes National Park (Baker, 1975; Giffin, 1978).
Despite its wide dietary range, captive studies show some food preferences in Hawaii pigs.
For example, Kikuta and Stone (1986) determined that feral pigs strongly preferred sweet foods,
such as papaya, sweet potatoes and Citobium tree fern trunks. In a Maui rainforest, Diong (1982)
noted that the feral pigs shifted from a diet composed primarily of native tree ferns (Citobium
spp.), a concentrated source of sugar and starch, to a diet comprised mostly of sugary strawberry
guava (P. cattleianum) fruits when they were seasonally available. Giffin (1978) observed a
similar shift in a rain forest on Hawaii Island, where diets dominated by tree fern starch shifted to
sweet banana poka fruits as they became seasonally available. These then made up the majority

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

of the diet. In contrast, Anderson (1994) did not find any significant relationships between feral
pig activity and the abundance of plant species used as food in the upper Hana rainforest on Maui.
Anderson (1994) attributed the lack of relationship between feral pig activity and the
abundance of potential food plants was due to an absence of sweet-fruited species for which pigs
have strong feeding preferences (alien plant species with sweet, succulent fruits). He did however
posit a link between earthworm density and pig activity in forests.
Diong (1982), through direct observation of pigs feeding, observed that they preferred young
shoots, leaves and fronds over older parts, and they fed in distinct ways on different plant species.
For example, when feeding on grass, leaves at the center of the plant are usually browsed,
whereas while feeding on flowering plants, as Eupatorium sp., leaves are stripped off the plant
and eaten in preference over the stems. Diong (1982) also observed that tree bark feeding
appeared to be the result of individual animals foraging, while trunk feeding of tree ferns was a
group foraging activity. Diong (1982) speculated that it took a group of pigs to push over a tree
fern; he did not however, obtain field evidence of tree fern felling by pigs, but suggested that pigs
weaken tree ferns through rooting around their bases, contributing to their fall.
3.2. Activity, home range and movement patterns
On Hawaii island, feral pigs activity peaks in late afternoon, night and early morning (Giffin,
1978). Diong (1982) followed the daily activity cycle of five feral pigs in Kipahulu Valley, Maui
and found a similar pattern. The first peak was around 06:00 h and the second peak between
06:00 and 09:00 h. These data suggest night and afternoon resting periods between active
foraging periods. Giffin (1978) suggested a circadian cycle of feral pigs was related to their age.
He found that younger animals often moved during daytime hours while older boars preferred to
move during dawn and dusk.
Giffin (1978) studied the movement patterns of feral pigs in both mountain-pasture and rain
forest habitats on Hawaii island and discovered that pigs in pastures move greater distances than
those in forests. He speculated that this occurred because water and cover are less abundant in
pastures. He also determined that boars travel less than sows. With the average daily distance
traveled by boars per day in the rain forest habitat being 0.9 km and for sows, 1.2 km. The
difference between males and females was greater in mountain-pasture (1.6 km per day for males
versus 2.2 km per day for females). In contrast, Diong (1982) found that boars travelled farther
than sows in his forested study area, with boars moving a mean of 1.2 km and sows 0.7 km over
24-h. However, in both studies, the animals have being chased for eradication when they were
tracked. This fact could disturb on pigs movement data.
Giffin (1978), using a mark-recapture technique estimated feral pig home ranges at 5.2
10.4 km2 in mountain-pasture habitat, and from 1.3 to 5.2 km2 in rain forest habitat. He reported
that feral pigs have overlapping home ranges that they do not actively defend. Diong (1982)
determined that the home range of boar (2.0 km2) was two times the home range of sows
(1.1 km2). Giffin (1978) did not find any indication of seasonal movements, but observed
movements in response to changes in food availability and a trend for pigs . . . to follow water
holes on the ranches, during several drought periods.
3.3. Social aggregation and reproductive behavior
Diong (1982) found both solitary and grouping behavior in animals in the Kipahulu Valley,
Maui. The solitary individuals were males, either an aged boar or a bachelor older than 1 year.

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

The 58 groups observed by this author consisted of family units and mixed groups of adult males
and females with an average of 2.6 and up to 9 individuals per group. The family groups consisted
of either a single sow and her most recent offspring or extended families, which incorporated
offspring from the previous season. The mixed groups were comprised of sub-adults and adults
with or without juveniles. On Hawaii island, Giffin (1978) found groups of up to 10 pigs resting
in the same nest. He suggested that these groups consisted of both related and unrelated animals
grouped temporarily, whereas a normal or usual group consisted of one or two sows and their
offspring. Groups could also consist of different litters of weaned young traveling and feeding
together.
Reproductive information for feral pigs in the Hawaiian Islands is derived mainly from Diong
(1982), who observed breeding all year round. The author noted two annual peaks in birthing, the
first from November to March and the second from June to September. The November to March
farrowing season coincided with long periods of heavy rains and late winter storms. The author
registered that five months was the minimum breeding age for sows and these nursed offspring
for only 3 months. Giffin (1978) observed that birthing was related to the winter-spring rainy
season at mountain-pasture on the island of Hawaii, and speculated that this was related to
greater food and water availability during this time of the year. In rainforest habitat, despite yearround water availability, the same author recorded a pronounced fall birthing season that
coincided with an increase in wild fruit production. Giffin (1978) reported that this was the only
habitat where sows farrowed twice a year. He estimated the minimum breeding age for sows at 10
months. In both habitats sows build elaborate nests when ready to farrow, placing dry grass or
ferns in the lee of bushes or rocks (Giffin, 1978). The piglets remained at these nests until 2 or 3
weeks old. The nursing piglets were weaned at around 3 or 4 months of age at which point they
either leave their mothers or remain with them for several more months.
Piglets normally develop a suckling order soon after birth and tend to nurse from the same teat
at each feeding (Pond and Houpt, 1978). This habit allowed Giffin (1978) to estimate the litter
size from the number of functioning teats on a sow; through this method the author found an
average of 4.3 and 3.4 young per sow at rainforest and mountain-pasture habitats, respectively.
3.4. Sensory behavior
Among pigs and other terrestrial mammals that occupy large home ranges and or display
fissionfusion social organization, vision can be a poorly developed sense (Drea et al., 2002). In
agreement with this generalization, Giffin (1978) and Diong (1982) found that feral pigs have
poor vision. They rely on acoustic signals and olfactory communication to maintain group
cohesion and social organization. Pig pheromones serve to regulate group behavior. These are
produced by the mammary and sub-maxillary glands in females and preputial and salivary glands
in boars (Pond and Houpt, 1978). The male pheromones serve both to attract sows in estrus and
mediate the boars social ranking within a group. Diong (1982) described two behavioral patterns
for boars when in groups with a sow in estrus: the boar slashed the bases of tree trunks with his
canines, marking them with foamy saliva or the boar held its snout directly upward, sniffing the
air and repeatedly snapping its jaws.
3.5. Population estimation from pig-sign
The amount of sign cannot be used to directly estimate the number of pigs since the amount of
sign a single pig will leave and the conditions that determine this amount are not known (Ohashi,

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

1988), this has implications for the use of sign in monitoring relative pig abundance. Pig-sign
however, is useful in determining trends in relative abundances and is often employed to measure
the effectiveness of feral pig control program. Hone and Stone (1989) developed a monitoring
method that uses these behavioral cues along with other pig-sign, as an indicator of relative
abundance. This technique uses the presence of fresh tracks, rooting, wallows, rubs, tusk marks,
beds and feces, recorded for 10 m intervals along transects, to produce a pig abundance index.
3.6. Feral pig control techniques in Hawaii
The current methods used to control feral pigs and the damage they cause are the same ones
applied more than 20 years ago (Cooray and Mueller-Dumbois, 1981; Diong, 1982; Ohashi,
1988). In general they involve trapping, hunting and fencing within protected areas.
Traps used to capture pigs have been either box or corral traps (Ohashi, 1988). Some animals, in
contrast, become trap wary over long-term live-trapping programs, especially older boars that
overturn box-traps or dig under corral traps. To avoid these problems, Diong (1983) designed a
metal box-trap with a dual trigger mechanism, which allows managers to counteract such trap
cautious behaviors. By alternating the root-bar-trigger system with the pull-pin-trigger, it is
possible to counter trap wary behavior in the field. Another innovation made by Diong (1983) was
the use of a multiple-catch door in order to increase the number of animals caught at one time.
Despite the advantages of box-traps, this method is limited to areas within reasonable
proximity to vehicular access (Ohashi, 1988) or by helicopter (Diong, 1983). Thus, leg snare
capture is the technique currently used by most wildlife managers and is particularly effective in
remote areas with dense vegetation (Anderson and Stone, 1993).
Several snares are put out in the late afternoon and captured pigs are shot the next morning.
There is concern for the welfare of leg-snared animals, challenging the usefulness of leg snares,
and leg snares have low capture rates. For example, Anderson and Stone (1993) used a density of
up to 118 snares per 100 ha to remove only 1.5 pigs. The effort expended to snare pigs ranged
from 0.2 to 10.0 person-days per pig, and the snare-night effort per pig increased from 940 to
6853 during the last 3 months of this control program.
Hunting with dogs may be a more efficient capture method. For example, Giffin (1978)
determined that hunting with dogs was the major factor responsible for low pig densities in state
Forest Reserves on Hawaii island. However, recreational/public hunting has not been shown to
be effective in remote areas (Ohashi, 1988). Furthermore, heavy hunting pressure results in less
game, which eventually results in lower interest by hunters and less hunting pressure. With fewer
hunters, the game population increases and hunter interest returns with subsequent higher
hunting pressure (Ohashi, 1988). Therefore, without adequate hunting pressure, which depends
on area accessibility and hunter interest, the required level of control may not be achieved.
Controlling feral pig damage in Hawaii is accomplished through fencing (Ohashi, 1988; Pratt
et al., 1999). However, extensive fencing is an unrealistic alternative to the long-term control of
pigs due to the high costs of fence building and subsequent maintenance (Ohashi, 1988).
The last report of the Hawaii Animal Control Research Consortium (Jenkins et al., 1994)
suggested four research options capable of producing improvements in ungulate control. One is
developing a radio telemetry system that signals when animals are caught in snares and traps
nowadays commercially available (Trap Monitor1, Innotech Communications). Another is to
employ more humane capture methods, in consideration of the well being of the animals. The
third option is to increase hunting with dogs. Lastly, researchers should obtain better statewide
game data that permit hunting management using a rigorous scientific approach.

S. Nogueira et al. / Applied Animal Behaviour Science 108 (2007) 111

4. Discussion
Feral pigs are not easy to observe in the wild. Nevertheless, Giffin (1978) and Diong (1982)
obtained reliable feral pig behavioral data for Hawaii. One of the most important aspects of
ecological behavior presented by these two authors was pig feeding behavior. Food habits of feral
pigs can alter the structure and dynamics of Hawaiian ecosystems (Giffin, 1978; Diong, 1982). In
one area, feeding pressure was directed at roughly 40 plant food species, most of them native
plants (Diong, 1982). Rooting associated with the search for earthworms (Diong, 1982) initiated
soil erosion, enhanced seed beds and promoted invasion of exotic weeds. It also removed moss,
liverwort and native herbaceous ground cover. Direct feeding on native trees reduces their
abundance and affected tree regeneration. Finally, feral pigs helped disperse strawberry guava
seeds (Diong, 1982).
Behavioral studies show that feral pigs aggregate seasonally to take advantage of seasonally
available foods (Giffin, 1978; Diong, 1982). This seasonal behavior, when coupled with vegetation
maps and movement data can facilitate more effective pig control by allowing hunters to
concentrate traps in important food areas (Stone and Taylor, 1984). Similarly, movements in
response to drought-period searches for water (Giffin, 1978) could further target trap placement and
increase capture rates. To accomplish such behaviorally targeted trapping, we need to increase our
knowledge of the feral pig food preferences and ecology of food plants in different habitats.
Most studies on feral pig behavioral ecology to date occurred in national parks on Maui and
Hawaii island (Spatz and Mueller-Dumbois, 1975; Jacobi, 1976; Cooray and Mueller-Dumbois,
1981). These studies documented detrimental effects to native biota. Ohashi (1988) noted that
most results of this nature have been used to justify intensive management programs within the
national parks, although those comprise only a very small percentage of important forest
communities in the Hawaiian Islands. Few if any behavioral studies have taken place in areas that
are not protected or isolated. Studies are needed in these areas, and methods other than
eradication must be developed to control pig populations. Additionally, there are topics that still
lack systematic study, such as feral pig social interactions, pheromone communication and in the
area of capturing, bait improvement.
The rough terrain in Hawaii makes feral pig control programs difficult to implement. Leg
snares are the method of choice to capture pigs in remote areas due to lightweight and portability
of the snares (Anderson and Stone, 1993). However, catch rates decrease with time in a given
area, and more young than adults are caught (Anderson and Stone, 1993). An explanation for this
development is that hunting pressure induces feral pigs to move away from the snaring area, at
least temporarily (Giffin, 1978).
This problem could be overcome through the joint use of leg snares and the box-trap with
double trigger mechanism (Diong, 1983) in remote areas (Ohashi, 1988). This combined
approach could be evaluated by monitoring pig-signs. This method also should be useful to
determine the path that the animals followed trying to escape the traps and indicate the best
places to locate the box-traps. Both leg snares and box-traps could be used together to harvest
animals from areas where they are considered noxious for their relocation into game or hunting
areas. In this way it is possible to efficiently decrease the number of feral pigs in protected areas,
and counteract concerns associated with feral pigs control programs.
Another reason to catch and remove pigs alive is the fact that they eat carrion (Baker, 1975;
Giffin, 1978). This feeding behavior was observed during the intensive feral goat and pig controls
in Hawaii Volcanoes National Park in 1970s. Thus, even if authorities use lethal control
methods pigs corpses should be removed, for sanitary and nutritional reasons, since corpses can

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spread disease and can serve as a source of protein for pigs. The increased food may help increase
the reproductive rates of remaining animals.
Knowledge of the reproductive cycles of feral pigs can help managers identify the best periods
to implement control programs. For example sows are more vulnerable, due to pregnancy, from
October through January in the rain forest on the island of Hawaii (Giffin, 1978), and from
September through December on Maui (Diong, 1982).
In Kipahulu valley Diong (1982) reported that group behavioral reactions against humans
might be countered by studying sensory functions in feral pigs. Pigs have been observed to smell
humans from nearly a half mile away. This acute sense also is used to detected leg snares on
Oahu, even when the hunters use gloves. Feral pigs can smell traps and avoid or jump over them
making trapping and control work more difficult (Nogueira-Filho, personal observation).
Anderson and Stones (1993) decreased rate of capture over time clearly shows that despite the
several advantages of the leg snare (Anderson and Stone, 1993) we still need to develop means to
reduce animal detection of traps.
Pigs communicate through smell in more complex ways than any other sense (Giffin, 1978;
Diong, 1982). For example, smell and hearing is very important in regulating group formation
and social behavior; these senses should be maximally exploited in the development of baiting
programs and other control techniques. It is important to develop highly efficient baiting
materials to develop alternative and more cost-effective methods of improving feral pig control.
A new idea is to use pheromones as bait for traps as proposed by Diong (1982) or to disguise
human scent left at traps. In other mammal species, the olfactory system is used to detect prey or
predators. Hyenas, for example, are crepuscular hunters and scavengers that compete against other
predators; spotted hyenas rely on their olfactory sense to locate prey and to warn of danger (Kruuk,
1972; Mills, 1990). Morven et al. (2005) reported on the discrimination of co-specifics by juvenile
domestic pigs, showing that juvenile pigs can discriminate between familiar group mates from
different litters using normal social cues. In this context, we believe that Hawaiian feral pigs
pheromone communication must be studied more systematically to provide better techniques for
their control. Olfactory-based cues might also help in the development of chemical repellents as a
potential non-lethal control technique (Baker et al., 2005; Littin and Mellor, 2005).
Knowledge of animal behavior can assist in all facets of pig management. Exploring the links
between behavioral ecology, feeding behavior, reproductive pattern, social structure, cognition,
communication and wildlife management science will make it possible to more effectively
control feral pig populations. There is a strong need for more behavioral studies addressing
management concerns and for publishing data in easily available, peer reviewed journals.
Acknowledgments
We thank Edwin D. Johnson, Statewide Hunting Coordinator, for giving us access to DOFAW
files. 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. This work was concluded through the project Koolau Mountains Watershed
Partnership funded by the DOFAW.
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