Biology and Impacts of Pacific Island Invasive Species. 1. A Worldwide
Review of Effects of the Small Indian Mongoose, Herpestes javanicus(Carnivora: Herpestidae)1
Warren S. T. Hays2 and Sheila Conant3
Abstract: The small Indian mongoose, Herpestes javanicus (E. Geoffroy SaintHilaire, 1818), was intentionally introduced to at least 45 islands (including 8
in the Pacific) and one continental mainland between 1872 and 1979. This small
carnivore is now found on the mainland or islands of Asia, Africa, Europe,
North America, South America, and Oceania. In this review we document the
impact of this species on native birds, mammals, and herpetofauna in these areasof introduction.

The small Indian mongoose has been
introduced to numerous islands, including
eight in the Pacific. Beyond its native range
in southern Asia, this species now occurs on
islands or mainlands elsewhere in Asia, Africa,
Europe, North America, South America, andOceania. Its negative effects on native biotaof these areas are a concern to natural-areamanagers.

1), typically has an adult body mass in therange of 300 to 900 g and a body lengthfrom 500 to 650 mm (Nellis 1989). It has
the slender body shape typical of the herpestid family, with short legs, short brown fur,and a tail that makes up roughly 40% of theanimal’s total length. Dentition is 3 : 1 : 4 : 2,with a wide carnassial shear region. Females
have 36 chromosomes and males have 35,
because the Y chromosome has translocatedonto an autosome (Fredga 1965).
A more complete species description may
be found in Nellis (1989).

name
Herpestes javanicus (E. Geoffroy Saint-Hilaire,
1818)
Synonym: Herpestes auropunctatus (Hodgson,
1836)
Common names: Small Indian mongoose,
small Asian mongoose, Javan mongoosediet
No study has yet been done on the diet of thesmall Indian mongoose in its native range(Figure 2). Table 1 summarizes studies ofsmall Indian mongoose diets in various areasof introduction. In all these studies, exceptthat from Korcula, Croatia, this species ofmongoose was found to be an omnivore thatis primarily insectivorous and secondarily ahunter of small vertebrates.
The most extensive of these studies wasthat of Gorman (1975), who examined 4,404
mongoose scats on Viti Levu, Fiji. He foundthat diets varied according to habitat. For
example, crabs were the main food item inmangrove forests, rats were most common in
cane-field scats, and cockroaches were mostcommon in scats from urban areas. On Korcula, the mongooses were found to be primarily frugivorous (Cavallini and Serafini

description
The small Indian mongoose, Herpestes javanicus (E. Geoffroy Saint-Hilaire, 1818) (Figure

1 Manuscript accepted 19 April 2006.
2 Department of Biology, Hawai‘i Pacific University,
Honolulu, Hawai‘i 96813 (e-mail: whays@hpu.edu).
3 Department of Zoology, University of Hawai‘i at
Mānoa, Honolulu, Hawai‘i 96822.

Pacific Science (2007), vol. 61, no. 1:3–16: 2007 by University of Hawai‘i Press
All rights reserved

3

4

PACIFIC SCIENCE . January 2007

Figure 1. The small Indian mongoose, Herpestes javanicus. Photo by R. J. Shallenberger.

1995). That study was conducted during December and January in a temperate environment, so the seasonally high abundance ofthe fruits of juniper ( Juniperus oxycedrus) andstrawberry tree (Arbutus unedo) and low abundance of insect prey may have affected foodchoice.
In all of the studies listed in Table 1,
mammal prey was mainly murid rodents:
the house mouse (Mus musculus), Norway rat(Rattus norvegicus), and roof rat (R. rattus);
however, Cavallini and Serafini (1995) foundthat mongooses on Korcula preyed uponevery available species of small mammal except shrews (Insectivora: Soricidae), which
are avoided by many species of predators. Insect prey was varied in most studies, including
substantial numbers of orthopterans, coleopterans, and lepidopterans. Mostello (1996)
found a preponderance of cockroaches (Orthoptera: Blattidae) among invertebrates inscats from two Hawaiian sites: Lualualei Valley, O‘ahu, and Mo‘omomi dunes, Moloka‘i.
Small Indian mongooses have been observed to eat toads (family Bufonidae), including the parotid glands, which many predators

find noxious (Nellis and Everard 1983).
By comparison, cats refuse to eat toads, anddogs that consume toads typically become
sick or even die. Mongooses also eat littoral
animals in Hawai‘i, including crabs (suborder
Brachyura) (La Rivers 1948, Baldwin et al.
1952). Nellis and Everard (1983) found themto refuse land crabs (Cardisoma guanhumi)
persistently on St. Croix, U.S. Virgin Islands,
though Wolcott (1953) found this species ofcrab in 4 of 108 examined mongoose stomachs from St. Croix and Puerto Rico.
La Rivers (1948) examined the contents of
23 mongoose stomachs from Pearl Harbor,O‘ahu, and reported that reptiles were a largepart of the diet. He also described two individuals working together to hunt crabs (Metapograpsus messor). One turned over stones andthe other attacked any crabs that emerged.
Nellis and Everard (1983) kept captive
mongooses for several years on a diet consisting of 50% pork liver and 50% ripe papaya,
with no water other than that contained in
the foodstuffs. They reported that mango
and papaya were favored foods of captive
and hand-raised animals.

Figure 2. Native range and routes of introduction of the small Indian mongoose.

PACIFIC SCIENCE . January 2007

6
TABLE 1

Diet of the Small Indian Mongoose as Determined by Scatological and Stomach Samples
Diet

St. Croix1

Puerto Rico2

Fiji3

Korcula 4

Hawai‘i5

Hawai‘i6

(36)
18%
5%
9%
59%
11%

(56)
3%
0%
6%
70%
11%

(4,404)
10%
4%
19%
44%
23%

(126)
26%
13%
0%
16%
45%

(86)
29%
8%
0%
45%
18%

(73)
39%
3%
2%
86%þ
8–15%

(Sample size)
Mammals
Birds
Herpetofauna
Invertebrates
Plants

1 U.S. Virgin Islands (Seaman 1952).
2 Pimentel (1955).
3 Gorman (1975).
4 Adriatic Sea (Cavallini and Serafini 1995). Many of the other data in this table were also recalculated by those authors.
5 Baldwin et al. (1952).
6 Mostello (1996). Unlike the other columns, these data reflect incidence, not percentage mass.

economic importance andenvironmental impact
Impact on Bird Populations
Two bird species, both of them groundnesters, were probably driven to extinction
by the small Indian mongoose. In Fiji, thebarred-wing rail (Nesoclopeus poecilopterus) was
described as common in 1875; in 1883, themongoose was introduced, and within a few
years the bird was extinct (Gorman 1975).
Similarly, the Jamaica petrel (Pterodroma caribbaea) was abundant in the late eighteenth
century; the mongoose was introduced to Jamaica in 1872, and this bird has not been seen
since 1893 (Collar et al. 1992). A third possible case of mongoose-driven extinction, theJamaican poorwill (Siphonorhis americanus), ismore poorly evidenced. This bird has notbeen seen since 1859, and Bangs and Kennard (1920) proposed that the mongoose
was responsible. However, later authors havedoubted this, because the bird seems to havevanished over a decade before mongooses
were introduced to Jamaica. Collar et al.
(1992) blamed deforestation and rats.
Stone et al. (1994) stated, without citing
sources, that the small Indian mongoose is a‘‘known or suspected’’ nest predator upon atleast eight federally listed endangered birds
in Hawai‘i. These include the nēnē or Hawaiian goose (Branta sandvicensis), the ‘alalā or
Hawaiian crow (Corvus hawaiiensis), the koloa
or Hawaiian duck (Anas wyvilliana), the alaeke‘oke‘o or Hawaiian coot (Fulica alai), theae‘o or Hawaiian stilt (Himantopus mexicanus
knudseni), the alae ‘ula or Hawaiian gallinule(Gallinula chloropus sandvicensis), the ‘ua‘u or
Hawaiian petrel (Pterodroma sandwichensis),and the ‘a‘o or Newell shearwater (Puffinus
auricularis newelli). Documentation of thispredation is, however, rare. Land managers
in Hawai‘i frequently find signs of mammalian nest predation on protected birds but
can rarely determine the species of predator.
Baldwin et al. (1952) found remains of Hawaiian petrels in 5 of 86 mongoose scats theyexamined from rocky pastures above 600 m
elevation on leeward Hawai‘i Island. Kingand Gould (1967) suggested, without citing
sources, that the mongoose was responsible
for the extirpation or reduction of the Newellshearwater on O‘ahu, Moloka‘i, Maui, andHawai‘i. Mongooses have also been blamed,again without citation of evidence, for local
extirpations of the Hawaiian duck (Anonymous 1968).
Four once-common bird species havebeen extirpated from all Fijian islands withmongooses but persist on mongoose-free islands: the banded rail (Rallus philippensis),sooty rail (Porzana tabuensis), white-browed
rail (Poliolimnas cinereus), and purple swamphen (Porphyrio porphyrio) (Gorman 1975). The
uniform crake (Amaurolimnas concolor) was extirpated from Jamaica sometime after 1881,
and Raffaele et al. (1998) opined that themongoose may have been a major factor.

Impact of Herpestes javanicus on Native Vertebrates . Hays and Conant

On St. Croix, the bridled quail dove (Geotrygon mystacea) was originally a ground nester and was thought extinct in 1921, 47 yr
after the introduction of the mongoose. Nellis and Everard (1983) reported, however,that this bird had actually become an arboreal
nester and had become fairly common. Allen(1911) blamed mongooses for the extirpationof a species of quail dove (genus Geotrygon)
from St. Vincent. Wetmore (1927) blamedmongooses for the decline in Puerto Rico of
five ground-nesting birds: two species ofquail doves, the short-eared owl (Asio flammeus), the West Indian nighthawk (Chordeiles
gundlachii), and the Puerto Rican nightjar(Caprimulgis noctitherus). The latter species
was believed extinct for about 30 yr before
1961 and now exists as an endangered species.
It is found both in areas that do and do notcontain mongooses, but is less abundant in
the former (Vilella and Zwank 1987, 1993).
The short-eared owl is also found sympatrically with the small Indian mongoose in
Hawai‘i, but the mongoose has not been citedas having a deleterious effect via predation on
populations of the Hawaiian subspecies (Asioflammeus sandwichensis).
On St. Lucia and Martinique, the mongoose and other introduced predators presumably have added to the pressure that has
reduced the populations of white-breasted
thrasher (Ramphocinclus brachurus). But Collar
et al. (1992) opined that these factors wereless important than habitat destruction andcompetition by the bare-eyed thrush (Turdus
nudigenis), an invasive species. After a longstudy of the effect of mongooses on Trinidad, Urich (1931) found that no birds had
been extirpated, including ground-nesting
species, during the 61 yr since the mongoose’s introduction.
The degree to which mongooses are responsible for the historical decline of bird
species is often hard to assess, because of exacerbating factors such as introduction of rats,
cats, dogs, and pigs, and habitat encroachment by human communities. It must be
noted that any bird species now living in
the presence of mongoose populations in
Hawai‘i has been doing so for over a century.
It has been suggested that ground-nesting

7

bird populations have established a predatorprey equilibrium with mongooses in the
Caribbean (Westermann 1953). This may
also be true in Hawai‘i, though it is surely
also true that the mongoose’s presence posesa substantial barrier to reestablishment ofground-nesting bird populations in their historical ranges.
Impact on Mammal Populations
Woods and Ottenwalder (1992) reported thatbones found in owl pellets demonstrate thatthe following four species of small mammals
were present on Hispaniola in 1930: theHaitian island shrew (Nesophontes hypomicrus),
the small Haitian island shrew (N. zamicrus),
the large Haitian island shrew (N. paramicrus), and the Hispaniolan spiny rat (Brotomys
voratus). All four species are now believed
extinct. The same authors proposed that themongoose might have been a contributing
factor to these extinctions, basing their opinion largely on the timing of the mongoose introduction, which they state variously as 1895
(p. 109) or 1907 (p. 87).
Mongooses were introduced to Cuba in
1886 (Nellis and Everard 1983), and Westermann (1953) believed that mongooses werelargely responsible for the extinction of the
Cuban solenodon (Solenodon cubanus) (Insectivora: Solenodontidae) around 1910. This species has since been rediscovered in eastern
Cuba, though it is still endangered (Nowak1991).
There is a common story in Hawai‘ithat small Indian mongooses failed to control
rats in areas of introduction because themongoose is diurnal and rats are primarily
nocturnal (Stone et al. 1994). Most publishedaccounts dispute this story, asserting that thesmall Indian mongoose served as an excellent
cane-field ratter (Pemberton 1925, Barnum
1930, Doty 1945), though it was eventually
made obsolete by the development of improved techniques of rat poisoning (Doty
1945).
On Trinidad, Urich (1931) found that rats
were rare in cane fields, though they had beena major pest before the introduction of themongoose in the 1870s. By 1882, a govern-

8ment botanist estimated that the mongoose in
Jamaica was saving the colony 100,000
pounds sterling (current value: $8.3 million)
per year (Espeut 1882). Spencer (1950, citedby Seaman [1952]), however, found that roof
rat populations were as high as 50 per hectare
in some parts of St. Croix, despite the presence of mongooses. Seaman (1952) wrotethat some cane fields on St. Croix still suffered 25% crop loss due to rats and believedthat rats were as much a problem as before
the introduction of the mongoose.
Another common story is that mongooses
drove rats to become arboreal nesters in areasof introduction (Nellis and Everard 1983).
On Hawaiian islands with mongooses, Polynesian rats (Rattus exulans) and Norway rats
are terrestrial nesters, whereas roof rats arearboreal nesters. This appears to be true on
islands with and without mongooses in Hawai‘i and throughout the world (Baldwin
et al. 1952). There is, however, evidence thatmongooses alter the relative abundance ofrats in favor of arboreal roof rats (Walker
1945). In Puerto Rico, Norway rats arecommon only in mongoose-free urban areas,
whereas roof rats are found in mongoose
habitat (Pimentel 1955). Hoagland et al.
(1989) made a census of populations of mongooses and rats on St. Croix and Jamaica, andfound more roof rats and fewer Norway rats
in mongoose habitat.
Nellis (1989) stated that mongooses ‘‘often
dominate over’’ cats (Felis catus [domesticus]),though the degree to which they limit theabundance of feral cats in areas of sympatry
has not been studied. Feral cats and wildmongooses peacefully share food at artificial
feeding sites on O‘ahu, feeding within centimeters of each other (W.S.T.H., pers. obs.).
More pertinently, on 3 June 1999, while
doing a radio-tracking study, one of us(W.S.T.H.) observed two large male mongooses pass together within 3 m of an adultferal cat, in a relatively undisturbed woodlot
and apparently by coincidence, without anyof the animals involved showing any sign ofexcitement or stress even while making eye
contact. This anecdotal observation suggeststhat adults of these species can coexist inpeaceful sympatry, at least under some condi-

PACIFIC SCIENCE . January 2007

tions, though it is also possible that they may
harry or prey upon each other’s young.
Seaman and Randall (1962) gave a description, including firsthand observations, of
predation of mongooses upon fawns of theintroduced white-tailed deer (Odocoileus virginianus) on St. Croix.
Impact on Reptile and Amphibian Populations
The reputation of the mongoose as an
extinction-driver for Caribbean herpetofauna
is partly due to exaggeration, though certainly
mongooses do reduce populations of some
reptiles and amphibians. Gosse (1851) reported that the Jamaica giant galliwasp (Celestus occiduus), an anguid lizard, was fairly
abundant in the mid-1840s. It is now widelybelieved to be extinct, though at least one
herpetologist has expressed doubts on thispoint (Crombie 1999). If this species is indeed extinct, the timing of its extinction is
unclear, so it is hard to link this extinction
firmly to the introduction of mongooses
in 1872. Henderson and Sajdak (1996) statedthat the mongoose was the primary causeof the apparent extinction of the Hispaniola
racer (Alsophis melanichnus). Westermann(1953) reported that the mongoose had
driven the black racer (Alsophis ater) extinct
on Jamaica, but in fact this snake still exists,
albeit in low numbers (Henderson 1992).
Westermann (1953) and Nellis (1979) also
blamed mongooses for the extinction of
the Saint Croix racer (A. sanctaecrucis) on St.
Croix in 1859 and the extirpation of theorange-bellied racer (A. rufiventris) on Nevis
Island in 1879, but mongooses were not introduced to those islands until 25 and 21 yr,
respectively, after the last official sightings ofthose snakes (Henderson 1992). Westermann(1953) further blamed mongooses for theextinction of several species of Dromicus
snakes in the Lesser Antilles, though infact these species were ‘‘never even close tobeing threatened’’ (Henderson 1992:6).
Urich (1931) believed mongooses might have
driven the skinks Mabuya aurata and M. agilis
extinct on Trinidad, but these species persist
today. He also blamed the mongoose for the
possible extinction of the black tegu lizard

Impact of Herpestes javanicus on Native Vertebrates . Hays and Conant(Tupinambis nigropunctatus) on Trinidad, but
this species is now common again, despite
the continuing presence of mongooses (Nellis and Everard 1983).
Mongooses apparently did, however, extirpate the Antiguan racer (Alsophis antiguae)from Antigua, the Martinique clelia snake(Liophis cursor) from Martinique, and theSanta Lucia groundsnake (L. ornatus) from
St. Lucia. Each of these species persists onmongoose-free satellite islands (Henderson
1992). Corke (1987) blamed the mongoose
for the extirpation of the windward clelia(Clelia clelia) and the skink Mabuya mabuya
on St. Lucia, but Henderson (1992) believed
the latter species too small and reclusive tobe judged extirpated on existing evidence.
The small Indian mongoose was introduced
to Martinique and St. Lucia (dates of introduction unknown) to control the fer-de-lance(Bothrops atrox), a venomous species of pit viper (de Vos et al. 1956). Some authors blame(or credit) the mongoose with the extirpationof this snake on those islands (Barbour 1930,
Nellis 1989), but it is uncertain whether mongooses actually attack these snakes in the wild.
Hinton and Dunn (1967) stated that mongooses are commonly killed by the fer-delance in ‘‘fighting pit’’ shows, but Nellis andEverard (1983) believed that the mongoose
usually wins.
In 1910, the small Indian mongoose wasintroduced to three Croatian islands to control the horned viper (Vipera ammodytes). In
the same year, it was also introduced to Okinawa to control the habu pit viper (Trimeresurus flavoviridis). The effects of these two
introductions on the local snake populations
have not been studied.
The mongoose was largely responsible forthe extirpation of the Saint Croix ground lizard (Ameiva polops) on St. Croix (Baskin andWilliams 1966). These lizards had coexistedwith mongooses for 90 yr, and were extirpated only when the sidewalks that were their
final refuge were torn up in the late 1960s.
Allen (1911) blamed the mongoose for the
decline of Ameiva species in Grenada, butthere have been no extirpations there (Henderson 1992).
Mongooses were probably largely respon-

9sible for the extirpation of the two largest
skinks, Emoia nigra and E. trossula, from thetwo largest Fijian islands, Viti Levu and Vanua Levu (Zug 1991). These species vanishedfrom those islands within a few years of themongoose’s arrival, and they are still foundon mongoose-free Fijian islands. Case andBolger (1991) made a census of diurnal lizard
abundance on seven Pacific islands with mongooses and 11 islands without mongooses by
walking transects in a variety of different habitat types. They found that there was nearly
100 times the abundance of diurnal lizards
on mongoose-free islands.
Small Indian mongooses prey upon at least
four endangered species of sea turtles. Nellisand Small (1983) documented 14 instances
within a 2-yr period in which mongooses
preyed upon the eggs or young of the hawksbill sea turtle (Eretmochelys imbricata) on Caribbean islands. A 1982 government study
on St. Croix found that mongooses killed
5% of leatherback turtle (Dermochelys coriacea) hatchlings (unpubl. study, quoted by Nellis and Small [1983]). Seaman and Randall(1962) reported strong evidence of mongoosenest predation upon green sea turtles (Chelonia mydas). Nellis and Small (1983) statedthat small Indian mongooses prey on nestsof loggerhead sea turtles (Caretta caretta).
The edible frog Leptodactylus pentadactylus
has been extirpated from the three Caribbean
islands of its original range to which mongooses have been introduced but persists on
two mongoose-free islands (Barbour 1930).
Gorman (1975) found frog remains in mongoose scats collected from Fijian rain forests,
suggesting that mongooses may have been
the cause of the observed decline of two
native ground frogs (Platymantis vitianus andP. vitiensis).
It is often hard to assess the degree ofblame that should be affixed to mongooses inherpetofaunal decline. Nellis (1979) documented effects on the ecological communityof a 5.5-ha Caribbean island when five pairsof mongooses were added experimentally.
Within a year, a ground lizard (Ameiva exsul )had been extirpated, and, apparently as a result, prickly pear moths (Cactoblastis cactorum)and geckos of the genera Spaerodactylus andPACIFIC SCIENCE . January 200710

Hemidactylus had become more common. The
flush in the moth population was accompanied by a crash of the prickly pear cactus(Opuntia) population. This experiment demonstrates that mongoose introductions candisrupt fragile island ecosystems and are capable of causing a herpetofaunal extirpation
under some circumstances.
Henderson (1992) proposed that the mongoose was to blame for a few herpetofaunal
extirpations in the Lesser Antilles but believed that habitat destruction was more toblame on the larger Caribbean islands. Aneven more conservative opinion was given by
Corke (1992:47), who, after examining sevenislands of the Lesser Antilles, stated, ‘‘Thereis no conclusive evidence for the widely reported role of introduced mongooses in the
decline of the islands’ reptiles.’’geographic distribution in thepacific region
The small Indian mongoose is currently
found on eight major Pacific islands. In Hawai‘i it is found on Hawai‘i, Maui, Moloka‘i,
and O‘ahu; in Fiji it is found on Viti Levu
and Vanua Levu; and in Japan it is found on
Okinawa and Amami.

On the Caribbean islands of St. Croix
and Trinidad, a comparison of population
estimates from various habitats has shownthat dry natural areas are preferred and rainy
areas are avoided (Nellis and Everard 1983).
Another Caribbean study, on St. John, also
found that small Indian mongooses weremore abundant in dry, low-elevation communities (Coblentz and Coblentz 1985).
In Hawai‘i, however, wet areas are exploited by dense populations of mongooses.
Baldwin et al. (1952) suggested that humid
areas are even preferred by this species in Hawai‘i and reported seeing an adult mongoosein deep forest in Kı̄pahulu Valley on Maui(annual rainfall: 750 cm per year); however,they added that this was a rare sighting. Tomich (1979) found a high density of mongooses in Waipi‘o Valley, Hawai‘i Island,
where rainfall is 200 to 450 cm per year, depending on exact location.
The small Indian mongoose has been reported in Nepal (part of its natural range) at
2,200 m (Prater 1936). Baldwin et al. (1952)
stated that it is found to elevations over
3,000 m in Hawai‘i, but Tomich (1986) expressed doubts that Hawaiian populations
exist much above 2,400 m elevation.

habitathistory of small indian mongoose
introductions

The small Indian mongoose is often reported
to have an aversion to rain and water, whichmight suggest that xeric habitats should be
preferred. This possibility is also supportedby the observation that trap success in removal studies falls to near zero in rainy
weather, even at usually successful sites (Nellis and Everard 1983) and even if the rain
continues for several days (Nellis 1973).
No study has determined habitat preferences of small Indian mongooses in their native range. Lekagul and McNeely (1977:600)
stated that in Thailand these animals ‘‘seem
to prefer grasslands and secondary growth todense forests.’’ According to Corbet and Hill
(1992), in the very dry northwestern portionof its native range the species is confined tomesic habitats and frequently is found aroundhuman habitation.

The small Indian mongoose has been introduced intentionally into wild habitats more
widely than any other mammal (Figure 2).
Indeed, all documented introductions of thisspecies have been intentional, because of its
reputation as a control agent upon rats andsnakes. It is also an efficient predator upon
many endangered and threatened vertebrates,
and has been considered by at least one land
manager to be ‘‘a vicious pest and wildlife’s‘enemy No. 1’ ’’ (Seaman 1952:188).
The introduced small Indian mongoose is
the only species of mongoose in the Western
Hemisphere and Polynesia. In 1872, a sugar
planter introduced four male and five female
small Indian mongooses to Jamaica fromeastern India (Espeut 1882). From Jamaica,
mongooses eventually were introduced to 29
islands throughout the Caribbean and also toImpact of Herpestes javanicus on Native Vertebrates . Hays and Conant

the northern coast of South America (Hoagland et al. 1989). An earlier New World
introduction of some unknown mongoose
species was made at Trinidad in 1870 (Urich[1914], cited in Hoagland et al. [1989]), but
this and all other original Caribbean mongoose introductions apparently failed, except
the one in Jamaica (Espeut 1882, Nellis andEverard 1983). Attempts were also made, before the introduction of mongooses, to introduce ferrets (Mustela putorius) to Jamaica, butthese attempts also failed, purportedly due tochigoes (‘‘burrowing fleas’’ [Tunga penetrans])(Hinton and Dunn 1967).
The range of the small Indian mongooseon the South American mainland is limited
to the developed coastal strip of Guyana, Surinam, and French Guiana (Nellis and Everard 1983), where it was introduced in 1900
(Hinton and Dunn 1967). The range of thismainland population appears to be limited,
for now, by a boundary of swamp and rain
forest (Nellis and Everard 1983). About 100
mongooses were also sent from Croatia to
Venezuela in 1926 (Tvrtkovic and Krystufek
1990) but apparently did not establish there.
In 1883, sugar planters imported the small
Indian mongoose from Jamaica to four Hawaiian islands (Hawai‘i, O‘ahu, Maui, andKaua‘i) and to the Fijian island of Viti Levu
(Gorman 1975, Nellis and Everard 1983).
For unknown reasons, the crate of mongooses was kicked off the dock at Kaua‘i, andto date the mongoose has apparently not established there, although a dead mongoose
was found in Kaua‘i in 1976 (Tomich 1986).
Mongooses were later introduced to the Hawaiian island of Moloka‘i and to the Fijian island of Vanua Levu.
The small Indian mongoose exists on Javaand Macau, and may have been introduced tothese islands hundreds of years ago. More recently, it has been introduced to other islandsof Indonesia and the Indian Ocean, including
Ambon, Mauritius, and Mafia (Nellis andEverard 1983).
This species has been introduced to three
Croatian islands of the Adriatic: Mljet (in1910), Korcula (in 1910), and Hvar (in 1970
[Tvrtkovic and Krystufek 1990]). It was introduced to the Peljesac Peninsula in Croatia

11

repeatedly in the 1920s, and a population existed there as late as 1936 but apparently diedout (Tvrtkovic and Krystufek 1990).
The small Indian mongoose was also
successfully introduced to Okinawa in 1910
(Ogura et al. 1998) and to nearby Amami Island in 1949 and/or 1979 (Abe et al. 1991).
A failed attempt was made in the late nineteenth century to introduce a mongoose into
Australia to control rabbits, but it is unknown
whether this was the small Indian mongooseor another species (Hinton and Dunn 1967).
physiology and behavior
Mean body temperature of small Indianmongooses is 39.5 C. The mean resting heart
rate is 252 beats per minute, and the mean
resting respiratory rate is 63 breaths per minute (Nellis and Everard 1983). Metabolic energy use for this species, as deduced fromoxygen consumption of restrained individuals, has been variously measured at 116 kcal/
kg/day (Lin and Kobayashi 1976) and 76
kcal/kg/day (Ebisu and Whittow 1976). The
expected energy use for mammals in the size
range of small Indian mongooses (300 to 600g) is 72 to 95 kcal/kg/day (Kleiber 1961). The
thermal neutral zone is narrow and very
high: 28 to 38 C (Ebisu and Whittow 1976).
Small Indian mongooses are, however, able tomaintain their body core temperature in therange of 0 to 41 C, though they show signsof extreme heat stress at 45 C (Nellis andMcManus 1974). Heat-stressed mongooses
pant and salivate on their chest and forelimbs
but do not lick them (Ebisu and Whittow1976). They can respond to cold by raising
their metabolic rate up to threefold (Nellisand McManus 1974).
The eye is emmetropic with at least 11 diopters of accommodation (Nellis et al. 1989),
and 25–40% of the photoreceptors are cones(Hope et al. 1982), suggesting that this species has unusually good vision for a small
mammal and can discern colors. The kidney
can achieve a urine concentration of >6,000
milliosmoles, suggesting adaptation to desert
conditions (Nellis 1989). There is a large,
appendix-like cecum (Palmiter and Horst
1982), but this appears to be lymphatic rather

PACIFIC SCIENCE . January 2007

12

than digestive in character. Most fat is stored
in the tail, which holds up to seven times thefat per mass of the body (Nellis and Everard1983).
This species is entirely diurnal (Kavanau
1975). Typical movements are a low slink anda gallop. Small Indian mongooses can swim
and climb trees (Nellis and Everard 1983) but
rarely do so. When killing vertebrate prey,small Indian mongooses deliver a bite to theback of the head, crushing the skull. Foraging
behavior has not been observed extensively or
described in this species, because it tends toforage under cover and is difficult to observe
for long, continuous periods. They form shallow burrows for use as dens.
Both male and female small Indian mongooses have a pair of pocket-shaped glands
lateral to an extensible pouch around theanus, rich in sebaceous and apocrine glands(Gorman et al. 1974). These structures accumulate lipids and proteins that support grampositive bacteria, which produce a mixtureof six saturated carboxylic acids. Mongooses
use a ‘‘handstand’’ marking behavior to place
this substance on vertical surfaces, supportingthemselves on their forepaws while marking
with the anal gland. Small Indian mongooses
are able to distinguish between the scentmarks of individual conspecifics in choicetests (Gorman 1976a), though the smell ofthese marks is undetectable by humans.
This species makes 12 distinct varieties of
vocalization (Mulligan and Nellis 1973),
including, for example, a contact call usedby young mongooses when following theirmother, a feeding call, and several agonistic
cries that are graded by intensity of stress.
Young small Indian mongooses show strong
following responses, and follow their mothervery closely until they are fully grown. Males
show some degree of sociality during thebreeding season (Hays and Conant 2003),
but the social system has not been fully
elucidated.
reproduction and population
dynamics
Pregnant females are found from February
through August in Hawai‘i (Pearson and

Baldwin 1953), Fiji (Gorman 1976b), and the
U.S. Virgin Islands (Nellis and Everard 1983),
all of which are between 17 and 20 degrees oflatitude either north or south of the equator.
Grenada, which is at 12 N, has a 10-month
breeding season (Nellis and Everard 1983).
Males produce sperm year-round in Hawai‘i(Pearson and Baldwin 1953), but the size ofthe testes changes seasonally: they are largest
from February through July and smallest in
October and November (Pearson and Baldwin 1953, Soares and Hoffman 1981). The
size of the prostate gland follows the same
cycle, as do serum levels of sex hormones(androgens, follicle-stimulating hormone, andluteinizing hormone) in males (Soares andHoffman 1981). Sperm are also produced
year-round in Fiji, where testes are largest
from September to December (equivalent toMarch through June in the Northern Hemisphere) (Gorman 1976b).
Females have a 4-day estrus at 20-day
intervals during the breeding season (Asdell
1964) plus postpartum estrus (Nellis andEverard 1983). Ovulation is induced by copulation (Asdell 1964). There is no delayed
implantation (Tomich and Devick 1970),
as there is in several other small carnivores(Sadleir 1969). Gestation takes 49 days, andpregnancy is not usually evident without
dissection (Nellis and Everard 1983). Meanlitter size is unknown, but examination of 255
wild-caught pregnant females on Grenada
showed them to have a mean of 2.2 embryos,
with a range of 1 to 5 (Nellis and Everard1983). The number of litters produced annually by a typical female in the wild has not yet
been determined with any certainty.
Pups begin accompanying their mother on
hunting trips at 6 weeks of age (about 200 gbody mass). Young mongooses follow theirmothers until 4 to 6 months of age, at leastin captivity, and this is also the range of ages
at which both genders attain sexual maturity(Nellis and Everard 1983). The youngest
wild-caught pregnant female with a confidently estimated age (determined by mass ofthe eye lens) was 4 months old (Nellis andEverard 1983).
Population densities have not been determined with certainty but may be extremelyImpact of Herpestes javanicus on Native Vertebrates . Hays and Conant

high in some regions of introduction, judging
by the small size of individual home ranges
found in a few studies. Nellis and Everard(1983) reported mean home ranges of 2.2 hafor two radio-tracked females and 3.6 ha forfive radio-tracked males on St. Croix. Hays
and Conant (2003) found mean home rangesof 1.4 ha for seven radio-tracked females
on O‘ahu. They also found that five radiotracked males had broadly overlapping homeranges in the breeding season, sharing a region of about 20 ha. All of these individual
home ranges are very small, in comparison
with those of typical small carnivores. Carnivores in the mass range of the small Indianmongoose (300 to 900 g) are expected tohave individual home ranges of 26 to 115 ha(Harestad and Bunnell 1979). No study has
been done of the home ranges of small Indian mongooses in their natural range, but it
may be speculated that populations are abnormally dense in many parts of the introduced range.
management
Despite its wide range of introduction and its
formidable reputation as a predator, the small
Indian mongoose can only be confidently said
to have driven three vertebrates to extinction
anywhere in the world: the barred-wing rail,
the Jamaica petrel, and the Hispaniola racer(and in each of those cases, there is room for
doubt). Considering the great number of islands the mongoose has been introduced to,and the great number of ecologically delicate
species it preys upon on these islands, thisfact may at first seem remarkable. Many othercommon anthropically introduced mammals,
such as cats, dogs, and rats, have probably
caused more than two extinctions apiece. But
this fact in itself may be a large part of the explanation of the relatively low number of extinctions directly attributable to mongoose
introductions: the mongoose was a latecomer
to its areas of introduction, and on mostislands simply added one further burden toalready-damaged ecosystems. By the time it
arrived, much of the destruction was alreadydone.
Nonetheless, the continued existence of13

the introduced mongoose is undoubtedly amajor impediment to the reestablishment ofmany extirpated or reduced vertebrate populations on dozens of islands. In most of theselocations, this is a role that the small Indianmongoose shares with other introduced predators, as well as introduced competitors
and habitat degradation caused by humanactivities.
Almost all efforts at mongoose population
management in areas of introduction havefocused on the use of traps to remove individuals from ecologically sensitive areas. These
areas include particularly the nesting sites ofrare, ground-nesting birds. Trapping programs are of limited use, because removing
the mongooses from a strip of habitat aroundthe sensitive area creates a region of empty
habitat that lures more mongooses into thearea. These programs are also very expensive,
because monitoring traps is a labor-intensive
procedure.
Diphacinone, an anticoagulant that kills byinternal hemorrhaging, first became available
for experimental use in 1988. It has proved tobe especially effective against mongooses, but
its use as a control agent is still primarily experimental (Keith et al. 1990, Smith 1998),
and it is not currently being widely dispersed
in areas inhabited by introduced mongooses.
prognosis
The small Indian mongoose is strongly entrenched in most of its areas of introduction.
It appears to have high population densities
on many large islands, is often found in habitat that would have been unsuitable for it
within its natural range, and in some places(e.g., Hawai‘i) has no natural predators andfew communicable diseases and parasites.
Only the most drastic measures, such as aerial
dispersion of diphacinone bait pellets, would
have any chance of eliminating such populations from even small islands, and the ecological costs of such efforts might outweigh theirbenefits in many cases. Perhaps the single
most crucial management issue is to preventthe further spread of the species. Fortunately,
new small Indian mongoose populations
are not likely to become established without

PACIFIC SCIENCE . January 200714

human agency, though this could occur
unintentionally (e.g., by release of pet mongooses).
acknowledgments
We thank an anonymous reviewer who supplied us with several useful references.
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