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Biology and Impacts of Pacific Island Invasive . 1. A Worldwide Review of Effects of the Small Indian , javanicus (: Herpestidae)1

Warren S. T. Hays2 and Sheila Conant3

Abstract: The , Herpestes javanicus (E. Geoffroy Saint- Hilaire, 1818), was intentionally introduced to at least 45 islands (including 8 in the Pacific) and one continental mainland between 1872 and 1979. This small is now found on the mainland or islands of , , Europe, North America, South America, and Oceania. In this review we document the impact of this species on native , , and herpetofauna in these areas of introduction.

The small Indian mongoose has been 1), typically has an adult body mass in the introduced to numerous islands, including range of 300 to 900 g and a body length eight in the Pacific. Beyond its native range from 500 to 650 mm (Nellis 1989). It has in southern Asia, this species now occurs on the slender body shape typical of the herpes- islands or mainlands elsewhere in Asia, Africa, tid , with short legs, short brown fur, Europe, North America, South America, and and a tail that makes up roughly 40% of the Oceania. Its negative effects on native biota ’s total length. is 3:1:4:2, of these areas are a concern to natural-area with a wide carnassial shear region. Females managers. have 36 and males have 35, because the Y has translocated name onto an autosome (Fredga 1965). A more complete species description may Herpestes javanicus (E. Geoffroy Saint-Hilaire, be found in Nellis (1989). 1818) : Herpestes auropunctatus (Hodgson, diet 1836) Common names: Small Indian mongoose, No study has yet been done on the diet of the small Asian mongoose, small Indian mongoose in its native range (Figure 2). Table 1 summarizes studies of description small Indian mongoose diets in various areas of introduction. In all these studies, except The small Indian mongoose, Herpestes javani- that from Korcula, Croatia, this species of cus (E. Geoffroy Saint-Hilaire, 1818) (Figure mongoose was found to be an omnivore that is primarily insectivorous and secondarily a hunter of small vertebrates. The most extensive of these studies was that of Gorman (1975), who examined 4,404 1 Manuscript accepted 19 April 2006. 2 Department of Biology, Hawai‘i Pacific University, mongoose scats on Viti Levu, . He found Honolulu, Hawai‘i 96813 (e-mail: [email protected]). that diets varied according to habitat. For 3 Department of Zoology, University of Hawai‘i at example, were the main food item in Ma¯noa, Honolulu, Hawai‘i 96822. mangrove forests, were most common in cane-field scats, and cockroaches were most Pacific Science (2007), vol. 61, no. 1:3–16 common in scats from urban areas. On Kor- : 2007 by University of Hawai‘i Press cula, the were found to be pri- All rights reserved marily frugivorous (Cavallini and Serafini

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 De- find noxious (Nellis and Everard 1983). cember and January in a temperate environ- By comparison, refuse to eat toads, and ment, so the seasonally high abundance of that consume toads typically become the fruits of juniper ( Juniperus oxycedrus) and sick or even die. Mongooses also eat littoral strawberry tree (Arbutus unedo) and low abun- in Hawai‘i, including crabs (suborder dance of prey may have affected food Brachyura) (La Rivers 1948, Baldwin et al. choice. 1952). Nellis and Everard (1983) found them In all of the studies listed in Table 1, to refuse land crabs (Cardisoma guanhumi) prey was mainly murid : persistently on St. Croix, U.S. , the (Mus musculus), Norway though Wolcott (1953) found this species of (Rattus norvegicus), and roof rat (R. rattus); in 4 of 108 examined mongoose stom- however, Cavallini and Serafini (1995) found achs from St. Croix and . that mongooses on Korcula preyed upon La Rivers (1948) examined the contents of every available species of small mammal ex- 23 mongoose stomachs from Pearl Harbor, cept shrews (Insectivora: Soricidae), which O‘ahu, and reported that were a large are avoided by many species of predators. In- part of the diet. He also described two indi- sect prey was varied in most studies, including viduals working together to hunt crabs (Meta- substantial numbers of orthopterans, coleop- pograpsus messor). One turned over stones and terans, and lepidopterans. Mostello (1996) the other attacked any crabs that emerged. found a preponderance of cockroaches (Or- Nellis and Everard (1983) kept captive thoptera: Blattidae) among invertebrates in mongooses for several years on a diet consist- scats from two Hawaiian sites: Lualualei Val- ing of 50% pork liver and 50% ripe papaya, ley, O‘ahu, and Mo‘omomi dunes, Moloka‘i. with no water other than that contained in Small Indian mongooses have been ob- the foodstuffs. They reported that mango served to eat toads (family Bufonidae), includ- and papaya were favored foods of captive ing the parotid glands, which many predators and hand-raised animals. Figure 2. Native range and routes of introduction of the small Indian mongoose. 6 PACIFIC SCIENCE . January 2007

TABLE 1 Diet of the Small Indian Mongoose as Determined by Scatological and Stomach Samples

Diet St. Croix1 Puerto Rico2 Fiji3 Korcula4 Hawai‘i5 Hawai‘i6

(Sample size) (36) (56) (4,404) (126) (86) (73) Mammals 18% 3% 10% 26% 29% 39% Birds 5% 0% 4% 13% 8% 3% Herpetofauna 9% 6% 19% 0% 0% 2% Invertebrates 59% 70% 44% 16% 45% 86%þ Plants 11% 11% 23% 45% 18% 8–15%

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 and ke‘oke‘o or Hawaiian coot (Fulica alai), the environmental impact ae‘o or Hawaiian (Himantopus mexicanus ), the alae ‘ula or Hawaiian gallinule Impact on Populations knudseni (Gallinula chloropus sandvicensis), the ‘ua‘u or Two bird species, both of them ground Hawaiian petrel (Pterodroma sandwichensis), nesters, were probably driven to and the ‘a‘o or Newell shearwater (Puffinus by the small Indian mongoose. In Fiji, the auricularis newelli). Documentation of this barred-wing (Nesoclopeus poecilopterus) was predation is, however, rare. Land managers described as common in 1875; in 1883, the in Hawai‘i frequently find signs of mamma- mongoose was introduced, and within a few lian nest predation on protected birds but years the bird was extinct (Gorman 1975). can rarely determine the species of predator. Similarly, the petrel (Pterodroma ca- Baldwin et al. (1952) found remains of Ha- ribbaea) was abundant in the late eighteenth waiian petrels in 5 of 86 mongoose scats they century; the mongoose was introduced to Ja- examined from rocky pastures above 600 m maica in 1872, and this bird has not been seen elevation on leeward Hawai‘i Island. King since 1893 (Collar et al. 1992). A third possi- and Gould (1967) suggested, without citing ble case of mongoose-driven extinction, the sources, that the mongoose was responsible ( americanus), is for the extirpation or reduction of the Newell more poorly evidenced. This bird has not shearwater on O‘ahu, Moloka‘i, , and been seen since 1859, and Bangs and Ken- Hawai‘i. Mongooses have also been blamed, nard (1920) proposed that the mongoose again without citation of evidence, for local was responsible. However, later authors have extirpations of the Hawaiian duck (Anony- doubted this, because the bird seems to have mous 1968). vanished over a decade before mongooses Four once-common bird species have were introduced to Jamaica. Collar et al. been extirpated from all Fijian islands with (1992) blamed deforestation and rats. mongooses but persist on mongoose-free is- Stone et al. (1994) stated, without citing lands: the banded rail (Rallus philippensis), sources, that the small Indian mongoose is a sooty rail (Porzana tabuensis), white-browed ‘‘known or suspected’’ nest predator upon at rail (Poliolimnas cinereus), and purple swamp- least eight federally listed endangered birds hen (Porphyrio porphyrio) (Gorman 1975). The in Hawai‘i. These include the ne¯ne¯ or Hawai- uniform crake (Amaurolimnas concolor) was ex- ian goose (Branta sandvicensis), the ‘alala¯ or tirpated from Jamaica sometime after 1881, (Corvus hawaiiensis), the koloa and Raffaele et al. (1998) opined that the or Hawaiian duck (Anas wyvilliana), the alae mongoose may have been a major factor. Impact of Herpestes javanicus on Native Vertebrates . Hays and Conant 7

On St. Croix, the bridled quail dove (Geo- bird populations have established a predator- trygon mystacea) was originally a ground nes- prey equilibrium with mongooses in the ter and was thought extinct in 1921, 47 yr (Westermann 1953). This may after the introduction of the mongoose. Nel- also be true in Hawai‘i, though it is surely lis and Everard (1983) reported, however, also true that the mongoose’s presence poses that this bird had actually become an arboreal a substantial barrier to reestablishment of nester and had become fairly common. Allen ground-nesting bird populations in their his- (1911) blamed mongooses for the extirpation torical ranges. of a species of quail dove ( Geotrygon) from St. Vincent. Wetmore (1927) blamed Impact on Mammal Populations mongooses for the decline in Puerto Rico of five ground-nesting birds: two species of Woods and Ottenwalder (1992) reported that quail doves, the short-eared owl (Asio flam- bones found in owl pellets demonstrate that meus), the West Indian nighthawk (Chordeiles the following four species of small mammals gundlachii), and the Puerto Rican were present on in 1930: the (Caprimulgis noctitherus). The latter species Haitian island shrew (Nesophontes hypomicrus), was believed extinct for about 30 yr before the small Haitian island shrew (N. zamicrus), 1961 and now exists as an . the large Haitian island shrew (N. paramic- It is found both in areas that do and do not rus), and the Hispaniolan spiny rat (Brotomys contain mongooses, but is less abundant in voratus). All four species are now believed the former (Vilella and Zwank 1987, 1993). extinct. The same authors proposed that the The short-eared owl is also found sympatri- mongoose might have been a contributing cally with the small Indian mongoose in factor to these , basing their opin- Hawai‘i, but the mongoose has not been cited ion largely on the timing of the mongoose in- as having a deleterious effect via predation on troduction, which they state variously as 1895 populations of the Hawaiian (Asio (p. 109) or 1907 (p. 87). flammeus sandwichensis). Mongooses were introduced to in On St. Lucia and , the mon- 1886 (Nellis and Everard 1983), and Wester- goose and other introduced predators pre- mann (1953) believed that mongooses were sumably have added to the pressure that has largely responsible for the extinction of the reduced the populations of white-breasted Cuban (Solenodon cubanus) (Insecti- thrasher (Ramphocinclus brachurus). But Collar vora: Solenodontidae) around 1910. This spe- et al. (1992) opined that these factors were cies has since been rediscovered in eastern less important than and Cuba, though it is still endangered (Nowak competition by the bare-eyed thrush (Turdus 1991). nudigenis), an invasive species. After a long There is a common story in Hawai‘i study of the effect of mongooses on Trini- that small Indian mongooses failed to control dad, Urich (1931) found that no birds had rats in areas of introduction because the been extirpated, including ground-nesting mongoose is diurnal and rats are primarily species, during the 61 yr since the mon- nocturnal (Stone et al. 1994). Most published goose’s introduction. accounts dispute this story, asserting that the The degree to which mongooses are re- small Indian mongoose served as an excellent sponsible for the historical decline of bird cane-field ratter (Pemberton 1925, Barnum species is often hard to assess, because of ex- 1930, Doty 1945), though it was eventually acerbating factors such as introduction of rats, made obsolete by the development of im- cats, dogs, and pigs, and habitat encroach- proved techniques of rat poisoning (Doty ment by communities. It must be 1945). noted that any bird species now living in On Trinidad, Urich (1931) found that rats the presence of mongoose populations in were rare in cane fields, though they had been Hawai‘i has been doing so for over a century. a major pest before the introduction of the It has been suggested that ground-nesting mongoose in the 1870s. By 1882, a govern- 8 PACIFIC SCIENCE . January 2007 ment botanist estimated that the mongoose in tions, though it is also possible that they may Jamaica was saving the colony 100,000 harry or prey upon each other’s young. pounds sterling (current value: $8.3 million) Seaman and Randall (1962) gave a de- per year (Espeut 1882). Spencer (1950, cited scription, including firsthand observations, of by Seaman [1952]), however, found that roof predation of mongooses upon fawns of the rat populations were as high as 50 per hectare introduced white-tailed deer (Odocoileus vir- in some parts of St. Croix, despite the pres- ginianus) on St. Croix. ence of mongooses. Seaman (1952) wrote that some cane fields on St. Croix still suf- Impact on and Populations fered 25% crop loss due to rats and believed that rats were as much a problem as before The reputation of the mongoose as an the introduction of the mongoose. extinction-driver for Caribbean herpetofauna Another common story is that mongooses is partly due to exaggeration, though certainly drove rats to become arboreal nesters in areas mongooses do reduce populations of some of introduction (Nellis and Everard 1983). reptiles and . Gosse (1851) re- On Hawaiian islands with mongooses, Poly- ported that the Jamaica giant galliwasp (Celes- nesian rats (Rattus exulans) and Norway rats tus occiduus), an anguid , was fairly are terrestrial nesters, whereas roof rats are abundant in the mid-1840s. It is now widely arboreal nesters. This appears to be true on believed to be extinct, though at least one islands with and without mongooses in Ha- herpetologist has expressed doubts on this wai‘i and throughout the world (Baldwin point (Crombie 1999). If this species is in- et al. 1952). There is, however, evidence that deed extinct, the timing of its extinction is mongooses alter the relative abundance of unclear, so it is hard to link this extinction rats in favor of arboreal roof rats (Walker firmly to the introduction of mongooses 1945). In Puerto Rico, Norway rats are in 1872. Henderson and Sajdak (1996) stated common only in mongoose-free urban areas, that the mongoose was the primary cause whereas roof rats are found in mongoose of the apparent extinction of the Hispaniola habitat (Pimentel 1955). Hoagland et al. racer ( melanichnus). Westermann (1989) made a census of populations of mon- (1953) reported that the mongoose had gooses and rats on St. Croix and Jamaica, and driven the black racer (Alsophis ater) extinct found more roof rats and fewer Norway rats on Jamaica, but in fact this still exists, in mongoose habitat. albeit in low numbers (Henderson 1992). Nellis (1989) stated that mongooses ‘‘often Westermann (1953) and Nellis (1979) also dominate over’’ cats ( catus [domesticus]), blamed mongooses for the extinction of though the degree to which they limit the the racer (A. sanctaecrucis) on St. abundance of feral cats in areas of sympatry Croix in 1859 and the extirpation of the has not been studied. Feral cats and wild orange-bellied racer (A. rufiventris) on Nevis mongooses peacefully share food at artificial Island in 1879, but mongooses were not in- feeding sites on O‘ahu, feeding within centi- troduced to those islands until 25 and 21 yr, meters of each other (W.S.T.H., pers. obs.). respectively, after the last official sightings of More pertinently, on 3 June 1999, while those (Henderson 1992). Westermann doing a radio-tracking study, one of us (1953) further blamed mongooses for the (W.S.T.H.) observed two large male mon- extinction of several species of Dromicus gooses pass together within 3 m of an adult snakes in the Lesser Antilles, though in feral , in a relatively undisturbed woodlot fact these species were ‘‘never even close to and apparently by coincidence, without any being threatened’’ (Henderson 1992:6). of the animals involved showing any sign of Urich (1931) believed mongooses might have excitement or stress even while making eye driven the Mabuya aurata and M. agilis contact. This anecdotal observation suggests extinct on Trinidad, but these species persist that adults of these species can coexist in today. He also blamed the mongoose for the peaceful sympatry, at least under some condi- possible extinction of the black tegu lizard Impact of Herpestes javanicus on Native Vertebrates . Hays and Conant 9

(Tupinambis nigropunctatus) on Trinidad, but sible for the extirpation of the two largest this species is now common again, despite skinks, Emoia nigra and E. trossula, from the the continuing presence of mongooses (Nel- two largest Fijian islands, Viti Levu and Va- lis and Everard 1983). nua Levu (Zug 1991). These species vanished Mongooses apparently did, however, extir- from those islands within a few years of the pate the (Alsophis antiguae) mongoose’s arrival, and they are still found from Antigua, the Martinique clelia snake on mongoose-free Fijian islands. Case and (Liophis cursor) from Martinique, and the Bolger (1991) made a census of diurnal lizard Santa Lucia groundsnake (L. ornatus) from abundance on seven Pacific islands with mon- St. Lucia. Each of these species persists on gooses and 11 islands without mongooses by mongoose-free satellite islands (Henderson walking transects in a variety of different hab- 1992). Corke (1987) blamed the mongoose itat types. They found that there was nearly for the extirpation of the windward clelia 100 times the abundance of diurnal (Clelia clelia) and the Mabuya mabuya on mongoose-free islands. on St. Lucia, but Henderson (1992) believed Small Indian mongooses prey upon at least the latter species too small and reclusive to four endangered species of sea turtles. Nellis be judged extirpated on existing evidence. and Small (1983) documented 14 instances The small Indian mongoose was introduced within a 2-yr period in which mongooses to Martinique and St. Lucia (dates of intro- preyed upon the or young of the hawks- duction unknown) to control the fer-de-lance bill sea turtle (Eretmochelys imbricata) on Ca- (Bothrops atrox), a venomous species of pit vi- ribbean islands. A 1982 government study per (de Vos et al. 1956). Some authors blame on St. Croix found that mongooses killed (or credit) the mongoose with the extirpation 5% of leatherback turtle (Dermochelys coria- of this snake on those islands (Barbour 1930, cea) hatchlings (unpubl. study, quoted by Nel- Nellis 1989), but it is uncertain whether mon- lis and Small [1983]). Seaman and Randall gooses actually attack these snakes in the wild. (1962) reported strong evidence of mongoose Hinton and Dunn (1967) stated that mon- nest predation upon green sea turtles (Chelo- gooses are commonly killed by the fer-de- nia mydas). Nellis and Small (1983) stated lance in ‘‘fighting pit’’ shows, but Nellis and that small Indian mongooses prey on nests Everard (1983) believed that the mongoose of loggerhead sea turtles (Caretta caretta). usually wins. The edible Leptodactylus pentadactylus In 1910, the small Indian mongoose was has been extirpated from the three Caribbean introduced to three Croatian islands to con- islands of its original range to which mon- trol the horned viper (Vipera ammodytes). In gooses have been introduced but persists on the same year, it was also introduced to Oki- two mongoose-free islands (Barbour 1930). nawa to control the habu pit viper (Trimeresu- Gorman (1975) found frog remains in mon- rus flavoviridis). The effects of these two goose scats collected from Fijian rain forests, introductions on the local snake populations suggesting that mongooses may have been have not been studied. the cause of the observed decline of two The mongoose was largely responsible for native ground (Platymantis vitianus and the extirpation of the Saint Croix ground liz- P. vitiensis). ard (Ameiva polops) on St. Croix (Baskin and It is often hard to assess the degree of Williams 1966). These lizards had coexisted blame that should be affixed to mongooses in with mongooses for 90 yr, and were extir- herpetofaunal decline. Nellis (1979) docu- pated only when the sidewalks that were their mented effects on the ecological community final refuge were torn up in the late 1960s. of a 5.5-ha Caribbean island when five pairs Allen (1911) blamed the mongoose for the of mongooses were added experimentally. decline of Ameiva species in Grenada, but Within a year, a ground lizard (Ameiva exsul ) there have been no extirpations there (Hen- had been extirpated, and, apparently as a re- derson 1992). sult, prickly pear moths (Cactoblastis cactorum) Mongooses were probably largely respon- and geckos of the genera Spaerodactylus and 10 PACIFIC SCIENCE . January 2007

Hemidactylus had become more common. The On the Caribbean islands of St. Croix flush in the moth population was accompa- and Trinidad, a comparison of population nied by a crash of the prickly pear cactus estimates from various habitats has shown (Opuntia) population. This experiment dem- that dry natural areas are preferred and rainy onstrates that mongoose introductions can areas are avoided (Nellis and Everard 1983). disrupt fragile island ecosystems and are capa- Another Caribbean study, on St. John, also ble of causing a herpetofaunal extirpation found that small Indian mongooses were under some circumstances. more abundant in dry, low-elevation com- Henderson (1992) proposed that the mon- munities (Coblentz and Coblentz 1985). goose was to blame for a few herpetofaunal In Hawai‘i, however, wet areas are ex- extirpations in the Lesser Antilles but be- ploited by dense populations of mongooses. lieved that habitat destruction was more to Baldwin et al. (1952) suggested that humid blame on the larger Caribbean islands. An areas are even preferred by this species in Ha- even more conservative opinion was given by wai‘i and reported seeing an adult mongoose Corke (1992:47), who, after examining seven in deep forest in Kı¯pahulu Valley on Maui islands of the Lesser Antilles, stated, ‘‘There (annual rainfall: 750 cm per year); however, is no conclusive evidence for the widely re- they added that this was a rare sighting. To- ported role of introduced mongooses in the mich (1979) found a high density of mon- decline of the islands’ reptiles.’’ gooses in Waipi‘o Valley, Hawai‘i Island, where rainfall is 200 to 450 cm per year, de- geographic distribution in the pending on exact location. pacific region The small Indian mongoose has been re- ported in Nepal (part of its natural range) at The small Indian mongoose is currently 2,200 m (Prater 1936). Baldwin et al. (1952) found on eight major Pacific islands. In Ha- stated that it is found to elevations over wai‘i it is found on Hawai‘i, Maui, Moloka‘i, 3,000 m in Hawai‘i, but Tomich (1986) ex- and O‘ahu; in Fiji it is found on Viti Levu pressed doubts that Hawaiian populations and Vanua Levu; and in Japan it is found on exist much above 2,400 m elevation. Okinawa and Amami. history of small indian mongoose habitat introductions The small Indian mongoose is often reported The small Indian mongoose has been intro- to have an aversion to rain and water, which duced intentionally into wild habitats more might suggest that xeric habitats should be widely than any other mammal (Figure 2). preferred. This possibility is also supported Indeed, all documented introductions of this by the observation that trap success in re- species have been intentional, because of its moval studies falls to near zero in rainy reputation as a control agent upon rats and weather, even at usually successful sites (Nel- snakes. It is also an efficient predator upon lis and Everard 1983) and even if the rain many endangered and threatened vertebrates, continues for several days (Nellis 1973). and has been considered by at least one land No study has determined habitat prefer- manager to be ‘‘a vicious pest and wildlife’s ences of small Indian mongooses in their na- ‘enemy No. 1’’’ (Seaman 1952:188). tive range. Lekagul and McNeely (1977:600) The introduced small Indian mongoose is stated that in these animals ‘‘seem the only species of mongoose in the Western to prefer grasslands and secondary growth to Hemisphere and Polynesia. In 1872, a sugar dense forests.’’ According to Corbet and Hill planter introduced four male and five female (1992), in the very dry northwestern portion small Indian mongooses to Jamaica from of its native range the species is confined to eastern (Espeut 1882). From Jamaica, mesic habitats and frequently is found around mongooses eventually were introduced to 29 human habitation. islands throughout the Caribbean and also to Impact of Herpestes javanicus on Native Vertebrates . Hays and Conant 11 the northern coast of South America (Hoag- repeatedly in the 1920s, and a population ex- land et al. 1989). An earlier New World isted there as late as 1936 but apparently died introduction of some unknown mongoose out (Tvrtkovic and Krystufek 1990). species was made at Trinidad in 1870 (Urich The small Indian mongoose was also [1914], cited in Hoagland et al. [1989]), but successfully introduced to Okinawa in 1910 this and all other original Caribbean mon- (Ogura et al. 1998) and to nearby Amami Is- goose introductions apparently failed, except land in 1949 and/or 1979 (Abe et al. 1991). the one in Jamaica (Espeut 1882, Nellis and A failed attempt was made in the late nine- Everard 1983). Attempts were also made, be- teenth century to introduce a mongoose into fore the introduction of mongooses, to intro- Australia to control rabbits, but it is unknown duce (Mustela putorius) to Jamaica, but whether this was the small Indian mongoose these attempts also failed, purportedly due to or another species (Hinton and Dunn 1967). chigoes (‘‘burrowing fleas’’ [Tunga penetrans]) (Hinton and Dunn 1967). physiology and behavior The range of the small Indian mongoose on the South American mainland is limited Mean body temperature of small Indian to the developed coastal strip of Guyana, Su- mongooses is 39.5C. The mean resting heart rinam, and French Guiana (Nellis and Ever- rate is 252 beats per minute, and the mean ard 1983), where it was introduced in 1900 resting respiratory rate is 63 breaths per min- (Hinton and Dunn 1967). The range of this ute (Nellis and Everard 1983). Metabolic en- mainland population appears to be limited, ergy use for this species, as deduced from for now, by a boundary of swamp and rain oxygen consumption of restrained individu- forest (Nellis and Everard 1983). About 100 als, has been variously measured at 116 kcal/ mongooses were also sent from Croatia to kg/day (Lin and Kobayashi 1976) and 76 Venezuela in 1926 (Tvrtkovic and Krystufek kcal/kg/day (Ebisu and Whittow 1976). The 1990) but apparently did not establish there. expected energy use for mammals in the size In 1883, sugar planters imported the small range of small Indian mongooses (300 to 600 Indian mongoose from Jamaica to four Ha- g) is 72 to 95 kcal/kg/day (Kleiber 1961). The waiian islands (Hawai‘i, O‘ahu, Maui, and thermal neutral zone is narrow and very Kaua‘i) and to the Fijian island of Viti Levu high: 28 to 38C (Ebisu and Whittow 1976). (Gorman 1975, Nellis and Everard 1983). Small Indian mongooses are, however, able to For unknown reasons, the crate of mon- maintain their body core temperature in the gooses was kicked off the dock at Kaua‘i, and range of 0 to 41C, though they show signs to date the mongoose has apparently not es- of extreme heat stress at 45C (Nellis and tablished there, although a dead mongoose McManus 1974). Heat-stressed mongooses was found in Kaua‘i in 1976 (Tomich 1986). pant and salivate on their chest and forelimbs Mongooses were later introduced to the Ha- but do not lick them (Ebisu and Whittow waiian island of Moloka‘i and to the Fijian is- 1976). They can respond to cold by raising land of Vanua Levu. their metabolic rate up to threefold (Nellis The small Indian mongoose exists on Java and McManus 1974). and Macau, and may have been introduced to The eye is emmetropic with at least 11 di- these islands hundreds of years ago. More re- opters of accommodation (Nellis et al. 1989), cently, it has been introduced to other islands and 25–40% of the photoreceptors are cones of and the , including (Hope et al. 1982), suggesting that this spe- Ambon, Mauritius, and Mafia (Nellis and cies has unusually good vision for a small Everard 1983). mammal and can discern colors. The kidney This species has been introduced to three can achieve a urine concentration of >6,000 Croatian islands of the Adriatic: Mljet (in milliosmoles, suggesting adaptation to desert 1910), Korcula (in 1910), and Hvar (in 1970 conditions (Nellis 1989). There is a large, [Tvrtkovic and Krystufek 1990]). It was in- appendix-like cecum (Palmiter and Horst troduced to the Peljesac Peninsula in Croatia 1982), but this appears to be lymphatic rather 12 PACIFIC SCIENCE . January 2007 than digestive in character. Most fat is stored Baldwin 1953), Fiji (Gorman 1976b), and the in the tail, which holds up to seven times the U.S. Virgin Islands (Nellis and Everard 1983), fat per mass of the body (Nellis and Everard all of which are between 17 and 20 degrees of 1983). latitude either north or south of the equator. This species is entirely diurnal (Kavanau Grenada, which is at 12 N, has a 10-month 1975). Typical movements are a low slink and breeding season (Nellis and Everard 1983). a gallop. Small Indian mongooses can swim Males produce sperm year-round in Hawai‘i and climb trees (Nellis and Everard 1983) but (Pearson and Baldwin 1953), but the size of rarely do so. When killing vertebrate prey, the testes changes seasonally: they are largest small Indian mongooses deliver a bite to the from February through July and smallest in back of the head, crushing the skull. Foraging October and November (Pearson and Bald- behavior has not been observed extensively or win 1953, Soares and Hoffman 1981). The described in this species, because it tends to size of the prostate gland follows the same forage under cover and is difficult to observe cycle, as do serum levels of sex hormones for long, continuous periods. They form shal- (androgens, follicle-stimulating hormone, and low burrows for use as dens. luteinizing hormone) in males (Soares and Both male and female small Indian mon- Hoffman 1981). Sperm are also produced gooses have a pair of pocket-shaped glands year-round in Fiji, where testes are largest lateral to an extensible pouch around the from September to December (equivalent to anus, rich in sebaceous and apocrine glands March through June in the Northern Hemi- (Gorman et al. 1974). These structures accu- sphere) (Gorman 1976b). mulate lipids and proteins that support gram- Females have a 4-day estrus at 20-day positive bacteria, which produce a mixture intervals during the breeding season (Asdell of six saturated carboxylic acids. Mongooses 1964) plus postpartum estrus (Nellis and use a ‘‘handstand’’ marking behavior to place Everard 1983). Ovulation is induced by copu- this substance on vertical surfaces, supporting lation (Asdell 1964). There is no delayed themselves on their forepaws while marking implantation (Tomich and Devick 1970), with the anal gland. Small Indian mongooses as there is in several other small are able to distinguish between the scent (Sadleir 1969). Gestation takes 49 days, and marks of individual conspecifics in choice pregnancy is not usually evident without tests (Gorman 1976a), though the smell of dissection (Nellis and Everard 1983). Mean these marks is undetectable by . litter size is unknown, but examination of 255 This species makes 12 distinct varieties of wild-caught pregnant females on Grenada vocalization (Mulligan and Nellis 1973), showed them to have a mean of 2.2 embryos, including, for example, a contact call used with a range of 1 to 5 (Nellis and Everard by young mongooses when following their 1983). The number of litters produced annu- mother, a feeding call, and several agonistic ally by a typical female in the wild has not yet cries that are graded by intensity of stress. been determined with any certainty. Young small Indian mongooses show strong Pups begin accompanying their mother on following responses, and follow their mother hunting trips at 6 weeks of age (about 200 g very closely until they are fully grown. Males body mass). Young mongooses follow their show some degree of sociality during the mothers until 4 to 6 months of age, at least breeding season (Hays and Conant 2003), in captivity, and this is also the range of ages but the social system has not been fully at which both genders attain sexual maturity elucidated. (Nellis and Everard 1983). The youngest wild-caught pregnant female with a confi- reproduction and population dently estimated age (determined by mass of dynamics the eye lens) was 4 months old (Nellis and Everard 1983). Pregnant females are found from February Population densities have not been deter- through August in Hawai‘i (Pearson and mined with certainty but may be extremely Impact of Herpestes javanicus on Native Vertebrates . Hays and Conant 13 high in some regions of introduction, judging the introduced mongoose is undoubtedly a by the small size of individual home ranges major impediment to the reestablishment of found in a few studies. Nellis and Everard many extirpated or reduced vertebrate popu- (1983) reported mean home ranges of 2.2 ha lations on dozens of islands. In most of these for two radio-tracked females and 3.6 ha for locations, this is a role that the small Indian five radio-tracked males on St. Croix. Hays mongoose shares with other introduced pred- and Conant (2003) found mean home ranges ators, as well as introduced competitors of 1.4 ha for seven radio-tracked females and habitat degradation caused by human on O‘ahu. They also found that five radio- activities. tracked males had broadly overlapping home Almost all efforts at mongoose population ranges in the breeding season, sharing a re- management in areas of introduction have gion of about 20 ha. All of these individual focused on the use of traps to remove individ- home ranges are very small, in comparison uals from ecologically sensitive areas. These with those of typical small carnivores. Carni- areas include particularly the nesting sites of vores in the mass range of the small Indian rare, ground-nesting birds. Trapping pro- mongoose (300 to 900 g) are expected to grams are of limited use, because removing have individual home ranges of 26 to 115 ha the mongooses from a strip of habitat around (Harestad and Bunnell 1979). No study has the sensitive area creates a region of empty been done of the home ranges of small In- habitat that lures more mongooses into the dian mongooses in their natural range, but it area. These programs are also very expensive, may be speculated that populations are ab- because monitoring traps is a labor-intensive normally dense in many parts of the intro- procedure. duced range. Diphacinone, an anticoagulant that kills by internal hemorrhaging, first became available management for experimental use in 1988. It has proved to be especially effective against mongooses, but Despite its wide range of introduction and its its use as a control agent is still primarily ex- formidable reputation as a predator, the small perimental (Keith et al. 1990, Smith 1998), Indian mongoose can only be confidently said and it is not currently being widely dispersed to have driven three vertebrates to extinction in areas inhabited by introduced mongooses. anywhere in the world: the barred-wing rail, the Jamaica petrel, and the Hispaniola racer prognosis (and in each of those cases, there is room for doubt). Considering the great number of is- The small Indian mongoose is strongly en- lands the mongoose has been introduced to, trenched in most of its areas of introduction. and the great number of ecologically delicate It appears to have high population densities species it preys upon on these islands, this on many large islands, is often found in habi- fact may at first seem remarkable. Many other tat that would have been unsuitable for it common anthropically introduced mammals, within its natural range, and in some places such as cats, dogs, and rats, have probably (e.g., Hawai‘i) has no natural predators and caused more than two extinctions apiece. But few communicable diseases and parasites. this fact in itself may be a large part of the ex- Only the most drastic measures, such as aerial planation of the relatively low number of ex- dispersion of diphacinone bait pellets, would tinctions directly attributable to mongoose have any chance of eliminating such popula- introductions: the mongoose was a latecomer tions from even small islands, and the ecolog- to its areas of introduction, and on most ical costs of such efforts might outweigh their islands simply added one further burden to benefits in many cases. Perhaps the single already-damaged ecosystems. By the time it most crucial management issue is to prevent arrived, much of the destruction was already the further spread of the species. Fortunately, done. new small Indian mongoose populations Nonetheless, the continued existence of are not likely to become established without 14 PACIFIC SCIENCE . January 2007 human agency, though this could occur the mongoose on St. John, US Virgin Is- unintentionally (e.g., by release of pet mon- lands. J. Mammal. 66:560–563. gooses). Collar, N. J., L. P. Gonzaga, N. Krabbe, A. Madrono Nieto, L. G. Naranjo, T. A. acknowledgments Parker III, and D. C. Wege. 1992. Threat- ened birds of the Americas. Smithsonian We thank an anonymous reviewer who sup- Institution Press, Washington, D.C. plied us with several useful references. Corbet, G. B., and J. E. Hill. 1992. The mammals of the Indomalayan region. Literature Cited Oxford University Press, Oxford, United Kingdom. Abe, S., Y. Takatsuki, Y. Handa, and H. Nigi. Corke, D. 1987. 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