Biology of the Mongoose (Herpestes javanicus) in a Rain Forest of Puerto Rico

Author(s): Francisco J. Viella

Source: Biotropica, Vol. 30, No. 1 (Mar., 1998), pp. 120-125
Published by: Association for Tropical Biology and Conservation
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BIOTROPICA 30(1): 120-125 1998

Biology of the Mongoose (Herpestes javanicus) in a

Rain Forest of Puerto Rico'
Francisco J. ViIeIIa2
U.S. Fish and Wildlife Service, Puerto Rican Parrot Field Office, P.O. Box 1000, Luquillo, Puerto Rico 00773

ABSTRACT
Ecological aspects of the mongoose (Herpestes javanicus) were studied in the Luquillo Mountains, a rain forest region
in eastern Puerto Rico. Information was obtained by removal trapping of mongoose from grids placed in tree plantations, colorado and tabonuco forests. Trapping efficiency was two mongoose per 100 trap-days, suggesting mongoose
abundance is low in wet montane forests of Puerto Rico. Sex ratio was biased (2.6:1) in favor of males. Body masses
of male mongoose inhabiting rain forests of the Luquillo Mountains were larger than those in dry forests at Guainica
(P < 0.0001). Stomach contents from 18 mongoose were examined, animal matter comprised 75 percent of the. total
food items encountered. Of these, 33 percent were from vertebrates. The food items most frequently encountered
were lizards (Anolis spp.), centipedes (Scolopendra spp.), and cockroaches (Blatellidae).

RESUMEN
Se investigaron varios aspectos sobre la ecologia de la mangosta (Herpestes javanicus) en la Sierra de Luquillo, una
region de bosque montano pluvial en el este de Puerto Rico. Se obtuvo informacion sobre la mangosta utilizando
trampeo de extraccion en un arreglo cuadriculado en plantacion de airboles, bosque de palo colorado y bosque de
tabonuco. La eficiencia de trampeo fue de dos mangostas por cada 100 dias-trampa, indicando que la abundacia de
mangostas es baja en los bosques montanos htumedos de Puerto Rico. La razon de sexo fue viciada (2.6:1) a favor de
los machos. La masa corporal de las mangostas macho en bosques htumedos de la Sierra de Luquillo fue mayor a la
de los capturados en el bosque seco de Guainica (P < 0.0001). Los contenidos estomacales de 18 mangostas fueron
examinados, el 75 por ciento compuesto de materia animal. De esta, 33 por ciento provenia de vertebrados. Los
articulos estomacales encontrados con mayor frecuencia fueron lagartos (Anolis spp.), ciempies (Scolopendra spp.), y
cucarachas (Blatellidae).

Key words: Anolis; Herpestes javanicus; mongoose; Moraceae; Puerto Rico; rain forest; Scolopendra.

(Wolcott 1953, Nellis & Everard 1983, Hoagland

THE SMALL INDIAN MONGOOSE (Herpestes javanicus)
et al. 1989, Vilella & Zwank 1993).

was introduced to the West Indies in 1872 in an

attempt to control rats on sugar cane plantations

In Puerto Rico, mongoose are known from ma-

(Espeut 1882). Today, the mongoose is found in a

ture dry forest, disturbed dry forest and scrub,

large number of islands and a wide variety of hab-

grasslands, cattle pastures, cane fields, and urban

itats (Hoagland et al. 1989). It has long been con-

areas (Pimentel 1955, Horst 1992, Vilella &

sidered responsible for the extirpation and extinc-

Zwank 1993). However, there is no information

tion of many terrestrial vertebrate species in the

for the mongoose on montane wet and rain forest

West Indies (Seaman & Randall 1962). However,

areas in Puerto Rico and very scant information for

recent studies suggest their effect may not equal

similar environments in the remaining West Indies

(Nellis & Everard 1983, Hoagland et al. 1989).

The objectives of this study were to obtain baseline
(i.e., feral cats) and habitat destruction (Henderson
information on the biology and food habits of the
1992). The information available on mongoose

that of other introduced mammalian predators

mongoose in rain forest regions of Puerto Rico.

food habits in the West Indies indicates they are

opportunistic feeders with a very generalized diet

STUDY SITE
1 Received 1 June 1995; revision accepted 7 October
1996.

The study was conducted in the Luquillo Experi-

2 Current address: Mississippi Cooperative Fish and

mental Forest (LEF; 18?10'N, 65?30'W) located

Wildlife Research Unit, Mail Stop 9691, Department of

Wildlife and Fisheries, Mississippi State University, Mississippi State, Mississippi 39762.

in the Luquillo Mountains of northeastern Puerto

Rico. The forest is wet with an annual rainfall of

120

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Biology of the Mongoose in Puerto Rico 121

2540 mm at lower elevations and 4320 mm at

venous injection of 0.2 ml deuthanasia-d in the

higher elevations. Trap grids (Fig. 1) were estab-

femoral vein. Carcasses were transported in an ice

lished within tree plantations, Tabonuco (Dacryodes

cooler and frozen for later examination. Sex,

excelsa) and Palo Colorado (Cyrilla racemiflora) life

lengths (total and tail), and body mass were re-

zones (Wadsworth 1951, Ewel & Whitmore 1973).

corded for each mongoose. Secondary sexual char-

Grid 1 was located at an elevation of 200 m

on timber stands adjacent to the U.S. Forest Ser-

acteristics, tooth wear, and pelage condition were

also recorded.

vice's El Yunque Ranger District. Vegetation com-

The entire digestive tract (esophagus to colon)

prised an overstory dominated by plantation spe-

was removed and contents suspended in a 5 per-

cies (i.e., Anthocephalus cadamba, Swietenia macro- cent buffered formalin solution. Contents were
phylla, Tectona grandis) with a dense undergrowth then washed with distilled water and run through
of vines, grasses, and shrubs. As a result of hurri2 sieves of various sizes. Contents were placed in a
cane Hugo (18 September 1989) debris and fallen

plastic petri dish and dried for 18 h at 35?C. Dried

logs were common throughout this grid (Brokaw

samples were dissected with a microscope and con-

& Walker 1991). Being on the edge of the reserve,

tents separated into categories for identification.

the grid was bordered by farmland, forest roads,

and the district offices.
Both grids 2 and 3 were located at higher ele-

Body masses for male mongoose captured at

LEF were compared with those trapped at Guianica

Forest (Vilella & Zwank 1993), a reserve of mature

vations (> 400 m) on the western parts of the LEF, coastal dry limestone forest located in southwestern
near the El Verde research area. In comparison to

Puerto Rico (18?00'N, 66052'W). The null hy-

grid 1, these grids were located in relatively interior

pothesis of no difference between the sample means
forest that had experienced little human impact.

(rain forest, dry forest) was tested using an unpai-

Grid 2 was dominated by Cyrilla racem#7ora, Mi-red t-test (Steel & Torrie 1980) with a test of ap-

cropholis chrysophylloides, and M garciniaefolia.

proximation for unequal variances. The Type I er-

Grid 3 was dominated by Dacryodes excelsa, Sloanea ror probability was set at 5 percent (t = 0.05).
berteriana, and Manilkara bidentata. For both grids
2 and 3, vegetative structure comprised advanced
secondary forest with little or no understory. Grids
were bordered by forest trails and by streams and

RESULTS
Mongoose were widely distributed in rain forests
of the Luquillo Mountains. Individuals were ob-

rivers.

served from the edge of the reserve (< 200 m) to

the elfin woodland regions located on the tallest

METHODS

peaks of the LEF (> 1000 m). Mongoose were

Removal trapping was conducted in the LEF from

commonly seen deep in the forest and away from

April to June 1993. Trap period 1 was from 20

any major roads or nature trails. They were fre-

April-13 May, and trap period 2 was from 18

quently encountered within nesting areas of the

May-10 June. Each grid consisted of 25 Havahart

live traps (model 1030, 50 X 17 X 17 cm) placed

Puerto Rican Parrot (Amazona vittata). However

in 5 rows at 50 m intervals (5 X 5 array), covering

ing parts of the year, by personnel working in re-

a 4 ha area (200 X 200 m). Traps were baited with

covery efforts for the parrot (FWS pers. comm.).

these are areas heavily visited, on a daily basis dur-

5 X 3 cm sponge squares immersed in a fish emul-

It is not uncommon for mongoose to move along

sion. All traps were opened shortly after sunrise,

trails and within areas heavily marked by human

checked at noon and checked again in the afternoon before closing them at dusk to prevent rats

scent (Coblentz & Coblentz 1985).

from entering. Mongoose are primarily diurnal

38 d (1400 trap-days) of trapping for a mean cap-

A total of 22 mongoose were trapped during

(Nellis & Everard 1983). Grid 1 was operated for

ture rate of two mongoose/100 trap-days (SD =
ten consecutive days during both trap periods, grids1.1). Four of the captured mongoose escaped dur2 and 3 were operated for nine d during both trap

ing handling, thus only 18 were available for ex-

periods. Trapping would terminate following no

amination. Relative densities of the sample by hab-

captures at any of the grids for three consecutive

itat type were 2.0 mongoose/ha for tree planta-

days.
Trapped mongoose were placed into a pillow

tions, 2.0 mongoose/ha for Palo Colorado Forest,

and 1.5 mongoose/ha for Tabonuco Forest.

case, immobilized with an intramuscular injection

of 0.2 ml ketamine, and euthanized by an intra-

favor of males. No females were captured in grid

Sex ratio of the sample was biased (2.6:1) in

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122 Vilella

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Biology of the Mongoose in Puerto Rico 123

TABLE 1. Food items identifiedfrom mongoose captured in the Luqillo Experimental Forest, Puert
Mongoose captured
Grid

Grid

Grid

Item 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16 17 18
Lizard
X
X
X
X
X
X
X
X
X
Bird
X
X
Tree
frog
X
X
Centipede
X
X
X
X
X
X
Crab
X
X
X
Shrimp
X
X
X
Coleoptera
X
X
X
X
X
Dictyoptera
X
X
X
X
X
X
Orthoptera
X
X
Spider
X
Moraceae
X
X
X
X
X
Flacourtiaceae
X
X
Melastomataceae
X
X
X
X
Unknown
plants
X
X
X

3. Trapped mongoose had an average total length

spp.), centipedes (Scolopendra spp.), and cockroach-

of 588 mm (SD = 51 mm, range 429-655 mm),

es (Blattelidae). Fruits and seeds (e.g., Miconia spp.)

an average tail length of 251 mm (SD = 33 mm,

made up the majority of the plant material.

range 150-285 mm), and an average body mass of

Body masses of male mongoose at LEF (Fig. 2)

739 g (SD = 184 g, range 397-1026 g). Alto-

were larger than those from the dry forests of

gether, males were larger than females. Average

Guanica (t = -5.72, df = 28, P < 0.0001). At

male total length was 612 mm (SD = 23 mm,

LEF body mass of males averaged 834 g (N = 13),

range 580-655 mm), average tail length was 264

whereas at Guainica forest males (N = 16) averaged

mm (SD = 10 mm, range 243-285 mm) and an

587 g (SD = 124 g, range 365-750 g). See Vilella

average weight of 834 g (SD = 108 g, range 624-

and Zwank (1993) for details on body character-

1026 g). Average female total length was 514 mm

istics of mongoose at Guainica forest.

(SD = 62 mm, range 429-580 mm), average tail

length was 217 mm (SD = 51 mm, range 150270 mm) and an average weight of 493 g (SD =
73.9 g, range 397-595 g).

DISCUSSION
Mongoose were widely distributed in the Luquillo

Five (28%) of the 18 mongoose examined had

Mountains, however, relative densities were very

noticeable teeth wear or canines and incisors either

low in all habitat types sampled during the study.

missing or broken. Pelage condition was predomi-

This may reflect a low abundance of mongoose in

nantly clean and well groomed. Four of the 13

montane humid and rain forest regions of Puerto

Rico. Studies in other islands of the West Indies

males caught were identified to be in breeding condition (wt > 600 g, scrotum enlarged and descend-

have reported lower densities of mongoose in high-

ed). Average scrotal male length was 620 mm (SD

er elevation wet forest sites as compared to drier

= 32 mm, range 570-655 mm), average tail length

areas near the coast. On Grenada, Nellis and Ev-

was 268 mm (SD = 10 mm, range 243-270 mm)

erard (1983) reported a density of 4.0 mongoose/

and an average weight of 853 g (SD = 146 g, range

ha in tree plantations of blue mahoe (Hibiscus ela-

624-1026 g). No captured females were in breed-

tus) at Les Avocats, and 4.7 mongoose/ha at Grand

ing condition.

Etang, an area of montane rain forest with over

Stomach contents were examined in 18 of the

4000 mm of rain per year. However, at Mt. Hart-

22 mongoose captured (Table 1). Of the 14 food

man, an area of dry scrub and woodland, mon-

items found, four (29%) were plant material and

goose densities were 10.4 mongoose/ha.

10 (71%) animal. Of the animal matter, 33 percent

Conversely, Hoagland et al. (1989) found den-

was made up by vertebrates and the remaining 67

sities of 14 mongoose/ha in moist forest of north-

percent was invertebrates. The animals encountered

western St. Croix. However, it should be men-

most frequently in the diet were lizards (Anolis

tioned that while this region is referred to by locals

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124 Vilella

Body mass (g
300

400

Rain

1 Dry Forest
Forest

401 - 500- /// // ///

501 - 600

601

700-____

701 - 8000

801 - 900901

1000-

1001 - 1500

Number of mongoose

FIGURE 2. Distribution of body mass for mongoose trapped in dry forest of Guainica (N = 16) and rain forest of
Luquillo (N 13).

as the "rain forest" portion of the island, annual

help to elucidate whether this difference in rain

precipitation (1300 mm), elevation and mean an-

forest mongoose populations is a result of local en-

nual temperature, classify this area as moist forest vironmental and climatic conditions.
The absence of reproductively active females
(Ewel & Whitmore 1973). There are no bioclimatic regions of wet or rain forest on St. Croix.

and skewed sex ratios contrasts those found in oth-

er studies conducted in Puerto Rico (Pimentel

those reported by other investigators for higher el- 1955, Vilella & Zwank 1993). However, these
could be an artifact of small sample size and of the
evation forested regions of the West Indies.
removal trapping method. Removal trapping usuFrequent periods of precipitation at LEF could
Notwithstanding, these densities are greater than

also have contributed to the low capture rates ob-

ally results in more males being captured initially.

served during the study. Starting on 29 April dur-

However, continuous removal results in a more

ing trap period 1, and ending on 10 June when all

equal distribution of the sexes (Hoagland et al.

trapping was terminated, there were only 8 days

1989).

with clear and sunny weather. The remaining were

Stomach contents of mongoose were similar to

cloudy days with either light constant rain through-those reported by Wolcott (1953). He found large
numbers of centipedes (Scolopendra spp.), lizards
out the day, and/or sporadic heavy downpours.
(Anolis spp.), crabs, and various species of insects
Only two mongoose were captured during all of
trap period 2, these being males caught in grids 2

in 98 mongoose stomachs from Puerto Rico and

and 3. Grid 1 had no captures during the second

St. Croix. At the LEF, lizards made up the majority

period of trapping. Heavy rains during trapping

of the vertebrate sample (Table 1). The most com-

could have affected captures, as mongoose activity monly encountered lizards on the understory vegdecreases during and following rainstorms (Pimen- etation were Anolis cristatellus and Anolispulchellus.
tel 1955). In order to optimize trapping efficency

Two samples from grid 2 included feather parts.

when sampling mongoose populations in montane

Feathers were dark brown in color and bone frag-

ments were soft, possibly from chicks or juvenile

rain forest environments, it may be better to schedule trapping periods during the drier months of theRuddy Quail-Doves, Geotrygon montana. Bones
identified as treefrogs (Eleutherodactylus spp.) were
year (November-February).
Body masses and body lengths were within lim-

present in 2 samples.

Indies (Nellis & Everard 1983, Hoagland et al.

Centipedes were common in stomach samples

from grids 1 and 3 and absent from samples from

1989). However, male mongoose at LEF were larg-

grid 2. Additionally, not only were centipedes fre-

er than those captured along trap transects at

Guinica Forest (Fig. 2). While this observation is

quently encountered in stomach samples, but these

were usually of very large specimens. Grid 1 was

based on small sample sizes, further research may

located in an area heavily affected by Hurricane

its previously reported for this species in the West

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Biology of the Mongoose in Puerto Rico 125

Hugo in 1989. It had a more structurally diverse

Plant material is known to be an important

vegetative understory and abundant debris that

source of nutrients for mongoose (Gorman 1975).

could possibly enhance centipede numbers.

The majority of the vegetative samples encountered

Three samples from grid 2 included freshwater were from the families Melastomataceae and Moshrimp (Atya spp.) parts as well as body parts of
raceae. Fruits and seeds from the melastome genus

the freshwater mountain crab Epilobocera sinuatif-Miconia were found in 4 samples. Additionally, five
rons (Table 1). Land crabs (Cardisoma guanhumi)
stomach samples had fig fruits and seeds possibly
have been reported in stomachs of mongoose collected from sugar cane fields (Wolcott 1953, Pi-

of the endemic Ficus sintenisii, which is commonly

found in the LEE Slightly over half of all the sam-

mentel 1955). During trap period 1, five of the

ples contained some kind of plant material (Table

nine days of trapping had constant rain. As a result
1). This may reflect a greater variety and abunmany creeks overflowed causing fresh water

dance of plant foods available for mongoose in rain

shrimps and crabs to be more mobile during day-

forest environments.

light and perhaps, susceptible to predation by mongoose. Another explanation could be that mongoose on grid 2 were feeding on the piles left by

ACKNOWLEDGMENTS

foraging night-herons (Nycticorax spp.), which wereThanks are due to Sonia Najera who conducted fieldwork
frequently seen near streams and rivers at the LEE

Mongoose are known to feed readily on available

carrion (Keith et al. 1985). Cockroaches were commonly encountered in samples from grids 2 and 3.

Grid 3 had a large area of rotten logs, and grid 2

had abundant leaf litter and fallen trees. These substrates provide good cockroach habitat.

as part of the Fish and Wildlife Service's Cooperative Student Program. Wilfredo Abreu and Gilberto Ortiz of the
Puerto Rican Parrot Field Office provided valuable field
assistance laying trap grids and monitoring captures. I am
grateful to Gerardo Camilo of the University of Puerto
Rico Terrestrial Ecology Division for assistance with identification of stomach samples. Special thanks to Roy
Horst and C. William Kilpatrick for excellent reviews
which greatly improved the manuscript.

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