B I O L O G I CA L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8

available at www.sciencedirect.com

journal homepage: www.elsevier.com/locate/biocon

Habitat use and spatial behaviour of the endangered

Southern river otter (Lontra provocax) in riparian habitats
of Chile: Conservation implications

Maximiliano A. Sepúlvedaa,e,*, José Luis Bartheldb, René Monsalvec,

Vicente Gómezd, Gonzalo Medina-Vogelc
a
Comité Nacional Pro Defensa de la Fauna y Flora, Programa Biodiversidad, Carlos Anwandter 842, Valdivia, Chile
b
Ines Gallardo 2129, Puerto Montt, Chile
c
Escuela de Medicina Veterinaria, Universidad Andrés Bello, República 252, Santiago, Chile
d
Instituto de Ecologı́a y Evolución, Facultad de Ciencias, Universidad Austral de Chile, Valdivia, Chile
e
College of Veterinary Medicine, North Carolina State University, 4700 Hillsborough St., Raleigh, NC 27606, USA

A R T I C L E I N F O A B S T R A C T

Article history: The spatial behaviour of 10 Southern river otters Lontra provocax was analysed using radio-
Received 29 September 2006 tracking data collected in the Queule River, Southern Chile. The estimated otter density was
Received in revised form 0.25 individuals km1. Females with cubs and adult otters displaying mating behaviours
21 August 2007 were the only social group recorded. Studied otters were solitary and observed behaviour
Accepted 27 August 2007 patterns suggest an intrasexual territoriality. No home range overlap among males existed
Available online 15 October 2007 while an average of 33% and 87% home range overlapped among females and between
sexes, respectively. In the situation of core areas, only one case of overlapping between
Keywords: an adult male and an adult female was observed. Average home range and core area
Territoriality lengths were 11.3 km and 0.9 km, respectively. Otters used rivers more frequently than
Home range expected, whereas they avoided small streams. Low intensity livestock farming, without
Ranging behaviour canalization of watercourses and riparian vegetation extraction practices, seems compat-
Radiotracking ible with the presence of Southern river otter populations since the spatial behaviour of
Lontra provocax the studied otters was not affected under these circumstances. Based on the present offi-
cial reserve sizes and distribution within the country, and also the increase of variant
anthropogenic land uses outside these areas that result with consequent habitat loss
and deterioration, we strongly recommend considering the importance of spatial dimen-
sion in quantity and quality habitats in further conservation planning for the Southern
river otter using an integrated catchment management.
 2007 Elsevier Ltd. All rights reserved.

1. Introduction servation of Nature and Natural Resources) (IUCN, 2006),

and recently classified for Chile as ‘‘endangered’’ between VI
The Southern river otter or huillı́n (Lontra provocax) is listed as and X Districts and ‘‘insufficiently known’’ for XI and XII Dis-
‘‘endangered’’ by the IUCN (International Union for the Con- tricts (República de Chile, 2007). It is confined mainly to the

* Corresponding author: Present address: College of Veterinary Medicine, North Carolina State University, 4700 Hillsborough St., Raleigh,
NC 27606, USA.
E-mail address: masepulv@ncsu.edu (M.A. Sepúlveda).
0006-3207/$ - see front matter  2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2007.08.026
330 B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8

Andean-Patagonian region of Southern Chile as well as some
lakes and fjords of Southern Argentina (Medina, 1996; Lari-
vière, 1999; Aued et al., 2003). Historically, the distribution of
the species was more extensive, including rivers, streams
and fjords from 34S to the Magellan Region (53S) in Chile.
Presently, populations in freshwater habitats are confirmed
in only a few river basins from 39S to 44S (Medina, 1996;
Aued et al., 2003). High concentrations of riparian vegetation,
woody debris, tree roots, gravel and natural riverbed sedi-
mentation, along with the availability of freshwater crusta-
ceans appear as decisive factors of the Southern river otter’s
distribution (Aued et al., 2003; Medina-Vogel et al., 2003). In
fact, land drainage of floodplains and canalization reduce
the carrying capacity of their habitat (Boon, 1992; Crook and
Robertson, 1999). Indeed, significantly less Southern river ot-
ter activity has been recorded in river sections where riparian
vegetation was removed, or the watercourse was canalized
(Medina-Vogel et al., 2003; Hauenstein et al., 2005). The Irriga-
tion Law passed in Chile in 1986 promotes investment in land
drainage for livestock farming and agriculture and conse-
quently encourages the removal of riparian vegetation and
the canalization of lowland rivers, particularly in those that
flow through swamp forest (Medina-Vogel et al., 2003). Fur-
thermore, in recent years increased demand of energy
sources by the growing Chilean economy promotes the con-
struction of large dams on several rivers where Southern river
otter populations exist. Despite the imminent threats and
critical conservation needs, knowledge about spatial behav-
iours, habitat requirements and relevance of anthropogenic
land uses for this rare species is scarce. These factors are crit-
ical in helping future management activities for this species
and it’s habitat, and is evidenced by studies of other carni-
vores (Sunquist and Sunquist, 2001). Therefore, the main
objective of this study was providing information to support
conservation efforts in the Southern river otter. The specific
questions were (1) what are the length of Southern river otter
home ranges and core areas? (2) are Southern river otters ter-
ritorial? (3) how is otter spatial behaviour affected by anthro-
pogenic pressure? (4) how could otter conservation be made
compatible with current policies or growth in Chilean
economy?
2. Methods
2.1. Study area
The study was conducted from February 2000 to December
2002 within the limits of the present northern distribution
of the species in Chile located in the upper section of the Que-
ule River within the Coastal Mountain Range (39S, 73W) and
in the Southern border of the IX Political Region of Chile
(Fig. 1). The topography consists of low mountains of up to
600 m above sea level and river valleys and open flood plains
25 m above sea level. Weather is mesotemplate humid, with
yearly average precipitations of 2139 mm and temperatures
of 12.5 C (Luebert and Pliscoff, 2005), with minimum levels

QUEULE WATERSHED
73° 10’ 73° 00’ 72° 50’

STUDY AREA

39° 10’

CHILE

39° 20’
N

0 5 10

Kilometers

Fig. 1 – Queule watershed area with square delimiting the study area in the upper part of the river.
 B I O L O G I C A L C O N S E RVAT I O N
of rainfall recorded in summer (January–March) and maxi-
mum levels during winter (June–August) (Campos, 1985).
The floodplains and areas next to streams have been used
mainly for exotic plantations of pine (Pinus radiata) and euca-
lyptus (Eucalyptus globulus), native forest, and cattle and goats
ranges with pastures (animal density approximately 10 ani-
mal km2). Percentage distributions in the study area were
33% for plantations, 53% for native forest and 14% for live-
stock farming areas. The native forest community consists
of patches of evergreen forest with Olivillo (Aextoxicon puncta-
tum), Ulmo (Eucryphia cordifolia), Guayún (Rhaphithamnus spino-
sus), Mañio de hojas largas (Podocarpus salignus), Tineo
(Weinmannia trichosperma) and deciduous forest of Roble (Not-
hofagus obliqua), Raulı́ (Nothofagus alpina) and Coigüe (Nothofa-
gus dombeyi). The riparian vegetation community consists of
the Temu Pitra swamp forest mainly composed by native spe-
cies such as Myrceugenia exsucca, Blepharocalyx cruckshanksii,
Luma gayana, Drimys winteri, Tepualia stipularis, Blechnum chi-
lense, Chusquea quila, Boquila trifoliata and Cissus striata (Hau-
enstein et al., 2005). Also, associated to river shores are
lined by a dense barrier of exotic Rubus constrictus. Based upon
the stream order system of Strahler (1952), the river basin in
the study area was subdivided into streams (first and second
order streams, 500 km) and rivers (third order stream, 230 km)
with ca. 275 km2 of total surface. The maximum flow of these
watercourses approaches 14 times the minimum flow. There-
fore, during times of elevated water levels, flooding of adja-
cent plains and riparian zones occurs. No river canalization
or drainage exists in these areas. Human presence is low with
scattered farm housing. The closest village is 10 km down riv-
er from the study location. This area is part of the Valdivian
Ecoregion, one of the most important in terms of biodiversity
and one of the most threatened in the world (Olson and
Dinerstein, 2000).
2.2. Capture of otters and data collection
Otters were live-trapped at latrines, which were previously
determined by fresh spraint and footprint detection. Trapping
was conducted each year from January to April using Victor
leg-hold traps (1.5 Soft catch, Woodstream Corp., Lititz, Penn-
sylvania 17543, USA).
For capture and insertion of radiotransmitters, otters were
anaesthetized using a combination of Ketamine (Imalgene,
10 mg/kg; Merial Aventis Pasteur S.A., Hernando de Aguirre
268, Piso 3. Providencia, Santiago, Chile) and Xylazine (Rom-
pun, 0.5 mg/kg; Bayer de Chile, Carlos Fernández 260, San
Joaquı́n, Santiago, Chile). Radio transmitters [40 g, 12 months
expected duration, 60 · 20 mm diameter cylinder, 150.00–
151.00 MHz (Sirtrack Ltd., Havelock North, New Zealand)]
were implanted by a veterinary surgeon within the abdominal
cavity in accordance with the same procedures described by
Hoover (1984). Pulse, respiration, and rectal temperature of
immobilized otters were carefully monitored throughout cap-
ture and immobilization. During transport and release proce-
dures the otters were placed in a well-ventilated transport
tube constructed from a 1 m section of 40 cm diameter PVC
pipe (Serfass et al., 1996).
Implanted otters were released on site, and all locations
within the first 5 days post release were excluded from study
1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8 331
to avoid potential confounding by the captivity on the otters’
spatial movements. Thereafter, otters were repeatedly lo-
cated, first by triangulation and then by homing from two mo-
bile stations. Each mobile station consisted of two observers
carrying one three-element yagi antenna, one receiver
(150.00–154.00 MHz., TR4, Telonics Inc., AZ., Mesa, AZ, USA),
a handheld compass and one Garmin GPS unit.
The range of the signal was up to 1600 m. Radiolocations
were estimated for each individual every month with a proto-
col between fixes of 12 h during 12 days. Each subsequent day
of study was started 1 h later in order to obtain the full daily
activity cycle. In the case of triangulation, radiolocation of an
animal was assessed by plotting the locations given by the
software LOAS (Ecological Software Solutions, www.eco-
stats.com) onto a UTM geo-oriented air photograph
(1:115.000). Radiolocation was given by GPS for homing.
The home range of an individual was defined as the total
length of watercourse between the furthest locations which
an otter used more than once during the study period (Mel-
quist and Hornocker, 1983; Bowyer et al., 1995). A range
asymptote is needed before one can be confident that the
range area is adequately described (Harris et al., 1990). By
plotting the number of fixes against cumulative range area,
we determined the otters reaching a home range asymptote.
Home range length was determined only for those otters with
adequate radiolocations corresponding to an asymptote.
In order to describe the core areas, the river drainage system
was divided into 350 m sections. Core areas were defined as
sections occupied by animals for at least 10% of the time during
the radio-tracking period (Melquist and Hornocker, 1983).
2.3. River size, adjacent area use and selection
Evaluating the impact of the various anthropogenic uses of
the land adjacent to the rivers inhabited by otters required
the following land use classifications: (1) open farm fields
(livestock farming), (2) pine forest plantations (plantations),
and (3) marshes, and native forest which include swamp for-
est (native). Habitat selection analysis between principal riv-
ers and streams was investigated to assess the importance
of river size. In avoiding biased interpretations, we analyzed
selection as described by Johnson (1980), evaluating second
order selection of habitat composition within home ranges
of individuals and third order selection of habitats in individ-
ual home ranges. During the study, radiotracked otters used
only riparian habitats. Based on this observation, we limited
the borders of the study area as the boundaries of the wa-
tershed (defined as the geometric space where all the points
drain to the same current and where the limits with other
continuous watersheds are marked by imaginary lines).
The distribution, limits and availability of these habitat
categories were confirmed by direct observation, UTM geo-
oriented air photographs (1:115,000) and Global Positioning
System (GPS), and then calculated using a Geographical Infor-
mation System (ArcView version 3.3, ESRI, 1996).
2.4. Data analysis
Data were separated by otter sex to compare home ranges and
core areas between the sexes. To test for the null hypothesis
332 B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8

X X
of no responses by the otter’s spatial behaviour to the catego- Ip ¼ Pki  Pkj
ries of anthropogenic use of adjacent lands and river size, we
estimated river size and adjacent area use and selection where Ip is the Simple Probability Index, and Pki and Pkj are the
according to methods used by Manfredi et al. (2006). We calcu- independent probabilities that at any arbitrary time otter i
lated the Levins’ index (B) (Levins, 1968) as and j are in the shared area (Powell, 2000).
X 1 All means are presented with ±1 standard deviation.
B¼ p2i

where pi is the fraction of a given habitat within the otters’ 3. Results

habitat. For this study, the index values ranged between 1
and 2 for river size and 1–3 for the adjacent area. Lower values
of this index indicate minimum habitat use diversity when
only one habitat use is occupied while higher values indicate
maximum diversity when all habitats are uniformly inhab-
ited. We measured selection/avoidance for each single habitat
using Ivlev’s (1961) Preference Index (E) as follows:
E ¼ ðUj  AjÞ=ðUj þ AjÞ
where Uj is the use of river size or adjacent area j and Aj is its
availability. The Preference Index values range from 1 (com-
plete avoidance) to +1 (complete selection). We also calcu-
lated the Selectivity Index (S) for each radiotagged otter,
P one cub in February 2001, and with two cubs in October 2002.
where S ¼ jUj  Ajj (Duncan, 1983; Lucherini et al., 1995). S
represents a measure of the overall degree of selection for
all river sizes and all adjacent areas combined, and range
from 0 (lack of selectivity) to 200 (maximum selectivity). Test-
ing the hypothesis of no territoriality among the study otters,
we calculated the spatial overlap of one animal on another as
the proportion of the latter’s home range overlapping the for-
mer’s home range (Macdonald et al., 1980). Data were sepa-
rated into three groups: females–females, males–males and
females–males and then the home range and core areas over-
lap were compared between the groups. Also, we performed a
static interaction analysis in which the sets of locations were
compared without reference to the temporal sequence of
locations, as we did not perform simultaneous observations
of animal locations (Powell, 2000; Kernohan et al., 2001; Fie-
berg and Kochani, 2005). Thus the probability that otters i
and j were in the overlapped area was computed as:
During this study, 12 Southern river otters were trapped,
radioimplanted, released and radiotracked. One male and
one female were recaptured, resulting in a total of 10 different
otters (Table 1). Otters were classified by sex: male (M), female
(F) and age: juvenile (J) or adults (A) based on body size, tooth
wear and staining.
During the study period, mating behaviours were observed
in untagged otters in October 2000, November 2001, and FJ3
was observed copulating with an untagged otter in December
2001. Family groups of otters composed of an adult otter and
two cubs were recorded in October 1999, November 2000, with
Radioimplanted individuals were never detected simulta-
neously at any location.
Otter density in our study site was 0.25 ± 0.04 otters km1
of river length based on the combined length of the home
ranges and the number of live-trapped and recaptured otters
(Table 2).
3.1. Home range and core areas length
Otters were radiotracked for a period ranging from 71 to 204
days, resulting in a total of 646 radiolocations with an average
of 53.8 ± 45.2 radiolocations otter1 (Table 1). The mean home
range length of the otters was 11.3 km ± 4.8 of river length
with two to four core areas per home range (range = 0.7–
1.4 km long) (Table 1). There was no difference in home range
size between sexes (Table 1).

Table 1 – Characteristics of body weight, body length, days of observations, home range, core areas, core areas proportion
and number of core areas by 10 radiotracked Southern river otters at Queule river, Southern Chile, between February 2000
and December 2002
Year Otter Body Total body length Length Number Home range Core area Core area Number of
weight (kg) including tail (cm) in days of fixes length (km) length (km) proportion (%) core areas

2000 FJ1a 5.5 104.0 189 17 11.3 1.0 9.3 3

MJ1a 6.2 115.0 204 18 – – – –
MA1a 12.6 117.0 178 15 9.6 0.7 7.3 2
2001 FJ3 4.8 97.5 159 41 12.0 0.7 5.8 2
MA2 10.0 119.0 199 147 22.3 1.4 6.3 4
MA4a 11.6 119.0 103 12 – – – –
FA1 10.0 112.0 97 105 7.4 0.7 9.5 2
FJ2 6.9 106.0 106 117 9.0 0.7 7.8 2
2002 FA3a 8.0 109.0 98 47 – – – –
MA5a 14.0 121.0 61 21 – – – –
MA3 12.0 118.0 71 51 8.5 1.4 16.5 4
FA2 8.0 107.0 100 55 10.1 1.4 6.9 2
M: males; F: females; A: adults; J: Juveniles.
Average body weights and lengths only for adults otters.
a
Otters with not enough radiolocations to determine either home range length, core areas or both.
 B I O L O G I C A L C O N S E RVAT I O N
Table 2 – Number of km of river length with presence of
otter studied, number of otter (radiotracked and direct
observations) in the study area and estimated density
Year Total river length Number of otters Otters
confirmed with observed in the (km1)
studied otters (km) study area
2000 20 4 0.20
2001 30 8 0.27
2002 10.8 3 0.28
Throughout the study, 14 core areas were identified (Fig. 3),
five of which were accessible. These core areas were contigu-
ous with the rivershore and were centered around otter dens
(holes dug into the river banks close to the water) and con-
tained dense patches of bamboo (C. quila) (50 m length · 10 m
width minimum surface) along with mature and fallen trees.
Of the 10 radiotracked otters, only three (FJ1, FJ2, and FJ3) had
core areas at one end of their home range (Fig. 3). Otter MA1
(2000 year) was recaptured and named MA2 (2001 year), main-
taining one of its core areas (Fig. 3). During 2001, one female
with a cub was observed around this core area. During 2002
this den was inhabited by FA2 and MA3. Coincidently, in that
year, MA2 was not trapped. FA2 and MA3 never used this den
simultaneously. All of the Southern river otters positioned
their core areas less than 350 m from a convergence of water-
Fig. 2 – Occupied areas in squares by eight Southern river otter individuals indicated by each letter in the map during years
2000–2002. The solid black circles in the map key illustrate the presence of otters MA4 in 2001 and MA5 in 2002.
1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8 333
courses, except for FJ2 which did not have this type of water-
course characteristic within the limits of its home range
(Fig. 3). Otter MJ1 was recorded at several locations, showing
a dispersion pattern. The otter soon left the study area after
its release and traveled 46 km downstream. Otters MA4 and
MA5 died 2 months after their release (Fig. 2), but the cause
of death for these otters was not apparent on necropsy be-
cause of the advanced state of carcass decomposition. FA3
moved 10 km downriver and settled in an area associated
with the village ‘Los Boldos’ making monitoring very difficult
because of electrical interference with the radio transmitter.
Only otters FJ3, MA2, FA1, FJ2, MA3, and FA2 were recorded
at enough radiolocations to create an asymptote curve at
42–96 locations (Fig. 3). No otter was located further than
10 m from the waterways.
We did not trap and monitor males simultaneously there-
fore there was no chance for home range overlap between
males. The home range overlap between females was
32.9 ± 22.9% and between sexes was 86.6 ± 8.5%. The Simple
Probability Index between the sexes was 0.65 ± 0.20 (n = 2),
and 0.08 ± 0.12 (n = 3) between females. Only one male and
one female adult otter had core areas overlap during year
2002, with 21.4% and 28.5% overlap, respectively (Table 3). In
2000, there was no core area overlap between animals MA1
and FJ1, but their core areas were contiguous (Fig. 3). Female
otters had no overlap of core areas, although otters FJ3 and
FA1 had contiguous core areas (Fig. 3, Table 3).
334 B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8

Fig. 3 – Home ranges as black lines and core areas as hollow circles for the following years and individuals: in 2000, MA1 and
FJ1; in 2001, MA2, FJ2, FJ3, and FA1; in 2002, MA3 and FA2.

Table 3 – Home range and core area overlaps among 10 radiotracked Southern river otters in the Queule river, Southern
Chile, between February 2000 and December 2002
Year Otters Home ranges Core areas
A–B Overlap (km) Overlap A (%) Overlap B (%) Overlap (km) Overlap A (%) Overlap B (%)

2000 MA1–FJ1 9.1 94.8 80.5 0.0 0.0 0.0

2001 FJ3–FA1 5.4 45.0 73.0 0.0 0.0 0.0
FJ3–FJ2 2.7 22.5 30.0 0.0 0.0 0.0
FJ2–FA1 1.1 12.2 14.9 0.0 0.0 0.0
MA2–FA1 2.7 12.1 36.5 0.0 0.0 0.0
MA2–FJ3 2.9 13.0 24.2 0.0 0.0 0.0
2002 MA3–FA2 7.9 92.9 78.2 0.3 21.4 28.5
Overlap is expressed as a proportion of the otter’s home range and core area length.

3.2. Selection of river size and adjacent area stream), otters most frequently used rivers (Figs. 3 and 4).
Almost all animals used rivers rather than streams with a
Only otters for which we could estimate home range size (n = 6) low Levins index (B), except for otters FA1 and FJ3 (Table 4).
were analyzed. In terms of size of the body of water (river/ With respect to adjacent areas, each otter used only one
 B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8 335

Stream
River
100

80
% of fixes

60

40

20

0
MA2 MA3 FJ3 FJ2 FA1 FA2

Fig. 4 – Habitat use (percentage of locations) of six Southern river otters in the Queule river, Southern Chile, between February
2000 and December 2002. M: male, F: female, A: adults, and J: Juveniles.

Table 4 – River size and adjacent area values of Levins index (B) and Selectivity index within the home range (S1) and study
area (S2) for each radiotagged Southern river otter in the Queule River, Southern Chile, between February 2000 and
December 2002
MA2 MA3 FJ3 FJ2 FA1 FA2

River size B 1 1 1.7 1 1.6 1.1

S1 8 2 8 0 52 10
S2 166 170 108 170 122 158

Adjacent area B 1.2 1.1 1.2 1 2 1.1

S1 46 16 20 10 44 34
S2 152 158 78 94 76 160

adjacent area (livestock or native), except for otter FA1 that
used both areas similarly (Fig. 5). Plantations were almost un-
used, with generally low diversity use of adjacent areas (Table
4). The Selectivity Index (S) indicated that there was a higher
selection for second order scale rather than third order for both
analyses, i.e. river size and adjacent area (Table 4). The analysis
of second and third order Preference Index (E) was similar for
all otters with a selection of rivers and avoidance of streams;
most pronounced at the second order scale (Table 5). Planta-
tions were avoided by all otters, whereas livestock farming
was selected by otters, with higher preference values (E) for
second order scale. Native adjacent area did not present a clear
selection pattern between different otters (Table 5).
4. Discussion
In this study, percent home range overlap and the Simple
Probability Index were higher between sexes rather than be-
tween females. The only overlap in core areas occurred be-
tween sexes. These results suggest an intrasexual
territoriality known in the subfamily Mustelinae and some
species of the subfamily Lutrinae (Powell, 1979; Dominic
et al., 2000). The fact that we could not trap more than one
male in any given study site during the same time period also
provides evidence of intrasexual territoriality.
The number of core areas per individual otter observed in
this study is similar to that of the comparably sized North
American river otter Lontra canadensis (Melquist and Hornoc-
ker, 1983). All accessible core areas corresponded with dens
where individuals spent long periods resting as observed in
the Cape clawless otter Aonyx capensis (Arden-Clarke, 1986),
which has a feeding strategy similar to the Southern river ot-
ter, preying mainly on freshwater crustaceans (Medina, 1998).
In this study, the presence of dens or resting sites appears to
be an important factor in the location of core areas. Since we
did not evaluate trophic offer at segment rivers, we were un-
able to evaluate its importance in location of core areas. Thus,
the core areas location seems to be associated at least with
presence of suitable dens or resting sites. Throughout this
study, the specific core areas were used repeatedly by either
the same individuals or by different otters moving into the
area. This pattern has been described for the Eurasian otter
(Lutra lutra) (Kruuk, 1995; Ruiz-Olmo et al., 2005) and may oc-
cur because suitable den location is a limiting factor (Beja,
1996; Kruuk, 1995; Yoxon, 2000). Therefore, the protection of
existing dens appears to be a crucial component for effective
336 B I O L O G I C A L C O N S E RVAT I O N 1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8

Plantation
Native
Livestock farming

100

80

60
% of fixes

40

20

0
MA2 MA3 FJ3 FJ2 FA1 FA2

Fig. 5 – Adjacent area use (percentage of locations) of six Southern river otters in the Queule river, Southern Chile, between
February 2000 and December 2002. M: male, F: female, A: adults, and J: juveniles.

Table 5 – River size and adjacent area composition (%) of the study area, river size and adjacent area composition (%) of the
home range (HR) and value of Preference Index within the home range (E1) and study area (E2) for six Southern river otters
in the Queule river, Southern Chile, between February 2000 and December 2002
Otter Study area (% of area) River size Adjacent area
River Stream Livestock farming Native Plantation

15 85 14 53 33
MA2 HR (%) 98 2 67 20 13
E1 0.02 0.5 0.15 0.54 0.53
E2 0.73 0.95 0.73 0.8 0.78
MA3 HR (%) 99 1 85 3 12
E1 0.01 1 0.04 1 0.26
E2 0.74 1 0.74 1 0.65
FJ3 HR (%) 69 31 18 82 0
E1 0.03 0.06 0.38 0.06 –
E2 0.64 0.47 0.27 0.27 1
FJ2 HR (%) 100 0 5 95 0
E1 0 – 1 0.03 –
E2 0.74 1 1 0.31 1
FA1 HR (%) 24 76 30 59 11
E1 0.21 0.35 0.27 0.1 1
E2 0.67 0.56 0.58 0.05 1
FA2 HR (%) 94 6 77 14 9
E1 0.03 0.29 0.1 0.75 0.38
E2 0.72 0.87 0.74 0.93 0.78
The values of the Preference Index range from 1 (complete avoidance) to +1 (complete preference). M: male, F: female, A: adults, and J:
Juveniles.

conservation of otter habitat. Additionally, conservation of
these den areas may serve as protection for other native wild-
life including endemic understory birds which use the bam-
boo (Sieving et al., 2000; Reid et al., 2004; Dı́az et al., 2005)
found in the otters habitat.
We observed that otters used rivers more frequently than
streams, possibly because higher order rivers provide a great-
er availability of water and therefore more aquatic prey which
comprise the otters’ diet (Medina-Vogel et al., 2003). Further-
more, all described dens were located on major rivers.
 B I O L O G I C A L C O N S E RVAT I O N
Streams had a summer depth of less than 20 cm (Sepúlveda
pers. obs.), which may have prevented typical otter swimming
and diving behaviour. Otter MA2, was observed walking in a
stream bed and seeking refuge in the woody debris along
the bank, instead of submerging in water, when it was star-
tled by people. A similar selection pattern was reported for
L. canadensis (Melquist and Hornocker, 1983) and A. capensis
(Somers and Nel, 2004) in that no otters lived entirely in
streams or first order tributaries.
Adjacent area and influence on otter movements showed
that livestock farming and native forest were used; likely be-
cause they were distributed in floodplains associated with riv-
ers which were selected (see above). Exotic plantations were
located on higher slopes associated with streams and were
subsequently avoided by otters. Higher values of selection
and avoidance were observed in the second order selection,
whereas the third order selection did not demonstrate a clear
pattern. This may be because more important variables were
present in the instream and riparian zones as riparian vegeta-
tion composition, riverbank structure, riverbed (Medina-Vogel
et al., 2003) and trophic offer not evaluated by our study. This
could explain why otters selected livestock farming areas and
why there was moderate avoidance of locations adjacent to
native forests.
The intensity of livestock farming in the study area did not
seem to affect the spatial behaviour of the radiotracked ani-
mals. This suggests that the species may be able to tolerate
some degree of human activities as described by Medina-Vo-
gel et al. (2003) in other Lutrinae species (Somers and Nel,
2004). This observation may be an important consideration
for conservation management decisions pertaining to areas
outside of natural reserves.
The vast diversity of environments (potamon and ritral riv-
ers, mountain lakes, and Southern fjords) used by the South-
ern river otter across its geographic range may mean that
spacing patterns will vary based on habitat type. The home
range sizes and densities reported in this study suggest that
large areas of freshwater habitat are required to sustain viable
otter populations, similar to other carnivore species (Simo-
netti and Mella, 1997; Sunquist and Sunquist, 2001). The need
for adequate otter habitat is concerning because of the high
levels of human impact throughout the majority of the fresh-
water distribution of the species (Medina-Vogel et al., 2003)
along with the disappearance of almost all riparian corridors
in the area (Smith-Ramı́rez, 2004). Furthermore, the otter’s
preferential habitat selection of lowland rivers could lead to
human-otter conflict because this is also prime area for agri-
culture and livestock. Integrated watershed management is
vital for the conservation of this semiaquatic species. The
present reforestation with exotic species as P. radiata and E.
globulus, the rapid deforestation of the chilean Temperate For-
est (Echeverrı́a et al., 2006) and the susceptibility to runoff
erosion from heavy rainfall (Smith-Ramı́rez, 2004) increase
the likelihood of flooding in the watershed lowlands. Conse-
quently, the human demand to drain the wetlands (to pre-
serve agricultural interests) is increased resulting in
deterioration and loss of primary otter habitat, i.e. the case
of the Temu Pitra swamp forest (Sepúlveda pers. obs.).
There are only three reserves in the Chilean System of Pro-
tected Areas (Carlos Anwandter Natural Sanctuary, Vicente
1 4 0 ( 2 0 0 7 ) 3 2 9 –3 3 8 337
Pérez Rosales Nacional Park and Chiloé Nacional Park) that
are located in the freshwater distribution where otters have
been confirmed (Medina, 1996; Sepúlveda et al. in prep.).
These areas have the minimal amount of otter habitat neces-
sary to preserve the populations. These reserves are isolated
within a mosaic of high human density and intense land
use for plantation forestry, farming and livestock (Armesto
et al., 1998). The reserves lack official corridors for the species
and the Carlos Anwandter Sanctuary (Ramsar Site) contains a
pulp factory 40 km upstream that has impacted the local bio-
ta (Artacho et al., 2007). Conservation strategies that encour-
age appropriate land use planning in a productive matrix
are considered a priority for carnivore species (Sunquist and
Sunquist, 2001) and specifically for the conservation of the
South American Temperate Forest (Armesto et al., 1998).
The creation of Southern river otter reserves that provide suf-
ficient quality and quantity of habitat, along with riparian cor-
ridors, are considered an urgent priority to enhance the
likelihood of survival of this carnivore as well as other coexis-
ting species of the chilean temperate forest.
Acknowledgements
We are grateful to the Earthwatch volunteers for field assis-
tance and hard work during trapping and radiotracking peri-
ods. We appreciate the support collaboration from the DID
Universidad Austral de Chile, the Earthwatch Institute and
the Frankfurt Zoological Society and G. Acosta, R. P. Brooks,
J. Rau, E. A. Silva-Rodrı́guez and two anonymous revisors that
reviewed and made comments of the manuscript. We thank
S. Molina for her important assistance with GIS and N. Bur-
dhayar, C. Briceño, M.J. Berecz and K.N. Wolf for English revi-
sion. This work was approved by the bioethical committee of
the Universidad Austral de Chile, and was done under the
restrictions imposed by the Subsecretarı́a de Pesca’s legal per-
mit No. 625 (April 19, 2000) for live trapping and radio tagging
Southern river otters.
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