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By using Global Positioning System technology, we documented the long-distance dispersal of a wolf (Canis lupus) from the northern Apennines in Italy to the western Alps in France. This is the first report of long-distance dispersal of wolves in the human-dominated landscapes of southern Europe, providing conclusive evidence that the expanding wolf population in the Alps originates from the Apennine source population through natural recolonization. By crossing 4 major 4-lane highways, agricultural areas, and several regional and provincial jurisdictions, the dispersal trajectory of wolf M15 revealed a single, narrow linkage connecting the Apennine and the Alpine wolf populations. This connectivity should be ensured to allow a moderate gene flow between the 2 populations and counteract potential bottleneck effects and reduced genetic variability of the Alpine wolf population. The case we report provides an example of how hard data can be effective in mitigating public controversies originating from the natural expansion and recolonization processes of large carnivore populations. In addition, by highlighting the connectivity between these 2 transboundary wolf populations, we suggest that documenting long-distance dispersal is particularly critical to support population-based, transboundary management programs. (JOURNAL OF WILDLIFE MANAGEMENT 73(8): 1300-1306; 2009)
Management and Conservation Note
Long-Distance Dispersal of a Rescued Wolf From the
Northern Apennines to the Western Alps
Dipartimento di Biologia Animale e dell’Uomo, Sapienza Universita
di Roma, Roma, 00185, Italy
WILLY REGGIONI, Parco Nazionale dell’Appennino Tosco-Emiliano, Cervarezza Terme, Reggio Emilia, 42032, Italy
LUIGI MAIORANO, Dipartimento di Biologia Animale e dell’Uomo, Sapienza Universita
di Roma, Roma, 00185, Italy
LUIGI BOITANI, Dipartimento di Biologia Animale e dell’Uomo, Sapienza Universita
di Roma, Roma, 00185, Italy
ABSTRACT By using Global Positioning System technology, we documented the long-distance dispersal of a wolf (Canis lupus) from the
northern Apennines in Italy to the western Alps in France. This is the first report of long-distance dispersal of wolves in the human-dominated
landscapes of southern Europe, providing conclusive evidence that the expanding wolf population in the Alps originates from the Apennine
source population through natural recolonization. By crossing 4 major 4-lane highways, agricultural areas, and several regional and provincial
jurisdictions, the dispersal trajectory of wolf M15 revealed a single, narrow linkage connecting the Apennine and the Alpine wolf populations.
This connectivity should be ensured to allow a moderate gene flow between the 2 populations and counteract potential bottleneck effects and
reduced genetic variability of the Alpine wolf population. The case we report provides an example of how hard data can be effective in
mitigating public controversies originating from the natural expansion and recolonization processes of large carnivore populations. In addition,
by highlighting the connectivity between these 2 transboundary wolf populations, we suggest that documenting long-distance dispersal is
particularly critical to support population-based, transboundary management programs. (JOURNAL OF WILDLIFE MANAGEMENT
73(8):1300–1306; 2009)
DOI: 10.2193/2008-510
KEY WORDS Canis lupus, connectivity, dispersal, Global Positioning System (GPS), Italy, long-range movements,
transboundary wolf management, wolf.
Dispersal strongly affects population dynamics, distribution,
gene flow, spatial and social organization, as well as
colonization and rescue effects (Howard 1960, Wolff
1977), and wolves (Canis lupus) are good candidates for
the study of dispersal (Fuller et al. 2003). Accordingly,
different aspects of wolf dispersal have been reported for
both North America (Gese and Mech 1991, Boyd and
Pletscher 1999, Fuller et al. 2003, Mech and Boitani 2003)
and northern Europe (Wabakken et al. 2001, 2006; Kojola
et al. 2006), but very limited information is available for
south-central Europe (Blanco and Corte
s 2007), where
higher human density and anthropogenic features may affect
wolf dispersal to a much greater extent. In addition, because
logistic and technical constraints did not allow, until
recently, detailed studies of long-distance dispersal by
wolves (Merrill and Mech 2000, Blanco et al. 2005), it is
difficult to fully understand wolf dispersal mechanics and to
predict landscape links (Fuller et al. 2003).
The advent of Global Positioning System (GPS) technol-
ogy made it possible to gain new insights into wolf dispersal
and connectivity among disjoint wolf populations (Kojola et
al. 2006, Wabakken et al. 2006). Analyses of detailed GPS-
revealed dispersal paths may provide information on how an
animal perceives and moves about through the landscape
(With 1994, Nams 2005). In particular, long-distance
dispersal trajectories might reveal which portions of the
landscape still provide connectivity between noncontiguous
populations (Graves et al. 2007).
This knowledge is particularly useful in human-dominated
landscapes, where spatially explicit management interven-
tions might enhance preservation and functionality of
existing links for the long-term viability of metapopulations
(Beier et al. 2006). For noncontiguous wolf populations
across international jurisdictions, documenting long-dis-
tance dispersal is also particularly critical, because it provides
evidence of their genetic and demographic connectivity,
therefore supporting the need for population-based, inter-
state management programs (Boitani 2003, Linnell et al.
In southern Europe, long-distance dispersal by wolves has
recently been inferred using noninvasive genetic data of the
naturally expanding wolf population across the Italian,
French, and Swiss Alps (Lucchini et al. 2002, Valie
re et al.
2003, Fabbri et al. 2007). Although these studies clearly
indicated that wolves in the Alps originated genetically from
the Apennine source population, they did not reveal
dispersal trajectories or sufficient evidence to resolve the
animated debate on the origin of the recolonizing wolves in
France (Spagnou 2003a). Whereas wolf opponents did not
rule out illegal release of captive wolves (Lucchini et al.
2002), some local experts were skeptical that wolves from
the Apennines could successfully travel through the narrow
and altered Ligurian Apennines to reach the Alps (Zunino
2003, quoted in Spagnou 2003b, in litteris).
By using GPS telemetry, we hereby document the first
report of long-distance wolf dispersal from the northern
Apennines in Italy to the western Alps in France. Although
based on a single event, this case provides evidence that the
transboundary wolf population in the western Alps could
have originated naturally from long-distance dispersers from
the Apennine source population. The wolf dispersal
trajectory we report directly demonstrates that a single,
1300 The Journal of Wildlife Management N 73(8)
narrow link still connects the 2 wolf populations across a
highly heterogeneous and human-dominated landscape.
We defined the study area by the outermost locations of the
dispersing wolf (Fig. 1), including the northern Apennines,
in Italy, and part of the southwestern Alps across the border
between Italy and France. Both areas were characterized by
rugged, mountainous terrain with altitudes up to 1,978 m in
the Apennines and 3,084 m in the Alps. Deciduous (mostly
beech, Fagus sylvatica) forests covered about 51% of the
Apennines, whereas the Alps featured 66% forest cover,
mostly composed by conifers (Abies alba, Larix decidua) and,
at lower elevations, deciduous trees (F. sylvatica, Acer
platanoides, Betula pubescens). Wild boar (Sus scrofa), roe
deer (Capreolus capreolus), and red deer (Cervus elaphus) were
locally abundant and heavily used by wolves, both in the
northern Apennines (Meriggi et al. 1996) and in the Alps
(Marucco 2003, Gazzola et al. 2005). Livestock, mostly free-
ranging cattle, was available to wolves all year, especially in the
northern Apennines. Wolf distribution in the Alpine portion
of the study area was continuous and expanding (Lucchini et
al. 2002, Marucco 2003, Valie
re et al. 2003) but it was
separated from the source Apennine population by a gap of
more than 200 km (Fabbri et al. 2007).
Following expansion of the wolf population in the central
Apennines during the past 30 years (Boitani and Ciucci
1993), the wolf range in the northern Apennines was
continuous up to about 44u409N latitude (Ciucci et al. 2003)
at the time of the study. However, it was believed to be more
discontinuous along the narrow, northwesternmost portion
of the Apennine chain (Meriggi et al. 2002). Snow cover
usually extended from November and December to April in
the Alps and from December and January through March in
the northern Apennines. Human density averaged 52.8
(6346.5 SD) and 50.1 (6307 SD) people/km
in the
Apennines and in the Alps, respectively, although its
dispersion varied considerably on a local scale. Mean
densities of permanent roads (highways, other paved roads,
and improved unsurfaced roads passable by 2-wheel drive
vehicles; Mladenoff et al. 1995) were 2.98 km/km
1.97 km/km
in the Apennines and in the Alps, respective-
ly. Agriculture and other anthropogenic cover types
accounted for slightly more than 50% in the Apennines,
but less than 10% in the Alpine range (Falcucci et al. 2007).
The Italian portion of the study area included 5 regional and
several more provincial administrative units, each with their
own land use and wildlife management jurisdiction, and
several protected areas (1,699 km
; 14% of the study area;
Fig. 1).
Figure 1. Global Positioning System (GPS)-estimated long-distance dispersal path of wolf pup M15, from the northern Apennines (Italy) to the western
Alps across the Italian–French border (11 Mar 2004–22 Jan 2005). Only selected towns (i.e., .10,000 inhabitants) and the main protected areas close to the
dispersal trajectory are shown. Open circles display sharp turning angles during directional dispersal, whereas the question mark corresponds to 4 days of
missing data (22–25 Sep 2004).
Ciucci et al. N Wolf Dispersal From Italy to France 1301
On 11 March 2004, we released in the northern Apennines
a 28-kg male wolf pup M15 (10 months old, age estimated
by tooth eruption and wear; Gipson et al. 2000) rescued on
24 February 2004 from a vehicle accident in the outskirts of
the city of Parma (Fig. 1). Immediate examination upon
rescue revealed limited external and internal bleeding, and a
slight limp at the front left leg (G. M. Pisani, Province of
Parma, personal communication). To avoid risks of human
habituation, in less than 24 hours, we moved the wolf to a
small and isolated stone hut within a nearby protected area
(Cento Laghi Regional Park) in the northern Apennines, as
the wolf gave signs of quick recovery. We fed M15 road-kills
(roe deer) and assessed its condition by observation at a
distance every 1 to 2 days (M. Andreani, Cento Laghi
Regional Park, personal communication). After 15 days, as the
wolf was quickly recovering, we released it in a secure place
nearby, located in the interstice between the territories of 2
wolf packs (Ciucci et al. 2003). Upon release, we sedated
(medetomidine and ketamine, antagonized upon release with
atipamezole; Kreeger et al. 2002) and fitted wolf M15 with a
Televilt (Lindesberg, Sweden) GPS-Direct collar. Although
the natal territory of wolf M15 was unknown, its 12-
microsatellite genotype matched a known genotype (E. Randi,
National Institute of Wildlife, personal communication),
whose corresponding fecal sample was previously (6 Dec
2003) collected at about 80 km southeast along the northern
Apennine chain (D. Pagliai, Alto Appennino Modenese
Regional Park, personal communication; Fig. 1). It is thus
plausible that wolf M15 was born from one of the packs in
that portion of the Apennines (Ciucci et al. 2003).
We retrieved data, including location, date, time, and
estimates of position quality (2D, 3D, and 3D+), from the
collar through the Global System for Mobile Communica-
tions (GSM). We allowed 180 seconds for each fix attempt,
and programmed the collar to acquire locations at 4-hour
(until 4 May 2004) and 12-hour (after 5 May 2004)
intervals. Field crews investigated clusters of GPS locations
during the postrelease period, at highway crossings, and
after settlement. We permanently lost GSM contact with
the collar by 22 January 2005, but only by 18 February 2005
did the field crew localize the carcass of wolf M15 by
homing in on the very high frequency signal. When found,
the wolf had probably died
10 days before and had been
entirely consumed by scavengers, making it impossible to
determine the cause of death.
We projected wolf M15 locations in ArcGis 9.2, assuming
inaccuracy of GPS position was negligible at the scale of our
analysis (expected range: ,30 m to , 99 m 95% of the time
for 3D and 2D positions, respectively; Dussault et al. 2001,
D’Eon et al. 2002). We analyzed the overall dispersal
trajectory using locations recorded at 12-hour sampling
intervals (n 5 484; 0730 hr and 1930 hr), including those
subsampled, at the same time intervals, from the 4-hour
sampling dataset (n 5 94).
We defined natal dispersal as the one-way movement from
the release (or presumed birth) site to an independent home
range, where wolf M15 would have presumably reproduced
if it had survived (Gese and Mech 1991, Boyd and Pletscher
1999, Wabbaken et al. 2006, Blanco and Corte
s 2007). We
quantified overall net displacement as the largest Euclidean
distance covered from the release site to the furthest location
along the dispersal trajectory. Differently, we quantified net
dispersal distance as the largest Euclidean distance from the
release site to the harmonic mean of the final home range
(Kenward et al. 2002). We approximated minimum
distances traveled as the sum of the Euclidean distances
traveled between successive 12-hour locations. Failed GPS
attempts (n 5 128) mostly comprised single locations (n 5
110), and only 14% included 2 successive locations. In case
one location was missing, we estimated it by linear
interpolation between successive locations (Ciucci et al.
1997, Stoner et al. 2007) to ensure a constant sampling
interval for the entire movement trajectory for movement
path analyses (see below).
To explore movement patterns during dispersal, we visually
inspected discontinuities in the cumulative net displacement
curve, because they reflected differences in the rate and extent
of geographical displacement (Fig. 2). By sequentially
demarcating patterns indicating use of the same general area
(i.e., little or no increase in net displacement) from those
reflecting a consistent travel from the release site (i.e., no
return to previously visited areas), we thus identified 11
dispersal phases, each featuring 1 of 4 different movement
patterns (Fig. 2): 1) local movements, with more or less
localized spatial behavior and recurrent use of the same
general area; 2) directional movements, with consistent
traveling in a predominant direction; 3) directional shifting,
an intermediate pattern between the previous 2, when wolf
M15 gradually shifted a restricted area of activity in one
direction; and 4) home range–like movements, similar to local
movements but more localized, reduced in extent, and for an
extended period of time. We considered the home range–like
movements as an indication of settlement (Gese and Mech
1991, Mech and Boitani 2003). To better characterize the
multistage pattern of dispersal (sensu Wabbakken et al. 2006),
we used linear, fractal, and circular metrics to describe
dispersal phases.
We computed fractals by the Fractal Mean method (Nams
1996) using FRACTAL (version 4.1,
staff/vnams/Fractal.htm, accessed 15 Jul 2009). We used
basic circular statistics (Zar 1999) to describe and test
directionality of travel, both within each dispersal phase
(first-order samples) and within movement patterns (i.e.,
directional vs. local movement phases; second-order sam-
ples). In particular, we measured angular dispersion of
traveling bearings by the mean vector r, a measure of angular
concentration that can vary from 0 (high angular dispersion
and no mean bearing) to 1 (all bearings have the same
direction), and used the Rayleigh’s z to test the null
hypothesis of no angular concentration. We used Moore’s
modification of the Rayleigh test (Zar 1999) to test the
hypothesis of no angular concentration in second-order
samples. In statistically comparing fractal and circular
metrics among dispersal phases, we assumed they were
1302 The Journal of Wildlife Management N 73(8)
independent, because wolf movements responded to differ-
ent environmental and social stimuli across the different
areas encountered during dispersal; however, we caution
against interpreting significance levels because all move-
ments pertain to a single wolf.
In 318 days since release (11 Mar 2004–22 Jan 2005), wolf
M15’s collar acquired 653 locations, with an acquisition rate
of 76.1%. Acquisition rate did not differ between the 4-hour
and the 12-hour GPS-schedules (G
5 1.02, P 5 0.31),
nor between the broadleaf (Jun–Oct) and the broadleaf-less
(Nov–May) seasons (G
5 2.19, P 5 0.14). At the 12-
hour sampling, acquisition rate of GPS locations was higher
in the evening (83.5%, at 1930 hr) than in the morning
(72.2%, at 0700 hr; G
5 9.34, P 5 0.002). Most
locations (66.6%) were of high accuracy (3D, 3D+), and
their proportion was not affected by sampling interval
5 0.11, P 5 0.74) or vegetative season (G
0.52, P 5 0.47). Since deployment, both measures of GPS
performance did not vary with increasing battery drainage
on a monthly basis (acquisition rate: F
5 0.08, P 5 0.78;
proportion of 3D locations: F
5 0.05, P 5 0.82).
Following release, wolf M15 spent about 2 months
roaming north, east, and southeast of the release site in an
area of about 514 km
, where we knew at least 3 other packs
existed (Fig. 1). During this period, at least 8 ground
investigations on clusters of
2 locations, partly aided by
snow, provided evidence that wolf M15 was traveling alone
and was feeding on roe deer, wild boar, and occasionally on
livestock carcasses (M. Andreani, personal communication).
Wolf M15 then abandoned this area and began traveling at
a faster pace in a west–northwesterly direction along the
Apennines, eventually reaching the French Alps by 2
October 2004, about 7 months after release. Wolf M15
then floated for about one month across the Italian–French
border, using an area of about 694 km
in the same general
locality where in 1993 the first noninvasive genetic sample
of a recolonizing wolf was collected (Valie
re et al. 2003). We
knew a minimum of 3 wolf packs resided in that area at the
time wolf M15 arrived (Wolf Alpine Group 2004). By 8
months after release, wolf M15 eventually began to restrict
its movements and settled on the Italian side of the Alps for
2.5 months, carving out a home range (95% fixed kernel) of
71.8 km
in an area between 2 resident wolf packs. Since
December 2004, a field crew investigated wolf M15’s tracks
in the snow (16 sessions, 44 km), and revealed that in 88%
of these sessions, wolf M15 was associated with another
wolf (F. Marucco, Piemonte Large Carnivores Project,
personal communication). Noninvasive genotyping later
confirmed that by 16 January 2005 wolf M15 had
permanently paired with wolf F70, a yearling female from
one of the resident packs (M. Schwartz, Rocky Mountain
Research Station, personal communication). The same
winter, wolf F70 was tracked alone from 9 February onward
(F. Marucco, personal communication), probably following
the death of wolf M15.
The furthest location wolf M15 reached during dispersal
was on the French side of the western Alps, for a maximum
net displacement of 217.3 km. Conversely, net dispersal
distance was 186.8 km, considering the release site, or
239.7 km with respect to the putative natal range (Fig. 1).
The net dispersal distance from the release site corresponded
to a minimum distance traveled of 958 km, which, corrected
by a factor of 1.3 (Musiani et al. 1998, Wabakken et al.
2006), yields an estimate of 1,245.3 km actually travelled.
Geographical displacement during dispersal was not
constant over time, and discontinuities in cumulative net
displacement indicated 11 sequential dispersal phases
(Fig. 2). Excluding directional shifting for which data were
incomplete, directional movements (n 5 4 phases) contrib-
uted the most to net (x
5 44.9, SD 5 37.8 km/phase) and
daily (x
5 8.6, SD 5 0.6 km/day) distance travelled,
whereas local movements (n 5 5, excluding home range–
like movements), contributed less (x
5 20.1, SD 5
10.4 km/phase, and x
5 3.4, SD 5 1.2 km/day for net
and daily displacement, respectively). Accordingly, phases
featuring local movements (n 5 5) were on average more
tortuous than directional movements (n 5 4; Fractal D: x
1.52, SD 5 0.21, and x
5 1.20, SD 5 0.06, respectively; t
522.96, P 5 0.021; Table 1). During localized phases
wolf M15 travelled minimum distances of 5.7–16.4 km/day,
indicating that it was not stationary but moved intensively in
the same general area, possibly searching for other wolves or
resources. Individually considered, directional phases had a
predominant traveling direction (0.20
0.81; 3.49
Rayleigh z
6.79, 0.001
0.05), whereas
localized phases did not reveal any particular directionality
0.15; 0.00
Rayleigh z
1.16, P .
0.05). Accordingly, directional phases had little angular
dispersion (r 5 0.81) and an overall bearing of 266.1uN (SD
5 37.1uN; Moore test: R
5 1.10, P , 0.05), whereas, in
contrast, localized phases did not display a preferred
traveling bearing (Moore test: R
5 0.47, P . 0.50).
Wolf M15’s dispersal trajectory extended through the
main Apennine chain, with an overall mean bearing of
Figure 2. Minimum daily distance and cumulative net displacement
traveled by wolf M15 from the release site during its Global Positioning
System–revealed dispersal from the northern Apennines (Italy) to the
western Alps (France and Italy; 11 Mar 2004–22 Jan 2005). We identified
11 dispersal phases on the basis of discontinuities in the cumulative net
displacement curve, and they are shown by alternate black and gray sections
(cf. Table 1).
Ciucci et al. N Wolf Dispersal From Italy to France 1303
264.5uN, at altitudes ranging from 270 m to 2,664 m.
During directional dispersal 3 sharp turning angles (phases 2
and 6; x
5 124.2uN, SD 5 24.8uN, in absolute values) were
on average greater (Watson and William test: F
5 14.89,
P , 0.001) than the overall mean of directional movements
5 44.8uN, SD 5 37.6uN, n 5 40), suggesting the wolf’s
attempt to redirect and maintain its traveling along the main
Apennine chain as it reached lower elevations or increasingly
developed areas (Fig. 1).
During dispersal, wolf M15 traveled across 2 national, 5
regional, and several provincial administrative units, and
went as close as 0.8–5 km to large towns such as Cuneo and
Genoa. The wolf navigated several potential barriers,
including 4 fenced 4-lane highways (traffic volumes in
Jul–Sep 2004 ranging 49,928–143,081 vehicles/day; AIS-
CAT 2004), several main railways, and many state,
provincial, and local paved roads. Wolf M15 crossed
highways with apparent ease (
12–24 hr) and, as from
field investigations, systematically used underpasses, which
are frequent along highways in these mountainous areas. As
an exception, wolf M15 clustered for 4 days at 700–1,100 m
east of highway A7 before crossing it. Although we cannot
exclude the presence of a carcass at the site, the juxtaposition
of the highway, an unfenced 2-lane state road, a railway, and
a river, all at the bottom of a steep valley flanked with
concrete banks .10 m high, might have presented wolf
M15 with a serious challenge, thereby delaying its
movements. At the edge of the northwestern Apennines,
while traveling in a southwesterly direction, wolf M15
turned northwestward before crossing highway A6, thus
reaching the westernmost portion of the Po River Valley at
altitudes as low as 300 m. Unfortunately, having lost GSM
data during this period (22–25 Sep 2004), we cannot assess
whether this change of direction might have resulted from a
failure in negotiating a more direct route to cross highway
A6 (Fig. 1). However, wolf M15 crossed highway A6
further north, utilizing the 900-m-wide, riparian vegetation-
rich drainage of the Pesio River running underneath the
highway. By following the same drainage for an additional
11 km, wolf M15 eventually reached the densely populated
outskirts of Cuneo, where it turned southward to finally reach
the Alps in less than 48 hours (Fig. 1). In this heavily
cultivated and developed area, wolf M15 traveled at a fast pace
during the night (15–16 km/night) and rested during the day,
using the thick and locally widespread corn plantations.
Being based on a rescued wolf, our study differs from other
telemetry-based wolf dispersal studies (Boyd and Pletscher
1999, Kojola et al. 2006, Wabakken et al. 2006, Blanco and
s 2007). We cannot therefore exclude that actual
dispersal distances were higher than those reported, or that
the prerelease events (vehicle accident, rescue, and recovery)
might have influenced to some extent wolf M15’s
subsequent dispersal behavior. Nevertheless, the observed
natural dispersal behavior supports the idea that limited
handling and no contact with humans during the brief
recovery were successful in avoiding conditioning or
habituation effects.
Although our results are based on a single case, they
provide clear evidence that wolves can disperse through the
human-dominated landscapes of the northern Apennines.
Wolf M15’s dispersal trajectory directly demonstrates that a
functional linkage still exists between the Apennine and the
Alpine wolf populations. This was previously inferred from
genetic studies, based on which unidirectional and male-
biased dispersal from the Apennine population could have
Table 1. Sequential phases of wolf M15 dispersal from the northern Apennines (Italy) to the western Alps (France) based on 484 Global Positioning System
(GPS) locations acquired at 12-hour intervals (Mar 2004–Jan 2005).
phase Date
Net displacement
Min. distance traveled (km)
Greatest Day
Mean SD Min. Max.
Local 1 11 Mar–7 May 98 25.6 0.4 173.2 3.0 2.8 0.1 11.8 1.45 0.05 59.7
Directional 2 8–14 May 7 39.9 5.7 54.8 7.8 2.9 5.6 13.5 1.23 0.76 262.7*
Local 3 15 May–13 Jun 55 13.9 0.5 92.2 3.1 2.6 0.1 9.9 1.81 0.14 295.8
Directional 4 14–17 Jun 7 24.6 6.2 28.5 7.1 3.7 3.2 10.7 1.16 0.81 331.0**
Local 5 18–27 Jun 12 8.0 0.8 29.8 3.0 2.4 0.6 5.7 1.34 0.01 111.0
Directional 6 28 Jun–4 Jul 10 15.5 2.2 42.0 6.0 5.0 0.6 15.2 1.27 0.20 251.9
shifting 7 5–26 Jul
23 12.8 0.6 20.1 0.9 0.7 0.1 2.1 1.24 0.32 268.2
Local 8 27 Jul–12 Sep 74 18.4 0.4 127.0 2.6 2.1 0.1 9.5 1.34 0.06 250.4
Directional 9 12–28 Sep
16 99.6 5.9 117.4 6.9 4.7 0.8 19.5 1.13 0.65 242.4***
Local 10 28 Sep–11 Nov 71 34.7 0.8 256.5 5.7 4.2 0.1 16.4 1.64 0.01 57.3
like 11 12 Nov–22 Jan 114 9.5 0.1 285.0 4.0 3.1 0.5 13.0 1.91 0.07 110.8
Greatest net displacement from the first location of current phase.
Cumulative Euclidean distances summed across successive locations.
By the Fractal Mean method (Nams 1996) using FRACTAL. Min. and max. scales were constrained by the average step size and one-third of total path
length, respectively (With 1994), and 10 divider lengths were used to measure the length of each movement path.
Marked values from angular distributions different from uniform expected (Rayleigh test; * P , 0.01; ** P , 0.001; *** P , 0.001).
Data from 8–14 Jul were lost due to downloading failure.
Data from 22–25 Sep were lost due to downloading failure.
1304 The Journal of Wildlife Management N 73(8)
occurred repeatedly at a rate of 1.25–2.5 wolves/generation
(Fabbri et al. 2007).
Although wolves may disperse as much as 390–1,092 km
(Boyd and Pletscher 1999, Wabakken et al. 2006), wolf
M15 traveled a dispersal distance higher than the average
reported for wolves in the more pristine landscapes of North
America (77–113 km; Gese and Mech 1991, Boyd and
Pletscher 1999, Mech and Boitani 2003) and northern
Europe (99 km; Kojola et al. 2006). This dispersal distance
is the highest so far documented by means of telemetry in
the human-dominated landscapes of southern Europe
(Spain: Blanco and Corte
s 2007; Italy: P. Ciucci, Sapienza
University of Rome, unpublished data).
Dispersing wolves seem to maximize breeding opportunities
rather than resource acquisition (Boyd et al. 1995, Wydeven et
al. 1995, Mech and Boitani 2003). Therefore, they may travel
long distances due to the low probability of finding a mate
(Boyd and Pletscher 1999, Wabakken et al. 2006). However,
not only conspecific attraction (Boyd and Pletscher 1999,
Blanco and Corte
s 2007), but also the rough and irregular
topography of the Apennines chain north of the release site
may have influenced both distance and direction of wolf
M15’s dispersal. Similarly to dispersing wolves in Montana,
USA, which used a narrow swath along the Rocky Mountain
chain where other wolves were present (Boyd et al. 1995), wolf
M15’s movements appeared to be funneled along the narrow
stretch of the northern Apennines, confirming a previously
postulated linkage effect of this tract of the Apennines
between the Apennine and the Alpine wolf populations
(Mech and Boitani 2003, Fabbri et al. 2007).
Wolf M15’s dispersal confirms the ability of wolves to
cross areas previously believed to act as barriers, such as
open, agricultural, and developed areas, or other linear
infrastructures (Mech et al. 1995, Merrill and Mech 2000,
Fuller et al. 2003, Valie
re et al. 2003, Blanco et al. 2005).
Elsewhere, however, highways with much lower traffic
volumes (4,000 vehicles/day) act as barriers to wolf
movements through direct mortality (Paquet 1993) and
reduced movement rates (Alexander et al. 2005). Although
we cannot infer population level responses from a single
event, we believe that in the Apennines highway crossing
may be facilitated by naturally occurring mitigation provided
by the many under- or overpasses largely negotiable by
wolves while traveling. Nevertheless, as exemplified by wolf
M15’s crossing of highway A7, the local juxtaposition of
several linear structures may represent a more difficult
obstacle, especially for dispersing wolves without prior
spatial knowledge of the area (see also Blanco et al. 2005).
Management Implications
Wolf M15’s dispersal conclusively demonstrates that wolves
from the Apennines can travel across the altered landscape of
the Ligurian Apennine chain to reach the Alps, supporting
previously inferred conclusions from genetic studies on the
natural recolonization of the Alps by long-distance–dispers-
ing wolves from the Apennines (Lucchini et al. 2002, Valie
et al. 2003, Fabbri et al. 2007). Because we made wolf M15’s
case public after its death, it was reported by most national
and local news media in France, which presented this case as
proof that wolves have returned to the Alps naturally (B.
Lequette, Mercantour National Park, personal communica-
tion). As a consequence, shepherds and farmers’ organizations
also ceased to openly support the artificial reintroduction
hypothesis. Hard data from a single wolf outweighed all
logical and biological inferences offered by scientists on the
natural expansion of the wolf range throughout northern Italy
and the Alps (Lucchini et al. 2002, Valie
re et al. 2003).
Acceptance of the natural recolonization process implied that
the wolves in the Alps are fully protected under the provision
of the Habitat European Directive and should be allowed to
establish a viable population. The ultimate evidence of the
habitat and population continuity across the Italian–French
boundary has been instrumental in prompting formal
meetings of the Italian, French, and Swiss authorities to
discuss a road map toward a common management plan of
the Alpine wolf population.
In this perspective, the functional connectivity between
the Apennine (source) and Alpine (colony) wolf populations
should be maintained, at least at the estimated current rate
(Fabbri et al. 2007) deemed sufficient to counteract serious
bottleneck effects for the Alpine wolf population. Our
results contributed to highlighting the landscape linkage
across the Ligurian Apennines for its future preservation
and mitigation of potential barriers.
M. Andreani, L. Grottoli, F. Marucco, L. Molinari, and L.
Orlando provided assistance during field investigations of
cluster of GPS locations. The Cento Laghi Regional Park,
the Province of Parma, and the Alpi Marittime National
Park provided logistic and administrative support for the
recovery, release, and monitoring of wolf M15. Funds were
provided by the Life-Natura Project LIFE00 NAT/IT/
007214 and the Department of Human and Animal Biology
of the Sapienza University of Rome. W. R. Clark and 2
anonymous referees provided useful comments and sugges-
tions on an earlier draft of this paper.
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1306 The Journal of Wildlife Management N 73(8)
... In Europe, some studies present dispersal rates similar to the one chosen in this study, as is the case with Finnish wolves who showed a dispersal distance of 98.5 km [80]. However, this number may be much higher, estimated at between 250 and about 400 km [81][82][83], and even reaching 800 km in the movements of the population of German wolves to Poland [12]. Ciucci et al. [83] determine the Italian grey wolf straight line dispersal distance (from the Northern Apennines to the Alps) as 240 km. ...
... However, this number may be much higher, estimated at between 250 and about 400 km [81][82][83], and even reaching 800 km in the movements of the population of German wolves to Poland [12]. Ciucci et al. [83] determine the Italian grey wolf straight line dispersal distance (from the Northern Apennines to the Alps) as 240 km. Similar data were presented by Ražen et al. [84], who set the straight line dispersal distance between the Dinaric-Balkan and Alpine grey wolf populations at 230 km. ...
Full-text available
Over the last few decades, much of the mountain area in European countries has turned into potential habitat for species of medium-and large-sized mammals. Some of the occurrences that explain this trend are biodiversity protection, the creation of natural protected areas, and the abandonment of traditional agricultural activities. In recent years, wolves have once again been seen in forests in the eastern sector of the Pyrenees and the Pre-Pyrenees. The success or failure of their permanent settlement will depend on several factors, including conservation measures for the species, habitat availability, and the state of landscape connectivity. The aim of this study is to analyze the state of landscape connectivity for fragments of potential wolf habitat in Catalonia, Andorra, and on the French side of the Eastern Pyrenees. The results show that a third of the area studied constitutes potential wolf habitat and almost 90% of these spaces are of sufficient size to host stable packs. The set of potential wolf habitat fragments was also assessed using the probability of connectivity index (dPC), which analyses landscape connectivity based on graph structures. According to the graph theory, the results confirm that all the nodes or habitat fragments are directly or indirectly interconnected, thus forming a single component. Given the large availability of suitable habitat and the current state of landscape connectivity for the species, the dispersal of the wolf would be favorable if stable packs are formed. A new established population in the Pyrenees could lead to more genetic exchange between the Iberian wolf population and the rest of Europe's wolf populations.
... Grey wolves, Canis lupus, are expanding their range throughout Eurasia and North America, almost entirely via natural dispersal (Ciucci, Reggioni, Maiorano, & Boitani, 2009). Although wolf social behaviour can vary, the general pattern of dispersal is that younger, nonbreeding adults leave their natal pack to seek out a mate and establish a new breeding territory elsewhere (Mech & Boitani, 2003). ...
... Under idealized circumstances, the majority of wolves eventually disperse, unless they attain breeding status within a pack. Many individual wolf dispersal events have been documented via radiotelemetry (Ballard et al., 1983(Ballard et al., , 1987Boyd & Pletscher, 1999;Fritts, 1983;Fritts & Mech, 1981;Fuller, 1989;Gese & Mech, 1991;Mech, 1987;Mech, Fritts, & Wagner, 1995;Peterson, Woolington, & Bailey, 1984;Van Camp & Gluckie, 1979) and GPS tracking (Ciucci et al., 2009;Jimenez et al., 2017;Kojola et al., 2006Kojola et al., , 2009Razen et al., 2016), including exceptional cases of over 1 000 km of linear distance (Wabakken et al., 2007). Broadly, these studies confirm that wolves establish territories to avoid areas occupied by other wolves and to avoid humans. ...
Full-text available
Dispersal is a fundamental ecological process that, on an individual level, is associated with intrinsic and fixed personality traits such as ‘boldness’. However, it is unknown whether personality traits or behavioural syndromes might change as a consequence of dispersal itself. We analysed 14 GPS-collared grey wolves, Canis lupus, in Finland that dispersed from their natal territories and settled in new territories, as well as 22 time-matched nondispersing controls. We used wolf avoidance of low-use forest roads, high-use primary roads and houses as proxies for boldness, and we estimated wolf responses to these features across spatial scales and across dispersal phases. Wolves, which already occupy portions of Finland with low human density, expanded into more human-impacted areas, but with a strong large-scale selection for relatively low human presence. At the finer scale, avoidance of human elements varied as a function of dispersal phase. During dispersal, avoidance of forest roads, primary roads and houses was substantially lower than in natal territories, reflecting the need to take more risks while moving in a small group (alone or with a single mate) in unfamiliar environments. After settling in new territories, the strength of avoidance of those elements continued to be lower than in natal territories, even after controlling for changes in availability. These results suggest that experiences gained across a major life-history transition, such as dispersal, might themselves influence personality traits.
... Ce constat est tiré de nombreuses études menées essentiellement en Amérique du Nord où les densités de populations lupines sont généralement plus élevées que chez nous [21] . [35,36,37] . Des phases de pause sont souvent observées durant cette période, ralentissant le rythme de dispersion de l'individu. ...
... Au sein de leur domaine vital, les individus peuvent parcourir des distances de l'ordre de 30 kilomètres par jour [36,40] . Les mesures de protection assurées par la Convention de Berne ont ensuite été confirmées en 1992 par la Directive européenne 92/43/CEE sur la protection des habitats naturels ainsi que de la faune et de la flore sauvages (dite « Directive Faune-Flore-Habitats » ou « Directive Habitats »). ...
Full-text available
First wolf plan for Wallonia : "Plan d'action pour une collaboration équilibrée entre l'homme et le loup en Wallonie".
... During this dispersal, when they can sometimes cover large distances, many solitary wolves are detected sporadically when their DNA is found on a prey carcass (Andersen et al. 2105;Ciucci et al. 2009;Ražen et al. 2016;Wabakken et al. 2007). Many wolves are killed during dispersal, mainly on roads, while some poaching also occurs (Reinhardt et al. 2019a). ...
Considering the spread of the grey wolf (Canis lupus) in Europe over the past 30 years, Luxembourg took some measures to prepare for the return of this apex predator, including the establishment of a management plan that notably addresses the issue of wolf depredation on livestock. Here we present the results of genetic analyses of putative wolf saliva, hair and scat samples collected from or near prey carcasses between 2015 and 2020. In two cases, the wolf was confirmed via DNA analysis: in July 2017 near Garnich and in April 2020 near Niederanven, both assigned to category C1 (hard evidence). A third case was classified as C2 (confirmed observation) based on prey carcass characteristics, while genetic analysis yielded no result. These are the first confirmed records of wolves in Luxemburg since 1893. Moreover, the two C1-cases originated from the Alpine (Garnich) and Central European (Niederanven) populations. Given similar developments in the neighboring countries and regions, we conclude that the area including the Benelux countries as well as Rhineland-Palatinate, Saarland and Northern France may well become a melting pot for wolves of the two aforementioned populations in the coming years and decades.
... Compared to the observations on wolf movement by Joly et al. (2019) and Mech and Boitani (2010), the distance might seem inconspicuous, however, studded with large rivers; the peripheries comprise dense human habitations. Ciucci et al. (2009) denoted a somewhat similar remark on wolf movement from the Apennines to the Alps traversing highways and settlements. On the contrary, the topography of the Sundarbans and adjacent regions is very different than the documented dispersal routes of wolves. ...
The Indian grey wolf, Canis lupus pallipes Sykes, 1831, is a small, cryptic subspecies and the only wolf living in arid plains and deserts of the Indian subcontinent. Since 1950, it has been considered extinct beyond 88° east longitude. Herein, we report an instance from Bangladesh after 70 years. A solitary male of C. l. pallipes was killed in retaliation in June 2019 as livestock predation events erupted and lasted for a month after a severe cyclone had swept coastal Bangladesh. The specimen was about 119 cm from nose to tail tip with a skull length of 26.23 cm. Two molecular markers, mt d-loop control region and 16S rRNA, and 54 cranial parameters consolidated the identity. Bayesian inference and maximum-likelihood analyses indicated its intraspecies position. The locality of conflict, 450 km eastward of the easternmost population of C. l. pallipes, is adjacent to the Sundarbans in the Ganges estuary that presents formidable tidal rivers as dispersal barriers. In 2017, another wolf was sighted from the Indian Sundarbans vicinity. The present incident and the sighting of 2017 remarkably appeared from the farthest corners of a 10,000 km 2 strong mangrove network that is rimmed by dense human settlements. The records surmise about the most challenging wolf dispersal route ever recorded. Additionally, the south-central coasts of Bangladesh, once home to wolves, bear old planted mangroves with open dunes but never surveyed for mammals. These facts necessitate a systematic camera-trapping in the coastal mangroves of Bangladesh exclusively intended for wolves.
... In areas of high prey availability, dispersal is postponed (Ballard et al., 1987;Blanco and Cortés, 2007;Jimenez et al., 2017) and is rather triggered by the onset of sexual maturity of young wolves (Gese and Mech, 1991;Messier, 1985;Packard and Mech, 1980) so that most wolves have dispersed from their natal pack by the age of 3 (Gese and Mech, 1991;Jimenez et al., 2017). Given wolves dispersal abilities, individuals may move from one population to another through long distance dispersal (Blanco and Cortés, 2007;Ciucci et al., 2009) in immigration and emigration processes. One of the main processes for dispersing wolves to reproduce is to form a new pack with a dispersing mate of the opposite sex (Mech and Boitani, 2003). ...
Full-text available
The occurrence of wolf populations in human-dominated landscapes is challenging worldwide because of conflicts with human activities. Modeling is an important tool to project wolf dynamics and expansion, and help in decision making concerning management and conservation. However, some individual behaviors and pack dynamics of the wolf life cycle are still unclear to ecologists. Here we present an individual-based model (IBM) to project wolf populations while exploring the lesser-known processes of the wolf life cycle. IBMs are bottom-up models that simulate the fate of individuals interacting with each other, with population-level properties emerging from the individual-level simulations. IBMs are particularly adapted to represent social species such as the wolf that exhibits complex individual interactions. Our IBM projects wolf demography including fine-scale individual behavior and pack dynamics based on up-to-date scientific literature. We explore four processes of the wolf life cycle whose consequences on population dynamics are still poorly understood: the pack dissolution following the loss of a breeder, the adoption of young dispersers by packs, the establishment of new packs through budding, and the different breeder replacement strategies. While running different versions of the IBM to explore these processes, we also illustrate the modularity and flexibility of our model, an asset to model wolf populations experiencing different ecological and demographic conditions. The different parameterization of pack dissolution, territory establishment by budding, and breeder replacement processes influence the projections of wolf populations. As such, these processes require further field investigation to be better understood. The adoption process has a lesser impact on model projections. Being coded in R to facilitate its understanding, we expect that our model will be used and further adapted by ecologists for their own specific applications.
... Dispersal is the primary way that maturing young gray wolves (Canis lupus lupus) potentially colonize new areas and maintain population genetic diversity (Fuller et al. 2003). Different aspects of gray wolf dispersal have been studied extensively in North America (Gese and Mech 1991;Boyd and Pleischer 1999;Fuller et al. 2003;Mech and Boitani 2003;Musiani et al. 2007;Treves et al. 2009;Jimenez et al. 2017) and in Europe (Wabakken et al. 2001(Wabakken et al. , 2007Linnell et al. 2005;Kojola et al. 2006;Blanco and Cortes 2007;Ciucci et al. 2009;Andersen et al. 2015;Byrne et al. 2018), but very limited information is available for North-East Asia, except of fragmented information from wolf dispersal research in China (Duan et al. 2016) and in the Gobi in Mongolia (Kaczensky et al. 2008;Joly et al. 2019). Wolves have been documented to disperse up to 1000 km with at least 3471 km traversed over 271 days (Wabakken et al. 2007), but more commonly 100-150 km from their natal pack (Linnell et al. 2005;Treves et al. 2009). ...
Using remote tracking (GPS+GSM module) we documented long-distance natal dispersals of two yearling wolves (Canis lupus) from The Daursky State Nature Biosphere Reserve, Russia. From the arithmetic center of natal home ranges the collared male and female traveled the straight-line natal dispersal distance of 280 km and 332.8 km, over 82 days and 34 days, respectively. Minimum distances of the entire tracking period were 3090.7 km (male) and 2056.7 km (female); the estimated actual travel distance of the entire tracking period was 9849 km and 4530 km, respectively. The travel speed of the wolves varied between phases (pre-dispersal, dispersal, and post-dispersal) and movement patterns (directional, nondirectional, and cluster). The mean travel speed of both wolves was the highest during dispersing (34.6 and 39.5 km/day), calculated as a minimum distance. It was one of the highest dispersal speeds among reported. The highest hourly mean travel speed was during pre-dispersing at dawn, moving directly (the male, 5.77 ± 4.25 km/h; the female, 4.09 ± 2.44 km/h). During pre-dispersing forays they returned several times to their home territories. During dispersal, yearlings crossed at least 5 territories of other packs. Wolves explored the steppe and forest-steppe in less modified habitats of the Russian part of the Dauria ecoregion and in the human-dominated Chinese part of the ecoregion.
... However, the few cases about the post-release behavior of rehabilitated wolves reported in the current scientific literature were mainly focused on animals without severe traumatic injuries. Indeed, only one study described the rehabilitation and successive release of two severely injured wolves in Northwest Portugal (RIO-MAIOR et al., 2016) and in only one other, the activities of a rehabilitated and released wolf were closely monitored over a long post release period (CIUCCI et al., 2009). ...
Full-text available
This case report describes the rescue of an eight-month-old male Italian wolf (Canis lupus italicus), the victim of a car accident that caused it a pulmonary contusion, a fracture of the shaft of right femur, and a metaphyseal fracture of the left stifle. A lateral surgical approach was performed to treat the animal’s multiple contusions and fractures. Afterwards the wolf was transferred to a wild animal recovery center for its rehabilitation, where it fully recovered. After 35 days in captivity the wolf was thus released into the supposed home-range of its original pack, and its movements were monitored by a GPS satellite collar. The collar worked correctly for 479 days. During that period the collar acquired a total of 1202 locations, indicating that the wolf had traveled at least 1590 km, with an average monthly distance (± SD) of 102 ± 40 km, exploring an overall area of about 270 km2. During the first 10 days after its release, the wolf remained in the area of its supposed native pack, whereas at about the age of 10 months the wolf began to make wide extraterritorial movements. The wolf’s last localization was acquired on 13th May 2018, about 17 months after its release, at a linear distance of about 65 km from the release site. This preliminary data showed that the wolf was alive and travelled long distances after its release, and demonstrates how a multidisciplinary management approach can support the recovery and successful release into nature of a rescued wild animal belonging to a flagship species with a notable ecological role, such as the Italian wolf.
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Aim Determining the drivers of movement of different life‐history stages is crucial for understanding age‐related changes in survival rates and, for marine top predators, the link between fisheries overlap and incidental mortality (bycatch), which is driving population declines in many taxa. Here, we combine individual tracking data and a movement model to investigate the environmental drivers and conservation implications of divergent movement patterns in juveniles (fledglings) and adults of a threatened seabird, the white‐chinned petrel (Procellaria aequinoctialis ). Location South‐west Atlantic Ocean. Methods We compare the spatial distributions and movement characteristics of juvenile, breeding and non‐breeding adult petrels, and apply a mechanistic movement model to investigate the extent to which chlorophyll a concentrations (a proxy for food resources) and ocean surface winds drive their divergent distribution patterns. We also consider the conservation implications by determining the relative overlap of each life‐history stage with fishing intensity and reported fishing effort (proxies for bycatch risk). Results Naïve individuals fledged with similar flight capabilities (based on distances travelled, flight speeds and track sinuosity) to adults but differed in their trajectories. Comparison of simulations from the mechanistic model with real tracks showed that juvenile movements are best predicted by prevailing wind patterns, whereas adults are attracted to food resources on the Patagonian Shelf. The juveniles initially dispersed to less productive oceanic waters than those used by adults, and overlapped less with fishing activity; however, as they moved westwards towards South America, bycatch risk increased substantially. Main conclusions The use of a mechanistic framework provided insights into the ontogeny of movement strategies within the context of learned versus innate behaviour and demonstrated that divergent movement patterns of adults and juveniles can have important implications for the conservation of threatened seabirds.
Natal dispersal (movement from the site of birth to the site of reproduction) is a pervasive but highly varied characteristic of life forms. Thus, understanding it in any species informs many aspects of biology, but studying it in most species is difficult. In the grey wolf Canis lupus, natal dispersal has been well studied. Maturing members of both sexes generally leave their natal packs, pair with opposite‐sex dispersers from other packs, near or far, select a territory, and produce their own offspring. However, three movement patterns of some natal‐dispersing wolves remain unexplained: 1) long‐distance dispersal when potential mates seem nearby, 2) round‐trip travels from their natal packs for varying periods and distances, also called extraterritorial movements, and often not resulting in pairing, and 3) coincidental dispersal by individual wolves from a given area in the same basic directions and over the same long distances. This perspective article documents and discusses these unexplained dispersal patterns, suggests possible explanations, and calls for additional research to understand them more clearly. Illustration depicts various movement patters of gray wolf Canis lupus natal dispersal. Although results and explanations are known for some patterns, this article discusses three dispersal patterns that lack an explanation.
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A LARGE, DARK WOLF poked his nose out of the pines in Yellowstone National Park as he thrust a broad foot deep into the snow and plowed ahead. Soon a second animal appeared, then another, and a fourth. A few minutes later, a pack of thirteen lanky wolves had filed out of the pines and onto the open hillside. Wolf packs are the main social units of a wolf population. As numbers of wolves in packs change, so too, then, does the wolf population (Rausch 1967). Trying to understand the factors and mechanisms that affect these changes is what the field of wolf population dynamics is all about. In this chapter, we will explore this topic using two main approaches: (1) meta-analysis using data from studies from many areas and periods, and (2) case histories of key long-term studies. The combination presents a good picture-a picture, however, that is still incomplete. We also caution that the data sets summarized in the analyses represent snapshots of wolf population dynamics under widely varying conditions and population trends, and that the figures used are usually composites or averages. Nevertheless, they should allow generalizations that provide important insight into wolf population dynamics.
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The relationship between the straight line distances (SLD), obtained from telemetry locations, and actual distances travelled by wolves Canis lupus (ADT), measured by snowtracking, was investigated in Bialowieza Primeval Forest, E Poland, in winter 1995/96. Radiolocations determined at 15-min time intervals approximated the ADT by wolves reasonably well. If wolves were relocated at 0.5- to 2-h intervals, SLD can be multiplied by a correction factor of 1.3 to obtain ADT. Within the range of SLD from 1 to 10 km, they could also be converted into ADT using a regression equation: ADT = 0134 + 1.19(SLD), with standard errors of prediction 0.13 to ± 0.3 km. The average travelling speed of wolves was 3.78 km/h (SD 1.23, range 1.6-6.1 km). Wolves walking the forest trails, roads and frozen rivers moved significantly faster than in the forest. Also, individuals travelling with other pack members moved faster than those walking singly.
THE FIRST REAL BEGINNING to our understanding of wolf social ecology came from wolf 2204 on 23 May 1972. State depredation control trapper Lawrence Waino, of Duluth, Minnesota, had caught this female wolf 112 km ( 67 mi) south of where L. D. Mech had radio-collared her in the Superior National Forest 2 years earlier. A young lone wolf, nomadic over 100 km2 (40 mi2) during the 9 months Mech had been able to keep track of her, she had then disappeared until Waino caught her. From her nipples it was apparent that she had just been nursing pups. "This was the puzzle piece I needed," stated Mech. "I had already radio-tracked lone wolves long distances, and I had observed pack members splitting off and dispersing. My hunch was that the next step was for loners to find a new area and a mate, settle down, produce pups, and start their own pack. Wolf 2204 had done just that."
We used VHF, GPS and satellite radiocollars to study details of long distance movements by four Minnesota wolves (Canis lupus). Number of locations during our tracking ranged from 14 to 274. Farthest distances reached ranged from 183–494 km, and minimum distances traveled (sums of line segments) ranged from 490–4251 km. Numbers of times wolves crossed state, provincial or interstate highways ranged from 1 to 215. All four of the wolves returned to or near their natal territories after up to 179 d and at least two left again.
We evaluated the accuracy and precision of tooth wear for aging gray wolves (Canis lupus) from Alaska, Minnesota, and Ontario based on 47 known-age or known-minimum-age skulls. Estimates of age using tooth wear and a commercial cementum annuli-aging service were useful for wolves up to 14 years old. The precision of estimates from cementum annuli was greater than estimates from tooth wear, but tooth wear estimates are more applicable in the field. We tended to overestimate age by 1-2 years and occasionally by 3 or 4 years. The commercial service aged young wolves with cementum annuli to within ±1 year of actual age, but under estimated ages of wolves ≥9 years old by 1-3 years. No differences were detected in tooth wear patterns for wild wolves from Alaska, Minnesota, and Ontario, nor between captive and wild wolves. Tooth wear was not appropriate for aging wolves with an underbite that prevented normal wear or severely broken and missing teeth.
1. I present a conceptual model that predicts the proximate and ultimate factors that determine whether a mammal species has the potential to be regulated by intrinsic (behavioural) or extrinsic factors. The model is based on three behavioural phenomena that purportedly regulate mammal populations: female territoriality, dispersal, and reproductive suppression. 2. The model predicts that intrinsic regulation can occur only in those species in which females are territorial, offspring-rearing space (or alpha status) is limited, and young females exhibit reproductive suppression. Female territoriality should occur in species that have altricial young and serve as a counter-strategy to infanticide from conspecific females. Female mammals that have precocial young or mobile altricial young that are not vulnerable to infanticide should not commit infanticide and should not be territorial. Thus, developmental state at birth would be an ultimate factor that determines whether offspring-rearing space potentially can be limited. 3. Dispersal should be density-independent in nonterritorial species and inversely density-dependent in territorial species and thus has limited potential to regulate population density in any species. 4. Behavioural reproductive suppression of young females is proposed as an adaptive mechanism to avoid inbreeding or to conserve reproductive effort in response to the threat of infanticide. 5. Intrinsic regulation should be most likely to occur in monogamous territorial species in which daughters grow up in the presence of male relatives (such as in canids and some primates). Polygynous species, in which females are territorial such as most rodents, have the potential for self-regulation; however, exposure to unrelated males and the fact that young females can breed on their mothers' territories usually preclude benavidural reproductive suppression. 6. Intrinsic regulation should not occur in species with precocial young, nonterritorial species, or in species in which daughters do not associate with male relatives, such as the ungulates, marine mammals, bats and marsupials. 7. The model predicts that female territoriality, the threat of infanticide, and the presence of male relatives in the natal home range are the proximate mechanisms for intrinsic population regulation in mammals. These factors apparently occur only under a limited set of conditions; therefore, most mammal populations are probably controlled by extrinsic factors. The model is presented with a series of a priori predictions from which hypotheses can be formulated and tested.
Dispersal is the movement an organism makes away from its point of origin to the place where it reproduces or would have reproduced if it had survived and found a mate. For the most part, the major dispersal movements are made by virgins about the time they attain puberty. Possession of the innate dispersal trait implies that such an animal is predisposed at birth to leave home at puberty and make one dispersal into surroundings beyond the confines of its parental home range. Such density-independent individuals have inherited an urge to leave home voluntarily. They often pass up available and suitable niches and venture into unfavorable habitats. Animals that make an innate dispersal movement are obsessed with a dispersal instinct. The "purposiveness" of the innate concept is not for the individual's welfare; rather, in spite of the high rate of mortality of innate dispersers, it has distinct survival value for the species. Innate dispersers are particularly important to a species because they, 1) increase the spread of new genes, 2) create wide outbreeding, 3) enable a species to spread its range rapidly as favorable habitats are created, 4) permit the species to have a discontinuous distribution, and 5) help the species quickly reinvade areas that may have been depopulated by catastrophes, such as floods, fires, or man's activities. Points that appear more or less to favor the existence of an innate dispersal concept include: 1) the distances of dispersal are, at least sometimes, significantly not random; 2) some introduced species spread their range too rapidly to be the result of population pressure factors; 3) reinvasion of a depopulated area does not commence at the edge and gradually overflow inward, but, instead, the density of the species builds up almost simultaneously over all of the area that is within the maximum limits of innate dispersals; 4) the rate at which innate dispersals are made seems to be density-independent; 5) the movements are made instinctively without any prior experience or instructors to imitate; 6) innate dispersers frequently cross or attempt to cross regions of unfavorable habitat, regardless of the availability of adjacent suitable habitats; and 7) the stimulus is of short duration, apparently being expressed only once, when the animal becomes sexually active for the first time. The presence of the environmental dispersal trait implies that the individual will remain where born or, by means of trial and error, eventually select a new home range usually within the confines of its parental home range. It will have a strong homing tendency and move only as far as forced by population pressure factors (intraspecific competition or density-dependent factors) such as parental ejection of young, voluntary avoidance of crowded areas, mating and territoriality, availability of food and homesites, or the presence of other organisms including predators. Minor shifts of homesites result in a dispersal, but these are all called environmental dispersals, even though a series of them by one individual might eventually result in a total dispersal distance that is quite extensive, even exceeding that of some innate dispersal movements. Environmental dispersal has only local significance, whereas innate dispersal is of geographic importance. To verify or refute the existence of an innate dispersal trait, the assistance of other investigators is urgently solicited, for findings of many workers will be necessary before we can thoroughly understand the dispersal behaviorism and dispersal pattern in different species.
Records are presented of three wolves that dispersed ≥200 km from northern Minnesota to the Minnesota-Wisconsin border, to southern Wisconsin, and to Michigan. This report documents that wolves cross major highways and other developed areas and that the recently recolonized wolf population in Wisconsin and Michigan could have originated from wolf populations in Minnesota other than those living along the Minnesota-Wisconsin border.