Division for Nature & Environment

Lars-Henrik Henriksson

Movement pattern of Moose (Alces

alces) in southwestern Sweden in relation to highway traffic intensity

Rörelsemönster av Älg (Alces alces) i sydvästra Sverige

relaterat till trafikintensitet vid motorväg

Biology D-level thesis

Date/Term: Supervisor: Per Widén Ass. Supervisor Mattias Olsson Examiner: Eva Bergman Serial Number: X-XX XX XX

Karlstads universitet 651 88 Karlstad Tfn 054-700 10 00 Fax 054-700 14 60

Abstract GPS telemetry is a method with good accuracy to determine animal movements in the terrain. It

is necessary to determine locations of free-ranging animals in order to understand movement

patterns and habitat use, and to understand the consequences of human impacts like highways.

This study aims to describe moose movement patterns and to evaluate the effect of highway

traffic intensity on moose movements across a highway.

Moose in Southwestern Sweden have different movement rates throughout the year. Increased

movement rate for females was observed during spring and summer. The breeding season (15

September -15 October) is the most important season for bulls. Our result shows that bulls

significantly increase their movement rate during the rut, compared to other times during the fall.

Movement rate increased twice compared with female movement rate during this period. No

difference was observed during the rutting period for females (15 September- 15 October)

compared with no rutting period during fall. During winter time, both sexes retain low

movements, mainly caused by energy saving actions. A distinct crepuscular rhythm was

exhibited during the summer and fall season, movements were more intense during dawn and

dusk hours. No distinct crepuscular rhythm was noticed during winter and spring seasons.

The traffic intensity at highway E6 in Southwestern Sweden increases during the morning hours

and reaches its maximum during midday. Moose in southwestern Sweden crossed highway E6

more often at night time than day time. Thus highway crossings by moose occurred at times of

peak moose movements, and traffic volume had lower importance.

Studier med hjälp av GPS är en noggrann metod för att bestämma djurs rörelser i terräng.

Det är av stor relevans att studera olika djurs rörelsemönster, habitat val och dess rörelser kring

motorvägar. Studien inriktar sig på rörelsemönster av älg kring motorväg relaterat till trafik

intensitet.

Resultatet visar att tjurar ökar sin rörelseaktivitet under brunst perioden jämfört med den övriga

höstperioden. Kor har ingen påvisad aktivitetsökning under brunstperioden. Under vintersäsong

minskar båda könen sin aktivitet, avgörande faktorer är att spara energi. Dygnsaktiviteten har

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tydliga aktivitetsökningar vid gryning och skymning under sommar och höst säsong.

Rörelsemönstret av älg varierar mycket genom alla årstider. Älgkor har en bevisligen högre

rörelseintensitet under vår och sommar. Brunst perioden (15 Sep-15 Okt) har tjurar högre

rörelseintensitet.

Trafik intensiteten på motorväg E6 har en kraftig ökning kring morgontimmarna och når sitt

maximum vid middagstid. Älgarna i studieområdet visar en tendens att passera motorvägen oftare

under natten jämfört med dagen. Studiens resultat visar att älgpassager sker oftast vid tidpunkter

då älgen har som högst rörelseaktivitet, trafikintensiteten har en mindre betydande roll.

Introduction During this century, roads and railroads have become important infrastructure networks,

contributing to fragmentation of wildlife habitats. Roads and highways also affects the

distribution of animal populations and limits the ability of individuals to disperse. The land area

is divided in smaller fragments and animal populations in smaller subpopulations, which may

cause higher vulnerability for catastrophic events and diseases (Jaeger and Fahrig 2004). Other

problems can be the management of small isolated populations, since the hunting may cause

demographic effects.

Population density is a principal factor that determines ungulate presence along roads, and

increased populations have been correlated with increased ungulate-vehicle collisions (Farell et

al. 2002). Other ungulate-vehicle collisions factors are seasonal behaviour, traffic speed and

traffic volume (Case 1978). In spring and early summer moose are vulnerable to collisions when

highway salts may attract them to roadsides. (Fraser 1979, Fraser and Thomas 1982). Roads have

great impact on local habitats and ungulates can be attracted to the roadside plantings (Case 1978,

Feldhamer et al. 1986, Warning et al. 1991).

Fencing has shown great effect reducing moose-vehicle accidents (Seiler 2003) and road fencing

has become a standard method in Swedish road management. Traffic accidents with moose has

increased rapidly since 1970 and today there are about 5000 police reported accidents each year

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in Sweden. Estimated annual number of collisions with ungulates in Europe is 507,000 (Groot-

Bruinderink and Hazebroek 1996). In Sweden, vehicle collisions with moose more often results

in human injury and death compared to other ungulate related collisions. Until 2001, more than

5000 km of highway have been fenced against large ungulates (O. Eriksson SNRA, pers. comm.).

The Swedish National Road Administration (SNRA) intends to increase fencing, when new

highways are upgraded and also to build wildlife passages to decrease the potential barrier effects

on moose and other animals. However, the main reason for ungulate mitigations is to increase

traffic safety (Seiler A, 2003).

Knowledge about the barrier effect of fenced roads on ungulates is limited. Experiments

indicated that exclusion fences decreased moose movements across roads by 90% (Skölving

1985, Nilsson 1987). Wheras a recent study in nothern Sweden (Seiler et al.2007) report a barrier

of 70%. Thus roadfencing considerably increases the barrier effect and fragmentation. In

addition, roadfencing may also result in local accumulations of moose along the fenced roads.

These accumulations along the road may cause forest damages (Ball and Dahlgren 2002).

The moose uses small units or patches for foraging and cover, in relation to their total home

range sizes. By definition, home range is the area an animal uses for its daily living (Burt, 1943;

Jewell, 1996). Moose movements strongly related to the forage-energy availability and forage-

amount and quality. Moose males use larger home-ranges than females (Cederlund and Sand

1994.). There are two explanations to this phenomenon. First the males have a bigger body size

and second, male moose have larger energetic requirements (Harestad and Bunnell, 1979).

Intersexual differences in range use and home-range size could be explained partly by the

dimorphism in body size. During the rut males are more mobile and large-bodied, large-antlered

mature males moves on larger areas than younger males.

In this study, I analyzed the daily and seasonal movement pattern of moose. The specific

objectives were to examine daily and seasonal movement patterns of moose in Southwestern

Sweden, including differnece between sexes, and to evaluate if movements across highway E6

are related mainly to traffic intensity or to natural movement patterns.

4

Material and methods The study was conducted in the south-western parts of Sweden north of Uddevalla. The study

area is divided by the highway E6, which stretches from Trelleborg in south to Svinesund, near

the border of Norway. The Swedish part of the highway E6 is under reconstruction and converted

from a two lane road without fence to a fenced four lane highway (M Olsson pers.com). The

eastern side of E6 was dominated by a homogenous coniferous forest (70% of the land area). The

western part consisted of a mixture of boreal forest (40%) and farmlands (30%). The proportion

of clear cuts was equal on both sides (10%). Norwegian spruce (Picea abies) and Scots pine

(Pinus sylvesteris) dominate the boreal forests, common hardwood species are alder (Alunus

glutinosa), Oak (Querqus robur), birch (Betula pendula, Betula pubescens) and mountain ash

(Sorbus aucuparia). Early successional stages after logging consisted of birch, aspen, willow and

other deciduous species. Logging has fragmented the forests into subunits of varying successional

stages.

5

- Forest

- Agriculture

- Urban areas

- Ocean & water

1 km

A

B

C

Figure 1. (M Olsson, review of studyarea). The highway E6 study area with wildlife passages (solid dots) from south to north: wildlife overpass completed June 2000, wildlife underpass completed June 2000, wildlife overpass completed June 2004. Highway reconstruction segments are indicated: A) finished in June 2000, B) finished in June 2004, C) finished in summer 2007. Open dots indicate capture locations of the 24 collared moose. Circles indicate mean annual home ranges of males and female moose calculated with minimum convex polygon method (MCP) and 95 % fixed-kernel method. From largest to smallest circle: MCP-male (48.6 km2), MCP-female (20.8 km2), 95 % fixed-kernel-male (19.1 km2) and 95 % fixed-kernel-female (12.1 km2).

During 2004, moose were counted by aerial survey. The result showed a difference in moose

density between the sides. East side of the highway the moose population was estimated to 9

moose/1000 ha and on the west side 5 moose/1000 ha (Svensk Naturförvaltning 2004). 30% of

the moose were bulls, 40% cows and 30% calves. The birth rate was estimated to be 0.9

calve/cow (M Olsson pers.com).

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In total, 24 individuals were GPS-collared, 6 adult bulls, 12 adult cows, 2 subadult cows and 4

subadult bulls from year 2002 to 2005. Moose positions were collected at 2-hour intervals and the

number of moose positions totaled 76170 during the study period of 46 months. Each moose was

collared up to 22 months before collars fell off due to preprogrammed drop-off mechanisms. Data

for individual animals were divided into four seasons based on thermal conditions (SMHI,

National database) and moose biology (Pulliainen, 1974; Cederlund and Okarma, 1988); spring:

16 March – 15 May, summer: 16 May – 30 August, fall: 1 september – 30 november and winter:

1 December – 15 March. All data were sorted based on id, date and time. Moose positions were

collected from 0000 to 2400, with time intervals of 2 hour. Data were managed in Microsoft

Excel. Rut period was selected for both gender during rut period from 15 September to 15

October. Movements are measured in meters/ time period.

We used ArcView GIS, version 3.3 (Environmental systems research institute, Redlands,

California, USA) for all GIS (Geographic system and Science)analyses. A road-grid was

imported to the GIS analyses including highway E6 and other local roads to be able to manage

and count the moose crossings. We defined a movement across the highway by two subsequent

locations documented on each side of the highway. Concern was taken to GPS errors and their

influence on location accuracy.

Differences in movement rates between males and females during all seasons were analyzed

using Wilcoxon Matched pair tests. Differences in movement rates during rut were analyzed

using Mann Whitney U-test.

We used multiple linear regressions to identify the relative importance of variations in traffic

volume on highway E6 and daily movement patterns of moose on the timing of overpass use by

moose (Statistica 7.1, StatSoft, Inc.). Specifically, the average distance moved by GPS-collared

moose during each of 12 2-hour time periods and mean traffic volumes during crossing hours

were regressed on the total number of overpass crossing by moose during respective 2-hour time

periods. Datasets were divided by seasons since the movement pattern differed throughout the

year. Analyses were considered significant at (P<0, 05).

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Results Seasonal movement pattern

spring

0 50

100 150 200 250 300 350 400

00:00 02:00 04:00 06:00 08:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00

Movement m/ 2h period

female male

Figure 2. Mean movement budgets of 2hour periods during spring season for males and females. Movements are

measured in meters/ 2-hour periods.

The daily activity pattern in spring (figure 2), shows two peaks, one at 0600 h and one at 2000-

2200 h. The evening peak was more pronounced than the morning peak. Differences in

movement rates between males and females was observed during spring (z=3.06; n=12;

p=0.002). Males have higher movement rate than females during all time periods in spring

season.

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summer

0 50

100 150 200 250 300 350 400

00:00 02:00 04:00 06:00 08:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00

Movement m/ 2h period

female male

Figure 3. Mean movement budgets of different time periods during summer season for males and females.

Movements are measured in meters/ 2-hour periods.

The daily activity pattern in summer season 16 may – 30 August (Figure 3), shows two peaks,

r

ifferences in movement rates between males and females was observed during summer (z=3.06;

one at 0400 h and one at 2200 h. The results indicate that moose are mainly active at crepuscula

hours and also during the short and light summer nights.

D

n=12; p=0,002). At peaks, males moved more intensively than females, compared to other times.

Minimum activity occurred from 0800-1800 h, with all male values under mean movement 100

meters/time period and female values under 75 meters/ time period. Female moose increased

their activity during night hours compared to daytime hours. Especially high movements-rates

were observed during 2200-0000 with a mean movement rate of 225 meters/time period.

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fall

0 50

100 150 200 250 300 350 400

00:00 02:00 04:00 06:00 08:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00

Movement m/ 2h period

female male

Figure 4. Mean movement budgets of different time periods during fall for males and females. Movements are

measured in meters/ 2-hour periods.

A distinct crepuscular rhythm was exhibited during the fall period (Figure 4). During fall males

almost have double activity rate than females and increases their movements during all time

periods of the day.

A difference between gender was observed during fall season (z=3.06; n=12; p=0,002).

Moose females use night hours, mean movement 100 meters /time period. A peak is reached

during 1800-2000 h, when females increases their movements, mean movement 150

meters/timeperiod.

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winter

0 50

100 150 200 250 300 350 400

00:00 02:00 04:00 06:00 08:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00

Movement m/ 2h period

female male

Figure 5. Mean movement budgets of different time periods during winter season for males and females. Movements

are measured in meters/ 2-hour periods.

During the winter period, 1 December – 15 march, moose seams to reduce their movements.

During time period 1600-1800 h moose movements increase and reach a peak, wich is the

normaly crepuscular time. During 0400-0800 moose movements seems to be short and mean

movement averages 50 meters/timeperiod. No differences between gender was observed during

winter (z=0.71; n=12; p=0,48).

Moose females have low winter movements during almost all time periods, except 1800-2000

when females increase their activity and reach a peak (Figure 5).

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Rutting movements of moose

females was observed during the rutting period (z=-4.17; n1=18; A difference between males and

n2=9; p<0,001). The results show that bulls in average moved 250 meters during each 2-hours

period, as to be compared to the females lesser movement rate of 100 m (Figure 6).

m f0

50

100

150

200

250

300

mov

emen

t

Figure 6. Differences between males and females during the rut.

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A difference was observed during the rutting period for males compared with no rutting period

during fall (z=-3.04; n1=9; n2=9; p=0,0012) (Figure 7). During the rutting period males increase

their movement rate from 150 m to 250 m per time interval. Results indicate a 60% higher male

mobility during rut season, compared with other times during the fall.

yes norut

0

40

80

120

160

200

240

280

320

mov

emen

t

Fig 7. Rutting behavior males.

No difference was observed during the rutting period fore females compared with other times

during the fall (z=-0.27; n1=18; n2=19; p=0,80) (Figure 8). The mean movement during the rut

season indicates 110 m movement during the 2-hour intervals; same as other times during fall.

yes norut

0

40

80

120

160

200

240

280

320

mov

emen

t

Fig 8. Rutting behavior females.

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Moose highway crossings in relation to traffic intensity

The traffic intensity increases during the morning hour and reaches its maximum during midday.

During the afternoon the traffic decreases, but is relative high at hour 2000-2200. During this

timeperiod moose have their highest highway crossings rates. Traffic intensity peaked during

summer period and decreased during autumn. Moose crossed the highway more often at night

time than during day time (Figure 9).

0,0 2,0 4,0 6,0 8,0

10,0 12,0 14,0 16,0 18,0 20,0

0 2 4 6 8 10 12 14 16 18 20 22Time of the day

Percent

Highway crossings Traffic intensity

Figure 9. The percent of highway crossings in relation to mean traffic intensity.

9 (7 males and 2 females) of the 24 collared moose have crossed the highway during the study

period. In total, 155 passages have been detected. Moose number 1 has crossed the highway at 95

(61 % of all crossings) occasions during the period 2002-2004. Moose female numbers 11 with

one calf has crossed the highway at two times during the winter season 2004. Additional

crossings are made by solitary females without calf.

We used multiple linear regression to test for movement rate and traffic volume effects on moose

crossing frequencies during all seasons. A model containing traffic volume and hourly movement

data for moose explained 53% of the variation in the timing of highway crossings by moose

(F(2,29)=16.44; r2=0.53; p<0.001). The number of moose crossings was negatively correlated to

mean traffic volume, but not significant at the p=0.05 level (Beta=-0.21+0.14 SE; n=32; p=0.16).

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The number of moose crossings was positively correlated with hourly movement rate for moose

(Beta=0.61+0.14 SE; n=32; p<0.001). Thus highway crossings by moose occurred at times of

peak moose movements, and traffic volume had lower importance.

Discussion The adaptive value of activity patterns has been well recognized (Aschoff, 1966). The daily

activity of an animal is a result from a complex trade-off between optimal foraging time, social

ecology, competitive and predator-prey interactions, environmental and physiological constraints,

and energy requirements. Other studies show that ungulates have especially daily and seasonally

routines. The Elk (Cervus elaphus) use their environment for daily activities like foraging, resting

and moving (Green 1990).

The double-peaked 24-hour rhythm of moose activity is common for a great number of species

(Aschoff 1966), and are found in other cervids (Cederlund 1989). Moose change their behavior

between periods of browsing and resting/ruminating. The moose individuals spend time at their

requirements both at day and night time but mostly at night time. The daily activity pattern is

synchronized between dawn and dusk (Miquelle 1989).

Behavioral rhythms play a major role in ecological relations of a species and form a part of its

evolutionary adaptation. Other important adaptations are related to seasonal and diurnal

variations in the environment (Cloudsley-Thompson 1961; Risenhoover 1986). Knowledge of

daily and seasonal activity patterns and time budgets are essential for evaluating foraging

strategies for herbivores (Cederlund et al. 1989). This provides us with useful information about

how ungulates spend their time and their living. Theoretically, allocation of time is usually

considered as an optimization process, whereby the time spent on an activity should be increased

as long as the resulting gain in time spent per unit of food exceeds the costs (MacArthur and

Pianka 1966).

Comparisons between the sexes and between seasons within a population, and between

populations, can clarify how animal’s strategies adapt it to different environmental conditions

(Zhang 2000). Ruminating animals have a serial foraging-resting-foraging activity pattern during

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the 24 h period. Most ungulates feed more in the early morning and late afternoon than other

times of day.

Interactions among the various elements of weather and other environmental and biological

conditions including day length, food supply, biological clock and reproductive conditions

controls the activity pattern of moose, (Garshelis et al. 1980).

My results indicated that moose responded to increasing light intensity during spring and

summer. Long daylight hours result in high degree of photosynthetic activity in plants.

There is evidence that females maintain large home range sizes in spring (including parturition in

late May to early June) (Markgren, 1969). High movements in spring season indicates high rate

of food patch allocating, which is distributed over the home-range area (Clutton-Brock et al.

1982). Plants during spring and early summer have low rates of fibrous tissue, which has low

grade of digestibility which are important fore lactating females (Bliss 1962).

A reduction in winter activity conserves energy and minimizes the negative energy balance

(Moen 1976). Usually ungulates take in less energy than they need during the winter and

consequently loose weight. Therefore, body reserves accumulated prior to winter are an

important aspect of nutritional strategy. One important factor that might impede movements is

snow depth (Phillips et al. 1973). Movements in snow accumulations results in increasing costs

of locomotion. Obviously moose try to conserve energy during wintertime.

Male moose have smaller home-ranges during winter compared to other seasons (Olsson et al.

unpublished). During winter moose females have lower metabolic needs, and choose areas with

great food availability (Houston 1968, Loisa and Pullianen 1968, Philips et al. 1973). Another

strategy to save energy might be that moose in wintertime keep close to the forest edges to avoid

wind exposure. Moose only walk as far as necessary into clear-cut areas to find food (Hamilton et

al. 1980).

My reults of spring period 16 March – 15 May, the local whether conditions may fluctuate and

the presence of snow conditions may decrease moose movements. However, snow cover during

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the study never exceeded the amount that would impede moose movements (Cederlund and

Okarma 1988). Later in the spring when temperature, light and food availability increases, moose have the

opportunity to compensate for their weight loss during the winter period. During spring and

summer, increased movement rates during crepuscular hours appear to be the main reason why

moose tend to increase their overall movements rates. In spring dispersing moose individuals

establish new home ranges. One possible reason is inbreeding avoidance, which is effectively

accomplished by female dispersal. Other reasons why moose females establish new home range

areas is related to food availability. Moose females separate with their offspring during late

spring and dispersing yearlings often establish their home range close to their mothers (Cormack

Gates et al. 2005). Among adult females, the primary determinant of home range establishement

appears to be access to nutritious food ( Mace et al. 1984). This may explain the spring movemts

in my study.

Movement patterns can be affected by several factors. Fragmented landscapes by clear-cuts,

which means shorter distances to adequate food patch, would reduce movement intensity

(Cederlund and Sand.1994) and increase site fidelity. American studies on moose indicate

generally larger home range sizes during summer season. Similar-sized areas were occupied by

cows with calves and those with out calves (Phillips et al. 1973).

There is a significant difference in movements among bulls and cows during the rut. My results

show that bulls increase their movements during the rut whereas cows do not. Strategies between

genders clearly differ. Bulls produce maximum number of progeny, and need to optimize their

sexual activities so they are available fore meeting a potential mating female. At same time bulls

need to maintain a good physical condition so that they can survive the winter. Bulls can loose up

to 20 % weight during the rut, females never lose more than 5 % (Lent 1974). Forage intake by

North American males decreased dramatically during the rut and especially larger bulls decreased

their foraging intake during the mid September (Lent 1974). Large-bodied, mature male moose

invest more than younger, smaller males in reproductive activities as in other male cervids

(Cederlund G. Sand H. 1994).Competitive ability and reproduction success is strong related to

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body size in male ungulates, younger bulls need to optimize their breeding possibilities by

foraging during the rut (Clutton-Brock et al. 1982).

My results show greater mobility by males during the rut period in all daytime hours.

Studies have shown that bulls decreases their forage intake during the rut, and for approximately

2 week in mid-September larger, older bulls almost stop feeding (Dale G and Miquelle, 1990).

Bulls obviously concentrate their movements and time budget to find a mating female. During the

breeding season bulls have their highest energy costs compared with all seasons (Maher and

Byers 1987). My studie indicates that mature bulls spend more energy costs on movements

during the rut, because of optimizing their breeding possibilities.

My data indicates that different moose individuals behave different in relation to highways.

During the study period, I followed an adult male that crossed the highway at 95 occasions. This

individual crossed the highway even during the holiday-midsummer traffic, with a traffic

intensity of 1400 cars/hour. Probably, this male learned how to cross the highway without getting

hit.

Movement studies on grizzly bears close to highways described that bears crossed more often at

night than other normal periods of activity. Grizzly bears were apparently choosing to cross when

they were less likely to encounter highway traffic. Adult female bears were most sensitive to

traffic, especially when accompanied by cubs, males appears to be least sensitive. (Waller et al.

2005).

My studies shows that moose crossed the highway more often at night, compared to other time

periods. The results indicate that moose prefer to cross the highway between 22:00 to 04:00.

The most dangerous time for moose to cross this highway might be during the time period 20:00

to 22:00; here traffic volumes are relatively high and moose crossings common. We have noticed

a tendency that moose are most active during time period 22:00 to 04:00. This result may be the

best indicator explaining why moose crossings are most abundant during the night hours. The

high frequency of highway crossings during periods with higher movements might be explained

by seasonal movements. Most crossings are detected during summer and fall, which coincide

18

with larger home-ranges during these periods. However, other factors like topographic

distribution, moose density, different age/sex classes, food-patch distribution or other influences

like habitat type (Chruszcz et al. 2003) have major effects. Moose females with calves prefer to

stay in habitats with less human disturbance. Moose movements across highly trafficked

highways appear to be most influenced of moose movements, and less to traffic volumes. Sounds

from vehicles and other disturbing factors close to roadsides do not prevent moose movements

especially during the crepuscular hours and night hours when moose movements generally is

high.

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Conclusion Males exhibit greater movement rates compared to females during all seasons, except winter.

Both males and females had distinct peaks of activity at crepuscular hours during spring, summer

and fall, but retained low movement rates through all hours during winter. Male movement rates

increased considerably during the peak period of rut, females had equal movement rates as other

time periods during the fall. Moose movements across highly trafficked highways appear to be

most influenced of moose movements, and less to traffic volumes. This indicates that the greatest

risk for collisions appears to be at times when moose move most, during crepuscular hours. This

theory should be evaluated further and compared to the timing of moose collisions during

different seasons.

Acknowledgments I would like to thank D-student Anders Olofsson and other friends for all willingness to help, and

support me during the study period. Special thanks to my assistant supervisor Matthias Olsson,

PhD student at Karlstad university, who have helped me a lot during this study. Last but not least

thanks to Per Widén, the supervisor for this project

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