788

Into thin air: vertical migration, body condition,

and quality of terrestrial habitats of alpine
common toads, Bufo bufo
M. Sztatecsny and R. Schabetsberger

Abstract: Since the end of the 1980s there has been increasing evidence of worldwide amphibian declines in high-
altitude regions. Moreover, amphibian conservation has so far mostly neglected the terrestrial habitat, which is essential
for effective protection measures. We determined the location and quality of terrestrial habitats of common toads (Bufo
bufo L., 1758) in a naturally fragmented alpine environment (Schlumsee, 1105 m a.s.l., Northern Calcareous Alps, Aus-
tria) characterized by a high diversity of available microhabitats. By radio-tracking 18 individuals during their post-
spawning migration, we located terrestrial habitats 130–1000 m horizontally away from and 85–390 m above the breed-
ing site. This is the first study to show vertical migration over several hundred metres in adult amphibians. Both adult
and juvenile toads completed the migration within 2–7 weeks and, on their way to the summer habitat, climbed 45°
scree slopes and ascended cliffs with slopes of up to 65°. Body condition indices were highest for individuals originat-
ing from the terrestrial summer habitat at the highest elevation, which is characterized by the highest vegetation diver-
sity, a high abundance of food, and the highest insolation, probably allowing the toads to extend their activity period
for food intake. Our study suggests that under demanding climatic conditions it can pay off for toads to undertake
costly migrations to reach high-quality habitats. For amphibian conservation, high-value habitat patches need to be lo-
cated and investigated to increase the effectiveness of protection measures.
Résumé : Depuis la fin des années 1980, il y a de plus en plus d’indications d’un déclin des amphibiens dans les ré-
gions de haute altitude à l’échelle globale. De plus, les mesures de conservation des amphibiens ont jusqu’à maintenant
pour la plupart négligé les habitats terrestres qui sont essentiels pour une protection efficace. Nous avons déterminé la
répartition et la qualité des habitats terrestres du crapaud commun (Bufo bufo L., 1758) dans un environnement alpin
naturellement fragmenté (Schlumsee, 1105 m au-dessus du niveau de la mer, Alpes Calcaires du Nord, Autriche) et ca-
ractérisé par une forte diversité d’habitats disponibles. En suivant par radiotélémétrie 18 individus lors de leur migra-
tion post-reproductive, nous avons trouvé leurs habitats terrestres situés à 130–1000 m horizontalement et 85–390 m
plus en altitude du site de reproduction. C’est la première fois qu’une étude montre une migration verticale de plu-
sieurs centaines de mètres chez un amphibien. Tant les adultes que les jeunes complètent cette migration en 2–7 semai-
nes et, dans leur déplacement vers leur habitat d’été, ils traversent des éboulis à inclinaison de 45° et grimpent des
falaises à pente pouvant atteindre 65°. Les indices de condition corporelle sont maximaux chez les individus récoltés
dans les habitats d’été de plus haute altitude qui sont caractérisés par la diversité de végétation la plus grande, une
forte abondance de nourriture et l’ensoleillement le plus important, ce qui permet probablement aux crapauds de pro-
longer leur période d’activité alimentaire. Notre étude indique que, dans des conditions climatiques exigeantes, il peut
être rentable pour les crapauds d’entreprendre des migrations coûteuses afin d’atteindre des habitats de haute qualité. Il
est donc important d’identifier et d’étudier ces parcelles d’habitat de grande valeur afin d’améliorer l’efficacité des me-
sures de protection pour la conservation des amphibiens.
[Traduit par la Rédaction] Sztatecsny and Schabetsberger 796

Introduction
Habitat loss and fragmentation due to anthropogenic land-
scape changes have been identified as key causes of local and
worldwide population declines (Barbault and Sastrapradja
1995), with amphibians being no exception (Alford and
Richards 1999; Stuart et al. 2004). Pond-breeding amphibi-
ans may be especially vulnerable to habitat degradation, as
they depend on more than one habitat type to complete their
life cycle (Wilbur 1980). Eggs and larvae develop aquat-
ically, whereas juveniles and adults lead a primarily terres-
trial life. Adults return to the aquatic breeding site only for
reproduction, and seasonal migrations between spatially sep-
arated habitats occur naturally in their life history (Sinsch
1990). The survival of juvenile and adult life stages has
strong effects on amphibian population persistence (Biek et

Received 19 November 2004. Accepted 25 May 2005. Published on the NRC Research Press Web site at http://cjz.nrc.ca on
21 July 2005.
M. Sztatecsny.1 Institute of Zoology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria.
R. Schabetsberger. Department of Organismic Biology, University of Salzburg, Hellbrunnerstrasse 34, 5020 Salzburg, Austria.
1
Corresponding author (e-mail: marc.sztatecsny@univie.ac.at).
Can. J. Zool. 83: 788–796 (2005) doi: 10.1139/Z05-071 © 2005 NRC Canada
Sztatecsny and Schabetsberger
al. 2002; Vonesh and De la Cruz 2002); knowledge of the
migration behaviour and terrestrial habitat requirements of
these stages is therefore highly relevant to amphibian con-
servation (Semlitsch 2000, 2002).
Some of the most alarming amphibian declines have been
reported from high-altitude sites (Barinaga 1990; Houlahan
et al. 2000). Alpine amphibian populations may differ mark-
edly from their lowland counterparts in life-history parame-
ters such as growth and longevity (Hemelaar 1988;
Schabetsberger and Goldschmid 1994; Miaud et al. 1999),
and little is known about their terrestrial ecology (but see
Matthews and Pope 1999; Pilliod et al. 2002; Schabetsberger
et al. 2004). Compared with lowlands, mountain terrain is
topographically diverse, highly fragmented, and only moder-
ately disturbed by human land use. The high geodiversity
and the compression of climatic zones along an elevational
gradient create a multitude of microhabitats across small
geographical scales (Körner 2002). Finding a high-quality
habitat might also be more crucial for amphibians at high el-
evations, as time for energy accumulation in order to survive
the long winter is limited.
The common toad (Bufo bufo L., 1758) is an explosive
breeder, with up to thousands of individuals gathering annu-
ally for only a few days to weeks at a breeding pond
(Heusser 1958, 1960). Its seasonal migratory behaviour in-
cludes a pre-spawning migration to the breeding site in early
spring, a post-spawning migration to the terrestrial summer
habitats, and an autumn migration to the hibernation sites
(Heusser 1968). The preferred terrestrial habitats are typi-
cally of high structural diversity and 55–1600 m away from
the breeding pond (Sinsch 1988a; Kyek et al. 1997); migra-
tion distances of B. bufo are greater than those known for
many other amphibians (Semlitsch and Bodie 2003).
It is unknown why common toads cover long distances
and what renders such demanding migrations advantageous.
However, such knowledge would help maximize conserva-
tion efforts by identifying those fragments of terrestrial habi-
tat that are most profitable to the population (Ovaskainen
and Hanski 2003). In demanding environments such as high
altitudes, effects of differences in habitat quality should be
pronounced and hence easy to detect. We therefore chose an
alpine study site where the diversity of available habitats is
greater than in lowlands and where the mountain terrain ad-
ditionally increases landscape resistance for migrating ani-
mals. The aims of this study were to (i) investigate, by
means of radiotelemetry, the abilities of common toads to
migrate in alpine terrain including vertical drop-offs and
cliffs, (ii) describe the migration behaviour of common toads
in a naturally fragmented environment with little human dis-
turbance, and (iii) locate and characterize the terrestrial habi-
tats of an alpine population of common toads. Based on the
knowledge gained from radio-tracking, we also investigated
(iv) the body conditions of toads in different terrestrial habi-
tats and (v) whether food availability drives migrations in al-
pine environments.
Materials and methods
Study area
The study was conducted around a temporary lake
(Schlumsee) in the Northern Calcareous Alps of Austria
789
(47°33′N, 13°06′E, 1105 m a.s.l.). Local climatic conditions
are cold and moist because of constant up-sloping effects
along the Alps’ northern edge (Cabela et al. 2001).
Schlumsee is situated in a karst depression (polje) with a di-
ameter of ca. 500 m and rims 100–800 m high, sealed with
fine clastic material consisting of ground moraine during the
last ice age (Klappacher and Knapczyk 1979). In early
spring, the bottom of this basin fills with meltwater from the
surrounding slopes and a headwater creek to form the lake,
which has a maximum area of about 0.02 km2 and a depth of
approximately 3 m. The water level gradually drops from
mid-May onwards, and the lake dries up by mid to late July,
forcing the amphibians to complete their aquatic develop-
ment within 8–10 weeks.
After drying out, the lake bottom turns into a meadow
partly covered with horsetail (Equisetum palustre L.). To the
southeast, the meadow gives way to a Norway spruce forest
(Picea abies (L.) Karst), tall forbs (mainly Cicerbita alpina
(L.) Wallr. and Veratrum album L.), and a mixed deciduous
forest (mainly Fagus sylvatica L. and Acer pseudoplatan-
us L.) (Fig. 1). In all other directions, scree slopes (25°–45°)
interspersed with large boulders and topped by rock faces
form the lakefront. The scree and boulders are only scarcely
covered with spruce and isolated larches (Larix decidua P.
Mill.). Via a pass at 1200 m a.s.l. at the northern edge of the
Schlumsee basin, a rift valley slopes to a wide valley at
about 500 m a.s.l.; in all other directions the terrain rises to
an extensive karst plateau (1500–2200 m a.s.l.).
Migration
From a pilot study conducted during May and July 2001,
we knew that the bulk of the toad population followed mi-
gration routes north and east of Schlumsee. We therefore
concentrated on these directions during radio-tracking in
2002. Toads used for radio-tracking were captured 0–200 m
from the lake from 18 to 30 May 2002. We implanted the
transmitters in the body cavity as described by Sinsch
(1988b). After measuring snout–urostyle length (SUL) to the
nearest millimetre using callipers and weighing the toads to
the nearest 0.1 g using an electronic mini-scale, we anaes-
thetized them in a 0.03% solution of MS 222 for transmitter
implantation. Because the toads recovered only several hours
after being anaesthetized, we kept them in plastic containers
partly filled with leaf litter overnight. We used two types of
transmitters (Holohil Systems Ltd., Carp, Ontario): 3 BD-
2GTHX (15 mm × 8 mm × 5 mm, 1.9 g) with an external
helix antenna, and 15 BD-2A with an internal helix antenna.
The BD-2A transmitters had been used before in an unrelated
study (Schabetsberger et al. 2004); we equipped them with
new button-cell batteries without removing the original batter-
ies and resealed them with body-compatible wax, increasing
the size of the transmitters from 8 mm × 7 mm × 5 mm to
20 mm × 8 mm × 6 mm and the mass from 0.8 g to 1.5 g.
A Televilt RX-98 tracking receiver (TVP Positioning,
Sweden; frequency range 150.000–151.999 MHz) and a
hand-held ZX-Yagi antenna were used to detect transmitter
signals. The signal detection range varied between transmit-
ter types, ranging from 15 m (BD-2A) to 300 m (BD-
2GTHX), and was lowest for toads hidden under boulders.
We could locate individuals with an accuracy of ca. 50 cm
independently of the transmitter type.
© 2005 NRC Canada
790
Fig. 1. Map showing the distribution of vegetation types in the
study area, the locations of sampling plots for pitfall traps, and
the locations of temperature data loggers.
We released the toads at the place of capture the morning
after the implantation procedure and radio-tracked them at
0700, 1400, and 2000 h from 18 May to 4 June 2002. There-
after, we visited Schlumsee for single detection efforts on 29
June, 20 July, and 20 August 2002. On 4 June, we removed
all BD-2A transmitters from toads that had not started their
migration, as the signal detection range of transmitters was
low. All other transmitters were removed when toads had
reached their summer habitats or, at the latest, on 20 August.
Toads were considered to have reached their summer habi-
tats when they had not moved more than 20 m within at least
3 weeks. Transmitter removal took the same form as the im-
planting procedure, and the toads were released at the points
of their final capture.
For distances of up to 300 m from the lake, we located
toads using a compass and a 20-m measuring tape; beyond
this distance, we used a global positioning system (GPS 12,
Garmin International Inc.), ignoring movements of less than
1 m. The steepness of slopes along migration routes within
300 m of the lake was measured every 10 m with a pendu-
lum and a goniometer attached to a pole. Beyond this dis-
tance we calculated total elevation gain from the GPS
positions. All capture locations were plotted on an aerial
Can. J. Zool. Vol. 83, 2005
photograph using geographic information system software
(ArcView® 3.2, Environmental Systems Research Institute,
Redlands, California). By integrating the aerial photograph
and altitudinal data from the Austrian Federal Office of Me-
trology and Surveying, we created a three-dimensional im-
age using ER Mapper 6.3 (ER Mapper, Australia).
Body condition index
To calculate the body condition index of toads from dif-
ferent habitats, we measured their SUL and body mass as
described before. We caught 70 females and 67 males be-
tween 16 and 18 May 2002, and 57 females and 176 males
between 12 and 13 May 2003, during their migration to the
summer habitats. To rule out effects of differences in their
reproductive cycle, gut contents, or degree of hydration, we
measured all toads immediately after spawning, a time when
they do not feed (Jørgensen 1992).
To compare the body condition of toads from habitats
north, east, and south of Schlumsee, we used the relative
mass (Wr) condition index, following Pope and Matthews
(2002). The Wr index was calculated separately for females
and males in 2002 and 2003 with the equation
W
Wr = 100
Ws
where W is the body mass of toads and Ws is the expected
length-specific mass predicted from a linear regression of
mass (log10W) on SUL (log10SUL). Ws was corrected for
bias introduced during conversion from logarithmic to arith-
metic units by Baskerville’s multiplicative bias correction,
exp(S⑀2/2), where S⑀2 is the error mean square of the linear
regression (Baskerville 1972).
Characterization of terrestrial habitats
To evaluate thermal conditions during the toads’ hiberna-
tion period inside and outside the Schlumsee basin, we re-
corded air temperature from 10 September 2002 to 10 May
2003 at the level of the lake and on the slopes north of the
lake (1250 m a.s.l.) (Fig. 1). We used Tinytag® data loggers
(Gemini Dataloggers (UK) Ltd.) attached to trees 2 m above
ground and recorded temperature at 4-h intervals starting at
0200.
Mean daily solar insolation was modelled following Rich et
al. (1995) using the software Solarflux for ArcView. The pro-
gram accounts for latitude and elevation, surface orientation,
shadows cast by surrounding topography, shifts in solar angle,
and atmospheric attenuation. The required digital elevation
model was based on grid data obtained from the Austrian
Federal Office of Metrology and Surveying. Mean daily in-
solation was calculated hourly, starting at 0000 h, on the
15th and the last day of each month from 28 February to 15
June. This time frame included the earliest recorded sight-
ings of B. bufo above 1000 m a.s.l. in Austria (Cabela et al.
2001) and the day closest to solstice, when the radiation
reaches its maximum. Data on vegetation were provided by
the Provincial Government of Salzburg and amended by a
detailed survey during field work in 2003 by M.S.
Food availability
To measure food availability, we quantified the abundance
and composition of potential prey organisms in survey plots
© 2005 NRC Canada
Sztatecsny and Schabetsberger
Fig. 2. Topographical relief of the Schlumsee basin showing the routes of radio-tracked common toads (Bufo bufo) in 2002. Contour
intervals of 20 m (dashed black lines) and 100 m (solid black lines) are shown.
inside and outside the Schlumsee basin. The plots measured
100 m × 200 m and were situated south of the lake and in
the terrestrial habitats north and east of Schlumsee as lo-
cated by radio-tracking (Fig. 1). Since we did not radio-track
animals migrating along the southern route, we were unable
to locate their exact terrestrial summer habitats; therefore,
we refrained from characterizing prey availability south of
Schlumsee.
Ten pitfall traps (glass jars 10 cm deep and 7.5 cm in di-
ameter, containing approximately 100 mL of ethylene gly-
col) for food items were randomly placed on a 10 m × 10 m
grid in each of the plots. Each trap was covered with a trans-
lucent plastic plate (10 cm × 10 cm), leaving a 2-cm opening
to the jar. The traps stayed exposed for 2 weeks, from 23
June to 7 July 2003. Captured organisms were assigned to
17 different categories, enumerated, blotted on absorbent pa-
per to remove excess moisture, and weighed to the nearest
milligram.
Due to the difficulties of finding toads in their extensive
terrestrial habitats, collecting data on gut contents was beyond
the scope of our study. However, we unintentionally captured
two male and six juvenile toads in the pitfall traps, which we
dissected. After removing the entire gut, we examined its con-
tents and identified prey items following Wheater (1986).
Results
Migration
We radio-tagged 14 female (76–90 mm SUL, 49–109 g)
and 4 male (65–71 mm SUL, 33–34 g) toads and followed
them for up to 94 days. The relative mass of transmitters
791
ranged from 1.4%–3.3% of body mass for females and
4.4%–4.5% of body mass for males. Six radio-tagged toads
migrated to the northern summer habitats and two toads mi-
grated to the habitats east of Schlumsee (Fig. 2). Total hori-
zontal distances covered by the toads were 130–500 m and
900–1000 m to the north and east, respectively; the maxi-
mum total elevation gain was 200 m to the north and 390 m
to the east (Fig. 2). On the eastern route, a female was local-
ized on a grassy ledge 5 m up a 65° rock face. Migration
lasted for 2–4 weeks to the north and 7 weeks to the east.
One female and one male, released 75 m and 150 m above
the elevation of Schlumsee on the northern and eastern up-
per rims of the basin, respectively, were captured during
their pre-spawning migration (Fig. 2). Both reached
Schlumsee within 24 h, travelling along direct routes with-
out avoiding vertical cliffs up to 2 m high. The median
steepness of slopes along their routes was 32.5° (min. 24°,
max. 90°) and 34.5° (min. 22°, max. 49°), respectively.
Transmitter signals from both toads were lost before they
started their migration to the summer habitats. Two females
and one male released at the lakefront did not leave the
Schlumsee basin, and transmitters had to be removed from
five toads before they started migration to avoid signal loss
(therefore not shown in Fig. 2). All radio-tracked toads
showed latent periods of 1–82 days before starting migra-
tion, during which they remained stationary.
Body condition
In 2002, female and male toads east of the basin had the
highest body condition (Fig. 3), with significant differences
between summer habitats for both sexes (Kruskal–Wallis
© 2005 NRC Canada
792
Fig. 3. Relative mass (Wr) condition index of common toads from the three terrestrial habitats north, east, and south of Schlumsee in
2002 and 2003. Regression equations give the overall relationship between snout–urostyle length (SUL) and body mass. The numbers
refer to sample sizes; the asterisks indicate statistically significant differences between the eastern habitat and the other two habitats.
females
160
Post-spawning Wr 2002
140
27
120
100
80
logWs= –1.45+1.647logSUL
females
Post-spawning Wr 2003
140
120
10
100
80
logWs= –2.942+2.443logSUL
North
Terrestrial habitat
test: females, χ2 = 9.975, df = 2, P = 0.007; males, χ2 =
6.027, df = 2, P = 0.049). In 2003, individuals east of the
basin again had the highest body condition; however, the
differences were not significant for females, probably owing
to the small sample size (Kruskal–Wallis test: females, χ2 =
3.394, df = 2, P > 0.5; males, χ2 = 25.595, df = 2, P <
0.0001).
Terrestrial habitats
The daily mean air temperature inside the Schlumsee ba-
sin was significantly lower than that outside the basin
(paired t test; t = 12.80, df = 242, P < 0.0001). Average tem-
perature recorded inside the basin was –0.2 °C (min.
− 22 .5 ° C, max. 24.5 °C), compared with 2.1 °C (min.
− 15 .8 ° C, max. 25.2 °C) outside. The first day with a mini-
mum air temperature below 0 °C was 25 September 2002
both inside and outside the basin; the last day with subzero
temperatures was 4 May 2003 inside the basin and 19 April
2003 outside the basin.
Mean daily insolation in the study area ranged from 2.5 to
23.3 MJ·m–2·day–1 (Fig. 4). The values for the terrestrial
summer habitats were more than two and three times higher
to the north and east, respectively, than in the Schlumsee ba-
sin. Areas inside the Schlumsee basin southwest of the lake
and on slopes northeast of the Schlumsee basin received the
least solar radiation.
The northern summer habitats lay entirely within decidu-
ous mixed forest dominated by beeches (F. sylvatica) and
sycamores (A. pseudoplatanus) with interspersed Norway
spruce (Fig. 1). The eastern habitats were less homogenous,
Can. J. Zool. Vol. 83, 2005
males
* *
23 23
24
20
20
logWs= –1.224+1.473logSUL
males
19 *
80
24
28 72
logWs= –2.082+1.948logSUL
East South North East South
Terrestrial habitat
including three vegetation types: larch forest formerly used
for cattle grazing; tall forbs (mainly Adenostyles alliariae
(Gouan) A. Kern. and C. alpina); and coniferous mixed for-
est made up of stone pines (Pinus cembra L.) and larches
with scattered spruce trees. The mixed stone pine and larch
forests form park-like, uncongested vegetation mosaics with
a rich photophile understory (Fig. 1). The tall forbs are re-
stricted to small areas with moist and nutrient-rich soil and
long-lasting snow cover. The area south of Schlumsee is
dominated by mixed beech and sycamore forest in lower re-
gions and mixed stone pine and larch forest on the slopes at
higher elevations.
Food availability
In terms of numbers of individuals, Collembola, Diptera,
Coleoptera (particularly Carabidae (ground beetles) and
Staphylinidae (rove beetles)), and Araneae dominated in all
three habitats (Fig. 5). Species of the order Opiliones were
abundant only in the northern habitat, and those of the sub-
family Formicinae (Hymenoptera) were completely absent
from the northern habitat and the basin. There were no sig-
nificant differences in the number of potential prey organisms
between habitats (Kruskal–Wallis test; χ2 = 5.43, df = 2,
P = 0.66), but the three habitats differed significantly in prey
mass (Kruskal–Wallis test; χ2 = 34.54, df = 2, P < 0.0001).
The highest masses were observed to the east, whereas the
lowest values again occurred to the north. Mollusca largely
dominated the invertebrate fauna to the east and in the basin
(Kruskal–Wallis test; χ2 = 30.471, df = 2, P < 0.0001),
whereas to the north, Carabidae dominated and Mollusca
© 2005 NRC Canada
Sztatecsny and Schabetsberger
Fig. 4. Map of the study area showing the solar insolation aver-
aged over 8 modelled days from 28 February to 15 June.
contributed only 2.3% to total mass. All categories except
Oligochaeta and Opiliones were found in the stomachs of
dissected toads (Table 1).
Discussion
Methodology
No animals died during radio-tracking, and wounds healed
well within 4–5 weeks. Surgical transmitter implantation
was the only practical way to radio-track common toads in
alpine environments and locate their distant terrestrial habi-
tats. Alternative methods, such as external attachment of
transmitters or insertion into the oesophagus, were unaccept-
able because of the short retention times and the high risk of
transmitter loss (authors’ observations; van Gelder et al.
1986; Oldham and Swan 1992).
Migrations
To our knowledge, this is the first record of vertical mi-
grations over several hundred metres for adult amphibians
(but see Funk et al. 2005 for juveniles). Common toads were
able to overcome considerable obstacles, surmounting scree
slopes with inclinations of up to 45° and climbing nearly
vertical cliffs to reach preferred habitats. Despite the eleva-
tion gain and the high friction (i.e., landscape resistance) of
the alpine habitat, distances between the breeding site and
793
terrestrial habitats were similar to those observed in
lowlands (Heusser 1968; Sinsch 1988a). Due to logistic dif-
ficulties, our sample size was small, but it still allowed us to
detect major migration routes predetermined by the relief
(Fig. 6). While surveying these routes in 2002–2003, we re-
peatedly found single female toads up to 1.2 km from
Schlumsee at altitudes ranging from 680 m (to the north) to
1500 m (to the south and east) (Fig. 6). Assuming that these
toads belonged to the Schlumsee population, which is likely
because it is the closest suitable breeding site, they had
gained up to 400 m in elevation to the east and southwest.
To the north, they first had to gain 100 m to leave the
Schlumsee basin via the pass at 1200 m and then descend
520 m. Metamorphosed juveniles observed in July 2001 and
2003 seemed to use the same routes and complete their mi-
gration to the terrestrial habitats in roughly the same time as
the radio-tracked adults.
Our findings suggest that distant terrestrial habitats should
be advantageous despite necessitating demanding migra-
tions. The boulder fields and scree slopes surrounding
Schlumsee are only scarcely covered with vegetation and
hence probably offer little food. The meadow forming the
lake bottom becomes accessible only after the lake has dried
out. Prey organisms were abundant on the meadow, making
a lack of food alone unlikely to drive the toads’ migrations.
However, to forage, toads depend on warm and moist condi-
tions (Kuhn 1994; Reading and Clarke 1995), which might
be rare inside the shaded and cool Schlumsee basin.
Anurans are capable of remarkable homing and orienta-
tion (Sinsch 1990). In lowland populations, most toads per-
form the post-spawning migration only once in their life
owing to high mortality rates of more than 50% per year
(Gittins et al. 1985; Kuhn 1994). However, in many amphib-
ians longevity increases with increasing altitude and latitude,
enabling them to breed repeatedly (Smirina 1994). The me-
dian age of Schlumsee toads is 10.5 years for females and
8 years for males (Schabetsberger et al. 2000). With females
and males having a minimum age for breeding of 7 and
6 years, respectively, it seems very likely that many individ-
uals return to the breeding site more than once.
Body condition and terrestrial habitats
Over two consecutive years, the condition index was high-
est for toads east of the basin. The body condition of an ani-
mal mainly reflects food uptake in the previous year and is
assumed to be indicative of habitat quality (Reading 1990;
Kuhn 1994; Pope and Matthews 2002). Snails, which domi-
nated the biomass of organisms captured east of the basin,
are part of the common toad’s diet (Eibl-Eibesfeldt 1951;
Wheater 1986; present study), and a single abundant prey
type rich in energy can have a significant effect on body
condition (Pope and Matthews 2002). However, a series of
samples collected throughout the season, amended by addi-
tional trapping techniques to avoid a bias in captures towards
ground-dwelling organisms, together with samples of toad
stomach contents, would have been required to compare
prey availability and temporal variation between habitats
(Donnelly 1991).
Besides the abundance and composition of food, the time
available for food uptake may affect body condition
(Jørgensen 1992). Toads are ectothermic animals, and tem-
© 2005 NRC Canada
794 Can. J. Zool. Vol. 83, 2005

Fig. 5. Abundance and mass of potential prey organisms in the two terrestrial habitats north and east of Schlumsee and inside the
Schlumsee basin (x axis scales differ between habitats).

North East Basin

Oligochaeta
Isopoda
Acari
Opiliones
Araneae
Chilopoda
Myriapoda
Collembola
Cercophidae
Carabidae
Staphylinidae
Other Coleoptera
Larval Coleoptera
Diptera
Formicinae
Other Arthropoda
Mollusca

2
4
6
8
.0

.0

2
4
6
8
0

0
25

20

15

10

0. 0
0. 5
0. 0
0. 5
0. 0
0

25

20

15

10

45

20
15

10

0
02
05
07
10
50

80

0.
0.
0.
0.

.
.
.
.
.

.
18

25

0
0
0
0
10

20
0.

Number trap–1 Mass trap–1(g) Number trap–1 Mass trap–1(g) Number trap–1 Mass trap–1(g)

Table 1. Gut contents (no. of organisms per taxonomic category) of six juvenile (Juv.) and
two adult male common toads (Bufo bufo) unintentionally captured in pitfall traps in the
terrestrial summer habitats east of Schlumsee and in the Schlumsee basin.
East Basin
Juv. Juv. Juv. Juv. Male Male Juv. Juv.
SUL (mm) 39.3 20 20 20.8 59.8 51 20 18.5
Taxon
Oligochaeta
Isopoda 1 2
Acari 4 3 3
Opiliones
Araneae 1 1 1
Chilopoda 2 1
Myriapoda 1 2
Collembola 1 1 2 2 1 1
Cercophidae 2
Carabidae 3 2 1 1
Staphylinidae 3 2 1
Other Coleoptera 3 2
Larval Coleoptera 1 1 1
Diptera 1 3 1 2 3
Formicinae 1
Other Arthropoda 3 3 1 1 1 1
Mollusca 1

peratures in the alpine environment of the Schlumsee study
area restrict their post-breeding activity to 4–5 months.
However, toads can regulate their body temperature
behaviourally by heliothermic heating, retreating to the
shade, and evaporative cooling (Carey 1978; Sinsch 1989).
The higher levels of insolation east of the basin may enable
the toads to maintain a favourable body temperature over a
wider range of air temperatures and hence prolong their
daily and seasonal activity. Furthermore, the greater diver-
sity of vegetation types to the east may provide higher
microclimatic variation, which may help toads, as well as
prey organisms, extend their activity period. Extending the
feeding period affects the body condition of toads, particu-
larly in alpine environments where, unlike in lowlands, body

© 2005 NRC Canada

Sztatecsny and Schabetsberger
Fig. 6. Aerial photograph of the study area showing the three
main migration routes of common toads. White squares indicate
free-ranging (i.e., untagged) common toads that were found dur-
ing surveys from 2001 to 2003. Contour intervals of 100 m are
shown.
reserves reach their maximum only at the end of the activity
period (Morton 1981; Jørgensen 1992).
The amount of snowfall is negatively correlated with the
body condition of anurans (Pope and Matthews 2002). We
do not have data on snow coverage; however, mixed beech
and sycamore forests are restricted to areas characterized by
large quantities of snow (Mertz 2000). This vegetation type
was dominant to the north and partly present to the south but
absent to the east.
Implications for conservation
Our study revealed differences among terrestrial habitat
types for common toads at high altitudes. In alpine environ-
ments characterized by high fragmentation but moderate lev-
els of human landscape alteration, it seems to pay off for
toads to perform extensive migrations to reach high-quality
habitats. Determining exactly what specifies habitat quality
was beyond the scope of this study; however, high vegeta-
tion diversity and solar insolation seemed to be important
factors fostering energy uptake during the short activity pe-
riod. Regarding the conservation management of amphibian
populations, this implies that spatially extensive investiga-
tions of terrestrial habitats are necessary to plan proper mea-
sures and to protect the most valuable habitat patches.
Because the most valuable habitat patches might occur far
from the aquatic breeding site, effective amphibian conser-
vation might be more demanding and costly than so far as-
sumed.
795
Acknowledgements
We are indebted to many people for help with fieldwork,
S. Kronreif for permission to work at Schlumsee, and I.
Althaler for providing vegetation data. We thank G. Fluch
and F. Schober for their support during transmitter refurbish-
ment, and I. Kleinbauer and R. Jehle for help with GIS ap-
plications. KIÖS of the Austrian Academy of Sciences
provided financial support during fieldwork. For many help-
ful comments on the manuscript we thank J.W. Arntzen, R.
Jehle, B. Schmidt, and the reviewers. During the preparation
of the manuscript M.S. was supported by FWF grant 14799
to W. Hödl. Permits were acquired from the Austrian Na-
tional Board for Experimental Animals (68.210/17-
Pr/4/2002) and the Provincial Government of Salzburg
(21301-RI-494/15-2002, 21301-RI-548/10-2003).
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