Oecologia (1988) 76:390-398
9 Springer-Verlag1988
Seasonal changes in the migratory behaviour
of the toad Bufo bulb: direction and magnitude of movements
Ulrich Sinsch*
Max-Planck-Institut f/Jr Verhaltensphysiologie, D-8131 Scewiesen, Federal Republic of Germany
Summary. The migratory behaviour of the toad Bufo bufo
was studied from February 1985 to April 1986 in the sub-
montane region of Bavaria, West Germany. Toads were
fitted with a mechanical tracking device to record individual
paths of migration. Three aspects of migratory behaviour
were quantified: orientation in relation to the breeding site,
straightness of path, and locomotory activity. The annual
activity period began with migration from the hibernation
sites to the breeding pond in April. The paths went straight
towards the breeding pond independent of the distance
(70-420 m). During the period of oviposition the preference
for the breeding site direction vanished and toads moved
away from the breeding pond, but in less straight paths
than before. In summer migratory activity decreased con-
siderably and was restricted to small areas, the home ranges,
at distances of 55-1600 m from the natal breeding pond.
The straightness of path was rather low, because toads often
returned to their starting points. During rainy nights toads
occasionally left their home ranges for extensive excursions.
In autumn most toads again migrated towards the breeding
pond, but paths were significantly less straight and direct
than in spring. However, toads stopped before reaching
the breeding pond and hibernated in holes or under the
leaf layer. The mortality rate of tracked toads was about
45%. The relative influence of 17 environmental variables
on locomotory activity was evaluated by principal compo-
nent analysis and stepwise multiple regression. Temperature
at night and rainfall variables accounted for significant
amounts of variance, whereas temperature by day, air hu-
midity, and atmospheric pressure showed no correlation.
Activity decreased if temperature approached 0 ~ C or after
long periods without rainfall. Within a certain range of
tolerance, however, the locomotory activity of the toads
was widely independent of environmental factors, indicat-
ing that endogeneous factors are more important sources
of variation in the migratory behaviour of these toads than
commonly assumed.
Key words: Bufo bufo - Migration - Locomotory activity
- Hibernation - Breeding
Important habitat resources (e.g. breeding site, food) of
many amphibians are separated in space and time. Periodi-
* Present address and addressfor offprint requests: Zoologisches
Institut der Universitgt Bonn, Poppeldorfer Schlol3, D-5300 Bonn,
FRG
Oecolbg/a
cal migrations from one resource to another characterize
the annual activities of the adults (Baker 1978). In regions
with pronounced seasonal variations of temperature, hiber-
nation during the winter months is followed by a spring
migration towards the breeding site. This part of the migra-
tory activity has been studied in various species of urodeles
(Taricha vulgaris, Twitty 1959; Ambystoma texanum, Pe-
tranka 1984) and anurans (Bufo americanus, Oldham 1966;
Bufo boreas, Tracy and Dole 1969). Even after passive dis-
placement most individuals of these species return to their
natal ponds (Twitty et al. 1964). When reproductive activity
has ended, the adults leave the breeding site (Bufo bufo,
Heusser 1967; Ambystoma talpoideum, Semlitsch 1985), and
spend the summer months in feeding areas, frequently sev-
eral kilometers from the breeding site (Bufo fowIeri, Stille
1952; Rana pipiens, Dole 1965). In autumn some species
enter into another migratory period before hibernation
(Rana sylvatica, Bellis 1962; Bufo bufo, Heusser 1967).
In the explosive breeders among the migratory amphibi-
ans the reproductive individuals of a population gather at
the breeding site within 1 or 2 weeks once a year (for review
see Wells 1977). This synchronization is even more remark-
able considering that (1) the activity of the ectothermical
amphibians depends at least partly on environmental condi-
tions and (2) the distance between feeding area and breeding
pond varies from a few meters to several kilometers for
individuals of the same population. However, the relative
influence of different environmental factors on the migrato-
ry behaviour of free-ranging amphibians is difficult to
quantify, because most factors are highly intercorrelated.
Univariate analyses of correlations between behavioural pa-
rameters and environmental factors remain questionable,
since effects may be masked or overestimated by the action
of neglected factors (Gittins 1983 a; Wisniewski et al. 1981).
The simultaneous consideration of all measured variables
by means of multivariate statistics represents a more reliable
approach to the problem (FitzGerald and Bider 1974; Smits
and Crawford 1984; Semlitsch 1985).
This study analyses quantitatively the annual pattern
of migration in the common European toad Bufo bufo and
its relation to season and climate. These toads are explosive
breeders which live solitarily most of the year, but meet
annually in large breeding aggregations at their natal pond
for a few days (Jungfer 1943; Eibl-Eibesfeldt 1950; Heusser
1960a, b, 1967, 1968). Their migratory behaviour includes
a prespawning migration to the breeding pond and a post-
spawning migration to the feeding areas in spring, locally
restricted summer movements, and a less conspicuous au-
 391

tumn migration to the hibernation sites. The daily paths Table 1. List of environmental variables considered in the multivar-
of individual toads were recorded throughout these four iate analyses as potential sources of variance in the locmotory
migratory periods, to investigate directional choice with re- activity of toads. All parameters were recorded continuously 10 cm
spect to the breeding site. In addition locomotory activity above ground by a Lambrecht Thermo-hygro-barograph
was related to 17 environmental variables by means of prin-
Independent variable Definition
cipal component analysis and stepwise multiple regression,
in order to evaluate quantitatively the influence of exoge- Air Temperature
nous factors.
(1) TEMPDMAX Maximum temperature between sun-
rise and sunset
(2) TEMPSS Temperature at sunset
Materials and methods (3) TEMPNMIN Minimum temperature between sunset
Study area and sunrise
(4) TEMPNAV Mean temperature between sunset and
The study was conducted in the sub-montane region o f sunrise
Bavaria near Seewiesen, West Germany, at altitudes be- (5) TEMPRATE Mean decrease of temperature between
tween 681 m and 718 m. The study population of Bufo inha- sunset and sunrise per hour
bited an area of about 6 km 2 mainly covered with pine Relative Air Humidity
forest and pasture. The breeding site was near the northern
(6) HUMIDMIN Minimum humidity between sunrise
shore of an artificial pond (3200 m 2 of water surface) within
and sunset
the study area. (7) HUMISS Humidity at sunset
(8) HUMINMAX Maximum humidity between sunset
General procedures. and sunrise
(9) HUMINAV Mean humidity between sunset and
Most toads were captured at night on gravel roads and sunrise
paths within the study area, others near the shores of the
Atmospheric Pressure
breeding pond. In situ every individual was fitted with a
track device on its back (Sinsch 1987a, b). The device con- (10) PRESSR Absolute pressure at sunrise
(11) PRESSS Absolute pressure at sunset
sisted of a sewing machine bobbin which was tied around
(12) P R E S - 2 4 Absolute change of pressure during
the toad's waist with an elastic tape. The bobbin contained 24 h before sunset
about 60 m of thread. Further details are described in Dole (13) PRES-- 12 Absolute change of pressure during
(1965). The loose end o f the thread was tied to a small 12 h before sunset
stake (referred to as "release point") and each toad was (14) PRES + 12 Absolute change of pressure during
released at the place of capture. The body mass of the toads 12 h after sunset
monitored ranged from 39 g to 46 g, to minimize unspecific
Rain
effects on the toads' behaviour.
On the morning of the following day two measurements (15) RAIND Hours of rainfall between sunrise and
sunset
were recorded for each individual: (1) locomotory activity
(16) RAINN Hours of rainfall between sunset and
loc as judged by the metres of thread unwound during the sunrise
diurnal activity period; (2) position of the animal in polar
coordinates, i.e. azimuth angle a and line-of-sight distance Soil Moisture
r relative to the release point. The hiding place of each (17) SOIL Hours after last rainfall
individual during its resting period was taken as the release
point the next day. All toads were tracked for at most
21 consecutive days, individually marked by toe clipping,
and then released at their last position. Newly captured
Statistical analysis
individuals from the adjacent area replaced those released.
The number o f toads tracked simultaneously varied from The migratory behaviour of the toads monitored was quan-
10 to 29. A total of 132 toads (112 males and 20 females) tified using three parameters: breeding site component cbs,
were monitored in the course of the study, 43 of them in straightness st of path and locomotory activity loc. Breeding
two tracking periods, 9 in three and 1 in four. Seventeen site component and straightness of path were calculated
environmental variables which might influence the locomo- separately for every individual using the equations: (1) Cbs=
tory activity of toads were continuously recorded within cos(abs), where abs is the direction of migration relative to
the habitat (Table 1). Records of the daily precipitation the breeding site; (2) st=r/loc, with r as line-of-sight dis-
made a rough estimate of soil surface moisture possible tance and loc as total locomotory activity (Batschelet 1981 ;
(as hours after last rainfall) because of the negative correla- Sinsch 1987a). The breeding site component ranges from
tion between the water content o f the surface soil and time - 1 to + 1 (Cos= + 1 means the toad moved directly towards
since the last precipitation. This variable represents an aver- the breeding site), the straightness of path from 0 to + 1
age value of the varying amounts of absolute moisture to (st= + 1 means the toad moved in a straight line). Both
be found in the toad habitat simultaneously due to local parameters were estimated only for toads which moved at
variation in vegetation and soil structure. The environmen- least 5 m during their diurnal activity period. Since the dis-
tal factors and toad activity were recorded during five study tribution of daily cb~ measurements frequently differed from
periods: in 1985:28 February-2 May; 10 June-6 July; 31 a normal distribution, the median values and the upper
July-19 August; 10 September-20 November; in 1 9 8 6 : 2 4 and lower quartiles (central 50% o f observations) were used
February-29 April. to describe the directional choice of the monitored toads.
392
hibernation reproduction m~mmer movements
[~
~:~ spring migration r-
+2C . - - _ a"-... M
7W ....
libernation r e ~ d u c t i o n
[~
sprln~ migration
+2c
~+10,
0 Fig. 1. Seasonal variation in ambient air temperature (recorded 10 cm above ground) in the
toad habitat in relation to the activities of the toad population studied. The upper line
-10- connects the daily temperature maxima, the lower line the the mimima. Broken lines indicate
1986 periods without records. The hatched area between maxima and minima lines represents the
MAR. I APRIL I MAY temperature range in which toads showed locomotory activity
Straightness and locomotory activity were presented as the
daily arithmetic mean and corresponding standard error.
Differences between distributions were tested by the Krus-
kal-Wallis H-test.
The association of environmental variables with the
mean locomotory activity of toads per day was estimated
by principal components analysis with orthogonal VARI-
MAX-rotation. This kind of rotation attempts to simplify
the columns of the factor matrix by making all values close
to either 0 or 1 and so facilitates the recognition of asso-
ciated variables within a principal component. The princi-
pal components analysis was based on the matrix of the
linear correlation coefficients of all variables. Extraction
of principal components was terminated when the eigen-
values was <_ 1. Those variables which loaded the same
principal component as locomotory activity were consid-
ered associated. All variables identified in this way were
submitted to a stepwise multiple regression analysis (proce-
dure: forward selection of variables) to obtain quantitative
models for the prediction of the locomotory activity. If
the variance explained by the next variable to enter did
not increase significantly the total of variance explained
(F-statistics, P>0.05), the regression analysis was termin-
ated. All calculations were performed by F O R T R A N 77
programs using the algorithms summarized in Flury and
Riedwyl (1983).
Results
During five study periods in 1985 and 1986 observations
of toad behaviour and records of the seventeen environmen-
tal variables were obtained for a total of 132 days (Fig. 1).
This data set was subdivided into four subsets for further
analysis: (1) prespawning migration in spring: 27 observa-
tions; (2) postspawning migration in spring: 25 observa-
tions; (3) summer movements: 53 observations; and (4)
autumn migration: 27 observations. The presentation of
the results follows this temporal classification of data with
an additional note on the hibernation behaviour (19 days
hibernatioz
autumn migration
"vv X
of observation). Breeding site component (directional
choice with respect to the breeding site), straightness of
path, and locomotory activity describe the migratory
behaviour of the toads monitored. Figures showing the time
course of these parameters do not present complete data
sets but sequences of 17-20 consecutive days. In addition
the distributions of all breeding site component and
straightness measurements within one migratory period are
summarized in frequency histograms. An analogous presen-
tation of the single records of locomotory activity does not
provide useful information because their magnitude is in-
fluenced by environmental factors (see results of multivar-
iate analyses). In all data sets principal component analysis
grouped the 17 measured environmental variables into five
principal components with eigenvalues > 1 (Table 2). PC1
represented temperature, PC2 moisture, PC3 nightly hu-
midity, PC4 absolute pressure, and PC5 change of pressure.
These principal components accounted for 80.1%-85.7%
of total variance.
Prespawning migration in spring
In 1985 the toads emerged from hibernation on 1 April
in 1986 on 30 March. The following migratory activity was
strictly nocturnal, beginning after sunset and lasting until
temperatures dropped to almost 0 ~ C, Migrating toads were
observed at distances of 70-420 m from the breeding pond.
Records of their paths are shown in Fig. 2. The toads, how-
ever, did not head exactly towards the small spawning area
near the northern shore but approached any shore of the
pond without preference. Some paths even led to a ditch
which terminated in the breeding pond.
The time course of the directional and locomotory pa-
rameters is shown in Fig. 3 (1985) and 6 (1986), and the
distribution of all single measurements in Fig. 7 (left). The
median breeding site components ranged between + 0.669
and + 0.946. The daily orientation of an individual towards
the breeding site did not depend on the distance to the
breeding pond (correlation coefficient r = 0.087, P > 0.05).
 393

Table 2. Association of the measured environmental factors (Table 1) with the five principal components (eigenvalues > 1) following
orthogonal VARIMAX-rotation. Names of the associated variables are given instead of the correlation coefficients because the absolute
values varied slightly within the four data sets

PC1 PC2 PC3 PC4 PC5

TEMPDMAX TEMPRATE HUMISS PRESSR PRES 24- -

TEMPSS HUMIDMIN HUMINMAX PRESSS PRES-12

TEMPNMIN HUMISS HUMINAV PRES + 12
TEMPNAV RAIND RAINN
RAINN
SOIL

Table 3. Quantitative models obtained by stepwise multiple regres-

sion (procedure: forward selection) to predict the mean locomotory
[m] activity of toads. Each of the four data sets on migratory activity
was analysed separately. R 2 is the coefficient of determination

Variable Coeffi- P Contri- Total

cient bution R2
to R 2
pond
(A) Locomotory activity during the prespawning migration in
spring
Intercept 10.54 <0.0001
TEMPSS 2.29 <0.0001 0.711 0.711
200
(B) Locomotory activity during the postspawning migration in
N spring
Intercept 12.83 <0.000t
TEMPNMIN 1.56 0.0004 0.370
SOIL -0.24 0.0302 0.124 0.494
lOO -~ ~ ~" ~%ditch
(C) Locomotory activity during the summer movements
Intercept 3.72 0.0001
RAINN 0.83 0.0010 0.I92 0.192
II
(

J~ (D) Locomotory activity during the autumn migration

0 Intercept - 2.40 0.1320
0 100 200 [m] 300 SOIL -0.04 0.0123 0.392
Fig. 2. Paths of seven migrating toads during the prespawning peri- TEMPSS 1.03 0.0014 0.157
od (753 April 1986). The capture site of each monitored toad RAIND 2.58 0.0023 0.152 0.703
is marked by a dot and the sex of the individual. The remaining
dots represent the places where the toads rested after the nightly
migrations accounted for 71.1% of variance in the locomotory activity
(Table 3).

Postspawning migration in spring

Since most (74%-100% individuals per day) of the moni-
tored toads approached the breeding pond by their daily
movements, the distribution of the 459 recorded breeding
site components was strongly biased: positive 88.7%, nega-
tive 11.3%. Intermediate frost periods did not affect breed-
ing site orientation. The paths of the toads were mostly
straight with 71.6% of all straightness measurements ex-
ceeding 0.8 (median: 0.86). The mean locomotory activity
ranged from 0 to 49.7 m/day, but was usually above 20 m/
day.
Principal component analysis of the mean locomotory
activity of the toads and the environmental variables of
the corresponding day showed that activity was associated
with P C I , the ambient temperature, but independent of
moisture, humidity, and pressure. Air temperature at sunset
(TEMPSS) fitted best of the four temperature variables and
In both years oviposition lasted only a few days: 9-14 April
in 1985; 2zb27 April in 1986. Within 3 days after the last
oviposition all toads had left the breeding pond. They dis-
persed up to 1.6 km away, as proved by the recapture of
a toe-clipped individual. Again all migratory activity was
restricted to the night. At the same time as the onset of
spawning 21 males that had not reached the breeding pond
turned away from the breeding site direction, while another
7 males and 4 females continued towards the breeding pond.
Therefore, the records of migrating toads during the period
of oviposition are not considered in this section.
The change o f direction preference during the spawning
activity is presented in Fig. 3 (1985). Out of 25 median
breeding site components, 17 were negative, and the total
range of medians covered --0.643 to +0.333. Conse-
394
1.0
[/l
o 0.0
-I.0
1.0
0.5
0.0
~j 40
a0
20
o 0
o
,---4
l0
0 i i i i i i i i i i i i i i i l l l l
2. 20.
April 1985
Fig. 3. Time course of breeding site component Cbs, straightness
st and locomotory activity of 10-20 toads during April 1985 (pre-
spawning and initial postspawning migrations). The median (hori-
zontal bar) of all individual breeding site components during 1
day is given as well as the upper and lower quartiles (broad vertical
bar) and the range (thin vertical bar). Straightness st and locomo-
tory activity loc are presented as mean __S.E. (vertical bar). In
some cases the standard errors were smaller than the radius of
the dots. The hatched area indicates the days with spawn deposition
quently, the distribution of the 353 sing!e measurements
was weakly biased: positive 39.1%, negative 60.9% (Fig. 7,
left). The paths of the migrating toads remained rather
straight (median: 0.74), but significantly less straight than
during the prespawning migration (H-test, P<0.01). The
mean locomotory activity was always lower than 20 m/day.
Principal component analysis showed that locomotory
activity was associated with PC1 and PC2, i.e. temperature
and moisture, whereas humidity and pressure did not corre-
late. Although associations with seven environmental vari-
ables were detected (Table 2), the best model of multiple
regression included only two, the nightly minimum temper-
ature ( T E M P N M I N ) and soil moisture (SOIL). However,
both variables together accounted for only 49.4% of the
variance in locomotory activity.
S u m m e r movements
Between May and June migratory activity decreased
strongly and finally the movements of every individual were
restricted to a small area, the home range. No net move-
ment in either direction was observed (Fig. 4). During the
study periods in June, July and August all toads monitored
showed this type of behaviour. In 6 out of 29 toads no
behavioural change occurred until hibernation in October,
while the rest left the home sites between late August and
October. The activity period during summer was limited
to the night hours.
1.0
r~
o 0.0
-i.0
1.o-
.+++ [4,+ ++ ,
0.5-
o.o-
40
30
20 + ,+ § + § §
o
10
0
12. 30. 2.
June/July 1985
Fig. 4. Time course of breeding site component Cbs, straightness
st and locomotory activity of 20-27 toads during June and July
1985 (summer movements). Presentation of data as in Fig. 3
On 45 out of 53 observation days the daily measure-
ments of breeding site components covered the whole range
from - 1 to + 1 in rather even distributions. Most median
breeding site components did not deviate significantly from
zero ( P > 0.05). The overall distribution of the 583 breeding
site components was not biased: positive 52.7%, negative
47.3% (Fig. 7, left). Straightness was significantly lower
than in the previous migration periods (median: 0.65, H-
test, P < 0.01), because 13.1% of the toads performed move-
ments that led them back to the start point (0<st<0.2).
Locomotory activity was generally low, but increased con-
siderably on rainy nights, when the toads made extensive
excursions outside their home ranges.
During the summer months locomotory activity loaded
exclusively PC2, i.e. moisture, whereas neither temperature
nor humidity or pressure showed any association. However,
the only environmental variable which accounted for a sig-
nificant part of variance (19.2%) was rainfall during night
(RAINN, Table 3).
A u t u m n migration
Most toads entered into another migratory period in Sep-
tember. However, migratory activity started earlier in toads
with home ranges far away from the breeding pond than
in those whose home ranges were closer. Therefore, the
line-of-sight distance between the start and the end position
of a toad following a complete tracking period was used
to decide whether the toad was still containing with the
summer movement pattern or whether it was participating
in the autumn migration. If the individual had moved more
than 100 m, the toad was considered to be migrating, be-
cause this distance far exceeds the extensions of a home
range (Sinsch 1987b). All migrating toads headed towards
 395

,,o 1.0-
"/

0'1 /
o 0.0
o oo
-I.0
I"ITIET',I
-1.0 Z
1
1.0- 1.0-

0.5
++++++ 0.5
r162162

0.0 0.0

"~ 40-

E 3o :
t......3
a0 }
O
20" §162 O
o
20
10
o
,,--,4 10-
~ 0 --
0 ? .... iT ,'7"T,;T ,",'iT
7. 25.
13. S e p t e m b e r 1 9 8 5 so. April 1986
Fig. 5. Time course of breeding site component cbs, straightness Fig. 6. Time course of breeding site component cbs, straightness
stand locomotory activity of 12-20 toads during September 1985 st and locomotory activity of 26~24 toads during April 1986 (pre-
(autumn migration). Presentation of data as in Fig. 3 spawning and initial postspawning migrations). Presentations of
data as in Fig. 3

the breeding pond, but migration was frequently inter- 60 1 Spring migrati~ J
rupted for a few days. None of them approached closer 40 i prespawning N
to the breeding pond than 70 m, although some would have
been able to reach the pond itself. In late October 1985
a week of night frosts ended the nocturnal movements. Out 60- Spring migration:
of 23 monitored toads 15 remained inactive and hibernated 0 postspawning
between 70 m and 760 m from the breeding site. After a
week of complete inactivity the remaining 8 toads shifted
to day activity and on sunny days they migrated throughout
~) (3. iLi,,,mmmi,,,,mt
80- Summer movements
the day. The migratory activity of these individuals ended
40"
on 10 November, after heavy snowfalls.
The following section refers exclusively to the nocturnal
part of the autumn migration, because data obtained during 60- Autumn migration
the daytime migration were too few for thorough analysis.
40
The time course of migratory activity in September is pre- 20
sented in Fig. 5. The migrating toads approached the breed- 0
ing pond, but not as directly as in spring. The median breed- -i 0 +1 0 +1
ing site components per day varied from + 0.07 to + 0.943. Breeding site component Straightness
Nevertheless, the distribution of the breeding site compo- Fig. 7. Distributions of the breeding site component Cbs and the
nents was clearly biased: positive 68.5%, negative 31.5% straightness of path st during the four annual periods of toad
(Fig. 7, left). The straightness of path increased again (me- activity. Data are summarized in histograms presenting the relative
dian: 0.78) and reached the level of the postspawning mi- frequency of observations in classes of 0.2 width. The absolute
gration in spring (H-test, P > 0.05), but remained lower than numbers of observations are: prespawning migration 459, post-
during the prespawning migration in spring (H-test, P < spawning migration 353, summer movements 583, autumn migra-
tion 288
0.01). The mean locomotory activity was mostly low.
Principal component analysis showed association of lo-
comotory activity with both PC1 and PC2, temperature
Note on hibernation
and moisture. The quantitative model obtained by multiple
regression included three environmental variables (Table The hibernation sites of 29 toads were observed sporadi-
3): soil moisture (SOIL), temperature at sunset (TEMPSS) cally in November, December, and March. Air temperature
and rain during the day (RAIND). The combined effect in the hibernation site and at 10 cm above ground were
of these variables accounted for 70.3% of variance in loco- recorded, Twenty-three toads hibernated at distances of
motory activity. 70 m-420 m from the breeding pond, and six within their
396
summer home ranges at 1060 m to 1230 m distance. Two
types of hibernation sites were used by the toads: holes
of any kind but mostly those of mice (n= 16) and the leaf
layer on the forest soil (n= 13). Both types gave the same
chance of survival (56% and 54% respectively). Tempera-
tures within the hibernation sites of surviving toads ranged
from +0.9 ~ C to +3,1 ~ C, even when the ambient air tem-
perature was lower than - 1 7 ~ C. In at least five sites the
death of the toad was caused by temperatures far below
zero. At the other ten sites the reasons for unsuccessful
overwintering remained unclear. Intervening temperature
increases (in November and December 1985) up to + 14~ C
did not interrupt hibernation.
Discussion
The annual sequence of migrations in Bufo bufo individuals
fitted with tracking devices was apparently the same as in
free-ranging toads: prespawning and postspawning migra-
tions in spring, summer movements, and autumn migration.
Thus, the tracking device does not influence the toads' ori-
entation behaviour or their willingness to perform migra-
tions (Sinsch 1987a, b). The speed of migration, however,
was reduced to about one-third of that of free-ranging toads
(Gittins et al. 1980; van Gelder et al. 1986). This reduction
of activity was due to (1) the additional mass of the tracking
device and (2) the local ground surface structure. The addi-
tional mass ranged from 14% to 17% of the toad's body
mass, so that its effect was assumed to be approximately
the same in all individuals. The paths of the toads were
usually marked by the thread becoming tangled with small
herbs or branches on the forest floor. The length of thread
unwound therefore also depended on the number of such
surface obstructions. Although most areas in which toads
were tracked had similar ground surface structures, this may
be an additional source of variance in locomotory activity.
Since these effects did not vary over time, the data on loco-
motory activity obtained by this method were suitable for
further comparative analyses. On the other hand, the use
of the models for quantitative prediction of the locomotory
activity is limited to the tracked population.
All annual migrations performed by adult toads were
significantly related to the direction of breeding pond, as
demonstrated by positive breeding site components in early
spring and autumn and by negative ones in late spring.
The orientation of the toads during the prespawning migra-
tion in spring depends mainly on olfactory and magnetic
cues, with visual control of straightness (Sinsch 1987a).
During the other migration periods the orientation cues
involved have not been investigated. The sudden lack of
breeding site orientation in most males simultaneously with
the onset of spawning might be triggered by spawn odours.
Whether toads return every year to the same home sites
after breeding cannot be judged from the available observa-
tions. However, mortality rates of more than 50% per year
(Gittins 1983b) suggest that most toads perform the post-
spawning migration only once in their life. The high rate
of winter mortality in the population studied (about 45%)
probably exceeds natural mortality because the tracking de-
vice certainly hindered the toads from entering safer hiber-
nation sites. During summer the toads exhibit a remarkable
fidelity to their individual home ranges as demonstrated
by frequent recaptures (Heusser 1968) and experimental dis-
placements (Sinsch 1987b). The homing ability relied
mainly on visual cues and familiarity with the adjacent areas
(Sinsch 1987b). A similar site fidelity outside the breeding
season has also been observed in Bufo valliceps (Grubb
1970) and in Bufo marinus (Carpenter and Gillingham
1987). Toads often returned to particular hiding places
within their home ranges indicating a limited site fidelity
as also observed in an enclosure (Pinston and Guyetant
1987).
Finally, the time course of breeding site orientation dur-
ing the autumn migration resembled that observed during
spring. The hypothesis that this migration serves to gather
the reproductive individuals of a population near the breed-
ing pond (" Warter/iume ", Heusser 1967, 1968) is supported
by the data presented. The wide range of distances
(70-1600 m) between individual home sites and the breed-
ing site decreased to a maximum of 420 m in the population
studied. The fact that six toads did not participate in the
autumn migration and hibernated at 1060-1230 m distance
supports the hypothesis that not all adults of a population
breed every year (Heusser 1967). Thus, only 2 or 3 weeks
of prespawning migration in spring are sufficient for most
reproductive toads to reach the breeding pond before onset
of spawning activity. To sum up, adult Bufo bufo individ-
uals move periodically between their natal pond and indi-
vidual home ranges with a breeding migration beginning
in autumn, interrupted by hibernation and continuing in
spring.
The seasonal changes in directional choice were accom-
panied by corresponding, but smaller, variations in the
straightness of path. The decrease of straightness following
reproduction was probably due to feeding for the rest of
the activity period, whereas toads starved during the pre-
spawning migration in spring (Heusser 1967). A further
decrease during the summer activity period caused by fre-
quent route reversals reflected the fidelity of some individ-
uals to certain sites within their home range,
The diel activity pattern of the studied population
broadly confirmed the observation that migratory activity
is nocturnal (Heusser 1967). However, toads (up to 28%)
of a Welsh population migrated during daylight as well
(Gittins 1983 b). In Bavaria too some toads.shifted to activi-
ty during daylight for about 3 weeks in late autumn, when
night frost did not permit nocturnal movements. Therefore,
it is likely that meteorological conditions rather than dark-
ness favour nocturnal migrations.
A variety of environmental factors that could influence
locomotory activity has been studied in Bufo spp., e.g. tem-
perature, rainfall, air humidity, atmospheric pressure, and
moon phases (FitzGerald and Bider 1974; Gittins 1983a;
Heusser 1967, 1968; Sinsch 1987b; Smits 1984; Smits and
Crawford 1984; Wisniewski et al. 1981). The present study
allowed a quantitative evaluation of environmental influ-
ences on migratory activity and its seasonal variation. The
proportion of variance in locomotory activity explained by
environmental variables ranged from 71.1% in early spring
and autumn to 19.2% in summer. Since temperature at
sunset alone explained 71.1% of variance during the pre-
spawning migration, a comparison with the results of simple
regression of nocturnal temperature on the number of im-
migrating male toads in a Welsh population is permissible.
In this case 50.1% and 64% (Wisniewski et al. 1981) or
20.9% (Gittins 1983a) of the variance was explained by
nocturnal temperature. The deviating result of Gittins is

probably due to the fact that his analysis is based on only
one breeding period (1981) in which temperature did not
reach critical limits. Principal component analysis and mul-
tiple regression confirm quantitatively that temperature at
sunset has the greatest influence on the number of migrating
toads (Heusser 1968) but reject an additional effect of rain-
fall. The minimum air temperature allowing locomotory
activity in toads is slightly greater than 0 ~ C as shown by
direct measurements. Threshold temperatures of 3 ~ 1 7 6
as calculated from the capture rates in fences (Gittins et al.
1980; Wisniewski et al. 1981) reflect the limits of the meth-
od rather than the true activity.
During the postspawning migration the effect of temper-
ature decreased to 37%, corresponding with the results in
Welsh populations: 28.1% (Wisniewski etal. 1981) and
8.1% (Gittins 1983 a). Soil moisture accounted for a further
proportion of variance, increasing the total enviromental
influence to 49.4%. Although the number of environmental
variables that had significant effects increased, their com-
bined effect declined probably in favour of endogenous fac-
tors. This tendency continued during the summer activity
period when only 19.2% of variance was accounted for
by rainfall. A preliminary analysis of the environmental
influences during summer suggested a greater contribution
(52.9%, Sinsch 1987b) but the data set investigated was
considerably smaller than that in the present study. In Bufo
americanus the combined influence o f rainfall and tempera-
ture was also low, amounting to 34.9% (FitzGerald and
Bider 1974). In the final migratory period of the year the
environmental influence increased to 70.3%, mostly due
to rainfall variables. All data demonstrate that the relative
importance of rainfall increased from the beginning o f the
annual activity period to its end, while that o f temperature
declined. Neither air humidity nor atmospheric pressure
were quantitatively associated with the magnitude o f move-
ments at any time.
The following hypothesis is presented to explain the sea-
sonally varying effect of temperature and rainfall: An envi-
ronmental factor affects the magnitude of locomotory activ-
ity only if this factor approaches the tolerance limits of
the toad. In fact, during spring and autumn temperature
was frequently near or below the critical limit of freezing
point. In the long periods of drought during summer and
autumn the water balance of the toads was probably sever-
ely affected. On the other hand the lack of influence of
temperature (range: 3 ~ C-17.5 ~ C) in summer indicates that
the toads are able to compensate for the effect of tempera-
ture on locomotory activity. Compensation could be based
either on thermoregulatory behaviour as in Bufo spinulosus
(Pearson and Bradford 1976) and/or on physiological adap-
tations o f metabolism as in Bufo terrestris (Smith 1976)
and other anurans (for reviews see Pough 1980, 1983). This
hypothesis, deduced from field data, could be supported
or rejected by further experimental studies under controlled
conditions in the laboratory. Nevertheless, this study dem-
onstrates clearly that in the ectothermal toad Bufo bufo
endogenous factors may explain a seasonally varying, but
significant, portion of the variation in locomotory activity.
Acknowledgements. I am very grateful to HG. Wallraff who recog-
nized various weak points in an earlier draft of this paper. For
further comments I thank I. Eibl-Eibesfeldt, D. Neumann and H.
Schneider. Thanks are also due to L. Sinsch and R. Wahl for
technical assistance in the field.
397
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Received January 8, 1988