ECOGRAPHY 23: 457–465.

Copenhagen 2000

Plasticity in age and size at metamorphosis in Rana temporaria –

comparison of high and low latitude populations

Juha Merilä, Anssi Laurila, Ane Timenes Laugen, Katja Räsänen and Maarit Pahkala

Merilä, J., Laurila, A., Timenes Laugen, A., Räsänen, K. and Pahkala, M. 2000.
Plasticity in age and size at metamorphosis in Rana temporaria – comparison of high
and low latitude populations. – Ecography 23: 457 – 465.

Effects of different combinations of stressors (viz. temperature, food level) on

growth, developmental and survival rates of Rana temporaria tadpoles from two
geographically widely ( :1500 km) separated populations were studied in a common
garden experiment. In both populations, low temperature and low food level lead to
lowered growth rates and delayed metamorphosis, whereas high temperature and
high food level had the opposite effect. Tadpoles from north metamorphosed earlier
and exhibited higher growth rates than tadpoles from south, suggesting local adapta-
tion to shorter growth period and cooler ambient temperature in north. Size at
metamorphosis did not differ between the two populations, but when the differences
in metamorphic age were accounted for, then the tadpoles from north were larger
than those from south. These results suggest considerable adaptive genetic differenti-
ation in growth rates, size and timing of metamorphosis between northern and
southern R. temporaria populations. In both populations, high food levels tended to
reduce tadpole survival rates and there was a negative correlation between growth
and survival rates across different treatments in both populations. In general,
tadpoles from north experienced high mortality rates in high food level – low
temperature treatments, whereas southern tadpoles experienced high mortality in
high food level-high temperature treatments. This suggest that there may be genetic
differences among different populations as how they would be influenced by high
nutrient loads, such as brought along for example by fertilization of forest or
agricultural soils.

J. Merilä ( juha.merila@zoologi.uu.se), A. Laurila, A. Timenes Laugen, K. Räsänen and

M. Pahkala, Dept of Population Biology, E6olutionary Biology Centre, Uppsala Uni6.,
Norby6ägen 18D, SE-752 36 Uppsala, Sweden (present address of J. M. and A. T. L.:
Di6. of Population Biology, Dept of Ecology and Systematics, Box 17, FIN-00014
Uni6. of Helsinki, Finland).

Geographic variation in morphological and life history
traits are of ubiquitous occurrence both in plants and
animals (Mayr 1963, Gould and Johnston 1972, Endler
1977, Linhart and Grant 1996). These differences have
been traditionally assumed to reflect adaptive, geneti-
cally determined differences caused by spatial variation
in local selective pressures. However, this view has been
challenged by demonstrations of occurrence of pheno-
typic plasticity, and few studies have actually demon-
strated a genetic basis for spatial differentiation. For
example, although there is an extremely large body of
data about geographic variation in morphological, be-
havioural and life history traits in birds (Zink and
Remsen 1995), only handful of studies have rigorously
tested genetic basis for these variations (James 1983,
Alatalo and Gustafsson 1988, Rhymer 1992, Berthold
et al. 1992, Strack et al. 1995). This shortage of evi-
dence for spatial genetic differentiation in quantitative
traits among different bird populations is understand-
able in the view of the logistic difficulties in performing
common garden or reciprocal transplant experiments
needed to demonstrate the genetic basis of observed

Accepted 21 November 1999

Copyright © ECOGRAPHY 2000
ISSN 0906-7590
Printed in Ireland – all rights reserved

ECOGRAPHY 23:4 (2000) 457

differentiation. Not surprisingly, the evidence for local April, and length of thermal growth season, defined as
genetic differentiation in quantitative traits in plants the number of days when daily mean temperatures
(Linhart and Grant 1996), insects (e.g. Parsons 1973, exceed 5°C, is ca 250 d (Odin et al. 1983). Breeding in
James and Partridge 1995, Nylin et al. 1996), other the northern population (Kiruna; 67°50%N, altitude=
invertebrates (e.g. De Meester 1996), fishes (e.g. 425 m) starts in late May – early June, and the thermal
Reznick and Bryga 1996) and amphibians (e.g. Berven growth season is ca 115 d long. Mean ambient temper-
et al. 1979, Berven 1982a, b) are much more common. ature in April in Lund is ca 5°C, whereas that in
However, as to the amphibians, the studies focusing on Kiruna is ca −5°C (July: 16°C and 12°C in Lund and
spatial genetic differentiation in quantitative traits are Kiruna, respectively). Consequently, the abiotic circum-
still few, and restricted mainly to few North American stances faced by the growing larvae in these two locali-
species (Berven et al. 1979, Berven 1982a, b, Berven and
ties are very different. From each of the two
Gill 1983, Harris et al. 1990, Bernardo 1994, but see
Lardner 1998).
The aims of this study were three-fold. Firstly, to
compare the degree of differentiation in larval life
history traits (viz. age and size at metamorphosis,
growth rate) between two geographically widely sepa-
rated populations of Rana temporaria in response to
variation in temperature and food availability. Sec-
ondly, to compare the degree of phenotypic plasticity in
larval life history traits between these populations to
see whether age and size at metamorphosis, as well as
growth rates show differences in their sensitivity to
different environmental factors, and thirdly, whether
this sensitivity differs among the two populations. Both
age and size in metamorphosis are important life his-
tory traits in amphibians because of their influence on
larval and juvenile survival, as well as on adult mating
success and fecundity (e.g. Collins 1979, Berven and
Gill 1983, Travis et al. 1985, Smith 1987, Semlitsch et
al. 1988)

Methods
The common frog R. temporaria is a widespread,
medium sized ‘‘brown frog’’ native to Palearctic breed-
ing in various kinds of small water bodies throughout
much of the western Paleartic (Gasc et al. 1997). Breed-
ing occurs in early spring, in cooler areas as soon as ice
start to disappear from lakes and wetlands. The eggs
hatch, depending on the water temperature, ca 1 – 3
weeks after fertilization, and the aquatic larvae meta-
morphose usually after 40–80 d from hatching. Al-
though adult life histories are known to vary along
latitudinal and altitudinal clines (e.g. Kozlowska 1971,
Elmberg 1991, Ryser 1996), little information is avail-
able on larval life history traits at different parts of the
distribution range. In particular, no study has ad-
dressed possible among population genetic differentia-
tion in life history, behavioural or morphological traits
using common garden or transplant experiments.
The animals used in this study originated from two
geographically widely separated populations (Fig. 1).
Breeding in the southern population (Lund; 55°40%N, Fig. 1. A map showing the approximate locations of the two
altitude =20 m) starts usually in late March – early Swedish populations of R. temporaria used in this study.

458 ECOGRAPHY 23:4 (2000)

populations we raised tadpoles from five different
full-sib families. In the case of the southern popula-
tion, we collected five male and five female frogs in 1
April 1998, which were artificially mated in labora-
tory in Uppsala. The eggs from each mating/family
were reared in 3 l volume of declorinated tap water
at 18°C until gill absorption was complete (stage 25;
Gosner 1960). After this, the larvae were transferred
to 0.7 l volume of water, and reared individually until
the metamorphosis. Individual rearings started 12
April 1998, which was taken as the starting date for
the experiments, and denoted as day zero. In the case
of the northern population, we collected five freshly
laid egg masses in 30 May 1998 which were brought
to laboratory in Uppsala. Eggs were allowed to de-
velop in ca 18°C until the Gosner stage 25 was
reached. After this, the larvae were transferred to 0.7
l volume of dechlorinated tap water and reared indi-
vidually until metamorphosis. Hence, the assumption
under laying all between population comparisons is
that the slight differences in pre-hatching conditions,
as well as the fact that larvae from southern popula-
tion originated from artificial crosses, whereas the lar-
vae from northern populations originated from
natural matings, did not influence the subsequent
growth performance of the larvae. However, the re-
sults from a similar experiment in which larvae were
obtained from artificial matings for both of these
populations suggest that the outlined differences in
pre-experimental treatment had no influence on subse-
quent performance of the larvae (Timenes Laugen et
al. unpubl.).
To study the influence of population of origin, tem-
perature and food level on different response vari-
ables (viz. time and size at metamorphosis, growth
rate and survival rate), full-sib tadpoles from five dif-
ferent families per population were raised in two dif-
ferent temperatures (14 and 22°C) and under three
different food levels (low, medium and high), with
three replicates per each family-temperature-food level
combination. Consequently, 18 animals per family
were utilised, and the total number of animals in this
experiment was 180. However, due to mortality, data
on metamorphosis dates and weights was available
only for 113 tadpoles. Within each temperature, the
animals were placed in three blocks, so that one
member of each family in a given food treatment was
present in all of the three blocks. In spite of signifi-
cant among block heterogeneity in the temperatures
(Two-way nested ANOVA: Temperature: F1,99 =
13774.62, p B 0.001; Population (nested within tem-
perature); F2,99 =1.16, p =0.32; Block (nested within
temperature): F4,99 =2.60, p =0.04; Population ×
Block: F4,99 =0.03, p = 0.99), block effects did not ex-
plain any significant amount of variation in any
response variables, and they are not considered in the
subsequent analyses. The lack of block effects may
ECOGRAPHY 23:4 (2000)
owe to small temperature differences among blocks
(max difference among blocks in 14°C: 0.9°C, in
22°C: 0.6°C).
Tadpoles were fed with fine ground 1:3 mix of Te-
traMin (Ulrich Baensch, Germany) and rodent pellets
(Hansson, Uppsala Sweden) every seventh day. The
initial food rations were 15, 45 and 180 mg for low,
medium and high food levels, respectively, after which
they were raised during the second week to 30, 90
and 240 mg, respectively. During and after the third
week, the food rations were kept in constant 60, 180
and 360 mg for low, medium and high food levels,
respectively. Water was changed once a week at the
same time when the larvae were fed. The date of
metamorphosis was determined by daily checks, and
the emergence of the first foreleg (stage 42; Gosner
1960) was taken as criteria for metamorphosis. At
this stage, the metamorphs were weighed to nearest
0.0001 g with an electronic balance. Growth rates are
defined as the metamorphic weight divided by the
number of days elapsed between hatching and meta-
morphosis ( = age).
The effects of different factors on age and size at
metamorphosis and growth rates were investigated
with linear models fitted with PROC MIXED in SAS
(Anon. 1996, Littell et al. 1996). In short, tempera-
ture, food level and population of origin were treated
as fixed effects, whereas the effect of family (nested
within populations) was considered to be a random
effect. To obtain correct degrees of freedom for ef-
fects in models, Satterthwaite procedure (Littell et al.
1996) was used. Analyses of all continuous variables
were performed on log-transformed variables to ho-
mogenize variances between different treatment
groups. If clearly non-significant (p \ 0.10), high or-
der interaction terms were excluded from all models.
Significant population × treatment interaction effects
would indicate genetic differences between popula-
tions for phenotypic plasticity. To illustrate and com-
pare the relative amount of plastic response in
different traits, we calculated coefficient of variation
of treatment means (CV = 100× standard deviation of
the treatment means/grand mean of treatments) fol-
lowing Schlichting and Levin (1990). Hence, the CV
is a quantitative measure of the relative response of a
given character to 6 treatments, and was computed
for each combination of population and trait. Ap-
proximate standard errors of CVs were obtained as:
SE= CV/2n, where n was the number of treatment
means used to estimate CV (Sokal and Rohlf 1981, p.
150). The CVs were considered to be significantly dif-
ferent if CV9 2× SE did not overlap between two
estimates. The survival analyses were performed with
generalized linear model using binomial errors, logit-
link function and type III sums-of-squares as imple-
mented in PROC GENMOD in SAS (Anon. 1996).
459
Table 1. Mixed model ANOVAs of age at metamorphosis and growth rate in Rana temporaria. var= variance component
estimate. NDF =numerator degrees of freedom, DDF = denominator degrees of freedom.

Random effects Age at metamorphosis Growth rate

var9SE z p var9SE z p

Family (population) 0.0002 9 0.0004 0.52 0.6046 0.0047 9 0.0036 1.29 0.0816
Residual 0.0060 9 0.0009 6.90 0.0001 0.0274 9 0.0039 6.89 0.0001
Fixed effects NDF DDF F p NDF DDF F p

Population 1 11.5 60.03 0.0001 1 9.9 10.69 0.0085

Temperature 1 101 2575.81 0.0001 1 98.3 71.63 0.0001
Food 2 99 28.81 0.0001 2 97.2 147.66 0.0001
Temperature×Population 1 100 4.49 0.0366 1 98.1 8.36 0.0047
Food×Population 2 99.8 11.38 0.0366 2 97.7 12.03 0.0001
Temperature×Food 2 98.6 41.14 0.0001 2 96.8 72.35 0.0001

Results lead to larger size at metamorphosis (Fig. 2b), but low

Age at metamorphosis
There was considerable plasticity in age at metamor-
phosis in response to temperature and food level treat-
ments, and these responses differed between the two
populations. The larvae from north metamorphosed at
significantly younger age than those from south (Table
1; Fig. 2a), but the largest impact on age at metamor-
phosis was exerted by temperature, lower temperatures
leading to delayed metamorphosis in both populations
(Table 1; Fig. 2a). Likewise, high food ratios advanced
the timing of metamorphosis (Table 1; Fig. 2a). How-
ever, as indicated by significant two-way interactions
between all treatment effects, the effects of temperature,
food and population were not independent of each
other (Table 1). The significant population× tempera-
ture interaction (Table 1) occurred because temperature
had larger effect on southern (mean age at 14°C/mean
age at 22°C=41%) than northern larvae (49%; Fig.
2a). Likewise, the significant population × food interac-
tion (Table 1) occurred because northern larvae were
more affected by variation in food level than the south-
ern larvae (Fig. 2a). The significant temperature× food
interaction showed that the effects of food treatment
were temperature dependent – higher food levels ad-
vanced metamorphosis only in 22°C, but not in 14°C
(Fig. 2a). Finally, the effect of family on age at meta-
morphosis was not significant, suggesting that genetic
influences on timing of metamorphosis were small rela-
tive to other sources of variation (Table 1).
temperature combined with the highest food level re-
duced size at metamorphosis in both populations (Fig.
2b; Table 2). Furthermore, significant population ×
food level interaction (Table 2) showed that the impact
of food treatment on body size was not similar in the
two populations because the size of the northern, but
not southern, larvae was reduced in the high food
treatment (Fig. 2b). There appeared to be a suggestion
of family effect on size at metamorphosis, but again,
this was weak and non-significant (Table 2).
The foregoing analysis dismisses the fact that the size
variation within and between the two populations are
likely to be associated with variation in the length of
the larval period, and therefore, we repeated the analy-
sis by introducing age at metamorphosis in the model
as a covariate (Table 2). When the positive main effect
of age on size at metamorphosis is accounted for, size
at metamorphosis is in fact larger in north (x=0.50,
SE= 0.02 g) than in south (x = 0.43, SE = 0.02 g; Table
2). However, the impact of temperature and food treat-
ment on (age adjusted) size are now much more compli-
cated, as for example indicated by significant three-way
interaction between temperature, food and population
(Table 2). However, the most interesting point of this
analysis relates to the significant age × population in-
teraction (Table 2), which shows that the relationship
between size and age at metamorphosis differs between
the populations – it is positive in south (b=1.0479
0.269, t97.9 = 3.89, pB0.001), but negative and non-sig-
nificant in north (b = −0.13490.298, t94.5 = − 0.45,
p= 0.65).

Size at metamorphosis
As a contrast to age at metamorphosis, there was no
main effect of population on size at metamorphosis
(Table 2; Fig. 2b), whereas the main effects of tempera-
ture and food treatments had again large impact on
development of the larvae (Table 2; Fig. 2b). In both
populations, low temperature and increasing food level
Growth rates
The northern larvae had significantly faster growth
rates than the southern ones (Table 1; Fig. 2c), and as
indicated by the significant population × temperature
interaction (Table 1), this effect was more pronounced
in the high than in the low temperature (Fig. 2c).

460 ECOGRAPHY 23:4 (2000)

However, the positive net effect of food level on growth
rates was fully realised only in the high temperature,
and the high food levels tended to slow down growth in
the low temperature (Temperature ×Food; Fig. 2c).
Furthermore, the significant population ×food level in-
teraction revealed that the high food levels had a
negative effect on growth of northern, but not of
southern larvae (Fig. 2c).
Survival
Sixtyseven of the total of 180 individuals (=37%) used
in the experiment died before metamorphosis, and this
mortality was not random in respect of population,
temperature and food treatment (Table 3; Fig. 2d). In
both populations, the tadpole mortality increased with
increasing food level, as revealed by significant main
effect of food treatment on survival probability (Table
3; Fig. 2d). Likewise, mortality tended to be higher at
14°C than in 22°C, but the highly significant popula-
tion ×temperature interaction (Table 3) revealed that
whereas the larvae from south suffered highest mortal-
ity in 22°C, the larvae from north suffered highest
mortality in 14°C (Fig. 2d). The population× food
level interaction further indicates that the effect of high
food levels on mortality tended to be much more
pronounced for northern than for southern larvae, in-
Fig. 2. Effects of population,
temperature and food level on
a) age at metamorphosis, b)
size at metamorphosis, c)
growth rate, and d) survival
of R. temporaria larvae
originating from two
geographically distinct
populations and reared in
laboratory. Horizontal bars in
(a – c) are 9SE (sometimes so
small that not visible).
ECOGRAPHY 23:4 (2000)
dependently of temperature (Table 3; Fig. 2d). Finally,
survival probabilities differed among families (Table 3),
suggesting that individuals from certain families had
higher than average propensity to die.
Since the survival was assessed only once a week, and
we did not measure the larvae before metamorphosis,
individual survival probabilities could not be related to
individual differences in growth rates. However, the
family specific survival rates were negatively correlated
with family specific growth rates within each of the
populations and temperatures (Lund 14°C: r = −0.08,
n = 15, p= 0.77; Lund 22°C: r = −0.66, n = 11, pB
0.03; Kiruna 14°C: r = −0.46, n =10, p= 0.18; Kiruna
22°C: r = − 0.44, n = 15, p= 0.10). The overall nega-
tive relationship was further verified by an ANCOVA
model where the effect of growth rate on survival
probability was modelled as being nested within each
population and temperature (Growth rate nested within
population and temperature: F4,44 = 3.76, p =0.0102).
Comparison of phenotypic plasticity between
different traits
The results presented above show that individuals from
two different populations respond differently to varia-
tions in temperature and food levels, but they do not
necessarily paint a clear picture how the plasticity in
461
Table 2. Mixed model ANOVA and ANCOVA of size at metamorphosis in Rana temporaria. var =variance component
estimate. NDF =numerator degrees of freedom, DDF =denominator degrees of freedom.

Random effects ANOVA ANCOVA

var9 SE z p var 9SE z p

Family (population) 0.0060 9 0.0041 1.44 0.0751 0.0048 90.0035 1.38 0.0816
Residual 0.0256 9 0.0037 6.90 0.0001 0.0228 90.0034 6.75 0.0001
Fixed effects NDF DDF F p NDF DDF F p

Population 1 9.8 0.41 0.5353 1 96.5 9.17 0.0032

Temperature 1 97.9 244.50 0.0001 1 96.3 1.15 0.2853
Food 2 97.4 101.35 0.0001 2 94.9 92.41 0.0001
Temperature×Population 1 97.7 3.57 0.0617 1 96.3 6.52 0.0123
Food×Population 2 97.4 5.18 0.0073 2 94.9 8.18 0.0005
Temperature×Food 2 94.7 36.06 0.0001 2 94.7 37.97 0.0001
Temperature×Food×Population – – – – 2 94.7 5.60 0.0050
Age – – – – 1 96.2 5.16 0.0254
Age×Population – – – – 1 96.2 8.65 0.0041

these populations and different traits vary in relation to
each others. In Table 4, we have compared the degree
of plasticity in age and size at metamorphosis in re-
sponse to temperature and food treatments in each of
the populations. These comparisons show that all traits
show roughly comparable levels of plasticity, and that
the degree of plasticity did not differ between the
different populations (Table 4).
Discussion
We found that frogs from northern Sweden (Kiruna)
metamorphosed at significantly younger age than frogs
from southern Sweden (Lund) when reared under iden-
tical conditions in laboratory. As age at metamorpho-
sis, or its reciprocal (1/age at metamorphosis) is a
measure of developmental rate (e.g. Berven et al. 1979),
our findings suggest that frogs from northern Sweden
have a genetic capacity to complete their development
faster than frogs from southern Sweden. This is what is
to be expected under the hypothesis that northern frogs
have adapted to short and cold summers, and in order
to achieve similar fitness, need to complete their devel-
opment faster than their conspecifics in south. In this
sense, our results parallel those observed in studies of
Rana syl6atica (Berven 1982a, b, Berven and Gill 1983,
Riha and Berven 1991) and Rana clamitans (Berven et
al. 1979), in which populations inhabiting cooler cli-
mate (high altitude) were shown to have higher genetic
capacity to complete their development faster, than
those originating from warmer climate (low altitude).
Similar tendency has also been observed in wide variety
of other organisms (review in Conover and Schlutz
1995), suggesting widespread adaptation to harsh envi-
ronments. However, we cannot entirely dismiss the
possibility that the observed among population differ-
ences reflect non-genetic maternal effects, which could
have been transmitted for example through egg size
differences between the two localities. Berven’s (1982b)
results from R. syl6atica suggest that a large proportion
of among population variation in developmental rates
can owe to among population differences in maternal
investment. However, we note that if such effects were
present, they could as well be genetic, and themselves
present adaptation to different environmental condi-
tions (Mosseau and Fox 1998). Further experiments are
needed to evaluate the role of maternal effects in ac-
counting among population divergence in different life
history traits.
Our analyses revealed several significant population-
by-treatment interactions suggesting that the pheno-
typic responses to temperature and food level
treatments differed among the two populations. This is
interesting as it suggests that similar environmental
changes, such as for example brought along with cli-
mate warming (e.g. Hulme et al. 1999), could influence
larval life histories in different populations in different
fashion. Inter-populational divergence in phenotypic
plasticity has been observed also between mountain and
lowland populations of Rana syl6atica (Berven and Gill
1983). Berven and Gill (1983) found that age at meta-
morphosis was very plastic in lowland for larvae, and
that it was more plastic than size at metamorphosis.
Mountain larvae exposed to similar conditions showed
more similar levels of plasticity in both age and size at
Table 3. Generalized linear model of survival probability of
Rana temporaria larvae. The model was tested using binomial
errors and logit link function with PROC GENMOD in SAS
(Anon. 1996).
Source DF x2 p
Population 1 0.13 0.72
Family (population) 8 15.92 B0.04
Temperature 1 5.80 B0.02
Food 2 21.73 B0.0001
Population×Temperature 1 31.10 B0.0001
Population×Food 2 6.48 B0.04

462 ECOGRAPHY 23:4 (2000)

Table 4. Comparison of plasticity in age and size at metamor-
phosis, as well as in growth rates, in two R. temporaria
populations in response to temperature and food treatments.
Values are coefficients of variation (CV 9 SE) estimated using
treatment means (see Methods).
Population Age Size Growth rate
South 36.7910.6 36.89 10.6 53.8 9 15.5
North 45.5 913.1 38.89 11.2 42.89 12.3
metamorphosis (Berven and Gill 1983). Although the
phenotypic responses to temperature and food levels in
all studied traits in this study differed between popula-
tions (cf. Tables 2 and 3), there were no obvious
differences in levels of plasticity among the two popula-
tions (cf. Table 5). Likewise, both size and age at
metamorphosis showed roughly similar levels of plastic-
ity when expressed at a comparable scale. However, the
sample sizes in these comparisons were small, and only
very large differences would have been possible to be
detected.
The relationship between size and age in maturity has
been in focus of several models (Stearns and Koella
1986, Reznick 1992) and empirical studies of life history
evolution (Stearns 1992). The common pattern among
amphibians is that size and age at metamorphosis are
positively correlated both phenotypically (e.g. Blouin
1992, Semlitsch 1993, but see: Kaplan 1985, Pfennig et
al. 1991) and genetically (Berven 1987, Blouin 1992).
Here, we found that the frogs from southern Sweden
had a positive relationship between age and size at
metamorphosis, whereas northern Swedish frogs did
not show any consistent relationship at all. Since both
large size and early metamorphosis are suggested to be
positively correlated with future fitness of metamorphs
(Smith 1987, Semlitsch et al. 1988, Berven 1990), this
suggest that the fitness of northern larvae may be less
limited by the constraint set by the size-age relationship
than that of the southern larvae. In other words, early
metamorphosis at large size seems not to be an option
available for southern larvae, whereas it could be that
for the northern larvae. In this sense, one could say that
there is more plasticity in development of northern, as
compared to southern Swedish common frog larvae.
The lack of positive correlation between size and age
at metamorphosis among the northern larvae could
also be explained by differences in time constraints
individuals in the two populations are likely to face. If
the northern larvae are more time constrained in their
development than the southern larvae (i.e. due shorter
summers), then it is possible that northern larvae ini-
tiate their metamorphosis, for example, after accumula-
tion of certain temperature sum independently of their
size, whereas timing of metamorphosis among the
southern larvae can be more strongly related to attain-
ment of certain size. This explanation is also consistent
with the fact that the northern larvae are faster growing
ECOGRAPHY 23:4 (2000)
than the southern larvae. In general, there seems to be
a tendency for animals in time constrained environ-
ments, even under considerable stress, to exhibit higher
growth rates than those living in less time constrained
environments (Ardent 1997).
One of the major tenets of life history evolution is
that since different components of fitness cannot be
maximised at the same time, negative correlations be-
tween different components of fitness should be of
commonplace occurrence (Stearns 1992). We observed
that the correlation between survival and growth rates
across different families was negative, implying that fast
growth rates were associated with increased mortality
rates. A number of mechanism could explain such a
trade off, but examples of costs for high growth rates
are still scarce (review in Ardent 1997). However, since
we did not have data on individual growth rates for
those individuals which died, this evidence for trade-off
is merely circumstantial. The negative correlation be-
tween survival and growth rates could also rise if high
food levels influenced growth rates and survival rates
independently, so as that there were no causal relation-
ship between growth and survival rates. In other words,
although the surviving larvae from highest food level
treatment might have enjoyed high growth rates than
those from low food level treatment, they might also
have suffered from poor quality of water (i.e. excess of
food). Consequently, if we had been able to measure
the growth rates of tadpoles that died before metamor-
phosis, it is possible that no negative relationship had
been observed. In fact, we observed that the highest
food level did not increase growth rates and size at
metamorphosis, except in the case of Lund individuals
at high temperature, but instead, tended to slow it
down and increase mortality rates. Nevertheless, it is
interesting that the survival rates in the two populations
responded differently to food treatments depending on
the temperature. Divergent responses of populations to
similar diet arrays suggest that they may be distinct in
their feeding ecology (cf. Steinwascher and Travis
1983), or alternatively, that some factor associated high
food levels, such as high ammonium or nitrate concen-
trations (e.g. Hecnar 1995), influenced the survival of
tadpoles differently in different populations. One possi-
bility is that northern populations inhabiting naturally
very oligotrophic waters cannot cope with high nutrient
loads associated with high food levels, whereas south-
ern tadpoles, naturally more often exposed to eutrophic
waters, are better able to deal with these conditions.
Future studies are required to differentiate between
these alternatives.
In conclusion, the results suggest that northern and
southern Swedish populations of the common frog have
diverged from each others in several larval key life
history traits, and the direction of this divergence is
consistent with the expectation that northern popula-
tions have adapted to the colder and shorter summers
463
they experience. Interactive effects of population of
origin, temperature and food level on survival and
growth rates further suggest that changes in key-habitat
parameters such as temperature and water quality,
might have different impact on life histories in different
populations of the common frog.
Acknowledgements – We thank Eevi Karvonen, Niclas Kolm,
Fredrik Södermann, and Geir Wickstrøm Løe for their help
with the laboratory work, and Stefan Andersson, Ingemar
Jönsson, Björn Lardner, Ralph Tramontano and Jon Loman
for their help in obtaining animals forming the parental gener-
ation for this study. This study was supported by grants from
the Swedish Natural Science Research Council (grant B-AA/
BU 12029-304 for JM), Maj and Tor Nessling Foundation
(project c 98053 to JM), Academy of Finland (to AL), as well
by grants from the Nordic Research Academy (NorFA; grants
to JM, AL and KR).
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