Biodiversity and Conservation 13: 409420, 2004.

2004 Kluwer Academic Publishers. Printed in the Netherlands.

Spatial scales for the management of amphibian

populations

JAIME BOSCH*, LUZ BOYERO and INIGO
MARTINEZ-SOLANO
Museo Nacional de Ciencias Naturales, CSIC, Jose Gutierrez
Abascal 2, E-28006 Madrid, Spain;
* Author for correspondence (e-mail: bosch@ mncn.csic.es)

Received 18 March 2002; accepted in revised form 22 January 2003

Key words: Amphibians, Biodiversity, Conservation, Ponds, Spatial scales

Abstract. Conservation programs must be based on a detailed knowledge of the distribution of

communities and populations, but this changes with the spatial scale of observation. In this context, we
examined the multiscale patterns of spatial variation of a montane amphibian assemblage in Central
Spain. The largest spatial scale examined (in a range of 1200 m) accounted for the maximum variability
in both species richness and larval abundance, while the smallest scale examined (pond) was responsible
for the maximum variation of larval abundances of many individual species. Habitat characteristics
seemed to be more related to spatial variation of the amphibian assemblage at a particular spatial scale (in
a range of 75 m), and distances among ponds were in part responsible for variation of larval abundance,
but not species richness.

Introduction

Patterns of variation of biological communities change with the spatial scale of

observation (Levin 1992). The understanding of these patterns at a variety of spatial
scales is necessary in order to avoid inappropriate sampling designs and experiments
(Downes et al. 1993; Boyero and Bailey 2001) and, moreover, it is critical to
prevent losses of biodiversity (Wiens 1989). As Noss (1992) noted, the fact that
ecological processes operate at different spatial (and temporal) scales has relevant
implications for practical conservation. However, management decisions regarding
conservation of biodiversity are often constrained by limited resources, which has
led to a search for optimization criteria. In this context, the choice of the appropriate
scales for biodiversity assessments could be helpful in minimizing costs and
maximizing efficiency in the management of natural populations.
Nowadays, amphibians are subject to great conservation efforts, as a consequence
of the multiple extinction episodes that have occurred in recent decades (Bradford
1991; Carey 1993; Richards et al. 1993; Lips 1998). Due to their dependence on
aquatic habitats, amphibians are generally distributed in aggregations near breeding
sites, which results in patchy distributions. Thus, the study of amphibian populations
at different spatial scales is likely to result in different conclusions and management
strategies. This directly affects the success of conservation policies and, as a
consequence, the maintenance and good status of amphibian species. In spite of that,
multiscale studies are scarce and usually involve comparisons of hierarchical
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subsets within large regions (see, for example, Hecnar and MCloskey 1997a).
However, finer scale approaches involving the analysis of scaling effects in
variables such as amphibian species richness and abundance in smaller areas (such
as common protected areas) might also be helpful for designing cost-effective
monitoring plans for long-term management of amphibian populations.
We studied amphibian diversity and abundance at different spatial scales in a
small protected area of Central Spain. Our main objective was to determine which
are the most appropriate spatial scales for conservation of the whole amphibian
assemblage and individual species in our study area. We also tried to determine
whether observed patterns in amphibian variation matched the spatial scales of
variation of their environment, or were best explained by distances among ponds,
and thus which factor could be best used as an estimator of amphibian diversity and
abundance.

Methods

Study area

Penalara Natural Park is a protected area (768 ha) located between 1800 and 2430 m
altitude in the Sierra del Guadarrama, Central Spain (408 509 N, 38 579 W). The area
consists mainly of granitic outcrops with scattered bogs and alpine grasslands at
higher altitudes, with heathlands dominated by Cytissus oromediterraneus and
Juniperus communis nana and forests of Pinus sylvestris at lower altitudes.
Within the Park, 242 ponds have been catalogued (Granados and Toro 2000), and
10 amphibian species presently breed: Salamandra salamandra, Triturus alpestris,
Triturus marmoratus, Bufo bufo, Bufo calamita, Hyla arborea, Alytes obstetricans,

Discoglossus galganoi, Rana iberica and Rana perezi (Martnez-Solano et al.
2003).

Variables

From June to September 1999 all catalogued ponds and water bodies in Penalara
Natural Park (n 5 242; Granados and Toro 2000) were each visited up to six times
during the reproductive period, to score larval abundances of all species present.
Additionally, a few temporal and shallow ponds that were completely dry at the
beginning of the sampling period in June 1999 were visited again in May 2001. The
surveys included diurnal visual transects around the perimeter of the ponds as well
as dip netting. Adult amphibians were also searched for in the vicinity of each pond.
Species richness (number of amphibian species per pond) was recorded. In addition,
for each species, larval abundance at each pond was scored using the following
categories: 0, 110, 1050, 50100, 1001000, .1000. For small abundances all
tadpoles were counted; in other cases abundances were estimated by visually
counting all larvae present in a portion of each microhabitat, and then extrapolating
to the whole pond according to the area occupied by that microhabitat. Counting
 411

larvae is not difficult in the area because most ponds (up to 60%) are smaller than 5
m 2 , the water is almost always crystal clear and vegetation growth is limited. Since
the goal of the sampling was to score all species reproducing and the maximum
larval abundances they reach at each pond, multiple counts were performed during
the study period and maximum observed abundances were considered.
Each pond was characterized by several variables including physical measure-
ments, vegetation, substrate and surrounding habitat (Table 1). The minimum
distance between each pond and its nearest neighboring pond was also considered
(minimum distances ranged from 2.0 to 117.0 m; mean 6 SE 5 10.2 6 1.1 m). All
ponds were characterized before the start of the breeding season, in the shortest
possible lapse of time (less than 2 weeks), in order to minimize differences among
ponds due to growth of the aquatic vegetation. No significant changes in vegetation
composition of ponds were detected during the sampling period.

Spatial scales

According to the dimensions of the studied area and the arrangement of ponds, four
spatial scales were considered: pond, group of ponds, sector and group of sectors
(Figure 1). Ponds that were close together (in a range of 75 m) were arranged into a
group of ponds (there was a total of 19 groups). In turn, groups of ponds close
together (in a range of 300 m) were arranged into a sector (with a total of eight
sectors). Finally, sectors that were close together (in a range of 1200 m) were
arranged into a group of sectors (with a total of two groups of sectors). Since at least
two elements were necessary to calculate within-group variation at each scale, some
ponds (30 from a total of 242) were excluded from this analysis because their spatial
position did not allow them to be arranged hierarchically into the considered scales.

Statistical analyses

For each data set (pond characteristics and larval abundances) a detrended corre-
spondence analysis (DECORANA or DCA; Hill and Gauch 1980) was performed,
using PC-ORD v. 3.17 software, in order to summarize the data into a minimal
number of variables. We used this statistical technique because it avoids the
common assumption of linearity of traditional statistics (i.e. principal components
analysis) and is able to detect unimodal relationships between species and external
variables. Therefore, for further analyses, we considered the scores of the DCA axes
of both pond characteristics and larval abundance data sets instead of the studied
variables. Three species were omitted from the DCA analyses because they occurred
only at a few sites: B. bufo, H. arborea and D. galganoi.
Variation of pond characteristics, larval abundances and species richness was
examined with nested ANOVA, which partitions overall variation into components
accounted by the different spatial scales. All scales were considered as random
factors in the analyses. A single analysis was performed for species richness, but
different nested ANOVAs were used for both pond characteristics and larval
abundances (where the axes resulting from DCA analyses were the dependent
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Table 1. Variables used to characterize the ponds included in the present analysis (including physical
measurements, vegetation, substrate and surrounding habitat).
Physical measurements
AR 29.6 6 6.7 (0.01025.0) Area (m 2 )
PE 23.2 6 2.6 (2.0391.0) Perimeter (m)
LE 6.8 6 0.5 (1.048.0) Length (m)
WI 3.7 6 0.3 (0.036.0) Width (m)
RA 2.1 6 0.1 (0.018.0) Radius (m)
AL 2082.7 6 4.1 (1952.02235.0) Altitude (m)
WD 33.2 6 1.1 (0.085.0) Water depth (cm)
MD 7.0 6 0.6 (0.050.0) Mud depth (cm)
SP 530.6 6 31.9 (82.01629.0) Spatial position relative to the entrance of the Natural
Park (m)
DD 217.5 6 3.3 (155.0365.0) Date at which the pond dries completely
SMD 129.4 6 1.4 (97.0155.0) Date at which the snow covering the pond melts com-
pletely
WF 0.5 6 0.1 (0.03.0) Water flux (visually estimated at four levels, from 0
no flux to 3 maximum flux)
TU 1.9 6 0.1 (0.05.0) Turbidity (visually estimated at 6 levels, from 0 no
turbidity to 5 maximum turbidity)
NBC 0.9 6 0.1 (0.04.0) Number of connections with brooks
Substrate and vegetation variables (visually estimated)
SR 3.4 6 0.8 (0.090.0) Proportion of shore with rocks
SG 49.5 6 3.2 (0.0100.0) Proportion of shore with grass
SM 35.7 6 3.0 (0.0100.0) Proportion of shore with moss
SMC 10.0 6 1.8 (0.0100.0) Proportion of shore with moss forming underwater
caves
BSLM 60.8 6 2.4 (0.0100.0) Proportion of bottom covered by sand, lime or mud
BAM 12.2 6 1.7 (0.0100.0) Proportion of bottom covered by aquatic macrophytes
BDVD 21.6 6 2.0 (0.0100.0) Proportion of bottom covered by dead vegetation or
dung
BG 5.4 6 1.2 (0.0100.0) Proportion of bottom covered by grass
SUG 2.5 6 0.8 (0.090.0) Proportion of surface occupied by grass
SAM 10.1 6 1.7 (0.095.0) Proportion of surface occupied by aquatic macrophytes
SADV 7.5 6 1.4 (0.0100.0) Proportion of surface occupied by algae and dead
vegetation
SWV 79.9 6 2.2 (0.0100.0) Proportion of surface without vegetation
SH Surrounding habitat, including:
79.2 Cervunal (alpine grasslands of Nardus stricta),
14.2 Cervunal with rocks,
6.6 Others
For continuum variables, mean 6 SE and ranges are shown. For surrounding habitat the proportion of
ponds in each category is shown. AR, PE, LE, WI, RA and AL are from Granados and Toro (2000).

variables in the ANOVAs). Additionally, univariate nested ANOVAs were per-

formed for larval abundance of each species separately in order to assess the
variation of abundances of the different species at different spatial scales. These
seven tests were considered as independent null hypotheses with separate estimates
of mean square error, so we did not correct probability levels. In all cases, the
percentage of the total variance explained by each spatial scale was estimated based
on the sums of squares (analysis performed with STATISTICA 5.5, StatSoft Inc.,
 413

Figure 1. Map of the studied area showing the different spatial scales considered: ponds, groups of ponds
(75 m), sectors (300 m), and groups of sectors (1200 m).
414

Table 2. Data on incidence of seven species of amphibians at the 212 studied ponds of Penalara Natural
Park, expressed as the percentage of ponds occupied for each spatial scale considered (the pond scale
represents the total number of studied ponds).
Species Group of sectors (1200 m) Sector (300 m) Group of ponds (75 m) Total (pond)
S. salamandra 100 100 84 58
T. alpestris 50 25 21 12
T. marmoratus 100 63 21 4
A. obstetricans 100 25 11 2
B. calamita 100 100 79 25
R. iberica 50 13 11 3
R. perezi 100 38 26 11

module Variance Components). Probability levels were not calculated for the pond
scale, because this scale constitutes the error term of the analysis, and thus only the
percentage of variance accounted by this scale can be assessed (Underwood 1997).
Finally, the minimum distance among ponds was related to pond characteristics,
larval abundances and species richness through simple (for species richness) or
multiple regression models.

Results

Data on incidence, i.e. percentage of ponds occupied (total and for each scale), are
given in Table 2. The first axis of DCA for pond characteristics explained 90% of
the variation in the data set, so only this axis was used as a dependent variable in a
nested ANOVA. The variables with higher contribution to this axis were SP, SAM,
BAM (positively) and SWV (negatively). Physical pond measurements (e.g. AR,
PE, LE, or RA) were clustered, as occurred with other related variables (e.g. AL,
DD, and SMD) (Figure 2). For larval abundances, the first three DCA axes
explained 34, 29 and 12% of the variance, respectively. Each axis separates one
species (S. salamandra, B. calamita and T. alpestris, respectively) from the rest of
the species (Figure 3). S. salamandra and B. calamita are the most abundant species
in the area, while T. alpestris is restricted to a very reduced section of the Park

(Martnez-Solano et al. 2003).
Nested ANOVAs for pond characteristics, larval abundance, and species richness
revealed different patterns of variation at different spatial scales (Table 3). In the
case of pond characteristics, significant variation occurred only at the scale of group
of ponds. The variance components of each scale showed that pond scale accounted
for a high proportion of variance (77%), but this scale included the error term of the
analysis, so we were unable to ascertain which percentage of variance was due to
real variation among ponds and which was due to measurement or other sources of
error.
Larval abundance and species richness showed significant variation at the group
of ponds scale, but also at the group of sectors scale. Pond scale (together with error)
accounted for a high percentage of variance (3348%), together with the group of
 415

Figure 2. The first two axes of DCA analyses for the variables (physical measurements, vegetation,

substrate and surrounding habitat) characterizing 212 ponds from Penalara Natural Park (see Table 1 for
variable codes).

sectors scale (3361%). Again, sector scale accounted for a low percentage of
variance (01%), showing no significant variation for any data set.
Variation of larval abundances differed among species at different spatial scales
(Table 4). Only B. calamita varied significantly at the group of sectors scale, and R.
perezi at the sector scale. At the group of ponds scale, both S. salamandra and B.
calamita varied significantly. The highest percentage of variance occurred at the
pond scale for every species. Moreover, this scale accounted for almost the total
variation of T. marmoratus, A. obstetricans and R. iberica (98, 99 and 94%,
respectively). In this case, the percentage of variance due to pond scale was
presumably very high, despite the error term being included in this scale.
Finally, minimum distance among ponds appeared not to be related with either
pond characteristics (multiple regression: F3212 5 0.407, P 5 0.7483) or species
richness (simple regression: F1211 5 1.567, P 5 0.2120), but it was strongly related
to larval abundances (multiple regression: F3212 5 24.650, P , 0.0001).

Discussion

Previous studies have already stressed the importance of considering spatial scales
in improving the efficacy of management plans, and identifying the effects of habitat
fragmentation (e.g. Schonewald-Cox et al. 1991; Doak et al. 1992; Lewis et al.
1996; Schwartz 1999). However, it may be argued that in small, well-preserved
416

Figure 3. The first three axes of DCA analyses for larval abundances of seven species of amphibians in

Penalara Natural Park. Ss: S. salamandra; Ta: T. alpestris; Tm: T. marmoratus; Ao: A. obstericans; Bc:
B. calamita; Ri: R. iberica; Rp: R. perezi.

areas like our present example at Penalara the benefit of considering scaling effects
in management plans might be negligible. Nevertheless, there are some biological
and environmental factors that operate (and thus exert their influence on popula-
 417

Table 3. Results of the nested ANOVAs showing variation at different spatial scales of pond characteris-
tics, larval abundances (both represented by the axes of the DCA analyses), and species richness.
Probability levels (P) and variance components (v.c., in %) for every spatial scale are also given (except
P values for pond scale, which constitutes the error term of the analysis).
Spatial scale
Group of sectors Sector Group of ponds Pond (including
(1200 m) (300 m) (75 m) error term)
Axis P v.c. P v.c. P v.c. v.c.
Pond characteristics 1 0.2202 3 0.4676 0 ,0.0001 21 77
Larval abundance 1 0.0099 33 0.4056 1 ,0.0001 18 48
2 0.0001 52 0.7334 0 ,0.0001 13 38
3 0.0001 61 0.4673 0 0.0018 6 33
Species richness ,0.0001 49 0.8669 0 ,0.0001 20 37

Table 4. Results of the nested ANOVAs showing variation at different spatial scales of larval abundances
of each species considered independently. Probability levels (P) and variance components (v.c., in %) for
every spatial scale are also given (except P values for pond scale, which constitutes the error term of the
analysis).
Species Spatial scale
Group of Sector Group of ponds Pond (including
sectors (300 m) (75 m) error term)
(1200 m)
P v.c. P v.c. P v.c. v.c.
S. salamandra 0.2202 21 0.4676 0 ,0.0001 23 56
T. alpestris 0.1139 8 0.0777 8 0.1375 4 80
T. marmoratus 0.2143 1 0.2959 1 0.7644 0 98
A. obstetricans 0.7888 1 0.5106 0 0.4238 0 99
B. calamita 0.0070 18 0.7621 0 0.0005 14 68
R. iberica 0.1455 3 0.0724 3 0.9331 0 94
R. perezi 0.1117 8 0.0262 9 0.4719 0 83

tions) at different spatial scales and whose effects may be detected even within such
relatively small areas. For example, there are factors intrinsic to species, which
include habitat selection at smaller scales, or dispersal abilities at larger scales that
may be relevant for management purposes; besides, we should also take into
account extrinsic stress factors, whose influence might be detected both at a local
scale (e.g. predation, pond temporality, organic pollution) as well as at a global scale
(e.g. climate warming, disease). Examples of the latter have already been reported in

our study area in Penalara. Thus, climate warming was tentatively associated with

the expansion of R. perezi in the study area (Martnez-Solano et al. 2002). Disease
has also had an important effect on the population of A. obstetricans at Penalara,
where a chytridiomycosis outbreak reduced the population size by more than 90%
(Bosch et al. 2001). Our data suggest that, for amphibian management purposes, the
two groups of sectors should be considered independently at Penalara Natural Park.
Thus, actions directed towards amphibian conservation should warrant the preserva-
418

tion of maximum amphibian diversity (in terms of species richness) in each unit
separately. The presence of stress factors acting differentially on these two groups of
sectors has already been documented. For example, the negative effect of fish
predation on amphibian populations is well known (see for example Brana et al.

1996; Hecnar and MCloskey 1997b). At Penalara, the salmonid Salvellinus
fontinalis has been historically introduced for sport fishing practices in very few
ponds and streams (Granados and Toro 2000). Avoidance of streams inhabited by
this predatory species has been suggested to explain the recent absence of A.

obstetricans and R. iberica from these otherwise favourable sites (Martnez-Solano
et al. 2003).
Within each group of sectors, conservation efforts should be focused on preserv-
ing only some groups of ponds. High variability in species richness and larval
abundances at this scale suggests that some groups of ponds might harbour more
amphibian diversity and abundances than other groups of ponds. Moreover, this
scale matches the scale of maximum variability of pond characteristics (physical
measurements, vegetation, substrate and surrounding habitat), which suggests that
groups of ponds with high diversity or abundances may share certain pond
characteristics. If this were true, efforts should not be focused on preserving the
maximum variability of pond characteristics, but on certain pond types with
characteristics that favour the presence of a maximum number of amphibian species
and individuals. Distances between ponds do not seem to be important for differ-
ences in pond characteristics or species richness. However, larval abundances
appeared to be similar in ponds that are close together, probably reflecting aggrega-
tion effects. Analysis of scaling effects on larval abundances of different species
suggests that special management plans should be developed for certain species with
particular and serious conservation problems, such as A. obstetricans here (Bosch et
al. 2001).
Long-term monitoring of amphibian populations provides valuable records of
amphibian diversity and abundance, but it also requires the availability of expensive
resources. However, detailed, exhaustive samplings (recording species diversity and
abundance at each pond) can be carried out in any relatively small study area during

one or several seasons (e.g. Martnez-Solano et al. 2003). The existence of a
consistent pattern of variation of amphibian abundance among ponds through time
might be used to validate the data obtained. This information could then be used to
design sampling programs which optimize the economic resources available and,
more important, which can quickly alert about possible conservation problems.
However, it must be taken into account that high abundance or densities do not
necessarily indicate high quality habitat (Van Horne 1982).
It must be considered that the specific spatial patterns relevant for amphibian
management found here cannot be directly applied to other areas. Comparative
studies regarding the same questions in other areas (especially human-altered areas),
or affected by other factors (for example, habitat fragmentation), are needed in order
to complete this picture. Nevertheless, we suggest that studies at multiple spatial
scales are necessary in order to obtain a complete understanding of the distribution
of amphibian populations, even in relatively small areas. We feel that such studies
 419

are essential for the development of good conservation strategies of amphibian

populations and may be especially helpful in designing cost-effective long-term
management strategies.

Acknowledgements

We thank the people working at the Penalara Natural Park for facilities provided
during the samplings. This work was funded by the project 07M / 0109 / 2000 of the

Comunidad de Madrid (PI: M. Garca-Pars). J.B. is supported by the Ramon y
Cajal program from the Ministerio de Ciencia y Tecnologa (Spain). L.B. is
supported by a fellowship from the Comunidad de Madrid (CAM, Spain) and the
European Social Fund. I.M.-S. is supported by a CAM-CSIC-MNCN predoctoral
de Medio Ambiente of the CAM funded the preliminary
fellowship. The Consejera

study Inventario de los Anfibios de Penalara and extended permits for field work.

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