Biological Conservation 116 (2004) 327–332

www.elsevier.com/locate/biocon

Distribution of the Mallorcan midwife toad (Alytes muletensis) in

relation to landscape topography and introduced predators
Robin D. Moorea, Richard A. Griffithsa,*, Alvaro Románb
a
The Durrell Institute of Conservation and Ecology, University of Kent, Canterbury, Kent CT2 7NS, UK
b
Associació per a la Recuperació del Ferreret, Mateu E. Lladó 34B. 1  C 07002, Palma de Mallorca, Balearic Islands, Spain

Received 18 January 2003; received in revised form 14 April 2003; accepted 24 April 2003

Abstract
The endemic midwife toad of Mallorca (Alytes muletensis) is restricted to a small number of breeding populations in the moun-
tainous northwest of the island. The decline of the species has been attributed to the impacts of introduced species such as the
viperine snake (Natrix maura) and green frog (Rana perezi), and toads may be surviving only in areas that are suboptimal for these
predators. The influence of landscape features (elevation, aspect and maximum slope) on the distribution of toads and associated
predators was therefore investigated using GIS. The presence of toads was positively associated with steep slopes. At sites where
they occurred with toads, the distribution of predators was negatively associated with elevation. Reproductive success within toad
populations was strongly associated with the number of pools at each site, while reproductive success within individual pools was
positively associated with elevation. These findings may be used to optimise the design and location of future reintroduction sites.
# 2003 Elsevier Ltd. All rights reserved.
Keywords: Endangered species; GIS; Introduced predator

1. Introduction Manly, 2002; Hall et al., 2002), although most studies to

According to Diamond’s (1989) ‘evil quartet’, the four
principal causes of species’ extinctions are habitat frag-
mentation, overexploitation, introduced predators and
chains of extinctions. Species on islands that have
evolved in the absence of any major predators or com-
petitors are likely to be particularly vulnerable to the
introduction of predators. Periods of coevolution
between predator and prey play an important role in
reducing dynamic responses of one species to the other
(Case and Bolger, 1991). However, novel predators can
severely impact populations of prey species because
there has been no opportunity for coevolution (e.g.
Savidge, 1987; Rodda and Fritts, 1992; Keisecker and
Blaustein, 1997; Fritts and Rodda, 1998; Short et al.,
2002). There are many cases of introduced species
negatively impacting native island fauna (e.g. Gaston,
1994; Wilson et al., 1998; Dowding and Murphy, 2001;
Jackson, 2001; Towns et al., 2001; Burbridge and
* Corresponding author. Tel.: +44-1227-823434; fax: +44-1227-
827839.
E-mail address: r.a.griffiths@kent.ac.uk (R.A. Griffiths).
0006-3207/03/$ - see front matter # 2003 Elsevier Ltd. All rights reserved.
doi:10.1016/S0006-3207(03)00202-7
date have focused on introduced mammals and there
are few documented cases of reptile introductions.
Exceptions include the introduced boa (Boa constrictor),
believed to threaten endemic fauna on Cozumel Island
in Mexico (Martinez-Morales and Cuaron, 1999) and
the brown tree snake (Boiga irregularis), introduced
accidentally to Guam in the 1950s and responsible for
the extinction of many species of native fauna (Savidge,
1987; Rodda and Fritts, 1992; Fritts and Rodda, 1998;
Rodda et al., 1999).
The endemic midwife toad of Mallorca (Alytes mule-
tensis) was once widespread over the island (Alcover et
al., 1984), but today persists only as a small number of
isolated breeding populations. These breeding popula-
tions are largely confined to the limestone gorges of the
Serra de Tramuntana mountains in the northwest of the
island, where populations may use one or more pools
scoured by ephemeral torrents. The decline of the toad
has been attributed by many workers to predation and
competition from species such as the viperine snake
(Natrix maura) and green frog (Rana perezi) (Tonge,
1986; Bloxam and Tonge, 1995), both of which were
introduced to Mallorca around 2000 years ago. Both of
328 R.D. Moore et al. / Biological Conservation 116 (2004) 327–332
these species consume tadpole and adult stages of A.
muletensis (personal observations); in addition, it is
possible that interspecific competition occurs between
the tadpole stages of A. muletensis and R. perezi
(Spence, 2002). While some endemic fauna went extinct
following the introduction of these and other predators
during Roman times (Alcover et al., 1981; Tonge, 1986),
A. muletensis was one of the few species to survive,
although the reasons for this remain unclear. One
hypothesis proposed by some authors is that toad
populations have persisted in gorges with steep sides,
because introduced predators find it difficult to colonise
and establish populations in such areas (Bloxam and
Tonge, 1995; Schley and Griffiths, 1998). A better
understanding of the factors influencing the distribution
of the toad and its associated predators is required
before any firm conclusions can be reached.
Recent advances in geographical information systems
(GIS) and remote sensing technologies have facilitated
the analysis of animal and plant distributions in relation
to spatial environmental attributes (e.g. Farina, 1997;
Corsi et al., 1999; Knutson et al., 1999; Howell et al.,
2000; Lenton et al., 2000). Relatively simple environ-
mental descriptors such as altitude, slope and aspect
have been used to explain distribution and abundance
patterns of taxa such as birds (e.g. Farina, 1997;
Detmers and Bart, 1999) and plants (Janet, 1998),
and it is likely that these variables strongly influence
the microclimate of a site (Stocks and Heywood,
1994). Knowledge of how such environmental features
influence the distribution of rare species and associated
predators may be an important consideration in the
management of the species.
This study used GIS to analyse the distribution of
Mallorcan midwife toads and their principle predators
in relation to landscape topography. We therefore tes-
ted the hypothesis that toad populations have persisted
in sites that are suboptimal for predators. In addition,
we determined which site characteristics were the best pre-
dictors of reproductive success within toad populations.
2. Methods
2.1. Census methods
Annual population surveys carried out at all known
sites since 1991 provided information on the status of A.
muletensis (Garcia and Buley, 1997; Román and Mayol,
1997; Román, 2001, 2002). As adults are cryptic and
spend most of their lives in cracks and fissures under-
ground, annual censuses focused on counts of tadpoles
in the breeding pools. These counts were made in July
each year, immediately following spring breeding activ-
ity. While tadpoles may take anything from several
weeks to over a year to complete development, tadpole
numbers were thought to be at their highest during
these census periods. Most breeding pools are clear and
devoid of vegetation, so it is relatively easy to make
direct counts of individual tadpoles (Schley et al., 1998)
and counts made by different observers are quite con-
sistent (Muñoz, 2000). Mean number of tadpoles coun-
ted in each pool at each site from 1991 to 2000 was
taken as a measure of reproductive success of each
population over this period. However, as the number of
breeding pools used by a population of toads at each
site varies between 1 and 19, reproductive success was
expressed in two ways. First, reproductive success per
population was calculated as the mean number of tad-
poles based on the total tadpole count in all pools at a
site. Second, reproductive success per pool per popula-
tion was calculated by dividing the mean number of
tadpoles counted per population by the number of pools
used by toads in that population.
During the annual censuses the presence or absence of
Natrix maura and Rana perezi was noted. Populations
were classified as having ‘predators present’ if predators
had been observed within 100 m of the population on at
least two separate visits.
2.2. Mapping procedure
Scanned paper maps of the Serra de Tramuntana on
1:5000 scale were obtained from the Conselleria de
Medi Ambient in June 2000. Torrents and 25 m con-
tours were digitised from these scanned maps using
Arcview 3.2 across the entire geographical range of A.
muletensis. From these contours a digital terrain model
(DTM) was constructed using Triangulated Irregular
Networks (TIN). DTMs such as this serve as an impor-
tant tool for deriving secondary spatial data sets such as
altitude, slope and aspect (Stocks and Heywood, 1994).
Slope and aspect surfaces with a grid size of 0.01 km
were generated from the TIN for analysis. The locations
of all toad populations were mapped using information
already available on the geographic location of all
known populations in the form of GPS readings and
paper maps (Garcia and Buley, 1997). To provide a
comparison with torrent habitats not occupied by toad
populations, 50 points were selected along torrents using a
random points generator extension for ArcView 3.2 and by
combining all torrents so that the localities were randomly
located throughout the range of the toad.
2.3. Statistical analysis
Stepwise multiple logistic regression (Hosmer and
Lemeshow, 1989) was used to test a number of topo-
graphical factors considered to be potentially important
in influencing the distribution of the midwife toad and
of the introduced predators N. maura and R. perezi. The
distribution of the two predators amongst toad sites was
 R.D. Moore et al. / Biological Conservation 116 (2004) 327–332
perfectly correlated (r=1.0) indicating that wherever N.
maura are present, R. perezi also occurs nearby and vice
versa. Logistic regression was considered the most
appropriate analytical procedure for assessing the dis-
tribution of toads and predators because it permits the
prediction of binary attributes such as presence/absence
(McCullagh and Nelder, 1983) and it has proved to be a
powerful technique for analysing species distributions
(e.g. Jokimäki, 1999).
The dependent or response variable in the regres-
sion—presence or absence of toads—was scored as 1 (all
torrent-dwelling toad populations; n=17) or 0 (ran-
domly generated points along torrents; n=50), respec-
tively. Because all the random points were generated
along torrents, any toad populations that inhabited
artificial water bodies outside natural torrents were
excluded from the analysis. The independent or expla-
natory variables were three topographical features
associated with each locality derived from surfaces con-
structed using GIS; elevation (m), aspect (in degrees,
where 0 =true north and 180 =true south) and max-
imum slope within a horizontal radius of 100 m (equal
to an area of 31,416 m2). Maximum slope—rather than
mean slope—was chosen as an independent variable as
it was thought that just one steep slope in the vicinity of
breeding site may be limiting for predators, in that it
would prevent emigration of predators from the site. A
radius of 100 m was chosen because this was considered
to provide an appropriate representation of the
immediate topography at each site. Scribner et al.
(2001), for example, also characterised habitat within a
100 m radius around each pond when investigating
environmental correlates of common toad (Bufo bufo)
abundance and genetic diversity. The number of inde-
pendent variables in the analysis was kept low due to
the limited number of sites being compared; this is in
accordance with the suggested minimum requirement of
at least four to five times more cases than independent
variables (Tabachnick and Fidell, 1983). Using the
stepwise logistic regression procedure, selection of vari-
ables for inclusion in the final model was based on the
Wald test (Hosmer and Lemershow, 1989; Jokimäki,
Table 1
Topographical characteristics of torrent localities containing toad populations (toads present) and randomly allocated torrent localities (toads
absent) and results of logistic regression to predict presence/absence of toads at each locality based on the same three topographical features
Variable Toads present (n=17) Toads absent (n=50)
Mean S.D. Mean
Max slope ( )a 69.6 11.4 46.1
Elevation (m)b 313.7 199.9 135.2
Aspect ( )b 149.4 34.1 135.2
df=1 for each analysis. S.D., standard deviation; b standardised regression coefficient; R, unstandardised regression coefficient; P, level of
significance.
a
Variable included in final model.
b
Variables excluded from final model.
329
1999). All variables were normally distributed and
therefore met the assumptions of the analysis.
Factors influencing the distribution of predators were
investigated by comparing landscape topography
between all mapped toad populations (17 inhabiting
natural torrent pools and four inhabiting artificial water
bodies). Seven toad populations with predators were
compared against 14 predator-free toad populations
and predator presence or absence (scored as 1 and 0
respectively) was regressed against the three indepen-
dent topographical variables described above.
In order to assess the factors influencing reproductive
success of A. muletensis over the period 1991–2000,
stepwise linear regressions were made for all toad
populations of (1) mean tadpole count per population,
and (2) mean tadpole count per pool per population
against three site characteristics (tadpole counts were
log-transformed to ensure normality). In the absence of
data on adult population sizes, adult fecundity could
not be estimated and larval counts were considered to
provide the best index of reproductive success between
populations. The site characteristics tested were eleva-
tion, number of pools per site (taken from Garcia and
Buley, 1997) and degree of isolation (calculated as mean
horizontal distance to the five nearest toad populations).
3. Results
3.1. Distribution of toads and predators in relation to
topography
Out of the three variables entered into the forward
stepwise logistic regression model, only maximum
slope—which was significantly positively associated
with toad presence—was included in the overall model
(Table 1). The model was significant (w21=16.2;
P < 0.001) and resulted in 76.1% correct predictions of
toad occurrence. A further stepwise logistic regression
on the same three topographical variables to predict the
presence or absence of predators at toad sites found that
only elevation—which was negatively associated with
Logistic regression based on toad presence
S.D. b R P
18.2 0.07 0.35 <0.001
59.7 – 0.14 >0.05
59.7 – 0.00 >0.05
330 R.D. Moore et al. / Biological Conservation 116 (2004) 327–332

predator presence—was included in the model (Table 2).
The overall model was significant (w21=6.4; P < 0.05)
and resulted in 71.4% of correct predictions of predator
occurrence.
3.2. Factors influencing reproductive success
Multiple linear regression showed that only the num-
ber of pools at each site was a significant predictor of
overall reproductive success of a population: this vari-
able showed a strong positive association with the mean
number of tadpoles counted in all pools at each site over
1991–2000 (Table 3). When reproductive success per
pool per population was regressed against the same
three independent variables, elevation emerged as a sig-
nificant predictor, and displayed a positive relationship
with the dependent variable (Table 3).
4. Discussion
Of the three topographical variables included in the
analysis of toad distribution, only maximum slope was
included in the final model. Toad presence was strongly
positively associated with this variable, indicating that
the immediate relief surrounding a torrent may be
important in dictating its suitability as a breeding site
for toads. Although A. muletensis is an excellent climber
and can negotiate vertical rock faces, it has been sug-
gested that steep sides may make sites inaccessible to
introduced predators (Tonge, 1986; Schley and Grif-
fiths, 1998). However, this theory was not supported by
the lack of a significant negative association between the
presence of predators and maximum slope. An alter-
native explanation is that the steepness of the immediate
terrain influences the microclimate of a site; steep lime-
stone rock faces surrounding a breeding pool may pro-
vide a more shaded and cooler environment that
favours longer hydroperiods and provides optimal con-
ditions for growth and development of tadpoles. The
mean preferred temperature and optimum temperatures
for development of A. muletensis tadpoles are 21.6  C
(Martens, 1984) and 21–24  C (Kadel and Hemmer,
1984) respectively, both of which are low relative to
other European amphibians including A. obstetricans
(Martens, 1984). Pools in open areas are more likely to
be exposed to the sun, making them prone to heating
and desiccation during the summer. Because of the low
optimum temperature for the growth and development
of tadpoles—and since the distribution of the toad is
likely to be constrained by water availability—environ-
ments favouring the persistence of cool plunge pools at
the base of ephemeral waterfalls are most likely to
favour the presence of the toad. Additionally, steep

Table 2
Topographical characteristics of toad populations where predatory snakes and frogs are known to occur (predators present) and those where
predators have never been seen (predators absent) and results of logistic regression to predict presence/absence of predators based on the same three
topographical features

Variable Predators present (n=7) Predators absent (n=14) Logistic regression based on predator presence

Mean S.D. Mean S.D. b R P

a
Elevation (m) 211.5 126.4 437.8 218.8 0.01 0.27 <0.05
Max slope ( )b 62.7 14.7 68.4 12.2 – 0.00 >0.05
Aspect ( )b 141.8 26.5 156.0 62.3 – 0.00 >0.05

df=1 for each analysis. S.D., standard deviation; Beta, standardised regression coefficient; R, unstandardised regression coefficient; P, level of
significance.
a
Variable included in final model.
b
Variables excluded from final model.

Table 3
Results of multiple linear regression of A. muletensis reproductive success (log means of annual counts 1999–2000) against elevation, number of
pools and degree of isolation (mean horizontal distance to the five nearest populations)

Variable Reproductive success per population Reproductive success per pool per population

Standardized Coefficient (b)/Partial Correlation t P Standardized Coefficient (b)/Partial Correlation t P

Elevation 0.37b 1.69 >0.05 0.42a 2.16 <0.05

No. pools 0.66a 16.67 <0.001 0.23b 1.02 >0.05
Isolation 0.05b 0.30 >0.05 0.01b 0.04 >0.05

Significance of overall models: reproductive success per population, R2=0.43; F1 19=14.4; P=0.001; reproductive success per pool per population,
R2=0.20; F1 19=4.7; P< 0.05.
a
Variable included in final model.
b
Variables excluded from final model.
 R.D. Moore et al. / Biological Conservation 116 (2004) 327–332
sides surrounding a site may provide more suitable
habitat for adults, which often take refuge in humid
cracks and crevices within the walls of the torrent
gorges (personal observation).
Within the geographical range of A. muletensis, the
distribution of two introduced predators N. maura and
R. perezi was virtually identical. This suggests that
snake populations may require the presence of R. perezi
to sustain high population numbers and that the pre-
sence of A. muletensis alone is not sufficient. One study
conducted on mainland Spain found that N. maura
preyed heavily upon adults and tadpoles of R. perezi
(Santos et al., 2000), and the presence of the frog may
be a significant factor in the successful establishment
and continued existence of the snake on Mallorca.
Moreover, there was a strong negative association
between predator presence and elevation at toad sites.
Although both the frog and the snake were found toge-
ther at elevations as high as 1000 m (personal observa-
tion), it is likely that the environmental and climatic
conditions associated with lower elevations are pre-
ferred by both predators. N. maura is restricted to lower
elevations in the northern part of its range; in Italy it is
constrained to elevations below 700 m and in the Iber-
ian Peninsula, where it reaches its highest elevation, it is
found up to 1700 m (Gasc et al., 1997).
Although R. perezi is known to occur up to elevations
of 1500 m in the Pyrenees and over 2000 m in parts of
Spain (Gasc et al., 1997), it may be constrained by a
lack of suitable habitat at higher altitudes on Mallorca.
A reduction in predation pressure and competition with
increasing elevation may be an important factor in the
persistence of toad populations in montane torrent gor-
ges and an important consideration in the design of
future reintroduction sites. The distribution of pre-
dators may additionally be influenced by human activ-
ity, and the creation of artificial reservoirs in the vicinity
of natural torrent habitats may increase the chance of
these areas being colonised by predators (Mayol, per-
sonal communication). Data are lacking on the ranges
of N. maura and R. perezi on Mallorca outside localities
occupied by A. muletensis, and this precludes a more
comprehensive analysis of biotic and abiotic factors
influencing their distributions.
The viability of isolated populations of a species
typically depends upon factors such as population size
and degree of inter-population connectivity (Hanski,
1999), which in turn are a function of both physical
distance and the characteristics of intervening habitat
(Forman, 1995). Of the three site characteristics con-
sidered—elevation, isolation and the number of pools
within a site—reproductive success of Mallorcan mid-
wife toad populations was associated most strongly with
the number of pools. This is an important finding
because it indicates that the viability of a population
may be greatly enhanced by increasing the number of
331
potential breeding pools within a gorge. Adult toads are
capable of travelling considerable distances from breed-
ing pools (a distance of 222 m has been recorded;
Román, 2000) and it is highly likely that torrent gorges
provide ideal corridors for the movement of adults.
Individuals seem more likely to move between pools
within the same torrent gorge than to traverse open
areas and mountain tops to reach new potential breed-
ing sites, so the number of pools within a torrent gorge
is likely to be an important factor determining the via-
bility of a population. When the number of tadpoles per
pool was regressed against the same three independent
variables, the latter showed a significant positive asso-
ciation with elevation. There are two possible explana-
tions for this finding. First, environmental and climatic
conditions associated with high elevations may enhance
the viability of toad populations, for the microclimatic
reasons described above. Secondly, because the presence
of introduced predators is negatively associated with
altitude, populations of tadpoles and adult toads may
suffer higher rates of predation at lower elevations.
The results of this study have a number of implica-
tions for the conservation programme of the Mallorcan
midwife toad. As the optimal habitat for the species is
high altitude torrent gorges containing many pools and
surrounded by steep sides, these factors should be con-
sidered in the design and location of reintroduction sites
for the toad. Additionally, the viability of toad popula-
tions may be significantly enhanced by the creation of
more pools within the vicinity of breeding populations,
since overall reproductive success appears to be strongly
associated with the number of pools available. More
data are required on the distributions of N. maura and
R. perezi on Mallorca to allow a better understanding
of the factors influencing the distributions of these
predators.
Acknowledgements
This work was carried out with the co-operation of La
Conselleria de Medi Ambient, who also supplied digi-
tised maps of the Serra de Tramuntana. We thank D.
Jay, V. Muñoz, C. O’Brien, C. Zayas and R. Barber for
logistical support in the field. J. Mayol, T. Beebee and
B. Davis provided helpful comments on earlier drafts of
the manuscript. The work was supported by the Natural
Environment Research Council.
References
Alcover, J.A., Moyà-Solà, S., Pons-Moyà, J., 1981. Les Quimeres del
passat. Els vertebrats fossils del Plio-Quaternari de les Balears i
pitiuses. Editorial Moll Palma de Mallorca.
332 R.D. Moore et al. / Biological Conservation 116 (2004) 327–332
Alcover, J.A., Sanders, E., Sanchiz, B., 1984. El registro fósil de los
sapos parteros (Anura, Discoglossidae) de Balears. In: Hemmer, H.,
Alcover J.A. (Eds.), Historia biologica del ferreret. Editorial Moll
Palma de Mallorca, pp. 109–122.
Bloxam, Q.M.C., Tonge, S.J., 1995. Amphibians—suitable candidates for
breeding-release programs. Biodiversity and Conservation 4, 636–644.
Burbidge, A.A., Manly, B.F.J., 2002. Mammal extinctions on Aus-
tralian islands: causes and conservation implications. Journal of
Biogeography 29, 465–473.
Case, T.J., Bolger, D.T., 1991. The role of introduced species in shap-
ing the distribution and abundance of island reptiles. Evolutionary
Ecology 5, 272–290.
Corsi, F., Duprè, E., Boitani, L., 1999. A large scale model of wolf
distribution in Italy for conservation planning. Conservation Biol-
ogy 13, 150–159.
Detmers, R., Bart, J., 1999. A GIS modelling method applied to pre-
dicting forest songbird habitat. Ecological Applications 9, 152–163.
Diamond, J.M., 1989. Overview of recent extinctions. In: Western, D.,
Pearl, M. (Eds.), Conservation for the Twenty-first Century. Oxford
University Press, New York, pp. 27–41.
Dowding, J.E., Murphy, E.C., 2001. The impact of predation by
introduced mammals on endemic shorebirds in New Zealand: a
conservation perspective. Biological Conservation 99, 47–64.
Farina, A., 1997. Landscape structure and breeding bird distribution in a
sub-Mediterranean agro-ecosystem. Landscape Ecology 12, 365–378.
Forman, R.T.T., 1995. Some general-principles of landscape and
regional ecology. Landscape Ecology 10, 133–142.
Fritts, T.H., Rodda, G.H., 1998. The role of introduced species in the
degradation of island ecosystems: a case history of Guam. Annual
Review of Ecological Systematics 29, 113–140.
Garcia, G., Buley, K., 1997. Report on the Mallorcan Fieldwork 1997.
Jersey Wildlife Preservation Trust (JWPT) Internal Report.
Gasc, J., Cabela, A., Crnobrnja-Isailovic, J., Dolmen, D., Grossenba-
cher, K., Haffner, P., Lescure, J., Martens, H., Martı́nez Rica, J.P.,
Maurin, H., Oliveira, M.E., Sofianidou, T.S., Veith, M., Zuiderwijk,
A., 1997. Atlas of Amphibians and Reptiles of Europe. Societas Eur-
opea Herpetologica. Muséum National D’Histoire Naturelle, Paris.
Gaston, A.J., 1994. Status of the ancient murrelet, Synthliboramphus-
antiquus, in Canada and the effects of introduced predators. Cana-
dian Field Naturalist 108, 211–222.
Hall, J.R., Woods, R.W., Brooke, M.D., Hilton, G.M., 2002. Factors
affecting the distribution of landbirds on the Falkland Islands. Bird
Conservation International 12, 151–167.
Hanski, I., 1999. Habitat connectivity, habitat continuity, and meta-
populations in dynamic landscapes. Oikos 87, 209–219.
Hosmer, D.W., Lemeshow, S., 1989. Applied Logistic Regression.
John Wiley and Sons, New York.
Howell, C.A., Latta, S.C., Donovan, T.M., Porneluzi, P.A., Parks,
G.R., Faaborg, J., 2000. Landscape effects mediate breeding bird
abundance in Midwestern forests. Landscape Ecology 15, 547–562.
Jackson, D.B., 2001. Experimental removal of introduced hedgehogs
improves wader nest success in the Western Isles, Scotland. Journal
of Applied Ecology 38, 802–812.
Janet, F., 1998. Predicting the distribution of shrub species in southern
California from climate and terrain-derived variables. Journal of
Vegetation Science 9, 733–748.
Jokimäki, J., 1999. Occurrence of breeding bird species in urban parks:
effects of park structure and broad-scale variables. Urban Ecosys-
tems 3, 21–24.
Kadel, K., Hemmer, H., 1984. Temperature dependence of larval
development in the Mallorcan midwife toad, Baleaphryne mule-
tensis. In: Hemmer, H., Alcover, J.A. (Eds.), Historia Biologica del
Ferreret. Editorial Moll Palma de Mallorca, pp. 169–174.
Kiesecker, J.M., Blaustein, A.R., 1997. Population differences in
responses of red-legged frogs (Rana aurora) to introduced bullfrogs.
Ecology 78, 1752–1760.
Knutson, M.G., Sauer, J.R., Olsen, D.A., Mossman, M.J., Hemesath,
L.M., Lannoo, M.J., 1999. Effects of landscape composition and wet-
land fragmentation on frog and toad abundance and species richness in
Iowa and Wisconsin, U. S.A. Conservation Biology 13, 1437–1446.
Lenton, S.M., Fa, J.E., Perez del Val, J., 2000. A simple non-parametric
GIS model for predicting species distribution: endemic birds in Bioko
Island, West Africa. Biodiversity and Conservation 9, 869–885.
Martens, H., 1984. Temperature selection in tadpoles of Baleaphryne
muletensis. In: Hemmer, H., Alcover, J.A. (Eds.), Historia Biologica
del Ferreret. Editorial Moll Palma de Mallorca, pp. 163–168.
Martinez-Morales, M.A., Cuaron, A.D., 1999. Boa constrictor, an
introduced predator threatening the endemic fauna on Cozumel
Island, Mexico. Biodiveristy and Conservation 8, 957–993.
McCullagh, P., Nelder, J.A., 1983. Generalized Linear Models.
Chapman and Hall, London.
Muñoz, V., 2000. Assessing Monitoring Methods for the Mallorcan
Midwife Toad, Alytes muletensis. MSc Dissertation, University of
Kent, UK.
Rodda, G.H., Fritts, T.H., 1992. The impact of the introduction of the
colubrid snake Boiga irregularis on Guam lizards. Journal of Her-
petology 26, 166–174.
Rodda, G.H., Sawai, Y., Chiszar, D., Tanaka, H. (Eds.), 1999. Prob-
lem Snake Management: the Habu and the Brown Treesnake. Cor-
nell University Press, Ithaca and London.
Román, A., 2000. Estudi dels mecanismes dispersius del ferreret (Alytes
muletensis). Memòria de beca de formació per a la investigació a la
zona 5.B. Conselleria de Treball i Formació. Govern Balear.
Román, A., 2001. Report on the Fourth International Meeting on the
Alytes muletensis Conservation. 22–23 February 2001.
Román, A., 2002. Recompte de les poblacions naturals i introduı̈des de
ferreret (Alytes muletensis) en totes les localitats (2001). Govern Balear.
Román, A., Mayol, J., 1997. Documents tècnics de conservació. La
recuperación del ferreret, Alytes muletensis. Proyecto life ferreret,
Palma de Mallorca..
Santos, X., Gonzalez-Solis, J., Llorente, G.A., 2000. Variation in the
diet of the viperine snake Natrix maura in relation to prey avail-
ability. Ecography 23, 185–192.
Savidge, J.A., 1987. Extinction of an island forest avifauna by an
introduced snake. Ecology 68, 660–668.
Schley, L., Griffiths, R.A., 1998. Midwife toads (Alytes muletensis)
avoid chemical cues from snakes (Natrix maura). Journal of Herpe-
tology 32, 572–574.
Schley, L., Griffiths, R.A., Román, A., 1998. Activity patterns and micro-
habitat selection of the Mallorcan midwife toad (Alytes muletensis) tad-
poles in natural torrent pools. Amphibia-Reptilia 19, 143–151.
Scribner, K.T., Arntzen, J.W., Cruddace, N., Oldham, R.S., Burke,
T., 2001. Environmental correlates of toad abundance and popula-
tion genetic diversity. Biological Conservation 98, 201–210.
Short, J., Kinnear, J.E., Robley, A., 2002. Surplus killing by introduced
predators in Australia—evidence for ineffective anti-predator adap-
tations in native prey species? Biological Conservation 103, 283–301.
Spence, D., 2002. A Comparison of Fitness in Founder and Descen-
dent Stock of Mallorcan Midwife Toad (Alytes muletensis) Tad-
poles. MSc Dissertation, University of Kent, UK.
Stocks, A.M., Heywood, D.I., 1994. Terrain modelling for mountains
In: Price, M.F., Heywood, D.I. (Eds.), Mountain Environments and
Geographical Information Systems. pp. 25–40.
Tabachnick, B.G., Fidell, L.S., 1983. Using Multivariate Statistics.
Harper and Row, NY.
Tonge, S.J., 1986. Collecting the Mallorcan midwife toad. Oryx 20,
74–78.
Towns, D.R., Daugherty, C.H., Cree, A., 2001. Raising the prospects
for a forgotten fauna: a review of 10 years of conservation effort for
New Zealand reptiles. Biological Conservation 99, 3–16.
Wilson, P.R., Karl, B.J., Toft, R.J., Beggs, J.R., Taylor, R.H., 1998.
The role of introduced predators and competitors in the decline of
kaka (Nestor meridionalis) populations in New Zealand. Biological
Conservation 83, 175–185.