A Recovery Program for the Natterjack Toad

(Bufo calamita) in Britain

JONATHAN S. DENTON,* SUSAN P. HITCHINGS,* TREVOR J. C. BEEBEE,*‡
AND ANTONY GENT†
*School of Biology, University of Sussex, Falmer, Brighton BN1 9QG, United Kingdom
†English Nature, Northminster House, Peterborough PE1 1UA, United Kingdom

Abstract: The natterjack toad ( Bufo calamita) is an endangered species in Britain and has been legally pro-
tected since 1975. This amphibian suffered a major decline during the first half of the twentieth century, due
partly to habitat destruction but mostly to successional changes in its specialized biotopes and anthropogenic
acidification of breeding sites. In addition to site and species protection, extensive autecological research over
the past 25 years has provided the foundations for an intensive, 3-year species recovery program funded by the
statutory nature conservation organizations (English Nature and the Countryside Council for Wales). This pro-
gram was based on habitat management and reintroductions to restored sites and followed similar but less in-
tensive efforts. Management of heath and dune habitats focused on restoration and maintenance of early
stages of seral succession, initially through physical clearance of invasive scrub and woodland vegetation and
later by the reestablishment of grazing regimes similar to those prevalent in earlier centuries. In some cases ex-
tra breeding pools were provided to either increase or stabilize natterjack toad populations that had become
reliant on one or very few pools at small sites or to promote range expansion within large habitat areas. By
1995 proactive conservation work had been carried out at 29 (69%) of the 39 sites with extant native popula-
tions, including 8 during the recovery program. Twenty reintroductions also had been attempted, including
nine during the program. At least six reintroductions resulted in the foundation of expanding new popula-
tions, and an additional eight have shown initial signs of success. Conservation methods developed for Bufo ca-
lamita should provide a useful precedent for long-term conservation of early successional habitats and species.

Programa de Recuperación para el Sapo Natterjack (Bufo calamita) en Gran Bretaña

Resumen: El sapo Natterjack (Bufo calamita) es una especie en peligro de extinción en Gran Bretaña y esta
sujeto a protección desde 1975. Este anfibio sufrió una declinación importante durante la primera mitad del
siglo veinte, debido en parte a la destrucción del hábitat pero principalmente a los cambios sucesionales en
sus biotopos y a la acidificación antropogénica de sitios de reproducción. Además de la protección de sitios y
de la especie, extensas investigaciones autoecológicas en los últimos 25 años proporcionaron las bases para
un programa intensivo de recuperación patrocinado por las organizaciones de conservación de naturaleza
estatutarias (English Nature y Countryside Council for Wales). Este programa se basó en el manejo del hábi-
tat y reintroducciones en sitios restaurados, como continuación de esfuerzos similares pero menos intensivos.
El manejo de brezales y dunas se enfocó en la restauración y mantenimiento de las primeras etapas sucesion-
ales, se inició con la remoción de hierbas y vegetación leñosa invasoras y posteriormente se restablecieron
regímenes de pastoreo similares a los prevalecientes en siglos anteriores. En algunos casos se proporcionaron
charcas para reproducción para incrementar o estabilizar las poblaciones o para promover la expansión del
rango en áreas con hábitats extensos. Para 1995 el trabajo de conservación se había desarrollado en 29
(69%) de los 39 sitios con poblaciones nativas, incluyendo 8 durante el programa de recuperación. También
se habían intentado veinte reintroducciones, incluyendo 9 durante el programa. Por lo menos 6 reintroduc-
ciones resultaron en la fundación de nuevas poblaciones y 8 más han mostrado señas iniciales de éxito. Los
métodos de conservación desarrollados para Bufo calamita deben constituir un precedente útil para la conser-
vación a largo plazo de hábitats en sucesión inicial y de especies.

‡ Address correspondence to T. J. C. Beebee, email T.J.C.Beebee@sussex.ac.uk

Paper submitted September 19, 1997; revised manuscript accepted February 4, 1997.
1329

Conservation Biology, Pages 1329–1338

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1330 Natterjack Toad in Britain Denton et al.

Introduction
In the past few years concern has increased about declin-
ing amphibian populations in several countries around
the world, especially with respect to species living at high
altitudes and in habitats not obviously vulnerable to direct
impacts from human activities (Blaustein & Wake 1990;
Wake 1991; Blaustein et al. 1994a; Pounds & Crump
1994). The extent to which amphibians are declining
more than other groups is the subject of contention,
partly because they seem to have fared no worse than
other taxa (i.e., reptiles in Europe [Beebee 1992] and ver-
tebrates in general [Smith et al. 1993]), but also because
of the inherent difficulty of distinguishing between long-
term declines and natural short-term fluctuations in popu-
lation sizes (Pechmann & Wilbur 1994). Nevertheless,
some species seem to be in trouble; causes include habi-
tat destruction (Cooke & Ferguson 1976; Dodd 1991),
pesticide pollution (Hazelwood 1970), spread of patho-
genic diseases (Laurance et al. 1996), increased road traf-
fic (Fahrig et al. 1995), introduction of alien species such
as Rana catesbiana in Spain (Arano et al. 1995), and en-
hanced UV-B irradiation (due to damage to the atmo-
spheric ozone layer) causing increased egg mortality
Figure 1. Distribution of natterjack toad Bufo calamita
(Blaustein et al. 1994b). It is unlikely, therefore, that am-
in Britain.
phibian declines around the world share a common cause
(Griffiths & Beebee 1992)—multiple factors will often
play a part. Similarly, multiple factors contribute to losses
at a local scale. The UV-B effects, for example, can act syn- anthropogenic acidification of breeding ponds; (4) the
ergistically with both low pH and with infection by patho- enhancement of natterjack populations, accounting for
genic fungi (Kiesecker & Blaustein 1995; Long et al. minimum viable population size (MVP) and metapopula-
1995). tion theory; and (5) reintroductions of natterjacks to re-
The natterjack toad (Bufo calamita) is a native of store the historically documented range.
western and central Europe, but in the northern part of
its range it is confined to a few specific habitat types,
particularly coastal sand dunes, saltmarshes, and inland Background Information
heaths (Beebee 1983; Fig.1). A massive decline during
the early part of the twentieth century resulted in the Extensive autecological studies were initiated during the
species disappearing from more than 75% of its histori- 1980s to determine natterjack habitat requirements and
cal sites in Britain, particularly from those on the heath- the causes of decline. These studies identified aquatic
lands of southern and eastern England (Beebee 1976). In and terrestrial habitat features of critical importance to
the mid-1970s only about 40 natterjack sites (defined as natterjacks. Bufo calamita is adapted to early succes-
discrete areas of terrestrial habitat with varying numbers sional stages that are inherently unstable, such as sand
of breeding ponds) remained. Numerous factors are dunes, upper salt marshes, and river flood plains. The
thought to have contributed to these declines. The spe- adaptations are manifest as two life-history features with
cies was protected by law in 1975, and by 1994 more habitat implications: (1) a specialized feeding behavior
than 80% of localities with natterjack populations also in which invertebrate prey are actively pursued over
had statutory habitat protection as sites of special scien- open ground and (2) small larvae that grow rapidly and
tific interest (Banks et al. 1994). metamorphose after only a few weeks in the pond
We report on the conservation work carried out both (Banks & Beebee 1988; Denton & Beebee 1994). Impor-
before (1970–1992) and during phase one of a species tant terrestrial habitat features for natterjacks include ar-
recovery program (phase one extended over the 3 years eas of bare ground (usually sand) on which to forage and
1992–1995) for natterjack toads in Britain. This included in which to burrow and hibernate; areas of short vegeta-
(1) the restoration of early successional habitats on tion (,30 cm high) sustaining prey invertebrates; and
heaths and dunes; (2) the maintenance of these habitat generally open (unforested) terrain promoting high sum-
stages by sustainable economic methods; (3) countering mer temperatures at ground level. Important aquatic habi-

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Volume 11, No. 6, December 1997

Denton et al.
tat features include shallow, exposed (usually ephemeral)
pools with warm water (.208C) to sustain rapid tadpole
growth rates; circumneutral pH (6.0–8.5); and few pred-
ators or tadpole competitors. Natterjacks also benefited
in the past from the anthropomorphic creation of habi-
tats with these properties, such as heathlands sustained
by timber cutting and livestock grazing.
Direct habitat destruction, including agricultural recla-
mation, afforestation with conifer plantations, and urban
construction, was substantive but not the most impor-
tant reason for the widespread disappearance of Bufo
calamita (Beebee 1977, 1983). Of greater significance
was seral succession on heathlands, and to a lesser ex-
tent on dunes, from the early stages typified by very
open habitat with dwarf shrub or grassy sward commu-
nities to later stages characterized by extensive scrub
and tree invasion. This alteration of habitat structure cre-
ated conditions suitable for the common toad (Bufo
bufo), which has clearly displaced natterjacks over large
areas that, apart from successional changes, are other-
wise little different from historical descriptions (Beebee
1977; Denton & Beebee 1994). Bufo bufo does not sur-
vive well in the early-successional, open conditions of
heaths or dunes, but it encroaches after scrub-mediated
alterations of microclimate. The early spring breeding of
B. bufo produces tadpoles that out-compete those of the
later-breeding B. calamita (Banks & Beebee 1987; Grif-
fiths 1991).
The deterioration of heath and dune habitats due to
successional processes has been extensively documented
in Britain and is associated with major losses of other rare
fauna and flora (Van der Meulen 1982; Marrs et al. 1986;
Webb 1986, 1990) dependent upon these distinctive ec-
osystems. Although coastal dunes represent one of the
few examples of primary habitat remaining in Britain,
most dune systems and probably all heathlands were
used for low-density livestock grazing until the end of the
nineteenth century. Apart from the most exposed loca-
tions, such as frontal dune ridges, this type of manage-
ment now seems essential for their perpetuation (Fuller
& Boorman 1977; Webb 1986). Indeed, the heathlands of
northwestern Europe are almost entirely anthropogenic
in origin and were grazed continuously (and occasionally
burned) after the major forest clearances that created
them some 5000 years ago (Webb 1986). Furthermore,
the oligotrophic and dystrophic pools that form on
leached heathland substrates are highly vulnerable to an-
thropogenic acidification. Natterjacks require circum-
neutral ponds for successful reproduction; in at least one
site a fall in pH due to recent atmospheric pollution ren-
dered the major breeding pool too acidic for natterjack
larval development (Beebee & Griffin 1977; Beebee et al.
1990). It seems likely on the basis of previous natterjack
records and recent pH measurements that acidification
has occurred elsewhere on heathlands and must have af-
fected other elements of the freshwater biota.
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Natterjack Toad in Britain 1331
About 36% of the surviving natterjack populations in
Britain averaged less than 100 adults over 10 years of
monitoring (Banks et al. 1994), giving rise to concern
that they may be below minimum viable population sizes
(Laan & Verboom 1990; Nunney & Campbell 1993). Frag-
mentation and isolation of surviving populations has also
increased, and recent investigation has revealed very low
levels of genetic diversity in British B. calamita popula-
tions (Hitchings & Beebee 1996), posing long-term risks
from environmental, demographic and genetic stochas-
ticity. Adult population density of B. calamita is directly
related to breeding pond density (Beebee et al. 1996),
and it is often possible to increase population size by dig-
ging more ponds. Also, amphibians may exist as metapo-
pulations in which source and sink breeding pools some-
times coexist within interconnected habitat patches (Gill
1978; Sjögren 1991). Creation of a network of ponds of
varying depths should improve prospects for maintaining
metapopulations because one or more ideal depth (deep
enough to persist until metamorphosis, but not so deep
as to harbor large numbers of predators) is likely to be
available every year irrespective of weather conditions.
Although work on natterjack toad conservation started
in the 1970s, the concept of undertaking recovery pro-
grams for threatened species in Britain was only recently
promoted by the statutory conservation organizations
(Whitten 1990). In 1991 the species recovery program
was launched by English Nature; by 1996 the program en-
compassed almost 100 species projects. The projects have
three phases: “pre-recovery” in which recovery feasibility
is assessed; “phase 1” in which intensive work to improve
status is carried out, usually over 3–5 years; and “phase 2”
in which follow-up work over at least a decade consoli-
dates and secures the gains made during phase 1. Bufo ca-
lamita was among the first suite of species included in the
recovery program because of the large amount of auteco-
logical information already at hand by 1991.
Rationale and Methods
Restoration of Early Successional Habitats on Heaths
and Dunes
In the absence of management or as a consequence of
anthropogenic stabilization of habitats (such as preven-
tion of dune accretion by beach cleaning), many natter-
jack sites have progressed to late stages of seral succes-
sion. A variety of evidence was used to assess what the
habitat had formerly been like at these sites and to guide
objectives for restoration. This included photographic
records of successional change, accounts by local natu-
ralists, and specific information about changes in natter-
jack populations or in numbers of competitor species.
On both heaths and dunes, restoration involved re-
moval of invasive scrub and woodland, mostly buckthorn
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1332 Natterjack Toad in Britain
(Hippophae rhamnoides) and birch (Betula spp.) on
dunes, and birch, Scots pine (Pinus sylvestris), and gorse
(Ulex europaea) on heathland. In dune systems this was
often achieved by means of heavy machinery, mostly
track vehicles with blades or grab-arms, leaving disturbed
grass and herb layers with extensive areas of bare sand
but without destroying the soil profile. These restorations
created habitat structures similar to those of early succes-
sional stages of the kind still maintained naturally by wind
action on exposed frontal dune ridges. Natural accretion
was encouraged, where possible, by the cessation of
beach-cleaning operations and by restricting access by
motor vehicles. On heathland in which the heather
(mostly Calluna vulgaris) understory survived, scrub
was usually removed by manual cutting to minimize dam-
age to the shrub layer. This was necessary because heath-
ers take several decades to reach maturity and are easily
killed by mechanical trampling, which leaves the habitat
unsatisfactory for many of the rarer species. In this habitat
areas of exposed substrate were created by rotovation of
patches or tracks interwoven among stands of undis-
turbed heath. Stumps of deciduous scrub or tree species,
if not removed, were treated with herbicides to prevent
regrowth. All cut vegetation was burned to remove as
many nutrients as possible from the ecosystem.
Maintenance of Heaths and Dunes by Sustainable
Economic Methods
Long-term maintenance of early successional habitats by
economically sustainable methods was an essential objec-
tive after completion of restorative management. The re-
introduction of management regimes that achieved this
aim in the past—notably low-density (,1 animal/3 ha)
livestock grazing to control or prevent scrub regenera-
tion—was the method of choice. Sheep or cattle were
used on different sites; some were grazed for all and oth-
ers for part of the year. Ease of establishing grazing re-
gimes varied. Some dune and upper salt marsh sites have
always been grazed, but others required prior erection
of stock-proof fencing and acquisition of animals, gener-
ally on a lease-lend basis with local farmers.
Countering Anthropogenic Acidification of Breeding Pools
Natterjacks require circumneutral pools for successful
development of their spawn and tadpoles. Anthropo-
genic acidification is considered a major threat on heath-
lands, where the oligotrophic pools are weakly buffered.
Direct evidence of acidification is expensive and time-
consuming to obtain, but it was available for one site
(Beebee et al. 1990). Elsewhere, inferences about likely
acidification were made on the basis of historical infor-
mation about water quality, especially records of plants
typical of circumneutral water and of previous successful
use by natterjacks within the past century. Two methods
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Denton et al.
were employed to restore acidified heathland ponds to
circumneutrality. At three sites the minimum quantities
of Ca(OH)2 necessary to raise pH to between 6.5 and 7.0
were added to ponds in powder form early in the spring
before toad breeding started. Amounts were kept as low
as possible to avoid the development of calciphile com-
munities, which would be atypical of heathlands, and it
was generally necessary to make additions every year or
every second year to maintain the circumneutral pH
necessary for natterjack development. A second method
at one large site exploited the shallow and ephemeral
nature of natterjack breeding ponds. In this case the thin
silt bed with its accumulated sulphate load was scraped
away when the large (10-ha) pond desiccated in sum-
mer. After refilling, this pond had a pH of over 5.5, com-
pared with less than 4.5 before scraping, for several
years afterwards (Beebee et al. 1990).
Enhancement of Population Size
Small, isolated populations are especially vulnerable to
extinction, which has significant management implica-
tions. Population size is related to availability of breed-
ing habitat (Beebee et al. 1996); consequently, enhance-
ment of populations as well as development of a functional
metapopulation may be possible through pond creation.
Minimum viable population size is not known for this spe-
cies, although colonies of less than 100 adults have per-
sisted for many generations, and this number was adopted
as a minimum target size for all populations pending fur-
ther study. Pond creation was therefore considered appro-
priate at all sites with small toad populations and where
there was evidence of lowered water tables, which re-
duces the breeding success of toads at existing ponds.
Natterjacks favor ephemeral pools with very shallow
margins that desiccate around midsummer in a year of
average rainfall. Pond excavation was usually carried out
by means of heavy machinery, with care being taken to
generate basins of appropriate shape and depth (less
than 1 m maximum depth in midwinter) and in particu-
lar to avoid overdeepening. Ponds created on heathland
sites were in some cases lined with concrete to counter
acidification and retain water above natural water tables.
New ponds were subsequently monitored both for use
as spawning sites and also to determine whether meta-
morphosis was successful. The former was quantified by
counting of spawn strings and the latter by estimation,
to within an order of magnitude, of toadlet production.
B. calamita females usually deposit a single spawn string
each year, laid in shallow water (,10 cm) and not inter-
twined with those of other animals (Beebee 1983). Count-
ing spawn strings is therefore a useful method for estimat-
ing adult numbers (sex ratios are usually about unity). The
long-term effects on adult population size, judged by total
spawn string production at a site, were also monitored
over the years subsequent to pond creation.

Denton et al.
Reintroductions to Restore Historical Range
An important objective of the recovery program was to
reestablish the species at a series of representative sites
that would, as far as possible, reconstitute its historical
range. This procedure falls within the definition of rein-
troduction accepted by the World Conservation Union
(1994). Priority was given to localities on heathlands in
southern and eastern England, following as close as pos-
sible to well-documented historical records. Sites were
chosen in which habitat was judged either already suit-
able or easily made so. Management was carried out as
necessary to restore both terrestrial and aquatic habitat
features, and then natterjacks were introduced as spawn.
The equivalent of two full strings, about 5000–6000 eggs,
but as samples from several strings to maximize genetic
diversity, were translocated per year for 2 consecutive
years. These were usually taken from the nearest native
population on the same habitat type. Survival to meta-
morphosis was monitored by estimating toadlet emer-
gence to within an order of magnitude. Re-introduction
success was measured against three criteria: (1) initial
success, when at least tens of toadlets emerged in at least
one of the two reintroduction years; (2) intermediate
success, when adults returned to breed within 3 years
(the age of sexual maturity) of the beginning of reintro-
duction; and (3) complete success, when breeding con-
tinued at the site for at least 5 years, adult numbers re-
mained stable or increased over that time, and toadlets
from second-generation spawn were produced. A failure
was ascribed to any attempt that did not result in adults
returning and breeding successfully within 10 years of
the first spawn translocation.
Results
Restoration and Maintenance of Terrestrial Habitats
Problems from successional processes were more acute
in some habitats than others (Table 1). All the few re-
maining heathland sites were affected, about 50% of
dune sites but less than 10% of salt marshes. The contin-
uation of livestock grazing on most salt marshes and on
many dune systems largely accounted for the lower lev-
els of scrub invasion, and conservation effort was tar-
geted at the heath and dune sites most at risk. These in-
cluded two of the three remaining heathland sites,
where in each case tens of hectares of pine, birch, and
gorse were removed. Common toad invasion at two of
the heathland sites was lower than on many others
where succession had progressed, apparently for site-
specific reasons (heavy grazing by wild rabbits [Oryctol-
agus cuniculus] at one, acidification of all the deeper
pools at the other), probably explaining why natterjacks
persisted there. By contrast, common toads were so nu-
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Natterjack Toad in Britain 1333
Table 1. Restoration and maintenance of natterjack toad sites in
Britain since 1970.*
Salt
Dune Heath marsh Other
sites sites sites sites Total
Total no. native sites 21 3 11 4 39
No. with substantial
scrub encroachment 11 3 1 1 16
No. with scrub clearance 6 (3) 2 (1) 1 1 10 (4)
No. always grazed 8 0 9 2 19
“Always grazed” sites
with B. bufo
incursion (%) 0 — 0 0 0
Ungrazed sites with
B. bufo incursion (%) 77 33 50 50 65
No. sites recently grazed
again 5 1 0 1 7
* Numbers in parentheses are of sites where natterjack numbers or dis-
tribution have increased commensurate with clearance operations.
merous at the third heathland site that manual removal
of adults and spawn was necessary for more than 10
years to allow any survival of natterjack tadpoles during
scrub removal to restore early successional conditions.
Scrub problems were less widespread on dunes, partly
because successional processes are inherently slower in
exposed coastal areas and partly because about 40% of
dune sites were still grazed by domestic livestock. Scrub
invasion on some dune systems became acute in recent
decades, however, and in one particular locality on the
Merseyside coast the alien shrub Hippophae rhamnoides
spread so rampantly that B. calamita was replaced by B.
bufo over more than 70% of the dunes between 1970 and
1985.
Although open habitats are provided by initial distur-
bance of the ground during scrub clearance, further
management is required to sustain the habitat structures
required by B. calamita and other key elements of early
successional fauna and flora. In the absence of grazing, it
was necessary to carry out clearances every few years.
B. calamita increased in numbers or extended its range
on four sites with extensive clearances. Trends at four of
the managed sites are not yet clear, although natterjacks
certainly persist, whereas at the other two managed
sites clearance was less complete and natterjacks have
continued to decline. By 1995 scrub clearance had been
carried out at 10 of the 16 sites deemed to require it, in-
cluding three large-scale exercises during phase 1 of the
recovery program. In view of the endangered status of
B. calamita, no sites were left deliberately uncleared as
controls because early work indicated the importance of
open habitats to this species (Beebee 1977), but natter-
jack populations remained precarious or were maintained
by artificial methods (such as removal of common toads)
at the six sites where scrub removal was not possible.
Sites where grazing was maintained appeared to be
much poorer habitats for common toads than were un-
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1334 Natterjack Toad in Britain
grazed sites (Table 1). Grazing has persisted in more
than 80% of salt marshes, and in this habitat natterjack
populations generally thrived best, needing little or no
scrub control. Over the past 10 years grazing regimes
have been reestablished at seven sites with a view to
maintaining early successional stages in the long term,
including two initiated during the recovery program.
One major area of natterjack habitat (a large section of
the Merseyside dunes) was targeted before and during
the recovery program. Clearfelling of a pine plantation
and reintroduction of sheep grazing was achieved on
part of the site, whereas scrub removal from another 40
ha was completed in preparation for grazing in future.
Much more time will be needed before it will be possi-
ble to make a proper assessment of the effects of the
new grazing regimes on natterjack toads, but studies
comparing grazed and ungrazed habitats have proved
encouraging with respect to both utilization by this spe-
cies and to decreased occupancy by B. bufo (Denton &
Beebee 1996) and the reappearance of other species
typical of early successional stages.
Restoration and Management of Aquatic Habitats
Anthropogenic acidification of breeding ponds was spe-
cifically identified at one of the surviving heathland sites
of B. calamita and has probably affected others from
which the species has disappeared (Beebee et al. 1990).
Although control of pollutant emission and consequent
relief of ecosystem acid loads must remain the long-term
goal, this is unlikely to be realized within the time neces-
sary for amphibian conservation at sensitive sites (Jug-
gins et al. 1996). During the 1980s, therefore, addition
of quicklime (annually) or removal of bottom sediment
(as a single exercise) were carried out on the acidified
site at two separate pools with pH of less than 4.5. Both
treatments restored pH to circumneutrality, and both
ponds were subsequently used for spawning by natter-
jacks. Tadpole mortality in the two ponds remained
higher than in naturally circumneutral pools, however,
apparently because numbers of invertebrate predators
became exceptionally large, and no metamorphic suc-
cess has yet occurred in either pond (Banks et al. 1993).
In contrast, neutralization of pools with quicklime at an-
other heathland site prior to a reintroduction during the
mid 1980s was successful, and a natterjack population to-
tally dependent on these pools has become established
there; lime addition has also created conditions that sup-
port natterjack development through to metamorphosis
in several previously acidic ponds in a third area.
A common management approach at many sites has
been to create totally new ponds (Table 2). This was of-
ten necessary to rescue populations because historical
breeding sites were damaged or destroyed, but another
frequent objective was to increase adult population
sizes. The intention in these cases also was to stabilize
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Denton et al.
Table 2. Pond creation at natterjack toad sites in Britain since 1970.a
Effects on population
Total no. No. sites with No
Habitat sites ponds added effect Increase Rescue
Dune 21 17 5 1 10
Heath 3 3 1 1b 2
Salt marsh 11 5 1 2 0
Other 4 1 0 1 0
Total 39 26 7 5 12
a
Sites at which the effects of ponds have not been well recorded were
omitted from the analysis.
b
At this site initial rescue with one pond was followed by population
increases after addition of extra ponds over a period of 20 years.
metapopulation structures and thus reduce the probabil-
ity of local extinction due to stochastic factors. Pond
creation was therefore usually targeted to localities
where population size was small (,100 adults) or en-
dangered by pond loss. Because most natterjack popula-
tions fell into one of these categories, over the past 25
years over 200 ponds have been created, spread be-
tween about 67% of the extant sites, including 15 con-
structed during the recovery program. At all sites at least
one and usually most of the ponds were used by natter-
jacks within 1 or 2 years of their creation. In about half
of the sites the new ponds rescued the populations from
probable extinction, as judged by the absence of suc-
cessful reproduction in any other water body. Overall
more than 65% of populations provided with new ponds
were rescued or increased (as measured by spawn string
counts) in the years following pond creation. This type
of management has proved successful on all habitat
types, but not every population responded with substan-
tive size increases. Failures could be attributed to poor
siting (two instances), inadequate terrestrial habitat man-
agement accompanying pond creation (two instances),
or overdeepening after misjudgement of water-table low-
ering (three instances). Efforts are currently being made
to rescue populations using ponds suffering from the wa-
ter-table problem by making the pools shallower. To be
successful, therefore, restoration or creation of ephem-
eral ponds requires careful planning and execution, in-
cluding measurement of water-table fluctuations over
several seasons.
Reintroduction
Returning natterjack populations to areas where they re-
cently have become extinct has proved increasingly suc-
cessful over the past 25 years. Out of 20 attempts at rein-
troduction, only 6 have definitely failed according to the
criteria we defined (Table 3), and 5 of these were the
first efforts of the early 1970s. All three reintroductions
to dune sites have succeeded within the limits of judge-
ment possible at the present time, although only one is

Denton et al.
Table 3. Reintroductions of natterjack toads in Britain since 1970.*
No. on No. on
Assessment dunes heaths
Total attempted 3 17
Intitial success 3 (100) 11 (66)
Intermediate success 2 (100) 5 (45)
Complete success 1 (100) 5 (45)
Definite failures 0 6
* Parentheses indicate percentage of success of those reintroductions
underway long enough for proper comparison.
more than 5 years old, and it (initiated in 1982) has es-
tablished one of the largest natterjack populations on
the east coast of England. The 6 unsuccessful efforts, all
on heathland, failed by the first criterion, and no suc-
cessful metamorphosis was recorded at any of them for
reasons subsequently related to poor water quality in
the ponds. Of the remaining 11 attempts on heathland, 5
have been completely successful and 6 are too recent
(,3 years old) for anything other than a preliminary as-
sessment, although all produced toadlets in their first
year and have thus satisfied the first criterion.
New ponds were created at all three of the dune sites
simply by excavating below the water table, whereas on
heathland the situation was more complex. All the fail-
ures utilized natural water tables, whereas only 4 of the
11 more successful ventures relied on this source. Three
of these 4 are 7 or more years old and have generated
self-perpetuating natterjack populations, although one
site requires occasional addition of quicklime to main-
tain circumneutral pHs in the two breeding ponds. One
of the other 7 sites, the first successful natterjack trans-
location (started in 1980), has created another very large
population in eastern England that relies entirely on a
single pool with a butyl plastic liner. The patterns of
Figure 2. Population size development of natterjack
toads at reintroduction sites.
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Natterjack Toad in Britain 1335
population growth at this site and at the first dune rein-
troduction site, as judged by spawn string counts, are
Total
no. shown in Fig. 2. The remaining 6 sites, including one
longstanding success, (.10 years) have concrete-lined
20
14 (70)
ponds. Supportive management for reintroductions has
7 (54) included scrub clearance at 2 of the 3 dune and 7 of the
6 (50) current 11 heath sites and the establishment of grazing
6 (30) at 1 dune and 2 heath sites.
Discussion
Conservation measures for natterjack toads in Britain
have improved in effectiveness over the past 25 years as
critical habitat features have been identified and the
much broader issues associated with habitat manage-
ment were successfully addressed. It is in the arena of
early successional habitat conservation that work on B.
calamita has proved particularly relevant. It is well
known that open habitats are often maintained by graz-
ing regimes (Dix 1959; McCune & Cottam 1985); many
such ecosystems in temperate zones arose following an-
thropogenic forest clearances and have been maintained
by regular disturbance ever since, although some were
developed naturally by large herds of wild herbivores
(Van Wieren 1995) or wind erosion on exposed coastal
sites. Even on these, however, grazing by domestic ani-
mals has played a part and has become more important
in recent decades following declines in wild herbivores
and the introduction to many sites of alien tree and
scrub species (such as Rhododendron ponticum and
Hippophae rhamnoides). The coincidence of these
plant introductions with the mass extermination of rab-
bits, the main natural herbivore of these habitats, during
the mid-twentieth century by myxomatosis infection led
to widespread overstabilization of dune systems in Britain
(Fuller & Boorman 1977). In addition, livestock farming
on such marginal land became increasingly uneconomi-
cal and mostly ceased by the early years of the twentieth
century. Conservation of heaths and dunes therefore re-
quires the clearance of a large backlog of scrub and tree
invasion, followed by reinstatement of management prac-
tices necessary to maintain early successional habitats,
most especially low-density livestock grazing. Fire, al-
though used elsewhere and occasionally used on heath-
land to restore early successional stages, has deleterious
long-term effects on vegetation structure and is generally
less appropriate than the use of livestock (Bullock &
Webb 1995). Although grazing is unlikely to be economi-
cal from the point of view of commercial agriculture, it
seems likely to be the cheapest as well as the most effec-
tive way of maintaining such habitats for conservation
purposes. After the initial costs involved in perimeter
fence installation and livestock purchase are met, mainte-
nance management by grazing should at least break even
and may even be marginally profitable (Kottmann et al.
Conservation Biology
Volume 11, No. 6, December 1997

1336 Natterjack Toad in Britain
1985). Comparable situations elsewhere in the world are
attracting similar solutions (e.g., wetland management for
bog turtles [Clemmys muhlenbergii; Tryon & Herman
1990]), but there is still considerable scope for experi-
mentation with grazing regimes. It will certainly be im-
portant to establish the most appropriate times of year to
use livestock and to avoid overgrazing and the range of
different problems (including vegetation damage and im-
poverishment of reptile fauna) that can result from it
( Jones 1981; Fleischerer 1994; Noss 1994).
At the species level, short-term measures for B. calam-
ita have mainly involved pond creation or restoration.
Mostly, ponds have been put in place following destruc-
tion of or serious damage to the original breeding sites,
including the lowering of the water table by land drain-
age, overgrowth with scrub or rank vegetation, and an-
thropogenic acidification. These were therefore essen-
tial rescue measures and have largely succeeded in their
aim of maintaining viable natterjack populations, at least
for a few years. A secondary objective was to increase
toad numbers at many sites, taking account of the need
to maintain assumed minimum viable population sizes
(of about 100 adults) and adequate metapopulation
structures in the face of long-term threats from environ-
mental, demographic, or genetic stochasticity (Nunney
& Campbell 1993; Harrison & Hastings 1996). Evidence
shows that natterjack population size is limited by pond
availability and that increasing pond numbers can have
positive effects on population size within a decade (Banks
et al. 1993; Beebee et al. 1996). But at more than half the
sites where pools have been created for this purpose
there is as yet no evidence of a significant change in toad
population size. As discussed earlier, reasons for these
failures include inappropriate installations and insufficient
terrestrial habitat management.
We believe that two particularly important points
have emerged from this recovery program: (1) it is es-
sential to take into consideration all aspects of a species’
ecological requirements and to ensure that management
is in place to address them and (2) the conservation of
threatened species requires a holistic approach at the
habitat scale. Thus, while certain small-scale habitat ma-
nipulations, such as the creation of breeding pools for B.
calamita, may be justified for the conservation of spe-
cific species, they are unlikely by themselves to be suffi-
cient to ensure the long-term survival of populations.
Our approach has been to integrate specific habitat man-
agement needs to allow the persistence of this species
within the objectives of managing for early successional
communities.
Reintroduction is a stringent test of autecological knowl-
edge, and success rates with B. calamita have improved
commensurate with ecological study. Reintroductions
have therefore constituted valuable bioassays of the con-
servation management methods described earlier. Be-
cause historical localities for amphibians are often not
Conservation Biology
Volume 11, No. 6, December 1997
15231739, 1997, 6, Downloaded from https://conbio.onlinelibrary.wiley.com/doi/10.1046/j.1523-1739.1997.96318.x by Consorci De Serveis Universitaris De Catalunya, Wiley Online Library on [19/04/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Denton et al.
known precisely, reintroductions were often made to a
general area from which natterjacks were recorded,
rather than, for example, to a specific pond or marsh.
Success rates of amphibian reintroductions have been in
contention (Dodd & Seigel 1991; Burke 1991), but in
general they have been higher for common rather than
rare species. Early efforts with B. calamita were justi-
fied primarily as rescue operations and used animals
from sites in the process of destruction; they failed be-
cause aquatic and terrestrial habitat requirements were
not well understood. Since 1980, however, reintroduc-
tions have mostly been successful by the criteria out-
lined in the section on rationale and methods; during
the recent recovery program confidence was high
enough to initiate 8 new sites. At all of the 15 reintro-
duction sites initiated since 1980, ancillary management
operations have also been carried out to restore or im-
prove terrestrial or aquatic habitats, including scrub
clearance (8 sites), pond creation (all 15 sites), and the
introduction of grazing regimes (3 sites). The intention
was to improve the quality of the entire ecosystem, and
thus benefit a wide range of species, as part of the nat-
terjack reintroduction program.
It will of course be necessary to develop and execute
conservation plans for B. calamita beyond phase 1 and
into phase 2 of the formal recovery program. Such de-
velopments are currently well advanced, and a full-time
supervisory officer has been employed to oversee an ex-
pansion of management methods, especially the restora-
tion of livestock grazing regimes. Partnerships between
many different people and organizations have been and
will continue to be essential for success. These include
statutory bodies, voluntary organizations, and private in-
dividuals. Other recovery programs world-wide have
demonstrated the value of intensive treatment for a rela-
tively short time that leads to longer-term strategies, in-
cluding ones for endangered amphibians (Alytes mu-
letensis; Bloxham & Tonge 1995).
Acknowledgments
More than 50 people assisted with natterjack toad con-
servation and the national recovery program and con-
tinue to do so. Special thanks are due to J. Buckley and
W. Boyd for data collection on translocation sites, and
English Nature, the Countryside Council for Wales, the
WorldWide Fund for Nature, the Ministry of Defence,
Sefton Metropolitan Borough Council, and the Herpeto-
logical Conservation Trust for financial support.
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