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John W. Wilkinson
Vol ume 11
Status of Conservation and Decline of Amphibians:
Published by Pelagic Publishing
PO Box 874, Exeter, EX3 9BR, UK
Amphibian Biology, Volume 11: Status of Conservation and Decline of Amphibians: Eastern
Hemisphere, Part 5: Northern Europe
ISBN 978-1-78427-016-2 (Pbk)
ISBN 978-1-78427-017-9 (ePub)
ISBN 978-1-78427-018-6 (ePDF)
Copyright © 2019 Pelagic Publishing
This book should be quoted as Heatwole, H. and Wilkinson, J.W. (eds) (2019) Amphibian Biology,
Volume 11: Status of Conservation and Decline of Amphibians: Eastern Hemisphere, Part 5: Northern
Europe. Exeter: Pelagic Publishing.
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British Library Cataloguing in Publication Data.
A catalogue record for this book is available from the British Library.
Cover image: The naerjack toad Epidalea calamita is a European endemic. The image shows a
calling male in the city of Jena, Thuringia, Germany. In highly industrialized countries with a
dense human population, the species often colonizes secondary habitats like quarry pits or waste-
land within large cities. Photo: Andreas & Christel Nöllert.
Table of contents of volume 11, Amphibian Biology:
Eastern Hemisphere, Part 5 (Northern Europe)
54 Status and conservation of amphibians in Luxembourg 1
Laura R. Wood, Edmée Engel, Richard A. Griths, Roland Proess and Laurent Schley
55 Status and conservation of amphibian species in the Federal Republic of
Richard Podloucky and Andreas Nöllert
56 Conservation and declines of amphibians in Poland 26
Maciej Pabijan and Maria Ogielska
57 Amphibian conservation in Switzerland 46
Benedikt R. Schmidt and Silvia Zumbach
58 Amphibian declines and conservation in Austria 52
59 Conservation and decline of European amphibians: The Czech Republic 58
Lenka Jeřábková, Martin Šandera and Vojtech Baláž
60 Amphibian declines and conservation in Slovakia 68
Ján Kautman and Peter Mikulíček
61 Conservation status of amphibians in Norway 76
Leif Yngve Gjerde
62 Conservation measures and status of amphibians in Sweden 92
63 Decline and conservation of amphibians in Finland 109
Ville Vuorio and Jarmo Saarikivi
64 Decline and conservation of amphibians in Estonia 115
65 Decline and conservation of amphibians in Latvia 122
Aija Pupina, Mihails Pupins, Andris Ceirans and Agnese Pupina
66 Amphibian declines and conservation in Lithuania 142
Giedrius Trakimas, Jolanta Rimšaitė and John W. Wilkinson
67 Decline and conservation of amphibians in Denmark 149
Kåre Fog, Lars Christian Adrados, Andreas Andersen, Lars Briggs, Per Klit Christensen,
Niels Damm, Finn Hansen, Martin Hesselsøe and Ue Mikkelsen
65 Decline and conservation of
amphibians in Latvia
Aija Pupina, Mihails Pupins, Andris Ceirans and Agnese Pupina
Abbreviations and acronyms used in the text and references
Bd Batrachochytrium dendrobatidis
Bsal Batrachochytrium salamandrivorans
IUCN International Union for the Conservation of Nature
GE genomic equivalents
RDB Red Data Book of Latvia
HD Habitats Directive
LEGM Latvian Environment, Geology and Meteorology Centre
MKN Ministru Kabineta Noteikumi (the documents, approved by the Cabinet of
Ministers of Latvia)
qPCR quantitative polymerase chain reaction or real-time polymerase chain reaction
I. Introduction: The country and its amphibian fauna
Latvia – ocially the Republic of Latvia (in Latvian: Latvija, Latvijas Republika) – is a country of
the Baltic region of northern Europe and lies on the east coast of the Baltic Sea and its Gulf of Riga.
Latvia has been a member of the European Union since 2004. The total area of Latvia is 64,589 km2;
its length from north to south is 210 km and from east to west is 450 km. Latvia has land borders
with Estonia in the north (343 km), with Lithuania in the south (588 km), with Belarus in the
southeast (161 km), and with Russia in the east (276 km) (Central Statistical Bureau Republic of
Latvia 2012). Latvia also shares its borders with Estonia, Sweden and Lithuania in the Baltic Sea.
A relatively large proportion of Latvia’s perimeter, relative to its surface area, transects various
natural habitats also shared by neighbouring countries. This ecological continuity with adjacent
countries is reected in similarities in their amphibian faunas.
The largest part of the territory of Latvia is at a height of not more than 100 m above sea level.
Latvia has a temperate seasonal climate that can also be described as oceanic/maritime (Central
Statistical Bureau Republic of Latvia 2012). Latvia has four well-dened seasons of approximately
equal duration. In winter the average temperature is –6°C, with minima of –20° to –30°C, usually
lasting for several weeks; winter conditions include a short day-length, a constant mantle of snow
up to 0.7 metres deep, total freezing of small ponds, and persistent ice 15–20 cm thick on larger
ones. Such conditions aect survival of amphibians overwintering either underwater or on land,
Decline and Conservation of Amphibians in
I. Introduction: The country and its
II. Threats to amphibians in Latvia
III. Declining species and species of
special conservation concern in
IV. Conservation of amphibians in Latvia
V. Monitoring of amphibians in Latvia
Decline and Conservation of Amphibians in Latvia 123
especially in their rst year of life, and declines occur during winter. The average air temperature
is 19°C in summer, with maxima of 25°–30°C commonly occurring, especially in the country’s
interior. It often rains, especially in the second part of the summer (Latvian Biodiversity Clearing-
House Mehanism 2015) (Figure 65.1), thereby creating numerous large and small ponds and other
wetland habitats used by amphibians.
There is a moderate marine climate in the western coastal regions of Latvia, with warmer win-
ters and cooler summers than in the eastern continental part of the country; this regional dierence
in climate inuences the distribution of the amphibian fauna within Latvia.
Of the total surface of Latvia, agricultural land covers 23, 909 km2 (44.0%) and forest 29, 996
km2 (55.2%); inland fresh waters cover the remaining 419 km2 (0.8%). The largest lake is Lubans
with a surface area of 80.7 km2; the deepest lake is Dridzis with a depth of 65.1 m. The longest river
in Latvia is the Gauja, which is 452 km long. The longest river owing through the territory of
Latvia from Belarus is the Daugava, which has a total length of 1,005 km of which 352 km are
within the territory of Latvia. There are many lakes, rivers, large and small ponds and other small
bodies of water used by amphibians as habitats and migratory routes.
The average density of the human population is 34.3/km2. There are ve economic/political
planning regions in Latvia: Kurzeme, Latgale, Riga, Vidzeme and Zemgale. They dier in relief,
climate and level of economic and agricultural development, as well as in urban saturation which
is important for the composition and distribution of the amphibian fauna.
Nowadays amphibians are represented by 13 permanently resident species in Latvia – 11 anu-
rans and two caudates; all have Least Concern IUCN Red List Status for Europe, but with some
species exhibiting changes in population density in Latvia (Table 65.1).
Latvia is the most northern country of the European Union in which the European re-bellied
Toad Bombina bombina resides, and the species reaches the northern extreme of its distribution
there (Kuzmin et al. 2008). Latvia is also the most northern extent in the EU of the European tree-
frog (Hyla arborea). Hyla arborea was ocially declared as extirpated in Latvia but was successfully
reintroduced into the country using zoocultures established from animals caught in Belarus
Four species – the green toad Bufotes viridis, the naerjack toad Epidalea calamita, the common
spadefoot toad Pelobates fuscus and the northern crested newt Triturus cristatus – reside in Latvia
near the northern borders of their European distribution. Epidalea calamita is encountered mainly
near the coast of the Baltic Sea and in central Latvia; over the past few years it has been observed
in the southeastern part of the country as well. The other three of these species are encountered
Fig. 65.1 Long-term monthly averages of air temperature and precipitation in Latvia (Latvian Environment, Geolo-
gy and Meteorology Centre 2015).
124 Amphibian Biology
throughout most of Latvia, except in the north, but their populations are mostly small and frag-
Four other Latvian amphibians are abundant over the entire country and occur in a wide range
of habitats: the smooth newt Lissotriton vulgaris, the common toad Bufo bufo, the common frog Rana
temporaria and the moor frog Rana arvalis. Pelophylax spp. are also widely abundant throughout
Latvia and are even encountered in urbanized habitats (Rimicans et al. 2010; Pupins and Pupina
2011a) (Table 65.2).
II. Threats to amphibians in Latvia
According to the ocial national requirements for development of Species Conservation Plans,
factors posing threats to the habitats, ecosystems and populations of the protected species in
Latvia can be divided into those of natural origin and those of anthropogenic origin. These can,
of course, act synergistically. Threats of either kind are of less concern for amphibians that are
widespread in Latvia, however, than they are for species that are represented by small or frag-
A relatively cold climate is a permanent natural cause of paucity and limited abundance of
amphibian species in Latvia, such as Triturus cristatus, Bombina bombina, Hyla arborea, Bufotes viridis,
Epidalea calamita and Pelobates fuscus. Of course, all species of Latvian amphibians are generally
adapted to the Latvian climate as conrmed by their persistence there over a long period of time.
However, the Latvian climate does pose challenges to amphibians in terms of achieving thermo-
optimality and nding suitable habitats for feeding, breeding (Pupina and Pupins 2009) and
overwintering. Therefore, the cold (–20° to –30°C) Latvian winters with relatively thin layers of
snow are dangerous for amphibians overwintering in terrestrial sites, such as burrows or among
Table 65.1 The species of amphibians present in Latvia, their IUCN status for Europe (IUCN 2014) and the trends
of their populations in Latvia.
Species English Name Latvian Name IUCN Red List
trend in Latvia
Lissotriton vulgaris (Linnaeus, 1758) Smooth Newt Parastais tritons Least Concern stable stable
Triturus cristatus (Laurenti, 1768) Northern
Lielais tritons Least Concern decreasing decreasing
Bombina bombina (Linnaeus, 1761) European Fire-
Least Concern decreasing decreasing
Hyla arborea (Linnaeus, 1758) European Tree
Least Concern decreasing increasing
Pelobates fuscus (Laurenti, 1768) Common
Least Concern decreasing decreasing
Bufotes viridis (Laurenti, 1768) European Green
Zaļais krupis Least Concern decreasing unk nown
Epidalea calamita (Laurenti, 1768) Natterjack Toad Smilšu krupis Least Concern decreasing decreasing
Bufo bufo (Linnaeus, 1758) Common Toad Parastais krupis Least Concern stable stable
Pelophylax lessonae (Camerano, 1882) Pool Frog Dīķa varde Least Concern decreasing stable
Pelophylax kl. esculentus (Linnaeus,
Edible Frog Zaļa varde Least Concern decreasing stable
Pelophylax ridibundus (Pallas, 1771) Eurasian Marsh
Ezera varde Least Concern increasing unknown
Rana arvalis (Nilsson, 1842) Moor Frog Purva varde Least Concern stable stable
Rana temporaria (Linnaeus, 1758) Common Frog Parastā varde Least Concern stable stable
Decline and Conservation of Amphibians in Latvia 125
roots, and may lead to high mortality. Additionally, during such winters small ponds freeze all
the way to the boom and, in deeper ones, the oxygen regimen worsens, thereby causing the death
of hundreds of amphibians overwintering there. Cool, short summers often occur in Latvia, with
the result that some tadpoles, especially of southern species like Bombina bombina, do not have time
to go through metamorphosis and gain sucient weight before overwintering (A. Pupina and
M. Pupins, unpublished data). In the relatively cool Latvian climate, amphibians mostly choose
small, stagnant ponds, well heated by the sun, for reproduction. Therefore, when a dry, hot sum-
mer occurs, such ponds often dry up, resulting in mass mortality of amphibian larvae (A. Pupina,
M. Pupins, unpublished data) (Figure 65.2).
Long-term anthropogenic transformation of habitats for agricultural use led to a reduction in
the number of ponds in Latvia, and in impairment of habitat suitable for amphibians. The dams
and ponds of beavers (Castor ber), which are optimal for amphibians in Latvia, also are often
destroyed by land-owners (Pupins and Pupina 2007). Drainage of wetlands, deforestation and
formation of dams along riversides, straightening of small rivers and their subsequent transfor-
mation into soil-reclamation canals have been traditionally carried out over several decades in
Latvia. These works were especially developed during the Soviet occupation of Latvia and led to
the disappearance of tens of thousands of bodies of water, including small ponds, especially in
the central part of Latvia. Phreatic decline, caused by melioration, and the wide use of fertilizers
in agriculture during the Soviet time led to rapid eutrophication and to many of the remaining
ponds becoming choked with weeds, which as a result became too dark and cold, and thus inap-
propriate for amphibians (Pupins and Pupina 2006; Pupina et al. 2012). Also, aquatic and terrestrial
habitats continue to be overgrown by bushes and canes nowadays because of deagrarianization,
which is becoming the main reason for degradation of amphibian habitats in Latvia.
Mortality on roads is an important negative factor for Latvian amphibians, especially during
the spring and autumnal mass migrations, and for small populations of declining species. For
example, in spring 2014 the authors counted 224 aened adult and young Rana temporaria in
Daugavpils on a 200-m stretch of asphalt road between breeding and wintering habitats. Also, 26
aened or injured adult Bufo bufo, most of them in amplexus (Figure 65.3), were found on a 50-m
section of unsealed road near the town of Akniste. There are no trac signs in Latvia warning
drivers of amphibian crossings, and barrier fences guiding amphibians to tunnels under the roads
Table 65.2 The species of amphibians present in Latvia and their distribution and status in the country (Caune
1992; Pupins and Pupina 2011a; Pupina and Pupins 2012).
Species Status in Latvia Distribution in Latvia
Lissotriton vulgaris common whole territory
Triturus cristatus rare whole territory
Bombina bombina very rare southern part of Latvia
Hyla arborea very rare western part of Latvia (reintroduced)
Pelobates fuscus rare south and central territory, excluding the north of Latvia
Bufotes viridis rare south and central territory, excluding the north of Latvia
Epidalea calamita very rare central and western part of Latvia, near the Baltic sea; southeastern
part of Latvia
Bufo bufo common whole territory
Pelophylax lessonae common whole territory
Pelophylax kl. esculentus common whole territory
Pelophylax ridibundus common south and central territory, excluding the north of Latvia
Rana arvalis common whole territory
Rana temporaria common whole territory
126 Amphibian Biology
Fig. 65.2 Drying Pelophylax sp. tadpoles in a pond where Pelobates fuscus, Triturus cristatus, and Bombina bombina
are also living (Katriniski, Demene, Daugavpils novads; 55° 43’ 10.56” N; 26° 32’ 54.51” E). Photograph by Aija Pupina.
Fig. 65.3 Mortality on a road: attened and injured adult Bufo bufo in amplexus (Akniste; 56°10’ 8.61” N; 25° 43’
16.25” E). Photograph by Mihails Pupins.
Decline and Conservation of Amphibians in Latvia 127
are not used; therefore, mortality on roads is very high and can lead to fragmentation or extirpa-
tion of small populations.
Burning of dried grass is widely practised in Latvia. Such burning is carried out in the early
spring on warm, sunny days and is very dangerous for amphibians that are migrating from over-
wintering sites to ponds for reproduction (Pupins and Pupina 2006). The authors have many times
observed such res near the edges of ponds inhabited by the declining species Triturus cristatus,
Bombina bombina, Pelobates fuscus and some others (Figure 65.4). The authors have a dozen times
found live amphibians with half-burnt bodies, injured by the burning of grass (Figure 65.5).
The degradation and disappearance of the majority of factories and agricultural rms in Latvia
after the end of the Soviet occupation led to the appearance of a great number of derelict buildings,
communication lines and shafts acting as mortal traps for amphibians. We managed to count 38
atrophied individuals of amphibians in just one shaft in Kumbuli, Daugavpils region.
There are a great number of illegal dumps around many Latvian villages, i.e. in forests, slashes,
drains, marshes and bodies of water, which are the main habitats of amphibians. These dumps
are the source of pollution of the environment of amphibians (Figure 65.6) wherein they are also
injured by cuts from broken glass (Figure 65.7).
Fig. 65.4 Burning old grass on the shore of a pond used by Bombina bombina, Pelobates fuscus, Lyssotriton vulgaris
and Pelophylax sp. (Daugavpils novads; 55° 50’ 5.62” N; 26° 29’ 3.43” E). Photograph by Agnese Pupina.
128 Amphibian Biology
Fig. 65.5 Eect of burning grass: adult Rana temporaria burned while migrating to a breeding pond (Daugavpils
novads; 55° 50’ 6.11” N; 26° 29’ 4.60” E). Photograph by Agnese Pupina
Fig. 65.6 Rubbish (empty bottle) and Bombina bombina in the breeding biotope of Rana temporaria, Triturus crista-
tus, Pelobates fuscus, Pelophylax sp., and Bombina bombina (Katriniski, Demene, Daugavpils novads; 55° 43’ 10.56” N;
26° 32’ 54.51” E). Photograph by Agnese Pupina.
Decline and Conservation of Amphibians in Latvia 129
Fig. 65.7 Adult Rana temporaria showing wound caused by broken glass in a rubbish dump (Kalkunes pagasts,
Daugavpils novads; 55° 50’ 14.04” N; 26° 29’ 27.25” E). Photograph by Mihails Pupins.
Fig. 65.8 This invasive sh Perccottus glenii found in a breeding pond of six amphibian species (Microreserve Ilgas,
territory Natura 2000, Skrudaliena pagasts, Daugavpils novads; 55° 41’ 32.88” N; 26° 46’ 14.61” E). Photograph by
130 Amphibian Biology
Invasive predators are dangerous for amphibians in Latvia, one of which is a sh, the Chinese
sleeper (Perccous glenii) (Figure 65.8). Having become established in small, warm bodies of water,
which are the main habitats used by amphibians for reproduction in Latvia, this species can fully
destroy or seriously damage the larvae of amphibians (Pupins and Pupina 2012a). Perccous glenii
was found by the authors in catchment basins of almost every large river and in many lakes of
Latvia (Pupins and Pupina 2012b); it was found in the territories of many populations of Bombina
bombina and Triturus cristatus, and continues to extend its distribution (Pupina and Pupins 2014;
Pupina et al. 2015).
Another invasive predator in Latvia regarded as dangerous for larvae of amphibians is the
widely spread sh Carassius auratus, whose natural range is eastern Asia. This species was imported
into Latvia in 1948 and released before 1990 into at least 180 (23%) of Latvian lakes and into many
other bodies of water (Andrusaitis 2003). We have discovered that in Latvia amphibians avoid
using bodies of water for reproduction if sh are present (personal observations). Introduction of
invasive and local sh into water inhabited by amphibians is carried out by local citizens and by
anglers; Perccous glenii is sold on the Internet (A. Pupina, M. Pupins, and Ag. Pupina, unpublished
There are seven species and subspecies of exotic freshwater turtles that we recorded in the wild
in Latvia: Trachemys scripta troostii, Mauremys caspica, Mauremys rivulata, Pelodiscus maackii, Trache-
mys scripta elegans (Pupins and Pupina 2011b), Trachemys scripta scripta, Graptemys pseudogeograca
kohnii (M. Pupins, A. Pupina, unpublished data); mainly single individuals of Trachemys scripta
elegans successfully overwinter and aempt oviposition (Pupins 2007; Pupins and Pupina 2012c).
These carnivorous species can be dangerous for small populations of declining species of amphib-
ians, especially in the south of the country (Pupins and Pupina 2006), and they can also introduce
Dangerous new predators for adult amphibians in Latvia are invasive mammals that are becom-
ing widespread in the country: the raccoon dog (Nyctereutes procyonoides) (29,200 individuals
registered in 2014), the American mink (Neovison vison) (23,200 individuals registered in 2014),
and probably the muskrat (Ondatra zibethicus) (5,500 individuals registered in 2014) (NeoGeo.lv
Research on the presence and distribution of local or exotic parasites in the Pelophylax kl. escu-
lentus complex was carried out in Latvia in 2017. One-hundred-and-forty-one adult and juvenile
specimens were collected for parasitological examination. The parasites found belong to the fol-
lowing systematic groups: Protozoa, Monogenea, Digenea, Nematoda and Acanthocephala. Fifty-
ve Pelophylax kl. esculentus complex tadpoles were examined. Three Trematoda species in their
larval stage: Alaria alata, Diplostomum sp., Opisthioglyophe ranae and Diplodiscus subclavatus in its
adult stage were detected in dierent organs of the tadpoles. One Nematoda, Thelandros tba, a
common parasite of frog tadpoles, was observed in the intestine (Kirjusina et al. 2018a, 2018b).
One-hundred-and-thirty-ve samples of Pelophylax kl. esculentus complex from 42 locations
were checked for the presence of Batrachochytrium dendrobatidis (Bd) and B. salamandrivorans (Bsal)
using qPCR in August 2017. Bsal was not detected in any sample. Bd infection was detected in 18
(43%) locations and in 53 (39%) samples. The infection intensity in the P. kl. esculentus complex
reached a maximum of 143.0 genomic equivalents of zoospores (GE) in a water basin near Jurmala
city (Kulikova et al. 2018).
Taking into account that Xenopus laevis and other tropical species of amphibians are widely sold
in pet shops in Latvia and by Internet (Pupins and Pupina 2011a), it can be assumed that Bd, and
other species of exotic parasites can be found in many populations of Latvian amphibians.
Decline and Conservation of Amphibians in Latvia 131
III. Declining species and species of special conservation concern
The Red Data Book (Andrusaitis 2003) is not the ocial nature protection document in Latvia;
rather that role is taken by a popular book. The status of the species in the Red Data Book of Lat-
via does not directly inuence practical actions towards species or their protection and can only
serve for educational or illustrative purpose.
There is a special document in Latvia approved by the Cabinet of Ministers “Ministru kabineta
noteikumi # 396” (Ministru kabineta noteikumi 2004) which contains a list of specially protected
species in Aachment 1, including six species of amphibians. These are Hyla arborea, Epidalea
calamita, Bufotes viridis, Triturus cristatus, Bombina bombina and Pelobates fuscus, all of which are
species of special conservation concern in Latvia and in the European Union (Table 65.3).
In 2003 (Andrusaitis 2003) Hyla arborea was considered as extirpated in Latvia. However, in
1987 the Laboratory of Ecology, led by herpetoculturist Dr. Juris Zvirgzds-Zvirgzdiņš, was founded
for remediation of this species at the Riga National Zoo. He considered widespread amelioration
and a complete extirpation of beavers, Castor ber, from Latvia to be the reasons for the decline of
the treefrog (J. Zvirgzds-Zvirgzdiņš, personal communication). Using Hyla arborea that had been
brought from Belarus for the formation of a breeding group and bringing 4,110 young individuals
that had been bred in zooculture for reintroduction into nature in 1988–1992, the Riga National
Zoo formed a stable population of Hyla arborea in Liepajas novads in southwestern Latvia (56° 30′
N) (Zvirgzds et al. 1995; Zvirgzds 2003). This population has been continuously increasing in
numbers and extending its area of distribution. Now the area of expansion occupies Kurzeme and
at least 1,000 km2 in Kalvene, Vaiņode, Aizpute and Priekuli (Zvirgzds 2003). However, the pres-
ent distribution of this species in Latvia does not mean that the danger of decline has disappeared.
There was also an opinion that Hyla arborea had never inhabited Latvia and that it had been intro-
duced into the country as an alien species (Caune 1992; A. Berzins, personal communication).
Epidalea calamita is a very rare species in Latvia, mainly inhabiting the coastal region of the
Baltic Sea and its Gulf of Riga and the central part of Latvia (Caune 1992; Berzins 2008; Pupins and
Pupina 2011a). Over the past few years it has been discovered in Dviete in the southeast of Latvia
(E. Racinskis, personal communication). A. Berzins (2008) wrote that
“… the Naerjack Toad inhabits mainly open, well-warmed landscapes with light,
sandy soils. There it lives in sand dunes, glades of pine forests, sand and gravel quar-
ries and meadows … … The reason for decline in the Naerjack Toad is complex and
not fully understood, … and it reects a continuous change in the cultural landscape
over the past hundred years. The former distribution in Latvia included many sandy
exposed coastal areas, but also inland sand or gravel pits with open ground and with
oligotrophic and shallow breeding waters. The open exposed areas on poor and sandy
Table 65.3 The species of amphibians of special conservation concern and their status in Latvia (Latvian
Environment, Geology and Meteorology Centre 2015; Ministru kabineta noteikumi 2004; Andrusaitis 2003).
Species Category in Red Data
Book of Latvia
legislation Bern Convention Protected species in
Latvia (MKN #396)
Triturus cristatus 2 (vulnerable species) HD II, IV II Yes
Bombina bombina 1 (endangered species) HD II, IV II Yes
Hyla arborea 0 (extinct species) HD IV II Yes
Pelobates fuscus 4 (undetermined species) HD IV II Yes
Bufotes viridis 3 (rare species) HD IV II Yes
Epidalea calamita 2 (vulnerable species) HD IV II Yes
132 Amphibian Biology
soils with a natural uctuating water regime have become rare and the ground often
becomes overgrown by dense vegetation, leading to changed microclimate …”.
Bufotes viridis occurs throughout the whole territory of Latvia except for its northern part (Caune
1992; Pupins and Pupina 2011a). Nowadays the northern border of its current European distribu-
tion is partly located in Latvia; the populations of the species are tessellated, and their number is
not large. It tends to live in anthropogenic landscapes and in the basins of large rivers in Latvia
(Rimicans et al. 2010). The main reason for its decrease in number is the rapid overgrowing of
terrestrial habitats and small ponds by bushes following the abandonment of unprotable farms,
thereby destroying habitat essential for this species. As an example, a large population of Bufotes
viridis, which earlier had bred on the ood meadows used by local citizens for haymaking and as
pasture for cows, disappeared in Niderkuni, Daugavpils, between 1987 and 2005 (A. Pupina and
M. Pupins unpublished data).
Triturus cristatus is rare in Latvia; the species is tessellatedly distributed through the whole of
the country (Caune 1992; Pupins and Pupina 2011a), also in the northern part; the populations are
not large and are fragmented.
Bombina bombina is the rarest species of amphibian in Latvia. Until 2004 only two populations
were known: at Islice (Bauska) and Ilgas (Silene), with up to ten vocalizing males in each popula-
tion (Andrusaitis 2003). In 2004 A. Pupina, Ag. Pupina, and M. Pupins discovered a new popula-
tion in Ainavas (Daugavpils region) (Pupins and Pupina 2006) and during the following years
they came across populations in Demene, Medumi, Ilukste, Aizkraukle and Eglaine (Pupina and
Pupins 2007; Pupins and Pupina 2011a).
All populations known in Latvia reside within 0–20 km from the southern and southeastern
border of Latvia (Kuzmin et al. 2008; Pupina and Pupins 2008). The main reasons for the decline
are the overgrowing of habitats by bushes and canes (Pupina et al. 2012); the occurrence of sh in
ponds, especially the invasive Perccous glenii; cleaning of soil reclamation canals; and destruction
of dams and bodies of water produced by beavers. Two of the present authors (A. Pupina and
M. Pupins) observed some cases of Pelophylax sp. actively preying upon juveniles of Bombina
bombina (Figure 65.9). They also noticed an increase in some Bombina bombina populations and an
Fig. 65.9 Pelophylax sp. eating
juvenile Bombina bombina
(Demenes pagasts, Daugavpils
novads; 55° 42’ 47.91” N; 26° 29’ 54.88”
E). Photograph by Aija Pupina.
Decline and Conservation of Amphibians in Latvia 133
expansion of the species to the north, possibly inuenced by climatic changes in Latvia (Pupina
and Pupins 2010, 2013a).
Pelobates fuscus is spread throughout the territory of Latvia (Caune 1992; Pupins and Pupina
2011a); this species is inclined to live in mellow and sandy soils, including agricultural ones (Rim-
icans et al. 2010). The populations are not large and are tessellatedly expanded and fragmented,
especially in the north of Latvia. The main reasons for the decrease in numbers are the overgrow-
ing of meadows by bushes following cessation of haymaking, the overgrowing of breeding ponds
by bulrushes, and the invasion of ponds by shes.
Bombina bombina, Triturus cristatus and Pelobates fuscus often are sympatric in breeding ponds
(Pupina and Pupins 2009). The main reasons for this species’ decrease in numbers are (1) drainage
of wetlands, (2) the overgrowing of breeding ponds by bushes, especially ponds in forests and
bordering forests, and (3) invasion of ponds by shes.
All of these species of amphibians are currently living in Latvia at the northern edge of the
species’ European distribution, or close to it (Caune 1992; Pupins and Pupina 2011a) and are vul-
nerable to climatic changes. The Latvian Climate Research Agency stated that
“… during the 20th century the average air temperature in Latvia has risen by 1 degree.
This increase is even more apparent in the years of the 21st century, which have already
passed. During the past 100 years, there have been uctuations in annual rainfall,
which tended to rise from the beginning of the second half of the 20th century” (Lat-
vian Biodiversity Clearing-House Mechanism 2015) (Figure 65.10).
Such changes of climate, namely increase of the temperature and humidity, can lead to changes
both in the habitats of rare Latvian amphibians (e.g. a more rapid invasion of water by vegetation;
changes in oral composition) and in distribution and number of amphibian species and its main
invasive threat Perccous glenii in Latvia, especially in those habitats on the northern border of the
species’ ranges (Nekrasova et al. 2018; Tytar et al. 2018).
IV. Conservation of amphibians in Latvia
All amphibians are protected in Latvia as an integral part of nature; there also are special measures
for conservation of particularly protected species. The conservation of amphibians in Latvia is
carried out on several levels: (1) the legislative level (adoption of laws, norms and requirements
by government and competent organizations, aimed at amphibian conservation); (2) the planning
level (development and approval of the Species Protection Plans for specially protected species of
Fig. 65.10 The annual average air temperature and precipitation amount in Latvia (Latvian Environment, Geology
and Meteorology Centre 2015).
134 Amphibian Biology
amphibians in Latvia and of the Nature Conservation plans for specially protected natural reser-
vations); (3) the level of territorial protection of nature (amphibian conservation on protected nature
reserves); (4) the project-actions level (implementation of projects for conservation, nanced by
local governments and by the European Union); and (5) the level of particular specialists’ and
landowners’ private actions. An ocial permit of the Nature Conservation Agency of Latvia is
required for any action aecting particularly protected amphibians or their respective protected
territories in Latvia. Such actions are assessed by a certied expert of the Nature Conservation
Agency in the category named “Amphibians and Reptiles” or “Amphibians”. There were ve of
these experts in Latvia in 2018 (Expert Register of Nature Conservation Agency 2018). In fact, all
the amphibian conservation measures in Latvia described below were developed and implemented
with the involvement of at least of one of these experts.
A. The legislative level
Latvia, as a member of the European Union, executes all its legislative norms and there are laws
and regulations that are directly or indirectly involved with the conservation of nature, including
amphibians. The most important Latvian laws are the Environmental Protection Law [Par vides
aizsardzibu], Law On Specially Protected Nature Territories [Par ipasi aizsargajamam dabas ter-
itorijam], General Regulations on Protection and Use of Specially Protected Nature Territories
[Ipasi aizsargajamo dabas teritoriju visparejie aizsardzibas un izmantosanas noteikumi], Regula-
tions on the Content and Elaboration Procedure of the Nature Protection Plan for Specially Protected
Nature Territories [Noteikumi par ipaši aizsargajamas dabas teritorijas dabas aizsardzibas plana
saturu un izstrades kartibu], Law on the Conservation of Species and Biotopes [Sugu un biotopu
aizsardzibas likums], and others.
The Regulations on the List of the Specially Protected Species #396 [Noteikumi par ipasi aizsar-
gajamo sugu un ierobezoti izmantojamo ipasi aizsargajamo sugu sarakstu Nr. 396] are the most
important ones aimed particularly at conservation of declining amphibians in Latvia wherein an
ocial list of six specially protected species is given: Hyla arborea, Epidalea calamita, Bufotes viridis,
Triturus cristatus, Bombina bombina and Pelobates fuscus (Ministru kabineta noteikumi Nr. 396; 2004).
The Regulations for the Establishment, Conservation and Management of Micro-reserves #45
[Mikroliegumu izveidosanas, aizsardzibas un apsaimniekosanas noteikumi Nr. 45] are also impor-
tant because they allow the development of micro-reserves for three species of the specially pro-
tected amphibians in Latvia: Epidalea calamita, Bombina bombina and Triturus cristatus.
B. The planning level
There are programmes and plans developed in Latvia that are indirectly or directly aimed at
amphibian conservation. The most important of the general plans are the National Programme on
Biological Diversity, the Latvian Environmental Policy Guidelines for 2014–2020.
A legislatively approved provision on the Plans for Management of Species exists in Latvia, i.e.
Species Conservation Plan. Such plans are a national strategy for conservation of a particular
species and describe the state of that species in Latvia, indicate the threats against it and its habi-
tats, and oer measures for conservation of the species. In 2018, there were two plans approved
for two amphibian species: Bombina bombina (Pupins and Pupina 2006) and Epidalea calamita (Ber-
zins 2008). Such plans are updated every ve to ten years.
There are many Nature Protection plans elaborated for particular specially protected territories
in Latvia. Such plans also include measures aimed at conservation of specially protected amphib-
ian species: as an example, Bombina bombina in microreserves Natura 2000 Ilgas, Islice, Katriniski
Decline and Conservation of Amphibians in Latvia 135
C. The level of territorial protection of nature
This level is related to the existence and operation of protected natural reserves. There are micro-
reserves in Latvia that were founded for the conservation of specially protected declining amphib-
ian species. The following micro-reserves and Natura 2000 territories were chosen for the
conservation of Bombina bombina: Katriniski micro-reserve, Strauti micro-reserve (Demenes pagasts,
Daugavpils novads) (Pupina and Pupins 2014b); Ilgas micro-reserve (Skrudalienas pagasts,
Daugavpils novads); Īslīce micro-reserve (Īslīces pagasts, Bauskas novads). For conservation of
Epidalea calamita: Puzes Reserve (Ventspils novads); Garkalnes Reserve (Dobeles novads); Karateru
Reserve (Limbazu novads); Ainazi Reserve (Limbazu novads); Adazi Rezerve. Hyla arborea: in
Embute pagasts, Liepajas novads (Berzins 2008). Other sympatric declining amphibian species
often inhabit the same micro-reserves; for instance, Pelobates fuscus and Triturus cristatus occur in
the micro-reserves of Bombina bombina (Pupina and Pupins 2009). Amphibian species inhabiting
all the specially protected territories are protected according to the norms of those territories.
D. The project actions level
Particular projects for conservation of specially protected amphibian species and their habitats are
implemented in Latvia with the aid of the EU European Commission and LIFE Programme; the
Ministry of Environmental Protection and Regional Development and its structural units (Admin-
istration of Latvian Environmental Protection Fund, Nature Conservation Agency), local govern-
ments (Daugavpils city council) and organizations (e.g. Riga National Zoo, Latgales Zoo, Daugavpils
University, Latvian Fund for Nature and Latgales Ecological Society).
Such projects can be international or national and usually foster complex actions: (1) conserva-
tion in-situ, (2) conservation ex-situ, (3) research (ecology, distribution, threats, habitats, herpeto-
culture), planning (Pupina and Pupins 2013b), and monitoring, (4) education of local inhabitants,
and (5) dissemination of results. For example, three projects for conservation of rare amphibians
have been implemented in Latvia and are co-nanced by the European Commission and LIFE
The 2010–2014 LIFE project “Conservation of rare reptiles and amphibians in Latvia” (acronym:
LIFE-HerpetoLatvia; LIFE09NAT/LV/000239) was co-nanced by the European Commission:
Country: Latvia. Target amphibian species: Bombina bombina. Two new microreserves and new
Natura 2000 territories “Strauti” and “Katriniski” were created in Demene parish; 27 ponds were
restored or created; more than 4,000 Bombina bombina juveniles were raised in amphibians’ aqua-
culture (Figure 65.11); and released for population enforcement in 2013–2014 (Figure 65.12). The
Project is very successful; both project sites are populated by Bombina bombina; also an additional
site “Ilgas” which was improved for Emys orbicularis (Pupina and Pupins 2014a, 2014b).
The 2004–2009 LIFE project “Protection of habitats and species in Natura Park Razna”
(LIFE04NAT/LV/000199) was co-nanced by the European Commission. Country: Latvia. Target
amphibian species: Bombina bombina. New ponds were created in Nature Park Razna and juvenile
Bombina bombina were introduced to establish a new population.
The 2004–2008. LIFE project “Bombina in the Baltic Region – Management of re-bellied toads
in the Baltic region” (LIFE-Bombina number LIFE04NAT/DE/000028) was co-nanced by the
European Commission. Countries: Germany, Denmark, Sweden and Latvia. Target species: Bom-
bina bombina. In two microreserves and Natura 2000 territories “Islice” and “Ilgas”, new ponds
were created and juveniles were released for population enforcement. The Project was very suc-
cessful (Brockmuller and Drews 2009).
Many conservation projects and research on amphibians in Latvia have been executed with the
nancial support of the Latvian Environmental Protection Fund and Nature Conservation Agency.
136 Amphibian Biology
Fig. 65.11 Breeding laboratory and recirculation system for amphibians in Rare Reptiles and Amphibians Breeding
Centre (LIFE-HerpetoLatvia Project, Latgales Ecological Society, Ainavas, Daugavpils novds; 55° 50’ 65” 5.65” N; 26°
29’ 6.60” E
Fig. 65.12 Releasing of Bombina bombina in restored pond where Pelophylax sp., Triturus cristatus, and Pelobates
fuscus also are breeding (LIFE-HerpetoLatvia Project, Katriniski microreserve, Demene, Daugavpils novads; 55° 43’
10.56” N; 26° 32’ 54.51” E).
Decline and Conservation of Amphibians in Latvia 137
V. Monitoring of amphibians in Latvia
Latvia undertakes long-time monitoring and scientic analyses of amphibian population trends.
The rst methodology for a state-nanced amphibian monitoring programme was developed in
2005 as part of a national biodiversity monitoring programme. It consisted of two major parts:
Natura-2000 monitoring and whole-state or so-called “background” monitoring. Biodiversity of
Natura-2000 sites were surveyed by formally approved monitoring methods during the EMERALD
Project realized by the Latvian Fund for Nature in 2001–2004. Among other protected species,
there were two Habitats Directive Annex-II amphibian species for which project datasheets were
prepared: great crested newt (Triturus cristatus) and re-bellied toads (Bombina bombina). Triturus
cristatus was registered in seven territories, but Bombina bombina only in two. The information
consisted mostly of presence-absence data with some evaluation of available habitats. The meth-
odology for Natura 2000 was developed in 2007 by an environmental consulting company, Estonian,
Latvian & Lithuanian Environment.
The methodology for “background” monitoring was approved in 2006 by a state agency: Latvian
Environment, Geology and Meteorology Centre. Unlike monitoring by Natura 2000, the methods
were applicable to all amphibian species. This monitoring was performed only once, in 2008, by
the company Estonian, Latvian & Lithuanian Environment. It consisted of two types of surveys.
The rst was the counting of vocalizing males in spring in three repetitions on the same transect
at intervals of two weeks. The other method was the seing of three-armed drift fences twice a
season – in late spring and in autumn. In each case traps were set for a period of about one week
during which they were checked three times. Trapped amphibians were removed and released,
but not marked. Using a random stratied sampling method, a total of nine squares, each 1 x 1 km,
in dierent places of Latvia were selected for monitoring. There was a transect ~0.5–1 km long for
counting vocalizing males (for periods of one hour) and one drift-fence trap (four buried buckets
and six metres of fence) in each. Results from vocalizing males and from the traps in spring gen-
erally were poor, but in autumn the results from traps were beer, with up to 30 amphibians
captured for each check of the traps. Only the most common species were recorded (Lissotriton
vulgaris, Bufo bufo, Rana arvalis, Rana temporaria, Pelophylax sp.). Results are available in the form
of a report accompanied by datasheets from the eld on the Nature Conservation Agency’s home
In 2013, a state-approved biodiversity monitoring programme and a methodology for moni-
toring amphibians were signicantly reworked and expanded by the Latvian Fund for Nature
during a state-nanced project.
The State monitoring programme for amphibians began in 2015 and was nished in 2018. It
consisted of a vocalizing anuran survey on 65 plots that covers the whole territory of the state and
included monitoring of the great crested newt at 21 sites where this species was recorded in
1980–2010. A survey of vocalizing anurans indicated the presence of green frogs (Pelophylax sp.)
in 100% of visited 5 x 5 km squares; common toad (Bufo bufo) – 93%; common frog (Rana temporaria)
– 83%; moor frog (Rana arvalis) – 80%; spade-foot toad (Pelobates fuscus) – 13%; treefrog (Hyla arbo-
rea) – 10 %; and re-bellied toad (Bombina bombina) – in 5%. Data from a survey by handnet of the
great crested newt (Triturus cristatus) conrmed its presence in 60% of historical sites.
We have been conducting personal monitoring of Bombina bombina since 2000 in the previously
known population at Ilgas, and since 2004 in newly discovered populations at Demene, Ainavas,
Medumi, Bauska and Verdini. Methods of monitoring include the counting of vocalizing males
and visual accounting, partly looking for tadpoles and eggs. The characteristics of ponds, the
presence of shes (especially the invasive Perccous glenii) and the degree of overgrowing by
vegetation were also estimated. We explored all known localities in Latvia 1–4 times per season
138 Amphibian Biology
until 2011; the localities have been selectively explored in 2012–2017 because of an increase in the
number of localities found to contain this species (> 300). The results have been entered in the
database and published (Pupina and Pupins 2008, 2010, 2012, 2013a; Pupina et al. 2008).
Latvia, with its large quantity of humid habitats and small ponds, non-freezing bodies of water
and other places for overwintering, undeveloped agricultural industry and decreasing human
population, generally corresponds to the needs of the amphibian species that inhabit it. However,
Latvia is a northern European country and a half of all the Latvian amphibian species – Triturus
cristatus, Bombina bombina, Hyla arborea, Bufotes viridis, Epidalea calamita, Pelobates fuscus, Pelophylax
sp. – reach the northern border of their European distribution in or near Latvia. Therefore, their
populations in Latvia are few in number and are fragmented. These species have special require-
ments for optimizing their habitats in Latvia, i.e. increasing the permanence and luminance of
non-freezing ponds used by amphibians for overwintering. The progressive overgrowing by
bushes, shading, and degradations of such habitats contribute to the decline of the amphibian
populations. The increasing threats of anthropogenic origin – mortality on roads, invasive species
of shes, invasive mammalian predators, invasive freshwater turtles and new parasites may lead
to the decline of the few amphibian species inhabiting Latvia, and accordingly their European area
of distribution will decrease. The climatic changes in Latvia can have both positive and negative
meaning for Latvian amphibians. Because of climatic change, research is urgently needed on (1)
the peculiarities of the ecology and distribution of declining amphibians on the northern border
of Latvia; (2) optimization of biotopes and enhancement of the endangered populations using
animals from zooculture; and (3) performance of the necessary measures aimed at protecting
limited populations and at uniting them into metapopulations by establishing population bridges,
including trans-border ones, for making genetic contact with stronger southern populations.
Conservation of Bombina bombina in 2010–2014 was executed with the support of LIFE Project
LIFE09NAT/LV/000239 “Conservation of rare reptiles and amphibians in Latvia” co-nanced by
the European Commission. The research in aquaculture was supported by the Rural Support
Service of Latvia, Project 16-00-F02201-000002, “Innovation”. The research on modelling was
supported by the National Research Program EVIDEnT, No 4.6. “Freshwaters Ecosystems Services
and Biodiversity”, Project LHEI-2015-19. The article was prepared and researches of amphibians
in Latvia have been executed with the nancial support of the Latvian Environmental Protection
We thank the Institute of Life Sciences and Technologies of Daugavpils University, Daugavpils
City Council, Latgales Zoo, Latgales Ecological Society, Nature Conservation Agency, Ministry of
Environment and Regional Development of Latvia, and the administration of the Latvian Envi-
ronmental Protection Fund for support of amphibian conservation in Latvia. We thank the editors
of the book series Amphibian Biology for their kind help, reviewing and suggestions. We mourn
the loss of our colleague and co-author, leading Latvian batrachologist Dr. Aija Pupina and are
grateful for her highly valued cooperation in research and conservation of amphibians in Latvia.
Decline and Conservation of Amphibians in Latvia 139
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