Status of Conservation and Decline of Amphibians:Eastern HemisphereWestern Europe

Edited byHarold HeatwoleJohn W. WilkinsonPUBLISHING

PELAGIC

Amphibian BiologyVOLUME 11

PART 3

PUBLISHINGPELAGIC

Edited by Harold Heatwole and John W. Wilkinson

Amphibian species around the world are unusually vulnerable to a variety of threats, by no means all of which are properly understood. Volume 11 in this major series will be published in parts devoted to the causes of amphibian decline and to conservation measures in regions of the world; this Part 3 is concerned with Western Europe (Britain, Ireland, The Netherlands, Belgium, France, Spain and Portugal).

Experts from each country contribute a chapter describing the ecological background and the conservation status of aff ected species, with an emphasis on native species. As well as infectious diseases and parasites (also covered in a general chapter), threats take the form of introduced and invasive species, pollution, destruction and alteration of habitat, and climatic change. These are discussed as they aff ect each species. All these countries have monitoring schemes and conservation programmes, whose origins and activities are described. Recommendations for action are also made.

Edited by leading scholars in the fi eld, Volume 11, when complete, will therefore provide a defi nitive survey of the amphibian predicament and a stimulus to further research with the objective of arresting the global decline of an entire class of animal.

Cover image: A pair of Rana iberica in amplexus. This species is regarded as of Special Concern and is the subject of a captive-breeding programme at the Pealara Breeding Centre in Spain, aimed at recovering populations aff ected by chytridiomycosis and the introduction of alien species. Photograph by Cesar Ayres.

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Amphibian Biology Vol. 11 Status of Conservation and D

ecline of Am

phibians: Eastern Hem

isphere, Part 3 Western Europe

Published by Pelagic Publishingwww.pelagicpublishing.comPO Box 725, Exeter, EX1 9QU

Amphibian Biology, Volume 11 Part 3Status of Conservation and Decline of Amphibians: Eastern Hemisphere

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Cover image: A pair of Rana iberica in amplexus. This species is regarded as of Special Concern and is the subject of a captive-breeding programme at the Pealara Breeding Centre in Spain, aimed at recovering populations affected by chytridiomycosis and the introduction of alien spe-cies. Photograph by Cesar Ayres.

Table of contents of volume 11, Amphibian Biology: Eastern Hemisphere, Part 3 (Western Europe)

31 Infectious diseases that may threaten Europes amphibians 1Trenton W. J. Garner, An Martel, Jon Bielby, Jaime Bosch, Lucy G. Anderson, Anna Meredith, Andrew A. Cunningham, Matthew C. Fisher, Daniel A. Henk, and Frank Pasmans

32 Conservation and declines of amphibians in Ireland 42Ferdia Marnell

33 Amphibian declines and conservation in Britain 48John W. Wilkinson and Richard A. Griths

34 Conservation and declines of amphibians in The Netherlands 58Anton H. P. Stumpel

35 Amphibian declines and conservation in Belgium 66Gerald Louette and Dirk Bauwens

36 Amphibian declines and conservation in France 74Jean-Pierre Vacher and Claude Miaud

37 Conservation and declines of amphibians in Spain 87Cesar Ayres, Enrique Ayllon, Jaime Bosch, Alberto Montori, Manuel Ortiz-Santaliestra and Vicente Sancho

38 Conservation and declines of amphibians in Portugal 92Rui Rebelo, Maria Jos Domingues Castro, Maria Joo Cruz, Jos Miguel Oliveira, Jos Teixeira, and Eduardo Crespo

34 Conservation and declines of amphibians in The Netherlands

Anton H. P. Stumpel

Abbreviations and acronyms used in the text and references:

CBS Centraal Bureau voor de Statistiek (Statistics Netherlands)EHS Ecologische Hoofdstructuur (National Ecological Network)IUCN International Union for the Conservation of NatureLNV Former name of the Netherlands Ministry of Economic Affairs, Agriculture

and InnovationNGO Non-governmental organizationRAVON (foundation) Reptile, Amphibian & Fish Conservation NetherlandsTRIM Trends and Indices for Monitoring Data.

I. IntroductionThe Netherlands is a small, densely populated country that lies in northwestern Europe at approx-imately 52 N and 5 E. It is bordered by the North Sea to the north and west, Germany to the east, and Belgium to the south. With 16 million inhabitants on a land area of 33,874 km2 (population density of 472/km2), the landscape is largely man-made, intensively cultivated, and urbanized. The western and northern parts are a river delta where the estuaries of the Rhine, Meuse, and Scheldt come together. Water is thus an important feature of the landscape. Reclamation of land began in the 13th century when man radically changed the landscape; dykes were built, and pol-ders with a dense network of ditches and canals were created. Apart from some marshes, no primary or pristine habitats were left. Where there was no natural freshwater on the surface, ponds were dug. Much land is low-lying, nearly a quarter of the total surface area being below sea level, the lowest at -7 m. The soil consists of various substrates, especially clay, sand, peat, marl, and gravel; there are no rocks. Most of the country is quite flat, the highest point being only 321 metres above sea level. The Netherlands has an Atlantic climate with an annual precipitation of 780 mm.

I. Introduction

II. Declining species and species of special conservation concernA. Bombina variegata

B. Salamandra salamandra

C. Hyla arborea

D. Pelobates fuscus

E. Triturus cristatus

F. Lissotriton helveticus

G. Alytes obstetricans

H. Bufo calamita

III. Conservation measures and monitoring programmesA. Conservation measures

B. Monitoring programmes

IV. Perspective

V. Acknowledgements

VI. References

Conservation and declines of amphibians in The Netherlands 59

Nowadays, most of the land surface has been drained for agriculture, construction of houses, industry, and for supply of drinking water. The landscape is drastically fragmented by many roads, canals, and railways. Both water and soil are strongly eutrophic due to airborne pollution.

The Netherlands has sixteen indigenous amphibian species and some of them reach the limits of their European range there. Salamandra salamandra, Alytes obstetricans, and Bombina variegata reach their northern limit in The Netherlands and are confined to the southeasternmost part of the country. Ichthyosaura alpestris, Hyla arborea, Pelobates fuscus, and Rana kl. esculenta reach their northwestern European extent. Rana lessonae does likewise, apart from a single isolated population in England. The Netherlands North Sea coast forms the western limit of the distribution area of Rana arvalis. Additionally, in The Netherlands Lissotriton helveticus is distributed only in the south-ern provinces, although this newt has a more northern distribution in Britain and Germany.

II. Declining species and species of special conservation concernIn The Netherlands, all indigenous amphibians have been legally protected under the Nature Conservation Act since 1973. As well as naming species, this act also designated some sites as Beschermde Natuurmonumenten (Protected Nature Monuments). The Flora and Fauna Act (Backes and Verschuuren 2001) provided an update of the Nature Conservation Act and included the provisions of the Bern Convention and the Habitats and Birds Directives. Nonetheless, the new Act has exemptions that provide loopholes, leading to a political tug-of-war when people apply for planning permission. This can lead to important habitats being destroyed. Recently, a new national Red List for reptiles and amphibians was published (van Delft et al. 2007).

Amphibians in The Netherlands are seriously under threat from the results of human activity and also from disease. Amphibians habitats, both terrestrial and aquatic, are being affected by large-scale land use, intensive animal husbandry, drainage, mechanical and frequent mowing and cutting, frequent crop rotation, overgrazing, and the use of fertilizers, herbicides, and pesticides (Stumpel 2004). Fragmentation and loss of habitat is also the result of the ever-increasing urbani-zation, with a landscape dissected by roads, canals, and railways. Moreover, there is little room for well-developed habitats for amphibians because of the practices of modern water management, whereby watercourses are straightened and frequent dredging occurs, and banks are mown. Ironically, also in protected areas, mismanagement sometimes happens. Lack of money, time, and skills lead to lack of management and to mechanical cutting, bad timing of measures, too frequent intervention, and overgrazing. In more general terms, amphibian decline can also be attributed to predation by pets, waterfowl, or fish that have been released in ponds or ditches, as well as to the pollution of waters used for breeding. Collectors also take their toll of the wild population (cf. Stumpel, 2004). Infectious diseases form a threat of a different order. People have only recently become aware of the present and potential threat of chytridiomycosis and the disease caused by Ranavirus (Spitzen-van der Sluijs et al. 2010). Much research is still needed for developing a con-servation policy that takes these threats into account (cf. Garner, this Volume).

All these factors have led to half of the indigenous species being on the national Red List (van Delft et al., 2007); their present status according to both Netherlands criteria (trend since 1950, rareness) and those from IUCN, is briefly indicated in Table 34.1. For more information, see the Atlas of the Herpetofauna of The Netherlands (Creemers and van Delft 2009).

60 Amphibian Biology

Table 34.1 The status of amphibians in The Netherlands, assessed at the national level (STATUS-NL) and according to IUCN criteria (STATUS-IUCN) (van Delft et al. 2007).

SPECIES STATUS - NL STATUS - IUCN

Bombina variegata Critically endangered Endangered

Salamandra salamandra Endangered Endangered

Hyla arborea Endangered Least concern

Pelobates fuscus Endangered Endangered

Triturus cristatus Vulnerable Least concern

Lissotriton helveticus Vulnerable Vulnerable

Alytes obstetricans Vulnerable Vulnerable

Bufo calamita Susceptible Vulnerable

A. Bombina variegataThe yellow-bellied toad (Bombina variegata) is confined to the southernmost part of the country. Only five populations, now inhabiting abandoned quarries, have survived the destruction and fragmentation of the species habitat. Recent introductions into newly developed habitats in those quarries have been successful. However, as long as the total population numbers less than 500 adult individuals, the long-term survival of the species remains uncertain. Population densities may fluctuate strongly between years. Appropriate habitat management for this toad includes the repeated creation of shallow, temporary waters shortly before their reproductive season.

B. Salamandra salamandraThe fire salamander (Salamandra salamandra) is restricted to the same area as B. variegata. It is a terrestrial ovoviviparous species of deciduous forests that contain streams and springs. The free-swimming larvae need clear, oxygen-rich water that is free of fish, and there must be plants, stones, or logs to serve as shelter. Threats originate from humans spoiling the habitat by clearing away aquatic plants, polluting the water, and straightening streams or making sluices in them. In addi-tion, some animals are caught by collectors and still others from foreign populations are set free, thus releasing alien genetic material into the native population.

C. Hyla arboreaThe European treefrog (Hyla arborea) is found in the southeastern half of the country. Its terrestrial habitat requires subtle transitions from one vegetation type to another, such that the quality of the terrestrial habitat determines whether a species is present in an area more than does suitable aquatic habitat. This species reproduces mainly in marshes and ponds. In the cultural landscape, habitats for reproduction, aestivation, and hibernation may be hundreds of metres apart so good connectivity among them is vital for survival. Also important is the position of a single habitat in the network of habitats of the metapopulation. Population densities may fluctuate strongly between years. Although H. arborea has been successfully introduced into newly developed habitats during the past 35 years, the species is still suffering elsewhere from the effects of drainage, habitat frag-mentation and from being collected. Moreover, the aquatic habitat remains under threat from infilling, lack of management, pollution, and release of fish.

D. Pelobates fuscusThe European spadefoot toad (Pelobates fuscus) is close to becoming classified as critically endan-gered. Being a burrowing amphibian, many of its terrestrial habitats in loose sandy soils have become unsuitable due to changed use of agricultural land. Apart from isolation and loss of

Conservation and declines of amphibians in The Netherlands 61

habitat, numbers are declining and populations are rapidly disappearing. However, a few small populations survive in the east of the country. Because spadefoots reproduce in old ponds and pools, they are susceptible to neglect of management programmes, including the removal of released fish, especially pumpkinseed sunfish (Lepomis gibbosus). Removal of plants both in and around the pool for management of the flora can also have a disastrous effect. Unfortunately, the species low rating in international conservation legislation has led to little interest from the gov-ernment and, consequently, appropriate management does not take place.

E. Triturus cristatusThe aquatic habitat of the great crested newt (Triturus cristatus) is similar to that of Hyla arborea and therefore the species is susceptible to the same threats. Their terrestrial habitat in such places as rough grassland, gardens, scrub, and deciduous woodland, need to be close to the aquatic sites used for reproduction. Consequently, the species is threatened by its aquatic and terrestrial habi-tats being isolated from each other, for example by roads. This leads to many newts becoming traffic casualties during their migrations in the early spring and in the autumn, and probably has an impact on local populations. The creation of new ponds near potential terrestrial habitat has been shown to be successful for this species.

F. Lissotriton helveticusThe palmate newt (Lissotriton helveticus) is limited to the southern part of The Netherlands, where it is mostly found in natural areas. This species is tolerant of a low pH in the heathland, moorland, and woodland pools where it reproduces. The drastic decline of this newt during the past 50 years can be attributed to large-scale reclamation of heathland and moorland for agriculture, forestry, and housing; the remaining habitats have become fragmented and drained. Air pollution causes the water in which it reproduces to become acidified to a pH below tolerance levels and has further contributed to its decline.

G. Alytes obstetricansThe midwife toad (Alytes obstetricans) has the same, limited distribution as Bombina variegata. It is a rare species in The Netherlands. It requires a loose, stony soil in a warm location. These condi-tions can be found on agricultural land that is traditionally managed, in quarries, and in old churchyards. A strong decline has recently been observed that cannot be linked to changes in habitat; perhaps the cause can be attributed to chytrid infection, as is described for Spain (Bosch et al. 2001).

H. Bufo calamitaThe natterjack toad (Bufo calamita) occurs in various types of habitat such as heathlands, coastal sand dunes, edges of salt marshes, marshes in river forelands, and quarries, as well as in temporary habitats left by man after excavating ground for building (Stumpel, 2004). Apart from heathlands, these habitats have dynamic environments; population densities may fluctuate considerably. This toad is an opportunistic breeder, making use of temporary, shallow waters. New habitats can be created rather easily and are quickly taken into use. Although populations in the coastal sand dunes remain stable, threats are posed by new ways of building, resulting in habitats on building sites not being available long enough for successful reproduction. Elsewhere, the bare soil in their terrestrial habitat disappears by vegetational succession. In general terms, the fragmentation of the landscape results in populations becoming isolated from each other.

62 Amphibian Biology

III. Conservation measures and monitoring programmes

A. Conservation measuresThe government is responsible for nature conservation and developing policies for that purpose. Governmental and non-governmental organizations carry out measures in the field, work that is often done by volunteers. The state subsidizes projects both for species and habitats. For some species, there have been national Action Plans. The National Nature Policy Plan, Natuurbeleidsplan (Ministerie, 1990), provides the basis for the Ecologische Hoofdstructuur (EHS) [National Ecological Network], which consists of core areas, nature development areas, and corridors. This network is still under construction and planned to be completed by 2018. There are 11 species that have been targeted: Salamandra salamandra, Ichthyosaura alpestris, Triturus cristatus, Lissotriton helveticus, Alytes obstetricans, Bombina variegata, Pelobates fuscus, Bufo calamita, Hyla arborea, Rana arvalia, and R. les-sonae. Under the auspices of the Habitats Directive of the European Union, The Netherlands is contributing to the European network of Natura2000 areas by designating protected areas. Amphib-ians, however, play a minor role in this process.

The main organizations concerned with nature conservation that also own land are Staatsbos-beheer (the state forestry service), Natuurmonumenten, Provinciale Landschappen, one for each of the twelve provinces, and the Waterschappen (the water boards). In addition, the Stichtingen voor Land-schapsbeheer organize landscape management for each province, focusing mainly on small landscape elements; these organizations do not own land. All organizations carry out management work that includes paying special attention to amphibians and their habitats.

Volunteers play an important role in the conservation of amphibians in The Netherlands. Numbering several thousand and organized into working groups, they conduct surveys, carry out management measures, monitor the situation in the field, and send in distributional data to the national database, which at present comprises some 440,000 records. Together with the RAVON foundation (Reptile, Amphibian & Fish Conservation Netherlands) that coordinates their activities, they are also engaged in promoting interest in amphibians among the general public.

Action Plans can be a useful tool for amphibian conservation. Such a plan for an amphibian species consists of finding out the number of its populations and their sizes, as well as the charac-teristics of the aquatic and terrestrial habitats and any connections between them. Working with the idea of basic habitats, sites were chosen where they could be developed, and the costs esti-mated. Depending on the resources available, plans were carried out, mostly phasing the work over several years. Once work had been completed, the habitats were monitored to follow their development and to see whether amphibians accepted them. Action Plans have been carried out countrywide for Alytes obstetricans, Bombina variegata, Hyla arborea, and Pelobates fuscus; Action Plans carried out by individual provinces dealt with additional species of amphibians.

The loss of ponds in the agricultural landscape was a major cause of the decline of amphibians. Thus, plans for ponds were developed that provided for the creation and restoration of waters for use as habitat for reproduction (Boothby 1999). The first pond project started in the province of Limburg in the early 1980s; other provinces followed, and as a result new ponds were dug coun-trywide. Unfortunately it was often not possible to create ponds at places with the best potential. Furthermore, there was no guarantee of continued maintenance. At first, only the common species benefited from the new and restored ponds; the rare and threatened amphibians for which they were meant did not. Later, measures became more tailored to the requirements of these rare spe-cies with the result that their populations were able to build up again locally, and extension of their distributional boundaries was made possible.

Conservation and declines of amphibians in The Netherlands 63

Conservation measures in the terrestrial habitat of amphibians are directed largely towards maintaining scrub patches, hedgerows, and other small elements of the landscape, such as heaps of stones and log piles, that can serve as refugia (Stumpel 1997). Limiting the growth of woody vegetation is also necessary for the upkeep of the vegetation mosaic. Among the special provisions for the movement of fauna are ecoducts, tunnels, and specifically-designed kerbstones, the last two more particularly for amphibians.

During the spring migration of amphibians, many volunteers help prevent large numbers end-ing up as traffic victims by closing off roads at dusk. Where barriers are not possible, warning signs are sometimes put up ordering vehicles to slow down. Mortality is reduced by bordering roadsides with fences containing pitfall traps along its length, so that amphibians can be captured and carried over the road in buckets.

After habitat has been restored, or new habitat created, a species may be reintroduced into an area to help build up a population. This has been done successfully for Hyla arborea and Bombina variegata by releasing both larvae close to metamorphosis and juveniles. A new approach to con-servation is the building up of a breeding stock from threatened populations for releases in new and restored habitats. These habitats can either not be reached by the amphibians in a natural way due to barriers in the landscape preventing their migration, or because the tenacity of a species to a site is so strong that they do not colonize new habitats readily. This stock is obtained by collect-ing eggs in the field and rearing the young in the laboratory; when the larvae are near metamor-phosis, they are set free in the target habitat. After successful projects with Hyla arborea, this is now being practiced with Pelobates fuscus, taking animals from sites where the species was threatened with extirpation.

Amphibians have also been released outside their natural range by people wishing to extend the species geographic distribution; the newt Ichthyosaura alpestris and the toads Alytes obstetricans and Bufo calamita are cases in point. The introduced species have persisted for decades and may have affected local populations of other amphibian species. Moreover, by confusing the original distributional pattern, and in some cases because the animals are not of indigenous stock, such introductions hamper research into the biogeography and genetics of native amphibians.

Exotic species have also become established as a consequence of releases. They often pose a threat to indigenous amphibians by competition, predation, and hybridization or because they sometimes carry diseases. This is the case with Triturus carnifex, which inhabits the same habitats as the indigenous T. cristatus and hybridizes with it. Rana (Lithobates) catesbeiana is competing for habitat with indigenous green frogs (Rana synkl. esculenta) and furthermore is a vector for chytrid infections (Bai et al. 2010).

Informing and advising nature wardens, farmers, and the general public forms an important part of nature conservation in The Netherlands. In this, the NGO RAVON plays a central role by creating platforms with all interested parties around the table, giving talks, developing websites, publishing magazines and folders, and keeping in contact with the volunteers in working groups all over the country. Not only has this organization created awareness of the need for protecting amphibians, it also manages the data bank with all records of amphibians, has an extensive docu-mentation centre and, together with students from universities and technical schools, carries out scientific research.

B. Monitoring programmesIn 1997, Meetnet Amfibien, the national amphibian-monitoring network was set up (Goverse et al. 2006). Hundreds of well-instructed volunteers (Groenveld et al., 2011) have since surveyed 338 areas that included over 2,500 breeding sites. Monitoring mostly takes place four times a year and

64 Amphibian Biology

this network is still growing. From the amphibian counts, trends in population development are calculated using a computer programme (TRIM) designed for monitoring data with missing val-ues (Pannekoek & van Strien 1998). The status of all species over this period has now been assessed, and shows a moderate increase or stable situation for twelve of the sixteen indigenous species. Although Witmer et al. (2002) found a continuing decline in amphibians during the second half of the 20th century; some species seem to be making a recovery during the past decade.

Short-term monitoring also occurs, usually for a period of five years. This is sometimes on the authority of the government that also funds it, or a volunteer working group might be interested in following the development of a local population.

IV. PerspectiveHalf of all amphibian species in The Netherlands are at present threatened in some way or another. It remains uncertain whether amphibians in general are recovering from a very rough time in the past. Numerous factors plead against this. Most habitats are suffering from drying out as a result of large-scale drainage and from eutrophication and acidification due to airborne nitrate pollution both from agriculture and traffic. Although nature reserves have been designated and new habitats created, many of them are too small and isolated to be able to sustainably support a population. Possible effects of climatic change have not yet been taken into account and remain open to inter-pretation. Evidence for causal relationships between amphibian decline and other environmental factors is largely lacking, thereby weakening public debate. Modern society is focused on short-term economic profit, which is always at the cost of nature. The fate of the amphibians is in the hands of politicians; only they have the power to turn the tide. It is hoped that the present volume convinces them that effective conservation of amphibians is urgently needed.

V. AcknowledgementsThanks are due to Jeroen van Delft, Edo Goverse, Claire Hengeveld, Annemarieke Spitzen-van der Sluijs, and Suzette Stumpel-Rienks for comments on the manuscript.

Conservation and declines of amphibians in The Netherlands 65

VI. ReferencesBackes, C. and Verschuuren, J. (eds), 2001. Natu-

urbeschermingsrecht 2001/2002. Richtlijnen, verordeningen, verdragen, jurisprudentie, Ned-erlandse wet- en regelgeving en toelichtingen, met name rond de Vogelrichtlijn (79/409/EEG), de Habitatrichtlijn (92/43/EEG) en de CITES-ver-ordening (338/97/EG). Sdu publishers, The Hague.

Bai, C., Garner, T. W. J. and Li, Y., 2010. First evidence of Batrachochytrium dendrobatidis in China: discovery of chytridiomycosis in introduced American bullfrogs and native amphibians in the Yunnan province, China. EcoHealth 7: 127134.

Bal, D., Beije, H. M., Fellinger, M., Haveman, R., van Opstal, A. J. F. M. and van Zadelhoff, F. J., 2001. Handboek Natuurdoeltypen. Expertise-centrum LNV, Wageningen. Second, fully revised edition.

Boothby, J. (ed), 1999. Ponds and pond land-scapes of Europe. Proceedings of the Interna-tional Conference of the Pond Life Project, Vaeshartelt Conference Centre, Maastricht, The Netherlands, 30th August2nd September 1998. John Moores University, Liverpool.

Bosch, J., Martnez-Solano, I. and Garca-Pars, M., 2001. Evidence of a chytrid fungus infec-tion involved in the decline of the common midwife toad (Alytes obstetricans) in protected areas of central Spain. Biological Conservation 97: 331337.

Creemers, R. C. M. and van Delft, J. J. C. W. (eds), 2009. De Amfibien en Reptielen van Nederland. Nederlandse Fauna, Series No. 9. Nationaal Natuurhistorisch Museum Naturalis, European Invertebrate Survey - Netherlands, Leiden.

Goverse, E., Smit, G. F. J., Zuiderwijk, A. and van der Meij, T., 2006. The national amphibian monitoring program in the Netherlands and NATURA 2000. Pp. 3942 in Herpetologia Bon-nensis. II. Proceedings of the 13th Congress of the Societas Europaea Herpetologica, ed by M. Vences, J. Khler, T. Ziegler and W. Bhme. Museum Alexander Koenig, Bonn.

Groenveld, A., Smit, G. and Goverse, E., 2011. Handleiding voor het monitoren van amfibien in Nederland. RAVON Working Group Monitor-ing, Amsterdam. Third edition.

Ministerie van Landbouw, Natuurbeheer en Visserij, 1990. Nationaal Natuurbeleidsplan. Sdu publishers, The Hague.

Pannekoek, J. and van Strien, A. J., 1998. TRIM 2.0 for Windows (Trends and Indices for Monitoring data). Statistics Netherlands (CBS), Voorburg. Research paper No. 9807.

Spitzen-van der Sluijs, A. M., Zollinger, R., Bosman, W., van Rooij, P., Clare, F., Martel, A. and Pasmans, F., 2010. Batrachochytrium dendrobatidis in amphibians in the Nether-lands and Flanders (Belgium). RAVON, Nijmegen.

Stumpel, A., 1997. Amphibians and reptiles in agricultural and urban landscapes in the Netherlands; design of and provisions for sub-habitats and corridors. Pp. 5364 in Opportunities for Amphibians and Reptiles in the Designed Landscape, ed by R. Bray and T. Gent. English Nature, Peterborough. English Nature Science Series No. 30.

Stumpel, A. H. P., 2004. Reptiles and Amphibians as Targets for Nature Management. Thesis Wageningen University, Wageningen.

van Delft, J. J. C. W., Creemers, R. C. M. and Spitzen-van der Sluijs, A. M., 2007. Basisrap-port Rode Lijst Amfibien en Reptielen volgens Nederlandse en IUCN criteria. RAVON, Nijmegen.

Witmer, M., Dirkx, J., Leneman, H., Notenboom, J., van Veen, M. and Sollart, K. (eds), 2002. Natuurbalans 2002. Kluwer, Alphen aan den Rijn.

Index 103

Aabnormalities 22Acanthocephala 21acidification 64Aor Mountains 93action plans 5052, 62, 70, 7980,

84, 88adenocarcinoma 89adenovirus(es) 4Aeromonas hydrophila 1011aestivation 60Africa 2, 87African clawed frog (also see

Xenopus laevis) 7677Agency for Nature and Forests

[Belgium] 69, 71aggregation 15agile frog (also see Rana dal-

matina) 5, 9, 53agriculture 22, 4243, 45, 4950,

55, 5962, 64, 7071, 76, 7880, 83, 88, 9597

agrochemicals 22, 88AHE (also see Herpetological

Association of Spain) 89AHEnuario 89Alderney 49alien species (also see exotic

species) 72, 88Alpine Geographical Region 75alpine newt (also see Ichthyosaura

alpestris; Mesotriton alpestris) 7, 20

alpine salamander (also see Salamandra atra) 77

Alsace 7778, 81Alytes cisternasii (also see Iberian

midwife toad) 13, 89, 93, 95, 97Alytes dickhilleni 89Alytes muletensis (also see

Mallorcan midwife toad) 13, 8889

Alytes obstetricans (also see midwife toad; common midwife toad) 7, 10, 13, 20, 54, 59, 6163, 67, 69, 71, 76, 8889, 93, 95, 9798

Alytidae 2Ambystoma tigrinum virus (also

see ATV) 4American bullfrog (also see

Lithobates catesbeianus; Rana catesbeiana) 7172, 7677

American red swamp crayfish (also see Procambarus clarkii) 96

Amphibian and Reptile Conser-vation (also see ARC) 51

Amphibian and Reptile Groups (also see ARG) 54

Amphibiocystidium sp. 9, 12, 1820Amphibiichlamydia salamandrae 10

Amphibiothecum 19Anatolian marsh frog (also see

Pelophylax bedriagae) 72Anglian Waters 52anophthalmy 22apicomplexan 21apody 15, 22Appenine yellow-bellied toad

(also see Bombina pachypus) 14ARC (also see Amphibian and

Reptile Conservation) 5152, 54

ARG (also see Amphibian and Reptile Groups) 54

ascomycetes 12Ascomycota 11Asia 2Atlantic Biogeographical Region

75Atlantic Ocean 94Atlas of the Herpetofauna of The

Netherlands 59ATV (also see Ambystoma tigrinum

virus) 4Australia 2, 6Azores 93

Bbacilli 11bacteria 1011Balkan Dinaric Alps 77Balkans 77Balkan water frog (also see

Pelophylax kurtmuelleri) 76Barcelona Zoo 88Basidiobolus ranarum 12Basidiomycota 11BAP (also see Biodiversity Action

Plan) 50, 5253Batrachochytrium dendrobatidis

(also see Bd) 6, 8, 1116, 52, 71, 76, 88, 97, 99

Bd (see also Batrachochytrium dendrobatidis) 12, 76

Belgium 7, 1011, 13, 20, 58, 6673Bern Convention 59, 6667Beschermde Natuurmonumenten

(also see Protected Nature Monuments) 59

Biodiversity Action Plan (also see BAP) 50, 52

Birds Directives 59BIV (also see Bohle iridovirus) 4black-spined toad (also see

Duttaphrynus melanostictus) 21Blastocladiomycota 11Bohle iridovirus (also see BIV) 4Bombina pachypus (also see

Appenine yellow-bellied toad) 14

Bombina variegata (also see yellow-bellied toad) 13,

5963, 6768, 7172, 76, 7980, 83boreal toad (also see) 11Boscais newt (also see Lissotriton

boscai) 7, 95Britain (also see UK) 4857, 59Brussels-Capital Region 6667, 69Bufo balearicus (also see green

toad) 75, 78, 90Bufo boreas (also see boreal toad)

11Bufo bufo (also see common toad)

7, 10, 1213, 15, 18, 2021, 49, 52, 67, 76, 81, 83, 88, 90, 95, 97

Bufo calamita (also see Epidalea calamita; natterjack toad) 4243, 45, 52, 49, 54, 6163, 67, 7072 83, 8990, 95

Bufo marinus (also see cane toad) 6 Bufo viridis (also see green toad;

Pseudepidalea viridis) 7778, 80, 83

bullfrog 78, 7172, 7677

CCabaeros National Park 89calcivirus(es) 4California 2, 11Calotriton arnoldi (also see

Montseny brook newt) 8889Calotriton asper (also see Pyrenean

brook salamander) 78, 82, 89cane toad (also see Bufo marinus) 6cannibalism 7captive breeding 88Caragh River 43carcinoma 910cardiac arrest 12carp pox 9Castlemaine Harbour 43catfish (also see Silurus glanis) 77,

96CBIO (also see Research Centre in

Biodiversity and Genetic Resources) 98

Cestoda 21CHACOScomBIO (also see

Research, Conservation and Outreach of the Biodiversity of Temporary Ponds 98

Chioglossa lusitanica (also see golden-striped salamander) 89, 93, 95, 9798

Chlamydia pneumoniae 10Chlamydia sp. 10Chlamydiales 10Chlamydophila abortus 10Chlamydophila pneumoniae 10Chlamydophila psittaci 10Chlamydophila suis 10choanocytes 18Chromista 17Chromomycosis 12

104 Amphibian Biology

Chytridiomycete 12chytridiomycosis 2, 1217, 59,

7879, 88Chytridiomycota 11chytrids 17, 5253, 61, 63, 71, 88,

90, 97climatic change 2, 55, 7879, 88,

90, 93, 9697, 99Clinostomum 21coccidian 21coccidiosis 20coevolution 5Coimbra 94Colorado 11common frog (also see Rana

temporaria) 7, 42, 45, 53, 81common midwife toad (also see

Alytes obstreticans; midwife toad) 13, 69, 95

common spadefoot (also see Pelobates fuscus) 910, 69, 76, 78, 83

common toad (also see Bufo bufo) 7, 13, 5253, 55, 71, 81, 88, 95

common treefrog (also see common treefrog; European treefrog; Hyla arborea) 95

competition 54, 71, 77concatamers 4Conservatoire des Epaces naturels 81Conservaton of Habitats and

Species Regulations 50Conservation of Natural

Habiatats and of Wild Fauna and Flora [European Union] 67

Conservation of Wildlife (Jersey) Law 52

consumption of frogs 77Continental Biogeographical

Region 75Corsica 75, 78, 80crayfish 77, 88, 9697, 99crested newt 9Croatia 7cryptococcosis 12Cryptococcus 12Cryptosporidium fragile 21Czechoslovakia 20

Ddeformities 22Denmark 7Department of Environment

[France] (also see Direction Rgional d Enviroment, de lAmnagement et du Logement) 80

Department of the Environment (Ireland) 44

Department of the Environment, Food and Rural Affairs (DEFRA) (UK) 51

dermatosepticaemia 11Dermocystida 18dermocystid(s) 1819Dermocystidium, rosette agent,

Ichthyophonus, and Psorosper-mium (also see

DRIP) 18Dermosporidium 19Digenea; digenean 21Discoglossus galganoi (also see

west Iberian painted frog) 89, 93, 95, 97

Discoglossus jeanneae (also see Iberian painted frog) 89

Discoglossus montalentii 80, 82Discoglossus pictus (also see

Mediterranean painted frog) 82, 87, 8990

Discoglossus sardus (also see Tyrrhenian painted frog) 13, 80, 8283

disease(s) (also see ID) 123, 55, 59, 63, 71, 7577, 88, 9697

disecdysis 13dispersal 71distributions 54DNA 4, 79, 13, 19DNA polymerase 4Doana National Park 89DRIP (also see Dermocystidium,

rosette agent, Ichthyophonus, and Psorospermium) 18

Durrell Zoo 53, 88Duttaphrynus melanostictus (also

see black-spined toad) 21

Eeastern red-spotted newt (also see

Notophthalmus viridescens) 19Echinostoma 21Ecologische Hoofdstructuur 62ectromelia 15, 22edible frog (also see Pelophylax kl.

esculentus; Rana esculenta) 14 EDP (also see Energias de

Portugal, Sociedade Anonima) 9899

electron microscopy 6, 20ELISA 6endemism 6, 79, 87, 93, 97endospores 19Energias de Portugal, Sociedade

Anonima (also see EDP) 99England 4954, 59Epidalea calamita (also see Bufo

calamita; natterjack toad) 6, 14, 18, 4243, 45, 52, 7072

epidemics 11epidemiology 7, 22

epizootics 11, 17eradication 54, 7677, 99erythema 5, 11erythrocytic virus(es) 4Etrela Mountains 93EU (also see European Union) 53,

80Eucalyptus spp. 96eukaryotic translational initiation

factor 2 5Euproctus montanus 82European Alps 77European spadefoot toad (also

see Pelobates fuscus) 6061European treefrog (also see

common treefrog; Hyla arborea) 60, 77, 95

European Union (also see EU) 16, 71, 95, 98

eutrophication 22, 64, 78exotic species (also see alien

species) 5354, 63, 68, 93, 9698

extinction 78, 12, 99extirpation 2, 11, 17, 44, 50, 5253,

6870, 75, 78, 97

Ffatty acids 10Favourable Conservation Status

(FCS) 51fertilizers 22, 59, 8788fire salamander (also see

Salamandra salamandra) 10, 13, 60, 70, 88, 95

fish 89, 5961, 7779, 88, 9697Flanders 6672flatworm 21flavivirus(es) 4Flemish Land Agency 69Flora and Fauna Act (The

Netherlands) 59flukes 21Ford Motor Company 9899France 10, 1314, 20, 68, 7586French Alps 75Frog Virus 3 (also see FV 3) 4fungi 1118, 5253, 76, 88, 90, 97FV 3 (also see Frog Virus 3) 4

GGambusia holbrooki 96Garonne River 75gene transcription 4Generalitat de Catalunya 88Gers National Park 96Germanasca Valley 78Germany 9, 1314, 5859, 68golden-striped salamander (also

see Chioglossa lustanica) 95goldfish 8Goussia spp. 20

Index 105

granulomata 10Great Britain (see Britain)great crested newt (also see

Triturus cristatus) 20, 5051, 54, 61, 71

green frog(s) (also Pelophylax perezi; Pelophylax sp.; Rana perezi; Rana synkl. esculenta) 7, 9, 72, 83, 88

green toad (also see Bufo balearicus; Bufo viridis; Pseudepidalea viridis) 20, 7778

Guernsey 49Gulf Stream 49Gull Valley 78

Hhabitat loss; habitat fragmenta-

tion; habitat destruction 5054, 5961, 7071, 7576, 78, 83, 88, 90, 95, 99

Habitats Directive [European Union] 43, 4546, 50, 59, 62, 6667, 7072, 75, 80

Habitat Suitability Index 54haemorrhage 45, 7, 11, 17harvesting of amphibians 2heat waves 96hemorrhagic disease 45, 7herbicides 59Herpesviridae 8herpesvirosis 34, 810herpesvirus(es) 4, 6, 810Herpetological Association of

Spain (also see AHE) 89hibernation 5, 9, 60histopathology 13, 22host population 16host-switching 16hotspot(s) 50hybridization 72Hyres Island 80Hyla arborea (also see common

treefrog; European treefrog) 5963, 6768, 7172, 7677, 83, 90, 93, 95, 97

Hyla meridionalis (also see Mediterranean treefrog) 77, 83, 90, 93, 95

Hyla sarda 83 hyperkeratosis 6, 12hyperplasia 12

IIberia 93, 95Iberian brown frog (also see

Iberian frog; Rana iberica) 7, 88Iberian frog (also see Iberian

brown frog; Rana iberica) 95Iberian midwife toad (also see

Alytes cisternasii) 95

Iberian painted frog (also see Discoglossus jeanneae) 89

Iberian parsley frog (also see Pelodytes ibericus) 95

Iberian Peninsula 8788, 93Ichthyophonida 18ichthyophonid(s) 18

Ichthyophonus 19Ichthyosaura 20Ichthyosaura alpestris (also see

alpine newt; Mesotriton alpestris) 7, 15, 1920, 53, 59, 6263, 67, 76, 82

ID (also see diseases) 23immunity 1314, 16immunohistochemistry 6, 13infectious agents 2Institute of Zoology 52introductions 16, 5354, 60, 72, 93invasive species 71, 99Iraty Forest 79 Ireland 4247Iridoviridae 4iridovirus(es) 4, 67, 88, 90Isle of Man 49Italian agile frog (also see Rana

latastei) 5Italian crested newt (also see

Triturus carnifex) 9, 77Italy 5, 79, 1314, 1920, 78IUCN 2, 5960, 82, 89, 95

JJDE (also see Jersey Department

of Environment) 5354Jersey 49, 5354Jersey Amphibian and Reptile

Group 53Jersey Department of Environ-

ment (also see JDE) 53Jersey Zoo 53Jura Mountains 75

KKochs postulates 11

LLake Geneva 77Lanzas alpine salamander (also

see Salamandra lanzai) 78Laune River 43Law on Nature Conservation

[Belgium] 71leech 19legislation 10, 43, 50, 53, 55, 6061,

66, 71, 89leopard frog (also see Rana

pipiens) 8Lepomis gibbosus (also see

pumpkinseed sunfish) 61, 77, 9697

Levantine frog (also see Pelophylax bedriagae) 72

LIFE 89, 98lipid 3Lisbon 93Lissotriton 20

Lissotriton boscai (also see Boscais newt) 7, 89, 93, 9597

Lissotriton helveticus (also see palmate newt) 14, 1920, 49, 53, 59, 6162, 67, 76, 8182, 89, 93, 95, 97, 99

Lissotriton helveticus sequeirai 93Lissotriton vulgaris (also see

smooth newt) 7, 9, 20, 4243, 45, 49, 53, 67, 82

Lithobates catesbeianus (also see American bullfrog; Rana catesbeiana) 7, 53, 63, 76, 83

Loire River 75Lorraine 7778Lous Mountains 93Luck frog kidney tumour 8Luxemburg 68

M macroparasitology 13Madeira 93Maine River 43malachite green 17Malaysia 21Mallorca 16Mallorcan midwife toad (also see

Alytes muletensis) 13, 88marbled newt (also see Triturus

marmoratus) 7, 20, 95marsh frog (also see Pelophylax

ridibundus) 4, 6, 72Massive Central 75Mediterranean Biogeographical

Region 75Mediterranean painted frog (also

see Discoglossus pictus) 87Mediterranean Sea 87Mediterranean treefrog (also see

Hyla meridionalis) 77, 95MeetnetAmfibien 63Mesomycetozoea;

mesomycetozoean(s) 1820Mesotriton alpestris (also see alpine

newt; Ichythosaura alpestris) 89metacercariae 2122metastasis 9metamorphosis 5, 20, 53, 63Metazoa; metazoan 21Meuse River 58, 68, 70MHC class 1a genotype 5microbiota 13microparasites 3, 10Micropterus salmoides 96

106 Amphibian Biology

midwife toad (also see common midwife toad; Alytes obstetricans) 7, 13, 61, 88

migration 63Million Ponds Project (MPP) 51Ministry of Ecology and

Sustainable Developmet [France] 80

mitigation 7, 5051, 55MNHN (also see Musum national

dHistoire naturelle) 8082Monchique Mountains 9394monitoring 5455, 6264, 7172,

7982, 84, 8889, 9899Monitoring of Amphibians and

Reptiles of Spain 89Monogenea; monogeneans 21Montes de Valsain Natural Park

89moor frog (also see Rana arvalis)

78, 83mortality 5, 711, 1314, 1623,

63, 76, 88, 96Moselle River 77mountain yellow-legged frog

(also see Rana mucosa) 11Mucor amphibiorum 11Musum national dHistoire

naturelle (also see MNHN) 80mushrooms 11Mycobacteria 10Myxozoa; myxozoan 21

NNational Amphibian and Reptile

Recording Scheme (also see NARRS) 54

NARRS (also see National Amphibian and Reptile Recording Scheme) 5455

National Atlas of Amphibians and Reptiles 93

National Ecological Network (The Netherlands) 62

National Environmental and Conservation Grant 98

National Environment and Rural Communities Act (also see NERC Act) 50

National Nature Policy Plan (The Netherlands) 62

National Parks and Wildlife Service (Ireland) 44

natterjack [toad] (also see Bufo calamita, Epidalea calamita) 6, 14, 4246, 52, 61, 70, 89, 95

Natura2000 62, 80Natural England 52Nature Conservation Ace (The

Netherlands) 59Natuurbeleidsplan 62Natuurpunt 69, 71

Nematoda 21Neotropics 2NERC Act 53Netherlands: see The NetherlandsNew World 2, 19NI Wildlife Order 43North Africa 87North America 2, 5, 78, 10,

1819, 2122North Sea 5859Norway 52Northwest Mountains (Portugal)

9293Notophthalmus viridescens (also see

eastern red-spotted newt) 19nucleic acid 3

OOceania 2oedema 5, 9Office national de des Forts 81OIE (also see World Organization

for Animals Health) 4, 12, 16oncovirus 8oocysts 20Oomycetes 1718Oospores 17Order on the Protection and

Management of Species [Belgium] 71

Oronectes limosus 77

PPacifastacus leniusculus 96, 99palmate newt (also see Lissotriton

helveticus) 9, 12, 14, 19, 61, 81, 95, 97, 99

pandemic 16parasite(s); parasitism 23, 6, 12,

1522, 76Parque del Sureste Natural Park

89parsley frog (also see Pelodytes

punctatus) 95parvovirus(es) 4, 6pathogen(s) 4, 6, 1114, 18, 22, 52,

90pathogenesis; pathogenicity 13,

20Paul do Boquilobo Natural

Reserve 94, 96PCR 67, 10Pellice Valley 78Pelobates cultripes (also see

spadefoot toad; western spadefoot toad) 77, 83, 8990, 93, 95, 97

Pelobates fuscus (also see common spadefoot, European spadefoot toad) 910, 18, 5963, 67, 69, 71, 76, 78, 80, 83

Pelodytes 97

Pelodytes ibericus (also see Iberian parsley frog) 93, 95

Pelodytes punctatus (also see parsley frog) 83, 90, 93, 95, 97

Pelodytes sp. 93Pelophylax (also see water frogs)

77, 79Pelophylax bedriagae (also see

Anatolian marsh frog; Levantine frog) 72, 83

Pelophylax kl. esculentus (also see Rana esculenta and Rana kl. esculenta) 7,

14, 18, 20, 67, 83 Pelophylax kl. grafi 83Pelophylax kurtmuelleri (also see

Balkan water frog) 76Pelophylax lessonae (also see Rana

lessonae) 1920, 49, 51, 54, 72, 83

Pelophylax perezi (also see green frog; Rana perezi) 83, 88, 95

Pelophylax ridibundus (also see marsh frog) 4, 53, 72, 83

Pelophylax sp. (also see green frog) 7, 9, 51

Pealara Breeding Centre 88Pealara Natural Park 89Peneda-Gers National Park 94 ,

99peptidoglycans 10Peronosporomycetidae 17pesticides 22, 59, 88pets 59pH 61phenology 89phospholipid 10Picos de Europa National Park 89Pinail Nature Reserve 77Pinus pinaster 96plants 3, 11, 17, 6061platyhelminth(s) 2, 2122Plethodontidae; plethodontid 2Pleurodeles waltl (also see ribbed

newt, sharp-ribbed newt) 14, 8889, 93, 95, 97

Po Valley 78pollution 2, 18, 22, 59, 61, 64,

7677, 83, 88polydactyly 22polymelia 22polymerization 4polypody 22pool frog 19, 5152POPAMPHIBIEN 81populations 48, 11, 14, 1617,

1922, 43, 5253, 55, 6061, 6364, 6769, 76, 8081, 8789, 90, 95, 9799

Port-Cros Island 80Portugal 7, 1213, 92102precipitation 92

Index 107

predation 53, 59, 71, 77, 93, 96, 99Procambarus clarkii (also see

American red swamp crayfish) 77, 9697, 99

Protected Natural Monuments (The Netherlands) (also see Beschermde)

Natuurmonumenten) 59Protection and Management of

Species [Belgium] 67Proteidae; proteid 2protein 3, 5, 17protein kinase 5protist(s) 2Protozoa; protozoan 2021Pseudepidalea viridis (also see Bufo

viridis; green toad) 20pumpkinseed sunfish (also see

Lepomis gibbosus) 61, 77, 97Pyrenean brook salamander (also

see Calotriton asper) 78Pyrenean frog (also see Rana

pyrenaica) 79Pyrenees Mountains 7576, 78

QqPCR 13 quarantine 21Quaternary 87

RRACE (also see Risk Assessment

of Chyridiomycosis to European Amphibian

Biodiversity) 76radiography 22RaHV-1 (also see ranid herpesvi-

rus 1) 89Rana arvalis (also see moor frog) 9,

18, 20, 59, 62, 67, 78, 83Rana catesbeiana (also see

Lithobates catesbeiana) 63Rana catesbeiana virus Z (also see

RCV-Z) 4, 10 Rana dalmatina (also see agile

frog) 9, 20, 49, 5354, 83, 89Rana esculenta (also see Pelophylax

kl. esculentus; Rana kl. esculenta) 7

Rana iberica (also see Iberian brown frog; Iberian frog) 7, 88, 90, 93, 95, 97

Rana kl. esculenta (also see green frog; Pelophylax kl. esculentus; Rana esculenta) 59, 63

Rana latastei (also see Italian agile frog) 5

Rana lessonae (also see Pelophylax lessonae) 59, 62

Rana mucosa (also see mountain yellow-legged frog) 11, 16

Rana perezi (also see green frog; Pelophylax perezi) 93, 95, 98

Rana pipiens (also see leopard frog) 89

Rana pyrenaica (also see Pyrenean frog) 79, 83, 89

Rana temporaria (also see common frog) 3, 57, 911, 18, 20, 4243, 45, 49, 53, 67, 81, 83, 89

ranavirosis 48, 11Ranavirus 46, 59, 76ranavirus(es) 48, 12, 71ranid herpesvirus 1 (also see

RaHV-1) 8RAVON Foundation 6263RCV-Z (also see Rana catesbeiana

virus 4Red Data Book; Red Data List;

Red List 45, 54, 59, 67, 72, 82, 8990, 93, 95, 97

red-eared slider (also see Trachemys scripta) 96

red leg 1011Regina ranavirus 4Rhine River; Rhine Valley 75,

7778, 83Rhone River 75reintroduction 16, 44, 63, 71, 89Reptile, Amphibian & Fish

Conservation Netherlands 62Reptiles 93Research Centre in Biodiversity

and Genetic Resources (also see CBIO) 98

Research, Conservation and Outreach of the Biodiversity of Temporary Ponds (also see CHACOScomBIO) 98

restoration 63, 7172, 98Rhabdias bufonis 21Rhine River 58ribbed newt (also see sharp-

ribbed newt; Pleurodeles waltl) 14, 8889

Ribeiroia 21Risk Assessment of Chyridiomy-

cosis to European Amphibian Biodiversity (also see

RACE) 76 RNA 4RoIs Wildlife Act 43Royal Decree (Belgium) 66

SSAC (also see Terrestrial Special

Area of Conservation) 80Salamandra algira 89Salamandra atra (also see alpine

salamander) 77, 82Salamandra corsica 82

Salamandra lanzai (also see Lanzas alpine salamander) 78, 82

Salamandra salamandra (also see fire salamander10, 13, 5960, 62, 67, 7071, 76, 79, 82, 88, 90, 93, 95, 97

Salamandra salamandra crespoi 93, 97

Salamandra salamandra gallaica 93, 98

Salamandra salamandra terrestris 15

Salamandridae; salamandrid 2salmonids 77Sambre River 70San Mamede Natural Park 94Santee-Cooper ranavirus (also see

SSRV) 4So Memede Mountains 93So Miguel Island 93SAP (also see Species Action Plan)

52Saprolegnia 90Saprolegniaceae 17saprolegniaceaen(s) 18Saprolegniales 17saprolegniosis 1718SARE (also see Monitoring of

Amphibians and Reptiles of Spain) 87, 89

Scheldt River 58Scotland 19, 49, 5152Segura River Basin 88Seine River 75serology 6Serra da Estrela Natural Park 94,

97, 99sexually transmitted diseases 15sharp-ribbed newt (Pleurodeles

waltl; also see ribbed newt) 95SHF (also see Socit Herp-

tologique de France) 76, 8082SIARE 87, 89Siberia 87Sierra Nevada 11, 16Silurus glanis (also see catfish) 96Singapore Grouper iridovirus 4Sintra Cascais Natural Park 98Sites of Special Scientific Interest

50Slovakia 4smooth newt (also see Lissotriton

vulgaris) 7, 9, 4243, 45snails 22Socit Herptologique de France

(also see SHF) 76southern marbled newt (also see

Triturus pygmaeus) 95spadefoot toad (also see Pelobates

cultripes) 89

108 Amphibian Biology

Spain 7, 1314, 18, 7677, 79, 8791, 94, 9697

Special Areas of Conservation 50Species Action Plan (also see SAP)

51Speleomantes strinatii 80, 82SSRV (also see Santee-Cooper

ranavirus) 4Staatsbosbeheer 62 State Forestry Service (The

Netherlands) 62Strait of Gibraltar 87sterols 10Stichtingen voor Landschapsbeheer

62stratum corneum 6, 12Sudoeste Alentejano and Costa

Vicentina Natural Park 94, 96Sweden 18, 52Switzerland 13, 1920synergism 90

Ttapeworms 21Tejo River; Tejo Valley 9294, 96Terrestrial Special Area of

Conservation (also see SAC) 80

Testudo iridovirus 4The Netherlands 7, 10, 18, 5865,

68tiger salamander 8togavirus 4toxicology 13Trachemys scripta (also see

red-eared slider) 96trade in amphibians 2, 1617translocations 52, 70, 98transmission [of disease] 78,

1416, 77Trematoda; trematodes 2122Triturus 20

Triturus carnifex (also see Italian crested newt) 9, 54, 63, 77, 82, 93

Triturus cristatus (also see great crested newt) 20, 4951, 5455, 6162, 67, 71, 80, 82

Triturus marmoratus (also see marbled newt) 7, 20, 54, 82, 89, 93, 9596

Triturus pygmaeus (also see southern marbled newt) 89, 93, 95, 97

Turkey 77turtles 88 Tyrrhenian painted frog (also see

Discoglossus sardus) 13

UUK (also see Britain) 68, 1314,

18, 20, 22, 4857

University of Kent 51University of Lisbon 98University of Porto 98urbanization 59, 76, 95USA 9

VValongo Mountain 98vectors of disease 53vIF2 5viraemia 6virion(s) 4, 78virulence 13, 16, 1819virus(es) 210Vosges Mountains 75

WWales 4951, 54Wallonia 6672waterfowl 59water frog (also see Pelophylax) 77,

79Waterschappen 62Western Alps 78Western spadefoot toad (also see

Pelobates cultripes; spadefoot toad;) 95

West Iberian painted frog (also see Discoglossus galganoi) 95

Wildlife and Countryside Act 50World Organization for Animal

Health (also see OIE) 4, 12

XXenopus 4

Xenopus laevis (also see African clawed frog) 53, 76, 83, 93, 99

Yyellow-bellied toad (also see

Appenine yellow-bellied toad; Bombina variegata) 60, 68, 76, 7980

Zzoosporangium(a) 6, 12zoospores 1112, 14, 17, 19Zurich 19