© Ben Birchall/PAThe Somerset Levels at the height of the floods. As communities begin to recover, the cost to wildlife is only now becoming apparent.
Seals, moles, hedgehogs, badgers, mice, earthworms and a host of insects and seabirds are among the unseen casualties of the floods, storms and torrential rains of the last few weeks, say wildlife groups.
As the waters started to subside across England, conservationists reported that about 600 guillemots, puffins, razorbills, kittiwakes and gulls have been washed up on the south coast and 250 seals drowned in Norfolk, Cornwall and the Channel Islands. A further 11,000 seabirds are reported to have been found dead on the French coast.
"The relentless storms hitting our coast have had a cumulative effect on animals, which can usually cope with bad weather, but are now on really low reserves and are dying in large numbers," said Niki Clear from Cornwall Wildlife Trust, which has reported that dozens of seal pups were washed up on beaches.
David Jarvis, director of British Divers Marine Life Rescue, said his organisation has been heavily involved in rescuing seals: "Since January we have had 88 callouts to rescue marine wildlife, the vast majority concerning seal pups. We would normally expect approximately 150 in the whole year."
Natural England, the government's independent wildlife advisory body, has reported damage to 48 of the UK's most important wildlife sites, including those of special scientific interest and national nature reserves. Tim Collins, Natural England's coastal specialist, said: "It is estimated that some 4,500 hectares of designated coastal nature conservation sites in England were flooded.
"The sites affected mostly comprise coastal grazing marsh, saline lagoons and reed beds. All of these sites are of national importance, and 37 are also of international importance.
"Sea walls protecting at least 80 conservation sites have been breached in Norfolk, Suffolk, Kent and the Tees estuary."
The scale and extent of the impact of flooding on many species is still being assessed, but it is expected that hibernating animals will have been the worst affected.
Marina Pacheco, director of the Mammal Society, explained: "Moles and field voles will have drowned if the flooding was rapid. If it was relatively slow, then they would have had a chance to move away, but that would have brought them into conflict with established populations so may result in injuries as they fight each other off."
Mark Jones, of Humane Society International, said many other animals have also been affected: "Some badger setts will almost certainly have been damaged or flooded out completely, meaning that whole families could be disturbed."
Matt Shardlow, head of the invertebrate charity Buglife, noted that much depends on the habitat: "Bumblebees, earthworms, snails, beetles and caterpillars living on floodplains have all been at risk in the floods and waterlogged ground.
"The biggest impact will have been in places that are not used to being flooded. Species living in floodplains will have adapted and can survive under water for considerable periods. It's been bad news for hibernating animals.
"Mould is the deadly enemy of insects and it does not stop growing. It means there could be fewer grubs for birds to feed on."
The floods will have carried away thousands of tonnes of bug-rich topsoil, but if they continue the impact could be far worse, said Shardlow.
"After several weeks under water, plants start to decompose and, with oxygen in short supply, give off noxious fumes. If the floodwater were to be contaminated with pesticides or other toxic industrial chemicals, it could be devastating."
Butterflies had a good summer in 2013 following some of the sunniest months in years, but many could have suffered from the wet. "We can expect fewer butterflies this year," said a spokeswoman for the Butterfly Trust.
Most natural sites are able to take occasional flooding, but any water polluted by overflowing drains, septic tanks or pesticides from farms and parks is likely to have exacted a toll on many animals. University of Reading microbiologists found highly elevated bacteria levels when they tested floodwaters in Somerset.
But it's not all bad news. Some species are expected to have thrived in the wettest months recorded in 250 years. Wading birds and wildfowl have been able to feed across much larger wetland areas and are now feeding in large numbers on flooded grasslands and taking advantage of drowned invertebrates.
But even fish species have suffered. Nearly 5,000 fish, for example, were found dead in fields near Goring-on-Thames in Oxfordshire after the river flooded then subsided quickly. "When floods come, you can also lose the fry - newly-hatched fish - which get swept away," said Martin Salter of the Angling Trust.
With very few frosty nights recorded so far this winter, and remarkably warm weather since Christmas, many conservationists now fear that a cold snap could prove disastrous for wildlife. "A month of ice and snow on top of a long period when animals could not hibernate would be very serious indeed," said Shardlow.
THE WINTER'S VICTIMS
BUTTERFLIES
Species such as the brimstone and small tortoiseshell, which thrived in last year's summer sunshine, are in danger because of the incessant rain and lack of frosts.
"The mild weather increases activity levels of pathogens, making the caterpillar, egg and pupae stages of the butterfly's life cycle all highly susceptible to diseases," says Liam Creedon of the Butterfly Conservation charity. "It makes butterflies vulnerable to increased predation by bats and birds."
SEALS
The December tidal surge which coincided with end of the seal breeding season was followed by a series of massive Atlantic storms. Together they wiped out several seal colonies and hundreds have been found along beaches that were dead, stranded, injured or too weak to survive. The worst-hit areas include Lincolnshire, Norfolk and Cornwall.
BIRD LIFE
The kingfisher, which lives by seeing then catching river fish, has been badly hit because the rains and floods have washed down so much silt that the waters have become too cloudy for them to feed. Wading birds such as snipe, redwing and redshank, which thrive on the Somerset Levels, will be in difficulty if the floods continue into their nesting season. Seabirds have perished in their thousands in the heavy seas.
TREES
Hundreds of ancient trees - including 300-year-old oaks, veteran ashes and beeches - have fallen in the last three months' storms. The New Forest in Hampshire has been badly hit, with record numbers of trees downed. The National Trust reports more damage in some areas than any time since the great storm of 1987. The Forestry Commission estimated that the St Jude storm in November may have resulted in the death of 10 million trees.
EARTHWORMS
Earthworms, which hibernate and breathe through their skin, could have been badly hit by the heaviest winter rains ever recorded in Britain. They depend on moisture to breathe, so they like damp soil, but are highly vulnerable to waterlogging and flooding. Tens of thousands may have suffocated in the floods, leaving shrews, moles, certain beetles and birds hungry.
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