Even the first sunshine of spring is not enough to prevent the lion enclosure at Copenhagen Zoo from looking forlorn. One female lion lounges on a branch basking in the rays, while the other lolls on the dusty ground beneath her. Otherwise the pen is empty.
"They're shifting around the pack a little - that's why those females are here alone," explains Martin, a zookeeper. "We have a new male lion in the holding facility and before we can introduce him to our females, they need to get comfortable."
He doesn't volunteer the fact that, as part of this process, the zoo last week slaughtered two older lions and two cubs. But when it's brought up, he's unapologetic.
"It's a necessary part of keeping a healthy population," he shrugs. "Because we don't bring in animals from the wild any more, we need to do this."
When the zoo announced that it was putting down the four lions, it argued that if it hadn't, the new male would have done the job himself in a much bloodier fashion.
"The new male in the pride would have killed the immature males as soon as he got the chance," the statement read. That's not all. The two older lions would have fought with the new male, and the older males would have killed any cubs fathered by the new male.
"This may of course seem harsh, but in nature it is necessary to ensure a strong pride of lions with the greatest chance of survival," the zoo explained.
It said it had tried and failed to find another institution willing to take the animals and this, along with the risk that the elder male might also impregnate his two daughters, left it with little choice.
"The zoo is recognised worldwide for our work with lions," chief executive Steffen Strade concluded in the statement. "I am proud that one of the zoo's own brood now forms the centre of a new pride of lions."
Since February, when the zoo put down a healthy giraffe called Marius, dissected it in front of children and then fed the carcass to the lions, Strade and his scientific director, Bengt Holst, have faced intense criticism from animal rights activists. Online petitions to save the giraffe, sack Holst and even close the zoo have gathered tens of thousands of signatures across the world.
"We have all had some hate mail and death threats and other nastiness," Carsten Grondahl, one of the zoo's vets, says. "But that's all emotional, and you can't argue with emotions."
"We have a low profile now," says Mette Nyborg, another zookeeper, when asked about future dissections. "There has been too much bad attention." In Denmark, however, at least judging by the 10 or so families at the enclosure on Saturday, almost everyone seems to support what the zoo has been doing.
"It's totally okay," says Mette Brendstorp, who is visiting with her daughters, aged 2 and 4. "If you talk about what's cruel, it's wanting to go to the zoo and look at all the animals, and then getting hysterical when the zoo takes responsibility to ensure that there is no inbreeding."
She approves of the zoo's decision to dissect the giraffe. "I'm a schoolteacher and I wouldn't blink twice about bringing my fifth grade class to see a dissection. They are not seeing it being killed; they are just seeing it being cut open."
"The public is very supportive," Martin, the young zookeeper, tells me. "They see that we have good intentions and we generally have very broad acceptance of the work that we do."
Grondahl believes that the uproar over the zoo's practices, which it had been quietly following without criticism for many years, has come about because people are now too distanced from the natural world.
"It's because we're far from nature now. I think that people don't realise that the meat they buy in the supermarket was once an animal."