– A pride of starving lions on the brink of death are being nursed back to health after being freed from their tiny cages in which they were slowly being driven mad, thanks to the generosity of MailOnline readers.

For the first time in their miserable lives, lioness Mary and her two cubs Zita and Geeta padded tentatively into more space than they had ever known when they were released into purpose-built compounds in the Caucasus Mountains, constructed with £30,000 of donations.

Finally they had room to move, to stretch, to see beyond the single row of steel rods which imprisoned them in what was nicknamed ‘the saddest zoo in the world’ in Armenia.

The cuts on their heads – suffered from repeated banging against the bars and sides of their prison – are now gone.

Mary has piled on a massive 100lb in weight since January and her ribs no longer protrude through her thick, glossy coat.

And their eyes shine bright thanks to good food, added vitamins and a love they have never experienced before.

MailOnline’s story of their suffering in cramped, filthy cages built from truck containers touched the hearts of readers worldwide.

Alan Knight OBE, the CEO of East Sussex-based animal welfare charity International Animal Rescue (IAR), said in all his years of fundraising he had ‘never known of such a swift response’ to an appeal for help.

But while the lions are safe and will never endure such suffering again, more money is needed to get them to the end of their journey: England.

IAR needs to raise an estimated £15,000 to fly them home while the UK’s The Nature Foundation is appealing for more than £200,000 to construct an enclosure in a Peak District sanctuary where the lions can live out their days in peace.

But so much has changed for the beautiful lionesses already. When MailOnline reported their plight in January, there were fears that any help would come too late because the wheels of animal charities can often turn with sclerotic – and lethal – slowness.

Mary, seven, and cubs Geeta, four, and Zita, two, were among the seven residents of the private zoo that had fallen into decay and squalor in the city of Gyumri – the place where the family of Armenian-American superstar Kim Kardashian and her sister Khloe stem from.

Bears Masha and Grisha from the same zoo were taken under the wing of Libearty Sanctuary in Romania, funded by Foundation Brigitte Bardot, where they are now being cared for under the auspices of the Armenian government’s Foundation for the Preservation of Wildlife and Cultural Assets.