Titian’s quivering dogs, baying for blood; Velázquez’s regal horses; the sly black cat at the end of the bed in Manet’s “Olympia”: what nuances of sensibility are revealed in a musée imaginaire of great artists painting animals. And then there is Francis Bacon, whose theme and subject is the bestial condition of humankind — instinctual, rapacious, stalked, caged. It goes without saying that Francis Bacon: Man and Beast, opening at the Royal Academy in London when government guidance allows and certainly among 2021’s most eagerly awaited shows, is an exhibition for now: Bacon’s enduring motif is man alone in a room, fearful and furious. But isolation, in art as in life, is beginning to pall, and the anticipated joy of this show is a new inflection to the modern master of gloom. With his animals, Bacon’s grand guignol menace is varied, inventive, sometimes even touched with comedy.

The perched ape in “Study for Chimpanzee”, depicted sketchily against brilliant cerise, is enjoying posing; he was based on performing monkeys in the 1950 adventure film “The Elusive Pimpernel”. “Study of a Baboon”, playfully interwoven with a tree, as if the animal is both imprisoned by its branches and incongruously sprouting horns, is softly delineated, almost tender. A pair of shadowy forms, darker than the night, sit hunched on a bright lapis lazuli ground in “Owls” — sinister yet touching. “Chicken”, its trussed feet a parody of a crown, is sourced from a Terence and Caroline Conran cookbook illustration, a smudge of blood added.

Bacon, son of a brutal racehorse trainer, grew up with animals, including the family’s red setters, but they triggered his severe asthma; life-long he was frightened yet fascinated by dogs. Set bizarrely in the benign landscape of Monte Carlo’s coastal palm-fringed road, a snarling canine spinning in a green circle — flowerbed? roulette wheel? — derives from Eadweard Muybridge’s photographs of a mastiff called Dread. In a magnificent grisaille follow-up, “Man with Dog”, the pair on a pavement by a drain cover, Bacon blended Muybridge’s source with Giacomo Balla’s futurist “Dynamism of a Dog on a Leash”. The man becomes a mysterious spectre, the dog a predator of the sewers, and even the smeared, out-of-focus pavement seems to move in Bacon’s dystopian urban version.

Bacon’s animals are not all about foreboding. The most serene painting he ever made is the broadly brushed “Elephant Fording a River”, expressing his pleasure, visiting South Africa, in watching big game. Bacon subsequently destroyed “Rhinoceros”, from the same time — 1952 — as too literal. But the feel of leathery skin is already visceral in “Head I” with which the Royal Academy show begins: a monkey with bared animal teeth dovetailed with Velázquez’s “Portrait of Pope Innocent X”. A papal tassel brushes the ear of this weird cross-species.

Bacon said he wanted to paint like Velázquez, but with the texture of hippopotamus skin. “Heads” 1-VI (1948-49) evolve from the animal-like to the first iconic, caged, screaming purple-robed Pope, face reduced to an open mouth, above which swings again the mocking phallic tassel. Also on display will be the sumptuous but unresolved, rarely shown “Pope and Chimpanzee” (1960), where a frenzied, overscaled turquoise ape attempts to upstage the pontiff imprisoned in a glaring pink block.

Animal-man combat excited Bacon: he thought bullfighting “a marvellous aperitif to sex”. An RA coup is to reunite 1969’s three “Study for Bullfight” paintings: the brown-mauve figures of the bull and toreador are fused, deformed, seeping ejaculatory white impasto in a dance of death eroding lines between flesh and meat, eroticism and violence. Another first is the inaugural UK display of “Study of a Bull” (1991), Bacon’s final completed painting. The bull, symbol of the life-force for Bacon as for Picasso, bows out: vaporous and ghostly, backed into a corner between a white mirror and a black void in a monochrome canvas incorporating particles of dust — self-portrait as a declining, threatened beast.

Man uninhibited, out of control, in a meaningless universe: there is an argument that Bacon always portrayed human experience as animal extremes of suffering or ecstasy. The suspended figure in “Fragment of a Crucifixion” is derived from a photograph of a barn owl carrying its prey. In “Two Figures in the Grass”, which provoked complaints to the police when shown at the ICA in 1955, coupling men resemble fighting animals. Bacon recorded his love affairs in imagery of hunter and hunted. “Man Kneeling in the Grass”, a bulky nude, buttocks prominent, crawling through grasslands evoked in long flicked strokes, was modelled on Peter Lacy, Bacon’s sadistic lover who thrilled the masochist artist by suggesting he chain him up in his stable (“you could sleep and shit there”). In another portrait, Lacy has the ringed, piercing eyes of a bird of prey.

In “Portrait of George Dyer Crouching” (1966), Lacy’s successor squats, naked, on a gang plank, like an animal about to jump; he stares down into an abyss. Dyer committed suicide two days before Bacon’s major Paris retrospective in 1971, but continued to appear in the paintings. In “Portrait with Bird in Flight” (1980), he is a caged, beautiful Ganymede, about to be abducted by Jupiter in the form of an eagle. As Jupiter gave Ganymede eternal life among the gods, Dyer acquires immortality through Bacon’s images.

Hybrid animal-human forms of classical myth launched Bacon’s oeuvre: “Three Studies for Figures at the Base of a Crucifixion” (1944), which he considered the beginning of his career, features bony, animal-toothed, long-necked quadrupeds with human hair and ears — Bacon identified them as the Greek Furies, goddesses of vengeance.

Forty years on they still permeate the triptychs, of which two outstanding late examples come to the RA. In the bullfight “Triptych” (1987), a swooping biomorph Fury, derived from a photograph of a diving brown pelican, flies low towards the bull, a fading, drooping figure, as if to pierce it with its sharp beak. A similar pelican/Fury, a coagulated knot of blood in its ear, opens, on the left panel, “Triptych Inspired by the Oresteia of Aeschylus”, and presides over the blood seeping under the door, a thick impasto trail, signifying murder.

“You don’t see them, you don’t, but I see them, they are haunting me down,” says Orestes of the Furies. Bacon gave those inner voices of fear and guilt, despair and entrapment, for which the Furies are ancient metaphors, grandiloquent modern expression. “Consciousness of mortality sharpens one’s sense of existing,” he said; his genius was to find a heightened pictorial language of animal sensation to evoke also that imagination and interiority which distinguish man from beast.

January 30-April 18, royalacademy.org.uk

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