The young male jaguar who went on a deadly rampage against some of his fellow animals inside the Audubon Zoo early Saturday appears to have facilitated his own escape, breaking through a steel cable barrier on the roof of his enclosure, a zoo spokeswoman said Monday as the toll of dead animals reached nine.

All told, the jaguar named Valerio killed five alpacas, three foxes and an emu after freeing himself from the “Jaguar Jungle” enclosure before 7:20 a.m. Saturday, a few hours before the park’s scheduled opening time.

[Update: Audubon Zoo's Jaguar exhibit closed until 'stronger materials' replace bitten-through steel wire]

The last of Valerio’s surviving victims, a fox named Rusty, died Monday morning, zoo officials said.

A tranquilizer dart ended Valerio’s killing spree. The 3-year-old jaguar, imported last year from the San Diego Zoo, remained in isolation Monday, zoo officials said. There are no plans to euthanize him.

Zoo officials initially said only that the jaguar escaped through what they found to be a "compromised" part of the exhibit's roof. But spokeswoman Katie Smith said Monday that preliminary findings of an investigation indicate that the breach “was caused by the animal.”

A spokesman with the zoo’s accrediting body, the Association of Zoos and Aquariums, said Audubon Zoo officials were taking the proper steps in reporting the incident to both the association and the U.S. Department of Agriculture.

The spokesman, Rob Vernon, said it would be up to Audubon officials to investigate the incident and issue a report to the association, which would then decide on any action.

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“Whenever something like this happens, which doesn’t happen very often, our standard procedure is for the member zoo to do exactly what Audubon Zoo is doing,” Vernon said.

The association’s jaguar program includes 115 jaguars held in 47 facilities, mostly in the U.S., according to Stacey Johnson, a San Diego Zoo official who coordinates the program.

The Audubon Zoo was last accredited following an inspection last fall, Vernon said, though he declined to release that inspection report. To pass, Vernon said, the zoo must show it meets or exceeds minimum standards. For jaguars, those standards are spelled out in a 128-page manual.

The manual includes specifics on the thickness and width of steel wire barriers where they are used, though it also says that, “If possible, institutions are recommended to completely enclose the top of any jaguar enclosure.”

Pound for pound, jaguars have the strongest bite force of any big cat in the world, as well as “excellent jumping and ambush capability,” according to the manual.

“A major risk to jaguars is the failure to use suitable materials to contain the animals. Jaguars often grasp mesh with their canine teeth, over time damaging or breaking them entirely. … Single-strand wire mesh types like chain link or welded wire can be compromised by animals biting and pulling them apart.”

Valerio, one of two jaguars at the zoo, appeared to use those abilities to full effect Saturday morning, making quick work of other animals as he ventured into nearby exhibits.

Jaguars are known as “apex predators” for their standing atop the food chain, with a lack of natural rivals where they typically live, from northern Argentina to the occasional sighting across the Mexican border in Arizona.

They are also eager predators with a broad palate, known to feast on dozens of animals, both big and small.

Still, Dr. Howard Quigley, the jaguar program executive director for Panthera, a wild cat conservation group, said Valerio may have been the first jaguar to kill an alpaca — let alone five of them.

Alpacas also dwell in South America, but at far higher elevations than jaguars.

Quigley noted that jaguars “aren’t usually a real preferred animal for many zoos, because they tend to be not very active, not very exciting for the viewing public.”

Quigley, who described jaguars as "versatile predators," expressed little surprise at how Valerio reacted once free of his enclosure.

“Any predator comes with a sort of hardwired behavior that has made them successful in the wild. They have a natural reaction to potential prey,” he said.

“This animal escaped for some reason. Whether he’s a Houdini or a mega-strong animal, I don’t know. But once they’re out, there is, as with most predators, a stimulus-response reaction. If an animal flees, they attack it and kill it," Quigley added.

“What happens with a lot of predators, when confronted with a lot of prey, they simply kill one, see another one, move on it, kill that one. It’s called ‘surplus killing,’ and most predators have this kind of natural response.”

Jaguars, though, rarely experience such a captive group of targets in the wild, he said, adding that jaguars tend to steer clear of humans.

“I’m more afraid of African killer bees or snakes,” said Quigley, who said he has been studying jaguars for 30 years.

“They’re probably, of the big cats, the one that is least likely to attack humans,” he said, describing the jaguar as “very powerful but timid about the use of its power, and especially timid about the use of its power against people.”

Most incidents of jaguar attacks on humans involve hunters “who have already shot the animal or cornered it. Not to say if a jaguar got loose at the zoo it wouldn’t be dangerous to people. It would be very dangerous.”

Rarely do they get free of their zoo enclosures. Jaguar attacks at zoos mostly involve handlers who enter their enclosures by mistake while the wild cats are loose.

For instance, three jaguars killed a zookeeper at the Vienna Zoo in Austria in front of a crowd of visitors in 2002.

In 2007, a Denver zookeeper opened the door to a jaguar’s cage and was attacked and killed, according to media reports.

In one case, in 1998, two jaguars escaped under a fence from their enclosure at a zoo in France and one of them killed a young boy. His father suffered serious head wounds trying to save his son.

Staff writer Ramon Vargas contributed to this report.


Follow John Simerman on Twitter, @johnsimerman.