The recent death of Cecil the lion has shone a spotlight on sport hunting, especially for threatened species. Advocates say responsible hunting helps preserve wildlife and wilderness, while critics note eco-tourism does, too — and with fewer casualties.

Yet hunters and tourists can both harm wildlife when they're reckless. Even many trophy hunters have denounced Cecil's killers, whose tactics are widely considered unethical and inept. Aside from lacking a permit to kill Cecil, they reportedly baited the pride leader out of a national park, shone a (literal) spotlight on him, failed to kill him with an arrow, then tracked him for 40 hours before ending his misery.

Irresponsible tourism may pose subtler threats, but it can also add to animals' existing stress from things like poaching or habitat loss, possibly altering behavior enough to affect survival. And while non-lethal eco-tourism makes certain animals directly more valuable to humans alive than dead, sometimes a few regulated deaths can also help a species by making its natural habitat more profitable.

It's generally better to save wildlife without sacrificing any lives in the process, but not all habitats appeal equally to non-hunting tourists. People may flock to see charismatic megafauna like elephants and whales, but what about ducks and quail? They aren't big-ticket tourist magnets, but in much of the U.S., they are a lucrative draw for hunting licenses. Duck hunters are legally required to buy duck stamps, for example, with 98 percent of all profits going directly to wetland conservation. And that's good for all kinds of native wildlife — the ducks just pay the bills.

But things are trickier when hunting a threatened species. Although some 200,000 lions existed a century ago, they're now down to about 20,000. Killing one lion in 2015 thus has more impact than killing a lone mallard, turkey or white-tailed deer, all of which number in the millions and have faster reproduction rates. So when is hunting actually helpful to wildlife, and when is it unsporting or unsustainable?

teddy bear cartoonThis 1902 cartoon by Clifford Berryman gave rise to "Teddy bears." (Photo: Clifford Berryman/U.S. National Park Service)

A 'peculiar charm'

Humans are natural hunters, having evolved bodies for long-distance pursuit and brains that can invent potent weapons like atlatls, bows and guns. Food was our original motivation to hunt, and many people still hunt for that reason, often with little or no criticism. From subsistence hunters to resourceful sportsmen, catching your own food is more about reconnecting with nature than trying to defeat it.

In places where people have disrupted natural ecosystems, hunting may also help control wildlife overpopulation. By removing wolves from most of the U.S., Americans ended up with too many white-tailed deer, which in turn cause problems like overgrazing and road collisions. Hunting licenses offer a way to keep deer populations in check, although this concept is more controversial when applied to less abundant animals, especially popular predators like wolves and bears.

(In many cases, the problem isn't really too many animals; it's that humans have built vulnerable homes, farms and ranches within their habitat. And there are ways to keep predators away from livestock without killing them.)

Beyond food and culling, some people hunt primarily for trophies. Big-game trophy hunting is divisive, but not all hunters are like those who killed Cecil. Ethical hunters pride themselves on quick kill shots from as close as possible, and value the experience of a hunt more than getting a trophy at any cost. U.S. President Theodore Roosevelt was an avid trophy hunter, yet he famously declined to shoot a subdued bear in 1902, inspiring the icon of "Teddy bears."

"In hunting, the finding and killing of the game is after all but a part of the whole," Roosevelt once wrote. "The free, self-reliant, adventurous life, with its rugged and stalwart democracy; the wild surroundings, the grand beauty of the scenery, the chance to study the ways and habits of the woodland creatures — all these unite to give to the career of the wilderness hunter its peculiar charm."

Chincoteague National Wildlife RefugeDucks and geese gather at sunset in Virginia's Chincoteague National Wildlife Refuge. (Photo: Steve Hillebrand/USFWS)

Land's sake

Research suggests regulated trophy hunting can help protect habitat for endangered animals, at least in some cases. But it's also capable of making things worse, as seen with Cecil. That's why some African countries like Kenya and Botswana have banned trophy hunting, instead banking on the power of eco-tourism alone. Australia also recently banned the import of trophy-hunted lions, and the U.S. is considering a similar move. Several major airlines have decided to stop carrying certain animal trophies, including lions, adding to international momentum against the practice.

Yet according to a 2005 study, the legalization of white-rhino hunting in South Africa spurred private landowners to reintroduce the animals, enabling a population boom from fewer than 100 to more than 11,000. (Some 20,000 southern white rhinos now exist, mostly in South Africa, but the northern subspecies is on the brink of extinction.) And as study author Nigel Leader-Williams wrote in a letter to Science magazine, legalized hunting had similar advantages for Zimbabwe's elephants.

"In Zimbabwe, implementing trophy hunting has doubled the area of the country under wildlife management relative to the 13% in state protected areas," Leader-Williams wrote. "As a result, the area of suitable land available to elephants and other wildlife has increased, reversing the problem of habitat loss and helping to maintain a sustained population increase in Zimbabwe's already large elephant population."

There is nuance involved, however. Any hunting needs a ceiling, since humans have a long history of hunting animals to extinction. But depending on the species, limits may be needed on the age and sex as well as the number of animals killed. Natural predators tend to kill the weakest members of a species, minimizing the dent in reproductive capacity, yet many human trophy hunters seek the biggest, most impressive males. Cecil, for one, was a pride leader with seven cubs. A rival male could now kill his progeny, compounding the impact of his death (although reports suggest Cecil's "brother," Jericho, may have adopted the cubs in his absence).

For hunting to benefit wildlife, the money it generates needs to flow directly toward conservation efforts. The U.S. duck stamp program is a good example of that, but the connection between hunting fees and conservation is often less transparent. Corrupt guides and government officials can take advantage of such murkiness — a 2013 report by Economists at Large found that only 3 percent of revenue from hunting companies in Africa stays in communities around the hunting areas.

"The vast majority of their expenditure does not accrue to local people and businesses," the report found, "but to firms, government agencies and individuals located internationally or in national capitals." The conservation value of hunting therefore depends heavily on how it's managed and regulated.

lion prideA group of lions kick back in Kenya, where eco-tourism is worth up to $4.1 billion (U.S.) per year. (Photo: Shutterstock)

Curiosity killed the cat

Some people consider hunting a type of eco-tourism, at least when it's done responsibly. But for many, the term implies a lower-impact tour through nature that's more about shooting photos than firearms. Either way, research suggests non-hunting tourists play a larger role in conserving Africa's wildlife.

"Nature-based tourism does play a significant role in national development, but trophy hunting is insignificant," the Economists at Large report states. "Across the investigated countries, trophy hunting revenue was only 1.8% of tourism revenues." A 2004 study by South Africa's Terrestrial Ecology Research Unit reached a similar conclusion, reporting that non-lethal eco-tourism in private preserves yielded "more than 15 times the income of livestock or game rearing or overseas hunting."

Although well-managed sport hunting can preserve habitat for certain species, the loss of even one animal can spur far-reaching problems for others. Before his death last month, Cecil was being monitored by scientists at Oxford University, whose previous research shows even legal hunting around the edges of Zimbabwe's Hwange National Park — where Cecil lived — is bad for local lion populations.

"Each removal of a male lion by hunters on the borders of the park created a 'territorial vacuum' which drew males from further inside the protected area into boundary areas, where they too became vulnerable to hunters," the scientists write, referring to a study conducted between 1999 and 2004. Zimbabwe responded to that report by banning trophy hunting around the park from 2005 to 2008, letting the researchers record "an almost exact reversal of the vacuum effect observed between 1999 and 2004. The lion population increased by 50% and the skewed population structure, caused by excessive off-takes of adult males, disappeared."

African elephantsThanks to eco-tourism, wild elephants are worth up to 76 times more alive than dead. (Photo: Tony Karumba/Getty Images)

Taking photos, not lives

It's worth noting that legal trophy hunters pose less danger to many of Africa's disappearing species than poachers do, especially rhinos, elephants and even giraffes. But whether the threat comes from poaching, habitat loss or any other human pressure, the key for conservationists is to help local people make more money by keeping their forests, fisheries and other ecosystems intact.

"I don't think conservation in a rural community will ever work unless the people are your partners," renowned primatologist and conservationist Jane Goodall told MNN in 2014. "Unless they get some benefit and get some pride."

Regulated hunting can bring in money for conservation, but it can also come with pitfalls like misspent funds or ecological fallout from individual deaths. Sometimes there's controversy, too, even among conservationists. The World Wildlife Fund and many other big environmental nonprofits cautiously support hunting in the name of habitat protection, but that kind of acceptance varies widely for different species. And although most wildlife already benefits humanity in hard-to-quantify ways — offering us indirect perks like pollination and prey management — non-lethal eco-tourism has begun to dramatically boost the direct value of some species to humans.

Each living elephant in Africa, for example, is worth $22,966 per year to its local economy by attracting eco-tourists, according to a 2014 report. And since elephants live for up to 70 years, that means an elephant can generate $1.6 million during its life span. Trophy hunters often pay tens of thousands of dollars to kill a single animal, and poachers can reportedly net $22,000 by selling a pair of tusks. But an elephant is clearly more valuable to humans alive than dead, possibly by a factor of 76.

(And that's not even counting the less tangible value inherent to any animal's life, of course. Especially one as clever and complex as an elephant.)

That doesn't mean eco-tourism is without risks; its revenue is also subject to misuse, and careless tourists can stress out wildlife or introduce invasive species. But when it's done right, eco-tourism offers a pragmatic way to help wildlife conservation outbid the competition. Responsible hunting of certain species can do that, too — but only if, like Teddy Roosevelt, we remember the source of its peculiar charm.

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Russell McLendon ( @russmclendon ) writes about humans and other wildlife.