CHARLESTON, W.Va. -- Angling pioneer James Henshall wrote in 1881 that smallmouth bass "inch for inch, pound for pound, [are] the gamest fish that swims."

Few West Virginians would disagree.

Any Mountaineer who has hooked a smallmouth has experienced the strength, the agility and the sheer never-say-die grit the species has become famous for. Fisheries officials consider the smallmouth the state's second-most important game fish, and then only because it inhabits fewer waters than its close cousin, the largemouth.

"There's a lot more fishing for largemouths, mostly because bass tournaments are usually held on waters where largemouths are dominant," said Mark Scott, Division of Natural Resources fisheries biologist for the state's southeastern counties.

"Smallmouths are river fish that prefer clear water and rocky habitat. Largemouths are found mostly in lakes and in large rivers. West Virginia has a lot more largemouth habitat than smallmouth habitat."

That doesn't mean the state is smallmouth-deficient. In fact, it's anything but. Four major rivers - the New, the Greenbrier, the South Branch and the Elk - are renowned and popular smallmouth fisheries that add significantly to the state's tourism bottom line.

A small but thriving industry has grown up around float fishing for smallmouths. On rivers such as the New and South Branch, both of which flow through remote, largely roadless canyons, outfitters offer guided float trips.

Because of the outfitting industry that has grown up around smallmouth fishing, Scott believes smallmouths are every bit as important economically as largemouths.

"I'd place the smallmouth right up there," he said.

West Virginia's early pioneers found smallmouths in the Ohio River and many of its tributaries. The species itself is native to the Mississippi watershed and the Great Lakes.

Interestingly, it was not native to the Potomac watershed.

A Wheeling resident, Gen. William Shriver, was the driving force behind today's thriving smallmouth fishery in the Potomac and its tributaries. According to Henshall, Shriver caught 20 smallmouths from Wheeling Creek, put them in a cage submerged in the tender of a Baltimore and Ohio Railroad locomotive, rode with them to Cumberland, Md., and released them into the C&O Canal.

Smallmouths prefer rock-bottomed rivers and lakes, so it should come as no surprise that the species' scientific name, Micropterus dolomieu, reflects that trait.

Bernard Germain de Lacépède, a French naturalist, noting the smallmouth's affinity for rocky environs, named the species in 1802 for his friend, French geologist Déodat de Dolomieu. Dolomite, a type of limestone, also gets its name from de Dolomieu, so he might be the only scientist in history ever to have both a rock and a fish named for him.

Smallmouths also prefer cooler waters than their largemouth cousins.

Spawning takes place when water temperatures reach 57 to 62 degrees Fahrenheit. In West Virginia that usually happens in late May or early June. River flows can have a dramatic effect on spawning success.

"If the water gets really low or really high, it has a negative impact on the spawn," Scott said. "In years when we have floods on the New, we lose an entire year's worth of spawn."

Anglers usually don't notice the bad spawn until two years later, when the smallmouths that never hatched would measure 8 to 12 inches in length.

"Most of the smallmouths fishermen catch are in that range, so when they're missing from the population, people notice," Scott explained.

After the females lay their eggs, the males move in to guard the nests. As is the case with many species, smallmouth females tend to be larger than males. Males usually max out at 2 to 3 pounds. Females can weigh 5 or more.

The heaviest smallmouth ever caught in West Virginia was taken in 1971 from the South Branch. The big female tipped the scales at 9 pounds, 12 ounces. The all-tackle world record, from Tennessee's Dale Hollow Lake, weighed 11 pounds, 15 ounces.

Diet studies have shown that smallmouths prey primarily on crayfish and minnows.

"Crayfish seem to be the dominant food source, minnows less so," Scott said. "Smallmouths also feed on hellgrammites, caddis flies and other aquatic insects, but mostly they eat crayfish."

Anglers usually fish for smallmouths with light spinning or bait-casting tackle. Plastic grubs, particularly in crayfish-like colors, are popular baits. So are crankbaits, spinnerbaits, buzz baits and soft plastic-twitch baits. Fly tackle can be effective too, especially for anglers adept at fishing streamers and slider-style balsa poppers.

DNR officials have imposed special regulations on parts of four popular Mountain State smallmouth streams.

Catch-and-release regulations are in effect for a 12-mile section of the New River, and on 8- and 9 1/2-mile sections of the South Branch. All black bass - smallmouth and largemouth - must be released immediately after being caught.

A 12-inch minimum size limit is in effect along the entire length of Wheeling Creek. Anglers must release any bass smaller than that.

On a six-mile section of the Greenbrier River, a slot limit is in effect. Anglers must release any bass that measure 12 to 20 inches in length.

Scott believes the future is bright for most of West Virginia's smallmouth fisheries. The lone exception is the South Branch, which in recent years has suffered from algae blooms and reports of smallmouths that exhibit both male and female characteristics.

Scientists are working to determine the causes of those problems. Some theories have been advanced, but so far none have been proven. Until they are, Scott believes the South Branch will lag behind the New as the state's preeminent fishery for "the gamest fish that swims."

Reach John McCoy at 304-348-1231 or johnmc...@wvgazette.com.