Three species of fishes, sharing similar habitat but worlds apart biologically, are sought by anglers with a dream of the 'inshore grand slam' - the legendary bonefish, tarpon and permit. Dr Julian Pepperell takes a look at what makes these three different-but-linked fishes tick. STORY: Dr Julian Pepperell photos: dean butler & peter morse

When we think of gamefish, our mind usually drifts out over the horizon to the wild blue yonder. Images of a dancing sailfish, a dashing marlin or a line-burning tuna in cobalt offshore waters all come to mind. However, there are other legendary gamefish that live much closer to shore, sometimes even within range of a good cast with a saltwater fly rod.

Three of these - the bonefish, tarpon and permit - occupy perhaps a unique joint pinnacle in the world of gamefishing: the inshore grand slam. And to catch all three in a day, or indeed, in a lifetime, is the dream of many an angler.

I first became really aware of the awe in which these fishes are held during my first trip to the Florida Keys. Staying at the famous Cheeca Lodge at Islamorada, I soon discovered the 'Light Tackle Bar' - a marvellous watering hole that reeked of fishing history.

The walls were covered with all sorts of fascinating fishing memorabilia, but the centrepiece was a set of three beautiful casts of the silver ghosts of the flats - a bonefish, a permit and a huge tarpon.

All three are wary stalkers of the sand flats in that part of the world, and it is off the Florida Keys and nearby Caribbean islands that it is possible to catch all three in a single day.
Possible, but not likely - and there's the challenge.

WHAT DO WE KNOW?
Given the importance of these three fishes to the area, it is not surprising that the great majority of what we know about them, sketchy as it may be, comes from the Florida region.
Even though they don't appear to be very similar at all, two of the three species covered in this article - the bonefish and the tarpon - are quite closely related. These two species are both members of the Elopiformes order of fishes (the third member of this group, the giant herrings or ladyfish, Elops saurus, also occurs in the same area, but is not as highly regarded as its more famous cousins).

Fishes of this group, including the tarpon and bonefish, are regarded as particularly 'primitive' in that they possess various characteristics nearer to ancestral fishes than more-recently evolved, 'modern' fish. These features include the position of the paired pelvic or ventral fins, which, rather than being located under the 'chin', are situated well back along the body under the single fin.

Both also have what are known as leptocephalic larvae - a feature only shared by the Elopiform fishes and the eels. These larvae, rather than being tiny, quickly grow into long (6cm), transparent ribbon-like fishes with very small heads. After weeks or even months in this state - during which time they apparently absorb nutrients directly from the water - they completely metamorphose, halving their length and finally coming to resemble small adults.

And lastly, both the tarpon and the bonefish possess a primitive swim bladder, which has a tube connecting it to the oesophagus. The swimbladder is also highly vascularised (supplied with many small blood vessels), and actually functions somewhat like a lung.

Bonefish and tarpon can therefore obtain extra oxygen by gulping air at the surface, allowing them to live in oxygen-poor environments in estuaries and, in the case of tarpon, freshwater rivers and lakes. In fact, it is very likely that tarpon must have access to atmospheric oxygen to survive.

BONEFISH
I have long been fascinated by the allure of the bonefish. Here is a fish that looks and acts like an Australian whiting, that only reaches a maximum size of about 8kg (although a 3kg specimen would be a 'good' fish), but which has a reputation that dwarfs gamefish many times this size.

The main species of bonefish that causes all the fuss in angling circles is Albula vulpes, literally translated as "the white fox" - an apt name if ever there was one. Until quite recently, it was thought that this single species was the only bonefish, distributed on tropical sand flats and atolls right around the world. However, DNA studies have shown that there are actually five species of true bonefish on the planet - a fact that causes some confusion in angling circles.

While the 'true' bonefish, Albula vulpes, is certainly widespread and does occur in all three major oceans, another widespread species with which it overlaps is the roundjaw bonefish, Albula glossodonta. This similar species occurs through the tropical Indo-Pacific, and is the most likely bonefish to be encountered in the Australasian region and the central Pacific, including Hawaii.

Unfortunately for the angler, these species are quite difficult to tell apart. As the name suggests, the roundjaw has a distinctly rounded lower jaw, while that of the 'true' bonefish is more pointed. However, probably the best feature to look for is the presence or absence of a small, black spot under the snout. The roundjaw normally possesses this marking, while the 'true' bonefish does not.

The other bonefishes are the sharpjaw, Albula neoguinaica, another with no common name the Albula argentea, as well as the threadfin bonefish, Albula nemoptera, which is confined to the Atlantic and is easily identified by its trailing rear extension of the dorsal fin (similar to that of the tarpon and giant herring).

To complete the picture, two other species of so-called bonefish should be mentioned. These are the longfin bonefish Pterothrissus belloci, and the Japanese gissu Pterothrissus gissu. Both occur in relatively deep water, and, while there are basic similarities between the two genera of bonefish, they certainly show major differences too - the main one being a single long dorsal fin running most of the length of the body in the two Pterothrissus species.

DISTRIBUTION
As noted, bonefish are cosmopolitan, meaning that they occur right round the world, mainly in the tropics. Their preferred habitat of shallow, sandy flats means that they are coastal when near continents, but they also occur patchily in similar habitats around islands and atolls, often in remote mid-ocean locales.

Well-known bonefish grounds include the Florida Keys, the West Indies, Christmas Island in the mid Pacific, and also in the eastern Indian ocean and the Seychelles in the northern Indian. In the Australian region, bonefish are more commonly encountered than realised, but are often mistaken for large whiting.

They occur around the northern half of Australia, as far south as Moreton Bay. There are always exceptions to the rule, however, and one highly-unusual recent capture of a bonefish in the Sydney area certainly raised a few eyebrows.

NSW Fisheries biologist Aldo Steffe, a keen angler, hooked what he thought was a good-sized jewfish while fishing with fish bait in Port Hacking. After a long tussle, he got the surprise of his life when he landed a 3.6kg bonefish!

A little further away from Australia, Fiji and Vanuatu both hold good populations of bonefish, as does New Caledonia. In the latter country, sportsfishing for bonefish has not been promoted, although I suspect that if it were it could really take off.

On a recent visit to the central fishmarkets in Noumea, I was amazed to see piles of huge bonefish for sale, apparently gill-netted in the relatively remote north of the main island.

SIZE AND GROWTH
Bonefish appear to grow to different maximum sizes in different regions. Off the southeastern United States and in the Caribbean, the largest fish reach about 77cm in length and a weight of just over 7kg.

This is an exceptional specimen, of course, and any fish weighing more than 3.5kg is regarded as a very good capture. Considerably larger fish, however, have been recorded around oceanic islands and atolls. Fish taken off Hawaii and Africa and western Africa have topped the 9kg mark.

The official world-record bonefish is an 8.61kg specimen caught off Zululand, South Africa way back in 1962. Another weighing 7.7kg was caught in the same region in 1976, while at least three fish weighing in excess of 7kg have been taken around Florida.

Two of these were taken on normal saltwater tackle, while the third - caught in 1997 (on 4kg tippet) and weighing 7.03kg - is, not surprisingly, the current world fly-fishing record. The largest bonefish recorded in Australia weighed just over 10lb (4.66kg) and was unexpectedly caught on a fish bait off the Swains Reefs in Queensland.

Growth rates of bonefish are not well understood. Best estimates of growth indicate that fish reach a size of about 23cm by two years of age, and that most are mature at about four years of age, by which time they have attained a length of about 45cm. They are a relatively long-lived species, apparently living up to 20 years old.

FEEDING AND BEHAVIOUR
Any description of the bonefish stresses its 'inferior' mouth, meaning that it is located underneath the head. This, together with the fact that the mouth is equipped with strong crushing teeth, indicate that this fish is a bottom feeder that is adapted to crushing shellfish.

Indeed, dietary studies have shown that bonefish primarily eat crustaceans, including prawns and crabs, as well as clams, mussels and oysters. They are not averse, however, to chasing the occasional fish, and toadfish are commonly found in the stomachs of larger fish.

Bonefish often forage in groups, from as few as three or four fish up to schools numbering in the hundreds. They move out over the sand flats, often in such shallow water that their dorsal fins break the surface - a sight sure to set the heart of any angler racing!

REPRODUCTION
Bonefish spawn close to the coast, but move into deeper water to do so. Off Florida and in the Caribbean, spawning takes place from about November to June, with little activity evident in the hotter months of the year.

As mentioned, bonefish possess a leptocephalus larval stage that reaches a maximum length of about 60mm. Metamorphosis to the more normal larval shape takes up to two weeks, and it is during this long larval period that bonefish probably disperse throughout their broad, cosmopolitan range.

TARPON
Of the three fishes under consideration, the tarpon is the one that may lay claim to being a true gamefish simply on the basis of its size alone.

This incredible-looking silver fish with its huge scales and mechanical-looking mouth is an amazing sight, and one of the most highly revered of all gamefishes. Pick up any book on gamefishing from the US, turn to the chapter on tarpon, and be prepared for an unbridled flow of superlatives in describing the "Silver King".

Identification and classification of tarpon is made easy by the fact that there are only two species worldwide - one in the Atlantic and one in the Indo-Pacific. Both are in the family Megalopidae, the giant tarpon being Megalops atlanticus, while its smaller cousin, the oxeye herring, goes by the name of Megalops cyprinoides.

The giant tarpon is found only in the Atlantic ocean, ranging from Canada to Argentina in the west to Senegal and Angola in the east. However, because they can tolerate freshwater, some stray tarpon have made it through the Panama Canal and a small population now exists on the Pacific coast of Panama.

The smaller tarpon cousin, the oxeye herring, is quite widespread, being found in South Africa, Zimbabwe, Madagascar, India, southeast Asia, Papua New Guinea the Queensland coast and some Pacific Islands including New Caledonia, Palau and the Marianas.

Showing similarities to the bonefish, both species of tarpon can tolerate water with low dissolved oxygen by gulping air at the surface and storing it in their lung-like swim bladders. This allows them to penetrate into quite turbid water including rivers and lakes some distance from the ocean, although fully adult oxeye herring tend to avoid such habitats, unlike its giant cousin.

SIZE AND GROWTH
In common with quite a lot of fish species, female giant tarpon grow considerably larger than males. Since 1956, the IGFA all tackle record stood at 128.46kg for a fish taken in a lake in Venezeula, South America.

This weight was tied by another fish caught off Sierra Leone (Africa) in 1991. However, in March 2003, this record was finally broken by a whopping 130kg tarpon caught off Guinee Bissau, west Africa. Two other specimens that appear in the line-class records weighed over 120kg and were also caught off Sierra Leone - obviously an area to target for large tarpon.
The largest American line-class record still in the charts is a 66.8kg fish caught on 4kg line, although the saltwater fly world record is considerably larger at 91.85kg for a fish caught off Florida in 2001.

There is a reference in a Cuban paper to a tarpon measuring 2.5m and weighing 161kg. However, since this is so much larger than the all-tackle record, some doubt must be attached to the claim.

The Indo Pacific species of tarpon, the oxeye herring, Megalops cyprinoides, is a much smaller fish. While sizes of 3–4kg are not uncommon, and fish up to 8kg have been caught in Australian waters, there are unconfirmed reports of the species reaching double this size - up to 18kg, in fact.

Again, while this sort of size is possible, it would always be better to have absolute proof of such large specimens.

Tarpon grow quite rapidly when young, reaching a length of about 1.2m in six to seven years, but they slow down considerably after that. The age of a 100-pounder (45kg), for example, is estimated to be about 15 years.

Tarpon survive well in captivity, so we have some data on actual longevity of individual fish. Male fish are known to live to at least 30 years old, while one particular female fish held in an aquarium in Chicago died in 1998 at the ripe old age of 63. As with all cases of longevity in aquarium fishes, however, it is a moot point whether fish in the wild would attain such an age.

FEEDING AND BEHAVIOUR
Tarpon feed mainly on fish, including mullet, pilchards, catfish and longtoms (needlefish). They are also not averse to crustaceans such as prawns and crabs.

The tarpon mouth is designed to create a strong suction when it is suddenly opened, and when a large fish performs this noisy feat under a bait, it is apparently a sight and sound not soon forgotten.

Some limited high-tech tagging of tarpon has been undertaken using pop-up satellite tags. Results have so far indicated that, in general, tarpon are more active during the day than at night - and interestingly that peak activities occur right after cold fronts pass over the area.

Other studies on the microchemistry of otoliths (earbones) indicate that, while some tarpon migrate between the marine waters of the Caribbean and the freshwater of Lake Nicaragua in Central America, others do not, which suggests that movements are not predictable and may even be individually unique.

REPRODUCTION
Being large fish, tarpon are highly fecund. It is estimated, that a 2m female would produce approximately 12 million eggs at a time.

Spawning peaks in summer months, although it is thought that some spawning occurs year round. Observation of the spawning of tarpon has never been verified, but one interesting observation made by two Florida-based biologists sheds light on possible pre spawning behaviour.

Derke Snodgrass of the National Marine Fisheries Service and John Baldwin of Florida Atlantic University described this behaviour as follows:

"The school of 12 to 16 tarpon was observed in four to eight feet of water on the ocean-side of Tavernier Key, Florida, at the start of the incoming tide under cloudy skies and light rain.

"The tightly-packed school consisted of a single, approximately 140lb, presumably female fish that was repeatedly and persistently accosted by eight to ten smaller, 15–30lb, presumable males. In addition, two to four larger fish, 60–80lb, were following in the school, but were not as aggressive in their affection for the large female as the smaller males.

"The small males would consistently bump and rub the female's vent region, at times even pushing her above the water's surface. The males maintained this close contact despite numerous directional changes by the female.

"At times the female would be resting/basking at the surface, seemingly motionless with the whole dorsal side of her body and tail out of the water. This was typically ended when she would erupt with a burst of speed and throw a huge boil of water and spray.

"During the 50 minutes which we were able to stay with the school as it paralleled the shore, no milt or eggs were observed being released, but certainly could have been."

The authors go on to say that direct spawning activity of tarpon has never been documented, and this has normally been thought to occur offshore after schools migrate from near-shore aggregating areas. They therefore speculate that these fascinating behavioural observations may represent the prelude to actual spawning activity.

Like the bonefish, tarpon larvae are leptocephalic, ribbon-like creatures during their first several months of life. The whole cycle, from transparent leaf-like waifs to juvenile fish measuring about 4cm in length, takes approximately six months - and during this extended period it is very likely that larvae could be transported over large distances.

PERMIT
The third member of the inshore grand slam is the permit. Not related to either the bonefish or the tarpon, this striking-looking fish is actually a member of the trevally or jack family, Carangidae.

There has always been some confusion in different parts of the world over what constitutes a permit and what is a pompano. This is relatively easy to clear up since they are both,
in effect, the same - that is, the permit is simply one particular species of pompano.

Globally, there are at least 20 species of pompanos all belonging to the genus Trachinotus. The true permit is Trachinotus falcatus, which is only found in the western Atlantic. A closely-related species, the Florida pompano, Trachinotus carolinus, is also found in the same region, and although it rarely grows to a size much more than 3kg (much smaller than the permit), separating the two species when small has always caused problems.

Keen-eyed anglers could count the soft rays of the anal fin, which number 20 or less in the permit compared to about 22 in the Florida pompano. However, a better method is to run one's finger over the tongue of the fish. The true permit has small teeth on the surface, while the pompano has a smooth tongue.

In Australia, there are four recognised species of Trachinotus, but here they are almost universally called 'darts'. I say "almost universally" since one species, Trachinotus blochii - the largest in the region - has long been known as the snub-nosed dart (or 'oyster cracker').

However, in recent years, some writers and anglers have begun referring to it as the Indo-Pacific permit. One problem with using the name 'permit' is that, as we have seen, it has traditionally only ever applied to one species of this largish tribe. All of the other members of the genus Trachinotus are called pompanos (although, just to confuse the issue, the fish called the "African pompano" is the only 'pompano' that is not in the genus Trachinotus, but is rather a species of the threafin trevally genus Alectis).

Having said all that, however, with the snub-nose becoming increasingly popular among the saltwater fly brigade, perhaps the name Indo-pacific permit will eventually be adopted and even accepted internationally.

The name 'pompano', by the way, is relatively easy to trace, coming from the word meaning "grape leaf" - roughly the shape of the fish (well, with a good imagination anyway). However, the derivation of the word 'permit' is a bit of a mystery.

Not surprisingly, the largest pompano in official record charts is indeed the permit, Trachinotus falcatus. Until recently, the champion specimen was one caught off Fort Lauderdale, Florida in 1997, which weighed 25.45kg. However, this record was broken in 2002 by a fish of 27.21kg caught off Brazil. And for those who wave the long wand, the saltwater-fly-record permit was caught off Florida in 1986 and weighed 18.82kg.

Only one other pompano is listed by IGFA - the African pompano. As indicated, this is not a true pompano, but for completeness it is included here. The record for that species is a 22.9kg fish caught off Florida in 1990.

Finally, one other large species of pompano should be mentioned - the southern pompano of South Africa, Trachinotus africanus. While the angling record for the species stands at 14kg, specimens held captive in Durban aquarium, South Africa have exceeded 25kg in weight.

The Australian claimant to the name 'permit', the snub-nosed dart, is not a listed species for records with the GFAA. However, it is reliably reported to grow to at least 10kg. Perhaps when and if the species is added to the list of recognised gamefish, a better idea of its maximum size will be gained.

WHERE TO CATCH THE GRAND SLAM
Obviously, Florida and the Caribbean regions hold the best possibilities for taking a grand slam. This is certainly true if we are talking about one of the three being the giant tarpon.
However, if we bend the rules a little to allow some variations on the 'true' inshore grand slam, we might include the oxeye herring, and the snub-nosed dart.

In that case, a bit of fun might be had looking for a variation on the inshore grand slam around, say, the Cape York region of Australia or the coast of New Caledonia.
Not quite the adrenalin rush of the real thing, but a challenge nevertheless!