The Largest Fish on Coral Reefs were the First to Go. "You don't know what you've got 'till it's gone," as we're only now finding out.

By Douglas Fenner, Ph.D.

Many reef scientists (including the author) have spent their entire careers diving on reefs that have few big fish, and have never questioned whether that was normal or natural. Seeing big fish like sharks, humphead wrasse, bumphead parrots, mantas, goliath grouper and giant grouper, is very exciting for divers, and a big attraction for dive operators. One dive operator in Australia estimated that a single shark that he could reliably bring divers to might be worth $25,000 to his operation. In part, they are so exciting because they are so rare today.

Big fish are rare on most reefs anywhere near people. I dove for years in the Caribbean, and only once saw a goliath grouper, which was just a juvenile. I've been diving for years in many places in the Indo-Pacific, and have only twice seen schools of bumphead parrots, both times of only about a dozen individuals. I've only once seen a full size adult humphead wrasse. It is very easy to assume that reefs have always been the way that we first saw them, and judge their future condition based on that. As reefs degrade, each generation uses a lower condition as the baseline to judge further losses. This is called the "shifting baseline" (Pauly, 1995; Sheppard, 1995).

Jackson (1997) wrote:

"The problem is that everyone, scientists included, believes that the way things were when they first saw them is natural."

Jackson et al. (2001) wrote:

"The shifting baseline syndrome is thus even more insidious and ecologically widespread than is commonly realized."

In the last few years, there have been a flurry of reports on the reef fish communities at very remote coral reefs in the Pacific, which are nearly pristine. First, there was a report on the Northwestern Hawaiian Islands published by Friedlander and De Martini (2002). Tourists can only go to the "main Hawaiian Islands" at the southeast end of the chain. The islands and reefs in the Northwest beyond Kaui are too small, although Midway briefly had a dive tourism operation. It turns out they are virtually swarming with big fish when compared to the main Hawaiian Islands.

The most common big fish in the Northwestern Hawaiian Islands is giant trevally (Caranx ignobilis), which reach 1.7 meters length (5 feet) and 68 kg (150 pounds) maximum. There are also lots of grey reef sharks and Galapagos sharks. These big predator fish are called "apex predators" because they are at the apex of the food chain. Amazingly, they compose around half of all the weight ("biomass") of all the reef fish on these reefs (Birkeland and Friedlander, 2001). In contrast, when large areas of reefs around the main Hawaiian Islands are surveyed, there are very few sharks at all. There are about 65 times as many sharks per unit of area in the NW Hawaiian Islands as the main islands.

The author has snorkeled a lot in the main Hawaiian Islands (and wrote a book on Hawaiian corals, "Corals of Hawaii"), but has only ever seen one or two sharks there. If you snorkel or dive in Hawaii, you will be surrounded by beautiful small fish, only. That is very different from a natural reef.

Occasionally in the main Hawaiian Islands, a tiger shark bites someone, and rarely the person dies from the wound. This is tragic. In the hysteria that follows, people go out and kill all the sharks they can find, perhaps around 200. Few if any of them are tiger sharks. (editor's note: and most people who work with tiger sharks will say that they are not aggressive with humans except in rare instances; one researcher in Australia regularly swims with them to video them, and lassos them around the tail to hold them on a ship's stern apron to do research.).

People in western cultures love to fear sharks, which is supported in the popular media, such as the movie "Jaws." Around the world, sharks kill about 5 people per year, but many more people are killed by lightning and bee stings, among other things. Humans kill tens of millions of sharks a year, so who is the bloodthirsty killer, shark or human?

The rarity of sharks in the main Hawaiian islands is typical of reefs near people. In the Philippines, in two years of about 10 dives a week, I saw about a total of about five sharks, and two of those were on the way to the market. A few years later I got to dive at Tubbataha reefs in the Philippines, which are remote and currently protected. Whitetip reef sharks were not uncommon there, though nothing like grey reef sharks on the truly remote reefs that have not been fished.

In Hawaii, large numbers of sharks were killed in 'shark control' programs, using long lines of hooks laid on the bottom at an average depth of 45 m. "In response to concerns over shark attacks on humans, large scale shark fishing programs were conducted in Hawaii from 1959 to 1976. During this period >4500 sharks were caught within the MHI..." (Papastamatiou et al. 2006). In spite of the very clear evidence that sharks have been heavily overfished to the point that they are endangered with local extinction in the main Hawaiian Islands, there has been no management action to protect them there. This may be because of pressure from fishermen. Fishermen are highly vocal there, and in politics, the squeaky wheel gets the grease. As a result, sometimes there is an attempt to base fishery management on special interest politics, not on science (Pala, 2007a ). In addition, sharks have a bad reputation with the public, and occasional shark attacks in Hawaii mean that sharks get no sympathy. Action to protect them would be very unpopular.

More recently, studies in the remote and unfished reefs in the Line Islands south of Hawaii, have found that there, too, about half of the fish biomass is in the big fish (Stevenson, 2006; Pala, 2007; Richie et al. 2008; Sandin et al. 2008).

Studies of reef fish in Guam and the Marianas found that fish larger than 50 cm in length were much more abundant around the islands at the northern end of the chain. Human population is concentrated on Guam and the southern islands, while the middle and northern islands are uninhabited. One fisherman's response to hearing that there were many more big fish at the northern end of the chain was "maybe that's where we should hold our fishing tournament next year!"

The fish expert, Dr. Gerry Allen, reports that in the remote and uninhabited Phoenix Islands (west of the Line Islands), in a one-hour dive, an average of about 15 sharks were seen. On many reefs near people, you may have to dive 100 or more dives to see a single shark. In the Phoenix Is., you might see about 1500 sharks in those 100 dives. 1500 times as many sharks. There are many differences between islands other than the intensity of fishing, which could produce these differences. However, in spite of all of the other differences, nearly every reef with people nearby has very few big fish left, and virtually every remote pristine reef that has been studied has high levels of these fish. In some locations the decline has been documented (as in the GBR, Fiji, Guam and Solomon Is. below).

In fisheries science, overfishing has been defined as any fishing that is greater than the "Maximum Sustainable Yield." (MSY). That is because at MSY, the long-term benefits to fishermen are maximized. (Maximum benefits to society are another question with additional considerations.) Most fisheries models say that MSY is at about 1/3 of the biomass of an unfished stock. So these big reef fish are not just overfished, at 1/1500th the biomass they are grossly overfished and approaching local extinction.

Modern fisheries science has tended to adopt a slightly more restrictive overfishing limit of "Optimum Yield." (OY). While OY is not defined exactly, it is a fish catch that is less than MSY to allow for uncertainties in the data and models, and high variability in annual recruitment of new fish. In addition, if there are deleterious ecosystem effects of fishing, OY would be set to a lower level of fishing to avoid those effects.

But no matter how you slice it, these big reef fish are grossly overfished. Overfishing benefits no one, including the fishermen, who can catch more on a sustainable basis if they fish less (at MSY). But each individual fisherman would catch more and make more profit in the short term if they were allowed to fish more, hence fishermen often push to be allowed to fish more than would be beneficial for them in the long term or for society as a whole. Essentially, to mine the fish out and let the future take care of itself. Legally, in the U.S., states must now manage public assets based on "Public Trust," the legal doctrine that the states manage public resources held in trust for all citizens. Thus, states legally must manage publicly owned assets for the benefit of all citizens, not just those who utilize those public assets for personal financial gain, like fishermen. Presently, the U.S. Federal Government is not restricted this way (Turnipseed et al. 2009).

Comparisons between fished and remote reefs are open to many interpretations due to the many differences between the reefs. Observations over time can be more powerful in countering shifting baselines. Pierre Laboute, author of many guidebooks to fish and invertebrates (e.g., Laboute, 2001; 2009) has been diving, observing, and photographing the reefs of New Caledonia, Madagascar, and French Polynesia for decades, and reports that when he first started, sharks and humphead wrasse were plentiful in these areas, with several large individuals seen on the average dive. Now, several to many dives are needed to see a single large shark or humphead wrasse.

A recent report from Australia indicates that the Cocos-Keeling Islands in the Indian Ocean (owned by Australia), which have no fishing, have sharks in abundance. The report also shows that areas of the Great Barrier Reef (GBR) that are open to fishing (which until recently was most of the reef) have much fewer sharks (Robbins et al. 2006). In the few little areas of the GBR where people are not allowed to go, sharks are abundant, like in Cocos-Keeling. Surprisingly, in areas where fishing is not allowed but people can go, sharks are in low abundance similar to in areas where fishing is allowed. Apparently, people are poaching sharks even in no-take Marine Protected Areas (MPAs). Only no-go areas provide sufficient protection for the sharks.

Robbins et al. were able to measure the rate at which sharks are declining on the GBR, and it is rapid. Fishing in Queensland (where the GBR is) is controlled by the Queensland Department of Primary Industries (DPI).DPI claims they have tightened up regulations (now each commercial fisherman is limited to possession of one grey reef shark or white tip reef shark) so it is well regulated. So recreational shark fishing is unrestricted, and each commercial fisherman is limited to possessing one reef shark per day, or 365 a year, and anyone can kill all they want and throw them back. The story is going around that fishermen who fish for coral cod (grouper) on the GBR and who make quite a bit of money off that, do not like to pull up just the head of a coral cod that a shark has eaten while it was on their line. So they deliberately catch sharks, kill them, and throw them back. All perfectly legal, unreported and unregulated.

Robbins et al. (2006) wrote,

"Our data suggest that for coral-reef sharks, immediate and substantial reductions in shark fishing will be required for their ongoing collapse to be reversed."

"Together, these findings indicate that extirpation of these species from fished coral-reef ecosystems is an immanent likelihood in the absence of substantial changes to coral-reef management."

"Inferred and projected declines such as ours appear sufficient to warrant "Critically Endangered" status under the IUCN Red List (A3d) criteria for this study area for both species."

"Moreover, the magnitude of the population decline is severe: Median rates of population decline are 7% per annum for whitetip reef sharks and 17% for grey reef sharks. If current population trends continue unabated, the abundance of whitetip reef sharks and grey reef sharks present on legally fished reefs will be reduced to only 5% and 0.1% respectively, of their present-day no-entry abundance levels within 20 years."

"The minimum change in mortality necessary to produce a median estimated population growth rate of 1.0 (i.e., population stability) was calculated for each species. Analyses indicate that reductions in annual mortality by one-third (36%) for the whitetip shark and one half (49%) for the gray reef shark would be required to halt these ongoing declines. However, with commercial catches of sharks nearly quadrupling on the Great Barrier Reef between 1994 and 2003, and recreational fishing also removing large numbers of sharks in Australia, the trend is strongly in the opposite direction."

"For instance, on coral reefs, food-web models indicate that trophic cascades initiated by overfishing of sharks may have contributed to the collapse of Caribbean coral-reef ecosystems."

The renowned coral reef scientist, J.E.N. "Charlie" Veron writes,

"When I first worked on the Great Barrier Reef, I always felt a moment of anxiety after rolling backwards off the side of a boat to go for a dive. We all felt that. We waited for the bubbles to clear just to make sure that there wasn't a big tiger among the sharks that always gathered around. Now, anywhere in the Asian region, I swim long distances over deep water without the slightest concern, for there are virtually no sharks left, big or small. I haven't even seen big fish in any numbers around an Asian reef in years. The plight of sharks is symptomatic of what is happening to reefs."

The vulnerability of sharks is highlighted in this quote from Nichols (1993):

"Sharks possess particular biological characteristics which render then especially susceptible to high fishing pressure, and as such, qualify them as a special case for management. As apex predators, they have few natural enemies. The biological characteristics of sharks - long lived, slow growth rates, low fecundity and reproductive rates (some species do not reproduce every year), long gestation period, relatively large size at first spawning, and strongly density dependent recruitment - result in shark fisheries being particularly sensitive to over-fishing."

Knowlton and Jackson (2008) wrote,

"The areas of biggest concern for the immediate future are apex predators at the top, because they are globally so rare, and corals at the bottom, because of their continuing decline, apparent vulnerability to even modest local human impacts, and extreme sensitivity to all aspects of global change. Both risk extinctions if nothing is done to halt their global downward trajectories."

McKleod et al. (2005) wrote,

"The key interactions among species within an ecosystem are essential to maintain if ecosystem services are to be delivered. Removing or damaging some species can dramatically affect others and disrupt the ability of the system to provide desired services. Small changes to these key interactions can produce large ecosystem responses. For example, the absence of large-bodied predators at the apex of marine food webs can result in large-scale changes in the relative abundances of other species."

Because huge numbers are taken in the live food fish trade (the trade is worth around US$1 billion per year), Humphead wrasse have been put on the CITES list, which is to protect them from international trade that would deplete them. But they are also taken by local fishers wherever there are people. Their abundance is inversely correlated with the abundance of people. Where the human population is greatest they are nearly absent, but where there are no people or fishing is not allowed, they are most abundant. The remote and uninhabited Phoenix Islands and Wake Island (a U.S. military base) have some of the most abundant populations known (Sadovy et al. 2003).

Sadovy et al. (2003) write,

"This species is particularly sensitive to fishing pressure. In most fished areas, density and body size have dropped substantially. It appears to be particularly heavily targeted and depleted in SE Asia and in some places faces extirpation." "Other giant reef fish share many problems..." "...humphead wrasse in recent years fetching as much as US$130/Kg at retail."

"Probable extirpations at edge of range sites signal the start of range reduction, an early step towards extinction."

"The prognosis for the persistence of exploited populations of C. undulatus, under current conditions and given the biology of the species is poor. Conservation and management are needed to ensure it persists in viable numbers wherever exploited."

"The humphead wrasse is particularly vulnerable to exploitation at anything other than the lowest levels of fishing pressure." "Moreover, the ecological role of such large fishes is not well understood and it could turn out that they are key species for long-term ecosystem stability."

At night, Bumphead parrotfish sleep in the same schools, either in the open or in holes that are not large enough for them to completely fit into. They tend to sleep in the same area each night. As a result, they are particularly easy to spear at night with a flashlight and SCUBA. A fisher that finds where a school sleeps can return night after night to the same spot and spear them until the entire school has been extirpated.

Populations once again are inversely related to human populations, with low populations of Bumphead parrotfish where there are lots of people and many more where there are no people (Bellwood et al. 2003). C. Birkeland and G. Davis report that big schools of bumphead parrots were common in Guam in the 1960's, but they were spearfished out in the 1970's, and now they are rare. Hensley and Sherwood (1993) confirm that they are now rare in Guam; Dulvy and Polunin (2004a) suggest they might be extinct on Guam.

In spite of this, night spearfishing on SCUBA remains legal in Guam. Guam has a highly vocal fishing community. In Fiji, interviews with people revealed that when night time SCUBA spearfishing came to an island, the markets were filled with bumphead parrots, they were half or more of all fish in the markets. Now, in those same areas, they are rare and not seen in the markets. On some islands they have actually gone locally extinct (Dulvy and Polunin, 2004).

In some areas of the Solomon Islands, Bumphead parrotfish currently dominate markets. In areas near people, populations have decreased and fishers go farther to find more abundant populations (Aswani and Hamilton, 2004). Professor Howard Choat reports that a small group of spearfishers can fill a large skiff with them in a single night.

Bumpheads spawn in groups, so it is possible that spawning is stimulated by the presence of a group (Hamilton et al. 2008). It may be that once populations are low enough that no groups are left, they have trouble spawning, and it is hard for them to restore their populations even if protected. Small populations could be doomed, so it may be important to not allow the last groups to be killed. It appears that bumphead parrots are particularly vulnerable to being extirpated by fishing, more so even than humphead wrasse.

Hamilton et al. (2008) write:

"Preliminary studies have identified B. muricatum as a group spawning species, a behavioural trait that is frequently associated with over-exploitation."

"The large size, schooling behaviour and nocturnal aggregation of B. muricatum make it a lucrative and highly prized target of many subsistence and artisanal nighttime spear fisheries in the Pacific."

Bellwood et al. (2003) write:

"It is a major contributor reef ecosystem processes and its absence highlights the potential for marked changes in ecosystem functioning." "The absence of parrotfishes will impair normal recovery processes and constitutes a loss of ecosystem resilience..."

"B. muricatum, in particular, is highly susceptible to spearfishing activity..."

Dulvy and Polunin (2004) write:

"This large conspicuous reef fish was formerly a prominent and abundant member of reef fish assemblages and catches, but is now encountered infrequently throughout large parts of its range." "...this species may be extinct at the Marshall Islands and possibly Guam..."

Giant groupers (Epinephelus lanceolatus) in the Pacific (also called Queensland groupers in Australia) can get to well over 2.7 meters (8 feet) long and 300 kg (660 pounds). They appear to be rare everywhere, including reefs without people. However, the equivalent species in the Caribbean, the goliath grouper (Epinephelus itajara), which can get to at least 2.4 m (7 feet) and 310 kg (682 pounds, and possibly 455 kg or 1001 pounds!), is a different story.

Although they are rare in the Caribbean, in Florida there are pictures of the trophy catches from tourist fishing boats called "headboats" that paint a different picture. The old photos show lots of huge goliath grouper, sometimes a whole row of them, from a single day's fishing by one tour boat. Today, the photo of the trophy board is likely to have mostly smaller fish than that (McClenachan, 2008).

But goliaths have been protected in the Florida Keys since 1990. Now if you dive there, you have a good chance of seeing a juvenile, maybe 3 feet and 100 pounds. Under protection, their numbers are increasing rapidly, though it will be some time until the giant sizes are reached. Meantime, some fishing companies have discovered that there are fishers who find it extremely exciting to hook a huge fish, even if the hook is barbless and the fish is released. So there are tour companies that specialize in catch and release fishing for goliath grouper (see http://www.floridalighttacklecharters.com/gallery_extremefishing.htm).

Meantime, problems are appearing. There are catch and release fisheries for other fish as well. When these smaller fish are released, goliaths and sharks quickly learn that the fish that is released is dazed and up in the water where there is no hiding place. So goliaths and sharks hang around some fishing boats and zoom in and eat the newly released fish. This does not please the fishers, they want to be able to catch them again. There are rumors of fishers catching sharks and/or goliaths and taking them off somewhere and killing them.

Fishing has long been known to usually remove the big fish first (e.g., Ricker, 1946; Jennings and Kaiser, 1998; Jennings et al. 1999; Pitcher, 2001; Dulvy et al. 2004). The incentive is for a fisher to go for the big ones: more to feed your family or more to sell. In American Samoa, a fisherman who speared a large terminal male humphead wrasse sold it for $120. If he had speared the most common fish, a surgeonfish, it would have brought about $1. So the big fish was 120 times more profitable per fish. It is usually more profitable to take the big fish (though there are specific fisheries for small fish, such as anchovies, herrings, and sardines if large numbers can be found).

You can even get a measure of fishing pressure by recording the sizes of fish present, the more fishing the fewer big ones (Graham et al. 2005). Over decades, fishing can begin with the largest fish, then once they are depleted move to the next size fish, and so on down to the smallest that are still profitable. This is called "Fishing down the food web." (Pauly et al. 1998; Pauly and Palomares, 2005) Think of the size range for reef fish- if reefs in an area have 600 species of fish, how many are large enough that people fish them, and how many are so small no one would fish them?

The most diverse families of fish on reefs are gobies and damselfish, and they are too small to be fished by any but the most desperately poor fishers. So at the small end of the size range on reefs, there are huge numbers of species that are too small to be fished. At the large end of the size range, there are just a few species, which are highly prized catches. Trophy catches are the largest fish, not the smallest. Fishing pressure increases with the size of the fish.

In addition, the numbers of individuals in a species decreases with the increasing size of the species. There are huge numbers of damselfish on most reefs, but even on unfished pristine reefs where half of the biomass is large fish, there are many fewer sharks, bumphead parrots, humphead wrasse and giant grouper than damselfish. The most abundant fish species on reefs where it occurs is a surgeonfish that reaches just 26 cm length. All this is because the larger the size of the individual, the more it takes to feed them. A reef can feed vast numbers of damsels, but only a limited number of sharks, humpheads, bumpheads and giant grouper. So, big fish are less abundant than small fish, and more heavily fished. The result is that they are much more rapidly depleted than small fish.

There is now a quantitative measure of vulnerability of fishing, which incorporates a variety of things about fish that make them vulnerable to fishing (Cheung et al. 2007). There is a website with a wealth of information about all the different kinds of fish around the world, called "FishBase" (www.fishbase.org). That website gives information on each species of fish. For each species, it now gives the "vulnerability index." The index has a range from 0 for no vulnerability to 100 for maximum.

Each of the different kinds of the largest reef fish, like sharks, humphead wrasse, bumphead parrots, and goliath grouper, all have vulnerabilities on the order of 75 (out of 100). Small fish have much lower vulnerabilities, often on the order of 25-35. The striped bristletooth (surgeon), Ctenochaetus striatus, is one of the most abundant reef fish most places where it is found in the Indo-Pacific (Lieske and Myers, 2001). It has a vulnerability less than 14.

The largest species of reef fish are highly sensitive to fishing, but the small fish are much more resistant to fishing, with the most abundant species being highly resistant. Sharks reproduce in a way that makes it particularly hard for them to recover quickly from fishing. Unlike bony fish, they produce a few large pups, instead of masses of tiny eggs. Reef sharks typically have about 1-5 pups, once a year or every other year. Thus, their ability to increase in population rapidly is extremely limited.

By contrast large female bony fish can release millions of eggs a year. The probability of survival of a single tiny fish egg (likely about 1 mm or 1/16 inch diameter) is minute compared to the probability of survival of a single shark pup. The larger the individual the better the chance of survival. But if conditions are just right, a large bony fish can have so many offspring survive they can replenish their population in one year, but that is quite impossible for a shark (or ray). Once depleted, large fish and sharks in particular, can be kept at low levels indefinitely by small amounts of fishing. Just the small amount of poaching in no-take areas on the GBR was enough to deplete the sharks there.

Fishing always removes fish, and almost always results in a decrease in fish abundance and biomass. The total biomass of fish on reefs is higher on more lightly fished reefs, and lower on more intensely fished reefs (Knowlton and Jackson, 2008). Much of those differences come from the removal of the big fish (Birkeland and Friedlander, 2001). If only large predators are removed, then their prey can actually increase in abundance (Graham et al. 2003). However, in most cases smaller fish are taken as well as the large predators, and smaller fish decrease as well as the large fish, because although they are released from predation by predatory fish, they are taken in even larger numbers by predatory humans.

The degradation of coral reefs around the world has only been recently recognized, but it has been going on for a long time. Only recently have studies of near-pristine reefs and historical records shown how degraded most reefs are, and how long this has been going on. The historical studies confirm that the big fish were depleted before the small fish (Pandolfi et al. 2003; 2005). Tropical megafauna once included amazing numbers of monk seals and sea turtles, millions of the latter. They were the first to go when westerners arrived (Jackson, 1997).

There is even archeological data showing the decline of reef fish stocks before westerners arrived (Wing and Wing, 2001). On land, humans have been implicated in the extinction of large mammals and birds, which often disappeared about the time humans arrived on a continent such as North America. While the megafauna disappeared, the smaller species survived. Mice and rats do really well.

Pitcher (2001) stated,

"For terrestrial mammalian megafauna, and associated specialized carnivores and scavengers, a pulse of late Pleistocene extinctions throughout the world coincides with the advent of cooperative hunting behavior and technology as modern humans spread around the globe." "In the past 100,000 years North America has lost 73%, South America 79% and Australia 86% of endemic genera of terrestrial megafauna."

Jackson et al. (2001) state that:

"Historical abundances of large consumer species were fantastically large in comparison with recent observations."
Jackson (1997) wrote:

"Megavertebrates are critical for reef conservation and, unlike land, there are no coral reef livestock to take their place."

How much is a big fish worth in the fish market? A hundred dollars or more? That's a lot to a poor fisherman in a developing country. But how much can a dive operator charge to take a diver to see that fish alive on the reef? $50 or more? Divers go nuts over really big fish, they are so exciting. How about a boatload of divers? How about a boatload of divers every day? How much total money do those divers spend on hotel room, dining, car rental, souvenirs and airfare? How many people are employed by all those businesses? A single, huge, famous fish can have divers spend over a million dollars a year to see it. Alive, that fish is made of solid gold. Dead, it's not worth much in comparison.

Mind you, you have to be in an area where you can attract divers, but diving is much more sustainable than fishing, and the goose can go on laying the golden egg year after year. Does it make sense to kill the goose?? If the hotel and the dive operation are owned by people from developed countries, then the local people may get little benefit from the big fish in their own country. So I prefer to stay in locally owned hotels and go with local dive operators.

"Save the Giant Reef Fish!!" Australia protects humphead wrasse, as does Niue. Palau has now protected all it sharks, plus its humphead wrasse and bumphead parrots. American Samoa has promised to protect all of its large reef fish species, including all sharks, humphead wrasse, bumphead parrots, giant grouper, and giant trevally. They will be illegal to take by any means, throughout the territory, at all times, for all sizes of those species, by anyone. They are being protected on the basis that they are uncommon or rare, and they are exploited, and thus there is a possibility that the exploitation could drive them into local extinction. It is much easier to demonstrate that a species is rare and exploited than to prove it is overfished, so this may be a rationale for protection that has wider applicability.

References 

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