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“How humans shift fish evolution” with Mikko Heino, Knowable Magazine , February 27, 2020. |
Fisheries-induced evolution (FIE) is the microevolution of an exploited aquatic organism's population, brought on through the artificial selection for biological traits by fishing practices (fishing techniques and fisheries management). [1] Fishing, of any severity or effort, will impose an additional layer of mortality to the natural population equilibrium and will be selective to certain genetic traits within that organism's gene pool. This removal of selected traits fundamentally changes the population gene frequency, resulting in the artificially induced microevolution by the proxy of the survival of untargeted fish and their propagation of heritable biological characteristics. This artificial selection often counters natural life-history pattern for many species, such as causing early sexual maturation, diminished sizes for matured fish, and reduced fecundity in the form of smaller egg size, lower sperm counts and viability during reproductive events. [1] These effects can have prolonged effects on the adaptability or fitness of the species to their environmental factors.
Fisheries-induced evolution differs from the Darwinian evolution model by virtue of the direct human factor. [2] For FIE, fishing enforces a greater selection pressure for traits, often through sheer effort and catch numbers, which can disparage natural selection pressures such as predator-prey interactions and environmental influences.
Fishing practices that permanently remove animals from their population (i.e. not catch and release) drive direct fisheries-induced evolution by removing the genetic materials of those animals from the population. Individuals that are untargeted, through the selection bias of fishing gears and/or legislation, are allowed to reproduce and proliferate their genetic materials. As fishing pressures persist, traits belonging to non-selected organisms are preserved through survival and become more dominant in frequency within the gene pool. Additionally, fishing on a targeted species incur knock-on effects to those around it by its disturbance of their natural interactions. In these situations, specific traits of the untargeted species may be favourable under the diminutive presence, or absence, of the targeted species, and therefore indirectly selected for.
The direct selection for biological traits through fishery practices is the result of fishery management regulations, and gear restrictions and selectivities. [1] The most obvious artificial selection for traits through management legislation can be observed in the imposed regulations on size (minimum landing size), sex, seasonality, and locations. There is also evidence that various types of fishing gears directly select on traits that make individual fish more likely to be captured, including aspects of their behavior (e.g. boldness, activity level) and physiology (e.g. anaerobic capacity, swimming ability). [3]
Catch size regulations vary with specificity to the targeted species and is often used to prevent exploitation during a specific part of the life cycle for the organism. Such regulations arose in response to the effects of FIE observed by the fisheries of Atlantic Cod ( Gadus morhua ). [4] Prior to the fishing technological revolution that has led to the species moratorium, the exploitation of Atlantic Cod have been selective towards larger-sized fish since the 1500s. Catch data for the species have quantitatively shown that this selectivity for larger fish for over 500 years have shifted the life-history patterns, resulting in earlier sexual maturation and smaller sizes at said maturation. [5] [6]
Sexual selectivity by a fishery works on the theoretical foundation that the preservation of females allows their reproductive input to offset the fishing mortality. Fisheries of species with low fecundity such as mud crabs (Scylla serrata) and blue swimmer crabs (Portunus armatus) often adopt this method and only allow the harvesting of males. [7] The direct selection for traits, with respect to sex selectivity, occurs when specific characteristics or behaviours increase the susceptibility of the organisms for harvest. For example, corking wrasses ( Symphodus melops ) are harvested as a biological control for sea lice within farmed salmon facilities of the northern hemisphere. Male fish dominate these wild fisheries in both catch per unit effort and weight owing to their strong nesting behaviour and territorial nature, which differentiate them from females and sneaker males. [8] Persistent fishing pressures over the years have reduced maturation age and size for these nesting males, in addition to increasing the density of sneaker males to the detriment of the localised population.
Indirect fishery-induced evolution occurs when a species with some level of ecological significance is targeted by a fishery, and their diminished presence within the ecology causes a flow on effect to other untargeted species. Keystone or umbrella species provide many ecological services to the environment which they belong to, ranging from the provision of habitat and food, to the control of biodiversity by preventing any one organism from dominating. Removal or reduction of these organisms often cause significant changes to the behaviours and physiology of the organisms which were once controlled.
Non-migratory reef sharks ( Carcharhinus melanopterus, C. amblyrhynchos, and Triaenodon obesus ) play the vital role in weeding out sick and ill-adapted individuals from within a population, in addition to controlling the abundance of larger size fish. Thus, these sharks are an ever-present mortality factor for many reef organisms. Fisheries targeting these apex predators inherently allow for the proliferation of medium-sized predatory fishes (e.g. barracudas, juvenile groupers, trevallies, snappers) at the expense of the cohabiting smaller species. When this happen, the FIE responses observed from smaller species have been consistent with their increasing egg production and shifted life-history for early sexual maturation.
Another example of FIE that is instigated indirectly can be seen in the ornamental or aquarium trade industry. In particular, corals and anemones are highly prized ornamental commodities and are often harvested at unsustainable rates for profit within the Malay archipelago. [9] Removal of anemones at rates higher than their resettlement into the reefs, as observed in the Philippines, is known to have caused drastic reduction in the localised population of clownfish or anemonefish ( Amphiprion sp. ). [10] Evolutionarily, anemone fish observed on reefs that were subjected to intense and prolonged anemone fishing were significantly smaller, even for mature adult pairs, than those found living with an anemone. [11] This size reduction is attributed to their need to hide in small coral crevices in the absence of a host.
Outside of observational data collected from active fisheries, the evidence for fisheries-induced evolution, or its symptoms, may be discerned through both commercial and recreational anecdotes. These stories often articulate the diminishing catch, reduction in their weight and length records, in addition to the mandatory increase to fishing efforts over time to attain similar fishing quotas to their historical references. However, experimental data was needed to clarify two fundamental questions regarding FIE, so that the aforementioned ramifications of smaller sizes and early maturation may not be attributed to non-anthropogenic factors such as population flux. The two evidentiary questions: a) Is fishing pressure capable of changing the gene frequency of a population to cause microevolution within such a short time frame? b) How can non-selective fishing gears (gill nets and trawlers) be selective for certain traits?
The implementation of aquaculture experimental designs afford the possibility of isolating different biological traits and observing their impacts and heredity within a population. [12]
A study by Solberg et al. (2013) [13] demonstrated that with sufficient fishing pressure and selectivity, a substantial change to a population genotype may be achieved without the geological time-frame often associated with evolution. In this study, Atlantic salmon ( Salmo salar ) were subjected to a modified 'common garden' design, and the researchers isolated stress as an abiotic independent variable to the growth rates of salmon. By selecting for fish with a lower biological response to stress, a domesticated population was created within 10 generations of breeding, which exhibited a threefold increase in growth rate over its wild counterpart. Thus, the difference between the absence of larger fish, which can be misattributed to overfishing, and FIE lies in the inherited biological characteristics that were untargeted by fishing.
Fishing techniques such as gill nets and trawling are often associated with a lack of selectivity within the environment which they are operated. [14] It is difficult to establish whether selectivity can occur under these practices, when fishing data suggest that an entire localised population may be caught or up to 80% of the effort to go to bycatches and not the targeted species. [15] An experiment by Biro and Post (2008) [16] shows that despite being mechanically unselective of gill nets, behavioural variation in certain fish species can indeed affect the likelihood of them being caught and therefore be a function for selectivity. Using a ‘selection experiment in the field’ model, two genotypes of rainbow trouts ( Oncorhynchus mykiss ) were transplanted into identical artificial lake habitats and subjected to matching gill net fishing pressures. The net in question were designed to capture all size variability within the simulated populations. Trouts exhibiting the genotypes for faster growth rates and therefore a more active lifestyle were found to have a 60% increase chance to be caught in comparison to those characterised by the more sedentary genotype. Therefore, for 'non-selective' fishing methods, the selecting factor rests not on the gear itself but on the inherent behavioural variations within the population.
Fisheries-induced evolution is a function of the fishing-induced genetic drift, and to some extent represent the reduction of genetic diversity within the targeted population. For many exploited population, this reduction in genetic diversity has been predicted to reduce their adaptability to both environmental variability and ecological competitiveness. [17] Indeed, the commonly observed fisheries-induced adaptation of younger and smaller size at sexual maturation counter specific life-history characteristics that would enable interspecies competition for resources. [18] A biological model by Kindsvater and Palkovacs (2017) [19] shows that FIE Atlantic cod, due to their smaller sizes, actually belong to a lower trophic level irrespective of their original position prior to the species moratorium. Furthermore, the FIE effect of reducing egg counts and their viability have weakened the number of recruitments back into the population after spawning, thereby reducing the localised population density. [20] [21] However, at the reduced population abundance and smaller sizes, the increase in per capita resource has been stipulated to counteract the effects of competitions between different species by reducing intraspecific interactions, which can allow for maximal density-dependent growth to happen. This also means that for harvesting to be sustainable, yearly or seasonal assessment of viable reproductive stocks and their recruitment rates must be performed to account for the compromised fecundity and recovery statuses.
There is limited data on the interactions between FIE affected species and their robustness to environmental fluctuations. [17] For many species, the additional mortality rate and the frequency of harvesting exerted by the fisheries make it difficult to elucidate how environmental factors such as temperatures, salinity and currents can be beneficial or detrimental for stocks, since their effects on deaths or recruitment gains are diminutive in comparison. However, with the rapidly evolving climates stemming from the anthropogenic inputs, an understanding of how FIE affects the adaptability of these aquatic population will be necessary, not only for future stock assessments, but also sustainable harvesting.
The Atlantic cod is a fish of the family Gadidae, widely consumed by humans. It is also commercially known as cod or codling. Dry cod may be prepared as unsalted stockfish, and as cured salt cod or clipfish.
Clownfish or anemonefish are fishes from the subfamily Amphiprioninae in the family Pomacentridae. Thirty species of clownfish are recognized: one in the genus Premnas, while the remaining are in the genus Amphiprion. In the wild, they all form symbiotic mutualisms with sea anemones. Depending on the species, anemonefish are overall yellow, orange, or a reddish or blackish color, and many show white bars or patches. The largest can reach a length of 17 cm, while the smallest barely achieve 7–8 cm.
The orange clownfish also known as percula clownfish and clown anemonefish, is widely known as a popular aquarium fish. Like other clownfishes, it often lives in association with sea anemones. A. percula is associated specifically with Heteractis magnifica and Stichodactyla gigantea, and as larvae use chemical cues released from the anemones to identify and locate the appropriate host species to use them for shelter and protection. This causes preferential selection when finding their anemone host species. Although popular, maintaining this species in captivity is rather complex. The Great Barrier Reef Marine Park Authority regulates the number of collection permits issued to aquarium fish dealers who seek this, and other tropical fish within the Great Barrier Reef Marine Park. The symbiosis between anemonefish and anemones depends on the presence of the fish drawing other fish to the anemone, where they are stung by its venomous tentacles. The anemone helps the fish by giving it protection from predators, which include brittle stars, wrasses, and other damselfish, and the fish helps the anemone by feeding it, increasing oxygenation, and removing waste material from the host. Various hypotheses exist about the fish's ability to live within the anemone without being harmed. One study carried out at Marineland of the Pacific by Dr. Demorest Davenport and Dr. Kenneth Noris in 1958 revealed that the mucus secreted by the anemone fish prevented the anemone from discharging its lethal stinging nematocysts. A second hypothesis is that A. percula has acquired immunity towards the sea anemone's toxins, and a combination of the two has been shown to be the case. The fish feed on algae, zooplankton, worms, and small crustaceans.
Selective breeding is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant males and females will sexually reproduce and have offspring together. Domesticated animals are known as breeds, normally bred by a professional breeder, while domesticated plants are known as varieties, cultigens, cultivars, or breeds. Two purebred animals of different breeds produce a crossbreed, and crossbred plants are called hybrids. Flowers, vegetables and fruit-trees may be bred by amateurs and commercial or non-commercial professionals: major crops are usually the provenance of the professionals.
Trawling is an industrial method of fishing that involves pulling a fishing net, that is heavily weighted to keep it on the seafloor, through the water behind one or more boats. The net used for trawling is called a trawl. This principle requires netting bags which are towed through water to catch different species of fishes or sometimes targeted species. Trawls are often called towed gear or dragged gear.
In population ecology and economics, maximum sustainable yield (MSY) is theoretically, the largest yield that can be taken from a species' stock over an indefinite period. Fundamental to the notion of sustainable harvest, the concept of MSY aims to maintain the population size at the point of maximum growth rate by harvesting the individuals that would normally be added to the population, allowing the population to continue to be productive indefinitely. Under the assumption of logistic growth, resource limitation does not constrain individuals' reproductive rates when populations are small, but because there are few individuals, the overall yield is small. At intermediate population densities, also represented by half the carrying capacity, individuals are able to breed to their maximum rate. At this point, called the maximum sustainable yield, there is a surplus of individuals that can be harvested because growth of the population is at its maximum point due to the large number of reproducing individuals. Above this point, density dependent factors increasingly limit breeding until the population reaches carrying capacity. At this point, there are no surplus individuals to be harvested and yield drops to zero. The maximum sustainable yield is usually higher than the optimum sustainable yield and maximum economic yield.
Overfishing is the removal of a species of fish from a body of water at a rate greater than that the species can replenish its population naturally, resulting in the species becoming increasingly underpopulated in that area. Overfishing can occur in water bodies of any sizes, such as ponds, wetlands, rivers, lakes or oceans, and can result in resource depletion, reduced biological growth rates and low biomass levels. Sustained overfishing can lead to critical depensation, where the fish population is no longer able to sustain itself. Some forms of overfishing, such as the overfishing of sharks, has led to the upset of entire marine ecosystems. Types of overfishing include: growth overfishing, recruitment overfishing, ecosystem overfishing.
Population ecology is a sub-field of ecology that deals with the dynamics of species populations and how these populations interact with the environment, such as birth and death rates, and by immigration and emigration.
Oncorhynchus is a genus of ray-finned fish in the subfamily Salmoninae of the family Salmonidae, native to coldwater tributaries of the North Pacific basin. The genus contains twelve extant species, namely six species of Pacific salmon and six species of Pacific trout, all of which are migratory mid-level predatory fish that display natal homing and semelparity.
Life history theory (LHT) is an analytical framework designed to study the diversity of life history strategies used by different organisms throughout the world, as well as the causes and results of the variation in their life cycles. It is a theory of biological evolution that seeks to explain aspects of organisms' anatomy and behavior by reference to the way that their life histories—including their reproductive development and behaviors, post-reproductive behaviors, and lifespan —have been shaped by natural selection. A life history strategy is the "age- and stage-specific patterns" and timing of events that make up an organism's life, such as birth, weaning, maturation, death, etc. These events, notably juvenile development, age of sexual maturity, first reproduction, number of offspring and level of parental investment, senescence and death, depend on the physical and ecological environment of the organism.
Premnas biaculeatus, commonly known as spine-cheeked anemonefish or the maroon clownfish, is a species of anemonefish found in the Indo-Pacific from western Indonesia to Taiwan and the Great Barrier Reef. They can grow up to be about 17 cm (6.7 in). Like all anemonefishes it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of the host anemone. It is a sequential hermaphrodite with a strict size-based dominance hierarchy; the female is largest, the breeding male is second largest, and the male nonbreeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male changes to female if the sole breeding female dies, with the largest nonbreeder becoming the breeding male. The fish's natural diet includes algae and zooplankton.
The sebae anemone, also known as leathery sea anemone, long tentacle anemone, or purple tip anemone, is a species of sea anemone belonging to the family Stichodactylidae and native to the Indo-Pacific area.
Bubble-tip anemone is a species of sea anemone in the family Actiniidae. Like several anemone species, E. quadricolor can support several anemonefish species, and displays two growth types based on where they live in the water column, one of which gives it the common name, due to the bulbous tips on its tentacles.
The Red Sea Clownfish, commonly known as the Red Sea or two-bandedanemonefish is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes. Like other species of the genus, the fish feeds on algae and zooplankton in the wild.
A fishery is an area with an associated fish or aquatic population which is harvested for its commercial or recreational value. Fisheries can be wild or farmed. Population dynamics describes the ways in which a given population grows and shrinks over time, as controlled by birth, death, and migration. It is the basis for understanding changing fishery patterns and issues such as habitat destruction, predation and optimal harvesting rates. The population dynamics of fisheries is used by fisheries scientists to determine sustainable yields.
In 1992, Northern Cod populations fell to 1% of historical levels, due in large part to decades of overfishing. The Canadian Federal Minister of Fisheries and Oceans, John Crosbie, declared a moratorium on the Northern Cod fishery, which for the preceding 500 years had primarily shaped the lives and communities of Canada's eastern coast. A significant factor contributing to the depletion of the cod stocks off Newfoundland's shores was the introduction of equipment and technology that increased landed fish volume. From the 1950s onwards, new technology allowed fishers to trawl a larger area, fish more in-depth, and for a longer time. By the 1960s, powerful trawlers equipped with radar, electronic navigation systems, and sonar allowed crews to pursue fish with unparalleled success, and Canadian catches peaked in the late-1970s and early-1980s. Cod stocks were depleted at a faster rate than could be replenished.
Salmon population levels are of concern in the Atlantic and in some parts of the Pacific. Salmon are typically anadromous - they rear and grow in freshwater, migrate to the ocean to reach sexual maturity, and then return to freshwater to spawn. Determining how environmental stressors and climate change will affect these fisheries is challenging due to their lives split between fresh and saltwater. Environmental variables like warming temperatures and habitat loss are detrimental to salmon abundance and survival. Other human influenced effects on salmon like overfishing and gillnets, sea lice from farm raised salmon, and competition from hatchery released salmon have negative effects as well.
Saltwater fish, also called marine fish or sea fish, are fish that live in seawater. Saltwater fish can swim and live alone or in a large group called a school.
The gill plate trade is the buying and selling of stingray gill plates for their use in traditional Chinese medicine. The gill plates are harvested from stingrays that are caught intentionally, or caught as bycatch. The plates are sold whole or in a powder form. A single kilogram of the gill plate can be sold for up to US$350, though the price varies by ray species.
Stephanie M. Carlson is the A.S. Leopold Chair in Wildlife Biology at the University of California Berkeley. Her research considers fish ecology, freshwater ecology, and evolutionary ecology.