Juha Merilä and Ary A. Hoffmann
Changing climatic conditions have both direct and indirect influences on abiotic and biotic processes and represent a potent source of novel selection pressures for adaptive evolution. In addition, climate change can impact evolution by altering patterns of hybridization, changing population size, and altering patterns of gene flow in landscapes. Given that scientific evidence for rapid evolutionary adaptation to spatial variation in abiotic and biotic environmental conditions—analogous to that seen in changes brought by climate change—is ubiquitous, ongoing climate change is expected to have large and widespread evolutionary impacts on wild populations. However, phenotypic plasticity, migration, and various kinds of genetic and ecological constraints can preclude organisms from evolving much in response to climate change, and generalizations about the rate and magnitude of expected responses are difficult to make for a number of reasons.
First, the study of microevolutionary responses to climate change is a young field of investigation. While interest in evolutionary impacts of climate change goes back to early macroevolutionary (paleontological) studies focused on prehistoric climate changes, microevolutionary studies started only in the late 1980s. The discipline gained real momentum in the 2000s after the concept of climate change became of interest to the general public and funding organizations. As such, no general conclusions have yet emerged. Second, the complexity of biotic changes triggered by novel climatic conditions renders predictions about patterns and strength of natural selection difficult. Third, predictions are complicated also because the expression of genetic variability in traits of ecological importance varies with environmental conditions, affecting expected responses to climate-mediated selection.
There are now several examples where organisms have evolved in response to selection pressures associated with climate change, including changes in the timing of life history events and in the ability to tolerate abiotic and biotic stresses arising from climate change. However, there are also many examples where expected selection responses have not been detected. This may be partly explainable by methodological difficulties involved with detecting genetic changes, but also by various processes constraining evolution.
There are concerns that the rates of environmental changes are too fast to allow many, especially large and long-lived, organisms to maintain adaptedness. Theoretical studies suggest that maximal sustainable rates of evolutionary change are on the order of 0.1 haldanes (i.e., phenotypic standard deviations per generation) or less, whereas the rates expected under current climate change projections will often require faster adaptation. Hence, widespread maladaptation and extinctions are expected. These concerns are compounded by the expectation that the amount of genetic variation harbored by populations and available for selection will be reduced by habitat destruction and fragmentation caused by human activities, although in some cases this may be countered by hybridization. Rates of adaptation will also depend on patterns of gene flow and the steepness of climatic gradients. Theoretical studies also suggest that phenotypic plasticity (i.e., nongenetic phenotypic changes) can affect evolutionary genetic changes, but relevant empirical evidence is still scarce. While all of these factors point to a high level of uncertainty around evolutionary changes, it is nevertheless important to consider evolutionary resilience in enhancing the ability of organisms to adapt to climate change.
Mark V. Barrow
The prospect of extinction, the complete loss of a species or other group of organisms, has long provoked strong responses. Until the turn of the 18th century, deeply held and widely shared beliefs about the order of nature led to a firm rejection of the possibility that species could entirely vanish. During the 19th century, however, resistance to the idea of extinction gave way to widespread acceptance following the discovery of the fossil remains of numerous previously unknown forms and direct experience with contemporary human-driven decline and the destruction of several species. In an effort to stem continued loss, at the turn of the 19th century, naturalists, conservationists, and sportsmen developed arguments for preventing extinction, created wildlife conservation organizations, lobbied for early protective laws and treaties, pushed for the first government-sponsored parks and refuges, and experimented with captive breeding. In the first half of the 20th century, scientists began systematically gathering more data about the problem through global inventories of endangered species and the first life-history and ecological studies of those species.
The second half of the 20th and the beginning of the 21st centuries have been characterized both by accelerating threats to the world’s biota and greater attention to the problem of extinction. Powerful new laws, like the U.S. Endangered Species Act of 1973, have been enacted and numerous international agreements negotiated in an attempt to address the issue. Despite considerable effort, scientists remain fearful that the current rate of species loss is similar to that experienced during the five great mass extinction events identified in the fossil record, leading to declarations that the world is facing a biodiversity crisis. Responding to this crisis, often referred to as the sixth extinction, scientists have launched a new interdisciplinary, mission-oriented discipline, conservation biology, that seeks not just to understand but also to reverse biota loss. Scientists and conservationists have also developed controversial new approaches to the growing problem of extinction: rewilding, which involves establishing expansive core reserves that are connected with migratory corridors and that include populations of apex predators, and de-extinction, which uses genetic engineering techniques in a bid to resurrect lost species. Even with the development of new knowledge and new tools that seek to reverse large-scale species decline, a new and particularly imposing danger, climate change, looms on the horizon, threatening to undermine those efforts.
Fisheries science emerged in the mid-19th century, when scientists volunteered to conduct conservation-related investigations of commercially important aquatic species for the governments of North Atlantic nations. Scientists also promoted oyster culture and fish hatcheries to sustain the aquatic harvests. Fisheries science fully professionalized with specialized graduate training in the 1920s.
The earliest stage, involving inventory science, trawling surveys, and natural history studies continued to dominate into the 1930s within the European colonial diaspora. Meanwhile, scientists in Scandinavian countries, Britain, Germany, the United States, and Japan began developing quantitative fisheries science after 1900, incorporating hydrography, age-determination studies, and population dynamics. Norwegian biologist Johan Hjort’s 1914 finding, that the size of a large “year class” of juvenile fish is unrelated to the size of the spawning population, created the central foundation and conundrum of later fisheries science. By the 1920s, fisheries scientists in Europe and America were striving to develop a theory of fishing. They attempted to develop predictive models that incorporated statistical and quantitative analysis of past fishing success, as well as quantitative values reflecting a species’ population demographics, as a basis for predicting future catches and managing fisheries for sustainability. This research was supported by international scientific organizations such as the International Council for the Exploration of the Sea (ICES), the International Pacific Halibut Commission (IPHC), and the United Nations’ Food and Agriculture Organization (FAO).
Both nationally and internationally, political entanglement was an inevitable feature of fisheries science. Beyond substituting their science for fishers’ traditional and practical knowledge, many postwar fisheries scientists also brought progressive ideals into fisheries management, advocating fishing for a maximum sustainable yield. This in turn made it possible for governments, economists, and even scientists, to use this nebulous target to project preferred social, political, and economic outcomes, while altogether discarding any practical conservation measures to rein in globalized postwar industrialized fishing. These ideals were also exported to nascent postwar fisheries science programs in developing Pacific and Indian Ocean nations and in Eastern Europe and Turkey.
The vision of mid-century triumphalist science, that industrial fisheries could be scientifically managed like any other industrial enterprise, was thwarted by commercial fish stock collapses, beginning slowly in the 1950s and accelerating after 1970, including the massive northern cod crisis of the early 1990s. In the 1980s scientists, aided by more powerful computers, attempted multi-species models to understand the different impacts of a fishery on various species. Daniel Pauly led the way with multi-species models for tropical fisheries, where the need for such was most urgent, and pioneered the global database FishBase, using fishing data collected by the FAO and national bodies. In Canada the cod crisis inspired Ransom Myers to use large databases for fisheries analysis to show the role of overfishing in causing that crisis. After 1980 population ecologists also demonstrated the importance of life history data for understanding fish species’ responses to fishery-induced population and environmental change.
With fishing continuing to shrink many global commercial stocks, scientists have demonstrated how different measures can manage fisheries for species with different life-history profiles. Aside from the need for effective scientific monitoring, the biggest ongoing challenges remain having politicians, governments, fisheries industry members, and other stakeholders commit to scientifically recommended long-term conservation measures.
Rewilding aims at maintaining or even increasing biodiversity through the restoration of ecological and evolutionary processes using extant keystone species or ecological replacements of extinct keystone species that drive these processes. It is hailed by some as the most exciting and promising conservation strategy to slow down or stop what is considered to be the greatest mass extinction of species since the extinction of the dinosaurs 65 million years ago. Others have raised serious concerns about the many scientific and societal uncertainties and risks of rewilding. Moreover, despite its growing popularity, rewilding has made only limited inroads within the conservation mainstream and still has to prove itself in practice.
Rewilding differs from traditional restoration in at least two important respects. Whereas restoration has typically focused on the recovery of plants communities, rewilding has drawn attention to animals, particularly large carnivores and large herbivores. Whereas restoration aims to return an ecosystem back to some historical condition, rewilding is forward-looking rather than backward-looking: it examines the past not so much to recreate it, but to learn from the past how to activate and maintain the natural processes that are crucial for biodiversity conservation.
Rewilding makes use of a variety of techniques to re-establish these natural processes. Besides the familiar method of reintroducing animals in areas where populations have decreased dramatically or even gone extinct, rewilders also employ some more controversial methods, including back breeding to restore wild traits in domesticated species, taxon substitution to replace extinct species by closely related species with similar roles within an ecosystem, and de-extinction to bring extinct species back to life again using advanced biotechnological technologies such as cloning and gene editing.
Rewilding has clearly gained the most traction in North America and Europe, which have several key features in common. Both regions have recently experienced a spontaneous return of wildlife. Rewilders on both sides of the Atlantic are aware, however, that this wildlife resurgence is not that impressive, given that we are in the midst of the sixth mass extinction, which is characterized by the loss of large-bodied animals known as megafauna. The common goal is to bring back such megafaunal species because of their importance for maintaining and enhancing biodiversity. Last, both North American and European rewilders perceive the extinction crisis through the lens of the island theory, which shows that the number of species in an area depends on its size and degree of isolation—hence their special attention to the spatial aspects of rewilding.
But rewilding projects on both sides of the Atlantic not only have much in common, they also differ in certain aspects. North American rewilders have adopted the late Pleistocene as a reference period and have emphasized the role of predation by large carnivores, while European rewilders have opted for the mid-Holocene and put more focus on naturalistic grazing by large herbivores.
Maria Cristina Fossi and Cristina Panti
A vigorous effort to identify and study sentinel species of marine ecosystem in the world’s oceans has developed over the past 50 years. The One Health concept recognizes that the health of humans is connected to the health of animals and the environment. Species ranging from invertebrate to large marine vertebrates have acted as “sentinels” of the exposure to environmental stressors and health impacts on the environment that may also affect human health. Sentinel species can signal warnings, at different levels, about the potential impacts on a specific ecosystem. These warnings can help manage the abiotic and anthropogenic stressors (e.g., climate change, chemical and microbial pollutants, marine litter) affecting ecosystems, biota, and human health.
The effects of exposure to multiple stressors, including pollutants, in the marine environment may be seen at multiple trophic levels of the ecosystem. Attention has focused on the large marine vertebrates, for several reasons. In the past, the use of large marine vertebrates in monitoring and assessing the marine ecosystem has been criticized. The fact that these species are pelagic and highly mobile has led to the suggestion that they are not useful indicators or sentinel species. In recent years, however, an alternative view has emerged: when we have a sufficient understanding of differences in species distribution and behavior in space and time, these species can be extremely valuable sentinels of environmental quality.
Knowledge of the status of large vertebrate populations is crucial for understanding the health of the ecosystem and instigating mitigation measures for the conservation of large vertebrates. For example, it is well known that the various cetacean species exhibit different home ranges and occupy different habitats. This knowledge can be used in “hot spot” areas, such as the Mediterranean Basin, where different species can serve as sentinels of marine environmental quality. Organisms that have relatively long life spans (such as cetaceans) allow for the study of chronic diseases, including reproductive alterations, abnormalities in growth and development, and cancer. As apex predators, marine mammals feed at or near the top of the food chain. As the result of biomagnification, the levels of anthropogenic contaminants found in the tissues of top predators and long-living species are typically high. Finally, the application of consistent examination procedures and biochemical, immunological, and microbiological techniques, combined with pathological examination and behavioral analysis, has led to the development of health assessment methods at the individual and population levels in wild marine mammals. With these tools in hand, investigators have begun to explore and understand the relationships between exposures to environmental stressors and a range of disease end points in sentinel species (ranging from invertebrates to marine mammals) as an indicator of ecosystem health and a harbinger of human health and well-being.