Climate change is not a new phenomenon. During the past 2 million years, there have been at least 10 cycles of global warming and cooling. When the polar ice caps melted during warming periods, sea levels rose to well above their earlier levels, and a larger portion of Earth experienced tropical climates. During cooling periods, the polar ice caps expanded, sea levels dropped, and tropical species’ ranges contracted. Sometimes these changes occurred gradually, which enabled the affected species to adapt. But the onset of some climate change periods was abrupt, causing major ecosystem disruptions and global mass extinction events. Yet, nature recovered every time; many of the species we see today are survivors of previous climate change events. It is thus fair to ask why today’s climate change is of such concern to us.
10.5.1 Climate change’s impact on people
History provides us with many lessons to illustrate the impact of climate change on human societies. These lessons start with the earliest well-documented example of a societal collapse—that of the Middle East’s Natufian communities roughly 10,000 years ago—which has been attributed to climatic changes (Weiss and Bradley, 2001). Since then, climate change has regularly contributed to the collapse of complex human societies across the world. Notable examples of such collapses include the Akkadian Empire (the world’s first empire) of the Middle East (Carolin et al., 2019), Egypt’s Old Kingdom (who constructed the pyramids), Central America’s Classic Mayan civilisation, the USA’s first English colony (deMenocal, 2001), several Chinese dynasties (Wang et al., 2010), and the Late Bronze Age societies along the Mediterranean Sea (Kaniewski et al., 2013). Also, in Southern Africa, the fall of the Mapungubwe Kingdom has been attributed to crop failures and declining grazing lands due to regional droughts and warming cycles (O’Connor and Kiker, 2004).
Unlike the unavoidable natural climatic shifts that led to the historical societal collapses discussed above, we have brought today’s climatic change impacts upon ourselves. Because of our general lack of response in addressing the drivers of climate change, thousands of people will suffer the consequences. Prominently, many parts of the world are already seeing higher temperatures and longer droughts (Engelbrecht et al., 2009). These conditions are compromising our quality of life (Watts et al., 2017) by leading to more intense wildfires (Jolly et al., 2015; Strydom and Savage, 2016), increased incidences of malaria (Siraj et al., 2014), increased crop failures (Myers et al., 2014; Medek et al., 2017), and increased competition for water (Flörke et al., 2018). Many coastal areas are also seeing storms increasing in intensity and frequency, exposing people living near large rivers, deltas, and estuaries to more frequent flooding (Figure 10.5.1) and storm surges (Fitchett and Grab, 2014). Sea level rise is expected to leave many low-lying oceanic islands uninhabitable within a few decades (Storlazzi et al., 2018). With all these impacts expected to increase the competition for space under an increasing human population, it would be wise for the world’s governments to start preparing for thousands of climate refugees that would need to be relocated in the near future (Merone and Tait, 2018).
To combat climate change, politicians of several countries have started to enact laws to reduce greenhouse gas emissions and habitat destruction. Many industries are also hard at work developing “greener” technologies to enable us to live more sustainable lives. Conservation biologists also play a crucial role in mitigating the negative impacts of climate change. In addition to highlighting the plight of the natural world to society at large, we could work towards reducing the loss of ecosystem services and preventing species extinctions. To accomplish this task, we need to identify which species and ecosystems are most sensitive to climate change and develop strategies that will ensure the continued persistence of as many sensitive species and their habitats as possible. The rest of this chapter is dedicated to methods we can employ to understand which species are sensitive, and how they may respond to climate change.
10.5.2 Climate change’s impact on terrestrial ecosystems
Aside from regional variations in temperature and precipitation, Earth’s surface will be a few degrees warmer in future than the temperatures we experience today. In effect, that means that today’s climatic zones will generally shift upslope in mountainous areas and towards the poles on lowlands, plains, and plateaus. To survive, climate-sensitive plants and animals will need to track these shifts so that they remain within their suitable climatic envelopes of temperature and precipitation. By altering regional climates, it makes habitats less hospitable to the species living in them. While increased carbon dioxide levels can help plants conduct photosynthesis more efficiently, they are threatened by harsh temperatures and extreme weather events. Additionally, with warmer conditions, moisture from snow melt arrives earlier in the season, lengthening the fire season.
In response to changing conditions, range shifts have also been observed in plants, butterflies, other insects, freshwater fishes, reptiles, amphibians, and mammals. Because individual plants cannot physically move to cooler regions, plant range shifts result from seed dispersal. Seeds are often dispersed in all directions away from a parent plant, but more of the seedlings that establish in northern locations or higher elevations survive, resulting in a gradual shift towards the poles or up mountains (Figure 10.5.2). However, species that cannot adapt to new conditions or shift their ranges quickly enough face extinction.
Climate change on mountains
Species that live on mountains are at particular risk from climate change. Because temperatures decrease by roughly 0.65°C for every 100 m in elevation rise (known as temperature lapse rates), a 1°C increase suggests that climate-sensitive species living on a mountain would be displaced by at least 150 m (1.5 m/year) upslope between the years 2000 and 2100. Species that live on the lower slopes of mountains and are mobile enough to make such an adjustment may have opportunities to move to higher ground. However, species that live on or near peaks may have nowhere else to go as the world heats up, resulting in what biologists call mountain-top extinctions. It is only a matter of time before vulnerable mountain specialists will follow the example of Costa Rica’s once abundant Monteverde golden toad (Bufo periglenes, EX), the first known amphibian extinction attributed to climate change (Crump et al., 1992).
Species that live on mountain peaks are vulnerable to climate change because they may have nowhere else to go as the world heats up.
Climate change in the lowlands
The response of species living in lowlands and on plains tend be more variable and complex than those living on mountains. While some species may only need to make minor range adjustments, researchers estimate that some taxa may need to move 500 km (Barbet-Massin et al., 2009)—maybe even 1,000 km (Hsiang and Sobel, 2016)—to keep up with climate shifts. For species that have already shifted their distributions by 200–300 km (Beale et al., 2013), adapting seems relatively easy thanks to their mobility and largely intact ecosystems. Unfortunately, the rate of climate change will likely outpace the ability for most species to adapt (Jezkova and Wiens, 2016; Wiens, 2016).
Species of tropical lowland forests and deserts are also highly vulnerable to shifting climates. Many tropical species have narrow tolerances for temperature and rainfall variation, while desert specialists may be at the limits of their physiological heat and desiccation tolerances (Figure 10.5.2). Consequently, even small changes in the climate of these two ecosystems may have major effects on reproduction, species distributions, and hence ecosystem composition (Box 10.5.1). One species already impacted is the nocturnal aardvark (Orycteropus afer, LC): a study in Southern Africa’s Kalahari Desert found over 80% mortality rates in this species during recent summers (Rey et al., 2017). The high levels of mortality in this species was attributed to above average temperatures, which subjected the animals to heat stress, leading to behavioral disruptions, declining body conditions, and eventually starvation. The impact of climate change on the aardvark is of concern because it is an ecosystem engineer: their burrows provide denning and refuge sites for multiple other species (Whittington-Jones et al., 2011).
Box 10.5.1 Desert Birds and Climate Change
Susan Cunningham1 and Andrew McKechnie2,3
1FitzPatrick Institute of African Ornithology, DST-NRF Centre of Excellence,
University of Cape Town, South Africa.
2DST-NRF Centre of Excellence at the FitzPatrick Institute,
Department of Zoology and Entomology, University of Pretoria, South Africa.
3South African Research Chair in Conservation Physiology, National Zoological Garden,
South African National Biodiversity Institute, Pretoria, South Africa.
Deserts, with their extreme temperatures and scarce and unpredictable rainfall, are among the most inhospitable environments on the planet. To survive and breed in arid regions, organisms must minimize their energy and water requirements, and avoid exposure to potentially lethal temperatures. Birds are generally small and diurnal; and are therefore among the groups of animals most vulnerable to even small increases in air temperatures associated with climate change. Studies of the effects of temperature on arid-zone birds can thus be highly informative in terms of identifying new conservation challenges posed by global warming, developing mitigation measures, and understanding the management interventions that may become necessary during the 21st Century.
Daytime temperatures in many deserts regularly exceed avian body temperature, creating conditions under which birds can avoid lethal heat stroke only by dissipating heat via evaporation. But rapid rates of evaporation increase the risk of birds becoming lethally dehydrated. Desert birds thus face life-or-death decisions between avoiding hyperthermia by evaporative cooling versus avoiding lethal dehydration by minimizing water losses. Mass mortality events occasionally take place during extreme heat waves when air temperatures exceed birds’ physiological tolerance limits. In Australia, for example, there are both historic and contemporary accounts of die-offs sometimes involving millions of birds. As Earth heats up under climate change, the risk of such die-offs in desert birds is expected to increase dramatically for the deserts of Australia and North America during the 21st Century (McKechnie and Wolf, 2010; Albright et al., 2017).
Arid regions are also experiencing significant temperature increases which are predicted to continue over the next several decades (Conradie et al., 2019). Under these conditions, the impact of air temperature on avian physiology can be mediated by behavior. Birds employ a trio of behavioral adjustments to manage heat load and keep their body temperatures within safe limits. These include shade-seeking, reducing activity to minimize metabolic heat production, and gaping the beak (panting, sometimes accompanied by gular flutter) to facilitate respiratory evaporative cooling (Figure 10.5.2). Although these behaviors can buffer birds against physiological costs of high temperature, they carry subtle but important costs of the own, notably via their impact on birds’ ability to forage.
For desert birds, foraging is critically important for maintaining both energy and water balance, as most species obtain all their water from food. Reduced activity almost inevitably means reduced food intake via impacts on time available for foraging. Seeking shade also carries costs: for some species, returns on foraging effort in shaded locations are significantly lower than in the sun (e.g. Cunningham et al., 2013). Finally, respiratory evaporative cooling can severely restrict the ability of actively-foraging birds to acquire food due to mechanical constraints on simultaneously gaping the bill and using it for prey capture and handling (e.g. du Plessis et al., 2012).
Under climate change, the implications of these behavioral trade-offs between foraging and thermoregulation are non-trivial. Inability to balance water and energy budgets mean birds progressively lose body condition during heat waves (du Plessis et al., 2012). Compromised foraging also affects birds’ capacity to provision offspring, resulting in reduced nest success and/or smaller, lighter fledglings which may struggle to survive and recruit into the breeding population (e.g. Cunningham et al., 2013, Wiley and Ridley, 2016).
Successfully balancing the trade-offs between foraging and thermoregulation, and between hyperthermia and dehydration, is the secret to success for birds in hot places. As the climate warms, achieving this balance will become ever more challenging. Sublethal behavioral costs of keeping cool kick in at temperatures cooler than those promoting mass mortalities. In some parts of the world, the loss of birds from desert ecosystems may therefore occur through the insidious whittling away of fitness and weakening of populations (Conradie et al., 2019) before we even witness the dramatic die-off events for which Australia is already infamous.
An additional concern for lowland ecosystems is that climate change will likely lead to the creation of novel (i.e. hotter) ecosystems unlike any others currently on Earth (Williams et al., 2007). These changes will lead to biotic attrition. The gradual impoverishment of biological communities of lowland ecosystems as species either go extinct or move away while tracking their climatic envelopes. What is not clear is how the niches left open by the net loss of species, and newly created niches in the novel ecosystems, will be filled. The most likely scenario is that more tolerant, generalist species will fill the empty niches. However, with the inevitable loss of some species, combined with the decoupling of important biological interactions (discussed below), some functions and services associated with lowland ecosystems are likely to eventually collapse. It is important to note that tropical lowland forests and deserts are by no means the only ecosystems vulnerable to biotic attrition.
Climate change and dispersal limitations
Across many diverse ecosystems, a great number of species are threatened by climate change because of their poor dispersal abilities. Because they lack appropriate dispersal mechanisms, species, such as slow maturing plants (Foden et al., 2007), mosses, and flightless insects may simply not be able to keep up with changing climatic conditions. The impacts of climate change on Africa’s dispersal-limited species can already be seen. For example, the once abundant Aldabra banded snail (Rhachistia aldabrae, CR) is today so rare that this Lazarus species (Figure 10.5.5) was once believed to be extinct due to climate change (Battarbee, 2014). Dispersal limitations will also greatly affect terrestrial species living on oceanic islands, which will find it near impossible to track their climatic niches as it moves over the ocean.
Climate change and biological interactions
Species that are highly mobile are not entirely spared from the negative impacts of climate change. Consider migratory species for a moment. The same way the musicians of an orchestra rely on a conductor to remain synchronized, migratory species rely on environmental cues, such as daylength and temperature, to decide when they need to start moving from one area to the next. But because different species rely of different environmental cues to time their life cycles (e.g. breeding), not all species will adjust to climate change at the same rate. There is consequently a high likelihood that climate change will disrupt these synchronous movements that the animal kingdom has developed over thousands of years (Renner and Zohner, 2018). This disruption of timed aspects of species’ life cycle, such as migration and breeding, is called phenological mismatch or trophic asynchrony. Researchers have already seen signs of phenological mismatch: some migratory birds have started to migrate to their breeding grounds at earlier dates than before (Both et al., 2006; Vickery et al., 2014). If these trends hold, they may soon start breeding before peak food availability, which could lead to lower fitness of offspring.
Resident species are also vulnerable to phenological mismatch. While these species might not be known for large-scale movements around the globe, they may still have to adjust their ranges to keep track of their climatic niches. Considering the improbability of different species will adapt at the same pace, there is thus a danger that important mutualistic relationships might be pulled apart these during range adaptations. This is of concern for species with specialized feeding niches, as seen in some pollinators. Extinctions arising from this decoupling of mutualistic relationships are referred to as coextinction (Koh et al., 2004), while a series of linked coextinctions is called an extinction cascade.
Climate change and reptiles
One may think that reptiles—often seen basking on sun-drenched rocks to obtain active body temperatures—may benefit from climate change. Yet, as a group, they are also expected to suffer under climate change. One reason is because many reptiles will also have to adapt their ranges to shifting climates (Houniet et al., 2009). Even more important, climate change will increase reptiles’ vulnerability to demographic stochasticity. Many reptiles—and some fish—have their sex determined by temperature during embryonic development, with warmer temperatures often leading to more females (Valenzuala and Lance, 2004). In general, females regulate their offspring’s sex ratios by fine-scale breeding site selection. Under climate change, however, it might be harder for the females to find breeding sites with suitable microclimates. Those species unable to adopt new mechanisms to control for offspring sex ratio bias may eventually go extinct, even under relatively small temperature shifts (Sinervo et al., 2010).
10.5.3 Climate change’s impact on freshwater ecosystems
With the world's freshwater ecosystems already strained by the demands of a growing human population, freshwater biodiversity will face several additional stressors associated with climate change. Climate change will impact water temperature, flow volume, and flow variability. Because these variables are three primary predictors of freshwater ecosystem composition (van Vliet et al., 2013; Knouft and Ficklin, 2017), it is expected that climate change will greatly affect freshwater ecosystem composition and functioning in the coming decades.
Warmer rivers and streams
Climatologists and hydrologists predict that freshwater ecosystems will generally experience temperature increases under climate change. Like their terrestrial counterparts, many freshwater species are sensitive to temperature shifts (e.g. Reizenberg et al., 2019). Warmer water also holds less dissolved oxygen, and increased pollutant toxicity (Whitehead et al., 2009). In addition, longer growing seasons and higher water temperatures will lead to a general increase in primary productivity and decomposition rates, which in turn will lead to increased nutrient loads, algae blooms, and eutrophication (Whitehead et al., 2009). All these factors will force many freshwater species—even those not sensitive to temperature shifts—to adjust their ranges to keep track of suitable conditions. Many of these adjustments will be impeded by habitat fragmentation, notably by dams and other human constructs that block suitable dispersal pathways. As an additional complication, many aquatic organisms cannot travel overland, so are naturally limited to adjust their ranges along the rivers and streams in which they live. But the orientation of these rivers and streams may not follow suitable thermal isolines: consider a cold-water species that needs to disperse to higher elevation—and hence upstream—as its climate niche moves higher up a mountain. For some freshwater species, the impediments to adjusting their ranges as necessary may be insurmountable.
Climate change will alter water temperature, flow volume, and flow variability, the three primary predictors of freshwater ecosystem composition.
Changing flow regimes
Changing precipitation levels will have several impacts on freshwater ecosystems, particularly as it relates to changes in their flow regimes (Thieme et al., 2010; Knouft and Ficklin, 2017). For example, areas that are undergoing decreased precipitation will experience decreased runoff and increased drying of wetlands and small streams, while areas with increased precipitation will experience increased storm surges and flushing. These changes, together with the impacts of increased water extraction rates and evapotranspiration under a warmer world, will cause significant changes in water levels, flow rates, sediment loads, water turbidity, and the structure of the physical environment.
Given these multiple stressors, there is a reasonable expectation that many freshwater species will go extinct or face significant population declines and range shifts over the next decades. These changes are of major concern especially in regions where so many people depend on fish and related natural resources for their livelihoods.
10.5.4 Climate change’s impact on marine ecosystems
Like tropical forests, the world’s oceans have historically provided a relatively stable environment in which marine organisms have evolved. While this stability promotes species diversity, it also leaves marine species more vulnerable to environmental changes. In fact, a recent study found that cold-blooded marine species are twice as vulnerable to the impacts of warmer oceans than their terrestrial counterparts (Pinsky et al., 2019). In addition to the impacts of storm surges (Figure 10.5.6) and ocean warming (which leads to rising sea levels and ocean deoxygenation), marine organisms also must deal with ocean acidification. These threats will likely have impacts like those faced in terrestrial and freshwater ecosystems, including range adjustments, biotic attrition, and decoupling or important interactions. Below we discuss the mechanisms that will lead to some of these changes in more detail.
As discussed earlier, human activities release massive amounts of CO2 into the atmosphere each day. Although forests and other plant communities get considerable attention for CO2 sequestration, the world’s oceans also play a key role in keeping Earth’s carbon balance in check. In fact, the world’s oceans absorb an estimated 20–25% of our current CO2 emissions (Khatiwala et al., 2009). Now, with more atmospheric CO2 available, oceans absorb more carbon, which dissolves in seawater as carbonic acid. While this absorption may slow climate change, it also increases the acidity (i.e. lowing the pH levels) of the world’s oceans. This process—known as ocean acidification—has several consequences that may directly and indirectly kill marine organisms. For example, it inhibits the ability of coral animals to deposit the calcium used to build their reefs’ structure (Mollica et al., 2018), and prevents shellfish from accumulating adequate amounts of calcium carbonate to develop shells strong enough for survival (Branch et al., 2013). Ocean acidification also disturbs predator-prey dynamics by impairing the senses of prey species (Leduc et al., 2013), and compromising the ability of marine creatures to communicate with conspecifics (Roggatz et al., 2016).
Climate change is causing sea level rise and increased seawater temperatures, with broad implications for marine ecology and people living in coastal areas.
Sea level rise
Over the past 30–40 years, ocean surface temperatures have warmed by about 0.64°C (NOAA, 2016).Ocean warming has several implications, the most well-known being sea level rise, caused by the thermal expansion of ocean water combined with the released water from melting glaciers and polar ice caps. As the oceans creep further inland, the extent of low-lying coastal ecosystems such as rocky shores or sandy beaches will shrink, and so also the sizes of the wildlife populations living in those areas. The extinction of Australia’s Bramble Cay melomys (Melomys rubicola, EX)—the world’s first documented mammalian extinction caused by anthropogenic climate-change—has been attributed to sea level rise (Gynther et al., 2016).
The incredible diversity of corals reef ecosystems is attributable to the relative stability of tropical oceans. Because of this stability, individual coral species have adapted to very specialized niches. Many corals thus tolerate only narrow ranges in temperature, sunlight levels, water opacity, and nutrient loads. Climate change is disrupting this stability, by changing the temperature (ocean warming), depth (sea level rise), sediment and nutrient loads (increased erosion and runoff) of the environments where corals live. These changes are leading to a breakdown of critical mutualistic relationships between photosynthetic algae and corals. In the process, corals also lose their vibrant colors, revealing the corals’ ghostly white skeletons, hence the name coral bleaching (Figure 10.5.7). This relationship breakdown deprives the corals of essential carbohydrates they obtain from the algae, causing the corals to starve to death if the stressful conditions continue for a prolonged time.
Marine fish and invertebrates rely on dissolved oxygen that enters the water either through the atmosphere, or by photosynthetic plankton. But because warmer water absorbs less oxygen, scientists predict that some areas of the ocean will see a 3–6% drop in dissolved oxygen concentrations under climate change (IPCC, 2014). This process, known as ocean suffocation or ocean deoxygenation (Ito et al., 2017), will leave parts of the ocean unsuitable for marine fishes and invertebrates. The impact of ocean deoxygenation will also be felt by economically important fisheries.
10.5.5 Climate change interacts with habitat loss
Habitat loss and climate change each cause negative impacts on biodiversity; however, these threats also interact to have an overall larger negative impact than the sum of these threats independently. Prominently, because of habitat loss, many species will be unable to adequately adjust their ranges to keep track of their shifting climatic niches. For example, some species might not be able to adapt their ranges because suitable habitat in their future ranges will be destroyed by human activity.
Climate change interacts with habitat loss, by impeding species’ ability to adapt, and by bringing dispersing wildlife into conflict with humans.
Range-shift gaps describes a habitat gap that prevents a species from dispersing from its current to future ranges (Figure 10.5.8). These gaps, which may occur naturally or because of habitat fragmentation, may also impede range adjustments under climate change. While the impact of range-shift gaps is an active area of research, it is expected that mountain-top species may be inherently vulnerable to range-shift gaps, particularly if they are unable to first disperse downslope before they can reach climatically suitable locations at higher elevation elsewhere.
Habitat loss and climate change are also expected to exacerbate human-wildlife conflicts. Losses of arable land will see even more natural ecosystems converted for agriculture which, in turn, will further increase competition among and between humans and wildlife for resources such as food, water, and suitable habitat (Serdeczny et al., 2017). As the human footprint expands across Earth, agriculture and infrastructure will impede the ability of specialist species to find food and adapt to changing conditions, while generalist species will be forced into agricultural lands and nearby human habitation as they search for resources and/or disperse across the landscape. Such a scenario will likely exacerbate human-wildlife conflict where predators living in fragmented ecosystems with diminishing natural prey populations are increasingly prone to wandering beyond protected area boundaries into ranching areas in search of food (Tuqa et al., 2014).