Climate change and birds: perspectives and prospects from southern Africa

Global climate warming, now conclusively linked to anthropogenically-increased CO₂ levels in the earth's atmosphere, has already had impacts on the earth's biodiversity and is predicted to threaten more than 1 million species with extinction by 2050. Climate change in southern Africa is expected to involve higher temperatures and lower rainfall, with less predictability and a greater frequency of severe storms, fires and El Niño events. The predicted changes to birds in Africa — the continent most at risk from climate change — have hardly been explored, yet birds and many other vertebrates face uncertain futures. Here, in one of the first focused analyses of the correlates of climate change vulnerability in southern African birds, we offer a wide-ranging perspective on which species may be most at risk, and explore which traits may influence the adaptability or extinction risk of bird species.

Our review suggests that small nomadic species with short generation times may be least at risk. While larger-bodied species may be physiologically buffered against environmental change, their longer generation times may make them less able to adapt evolutionarily to climate change. Migrant species, and those with specialised feeding niches such as pollinators, are also predicted to be at risk of population declines, based on low ability to adapt to new environments when introduced there as aliens. Species with small ranges (<50 000km²) restricted to the two southern African biodiversity hotspots most at risk from climate change — the Cape Floral Kingdom and the Succulent Karoo — are ranked according to low, medium or high risk of extinction. Those restricted to mountain slopes, mountain tops or islands, and those occurring mainly at the southern or western extremes of these biomes, are ranked as highest risk. These include endemic sunbirds, warblers and rock-jumpers — none of which are currently recognised Red Data species. Using climate envelopes we modelled the possible range shifts by 2050 of three pairs of species found in habitats considered to be at risk: fynbos, mountain and arid Karoo. All six species lost substantial portions of their range (x = 40%), with the montane Drakensberg Rock-jumper Chaetops aurantius losing most (69%). Significant reductions of available climate space in all species may interact with life history characteristics to threaten many southern African bird species unable to shift geographic range or adapt to novel resource conditions. We conclude with a list of research priorities and testable hypotheses which may advance our understanding of the complex influence that climate change is likely to have on African, particularly southern African, birds.     

Climate change and the optimal arrival of migratory birds

Recent climate change has sparked an interest in the timing of biological events, which is a general problem in life-history evolution. Reproduction in many organisms breeding in seasonal environments, e.g. migratory birds, is dependent on the exploitation of a short but rich food supply. If the seasonal timing of the food peak advances owing to climate change, then one would expect the bird to track those changes, hence, initiate migration and breeding earlier. However, when there is competition for territories and a risk of pre-breeding mortality, the optimal response to a shifting food distribution is no longer obvious. We develop a theoretical model to study how the optimal arrival time depends on the mean and variance of the food distribution, the degree of competition for territories and the risk of mortality. In general, the optimal shift in arrival date should never be as extreme as the shift in food peak date. Our results also show that we should expect the high variation of trends in arrival date observed among migratory birds, even if migration and information about climate change were unconstrained.     

Climate change increases the risk of malaria in birds

Malaria caused by Plasmodium parasites is one of the worst scourges of mankind and threatens wild animal populations. Therefore, identifying mechanisms that mediate the spread of the disease is crucial for both human health and conservation. Human‐induced climate change has been hypothesized to alter the geographic distribution of malaria pathogens. As the earth warms, arthropod vectors may display a general range expansion or may enjoy longer breeding season, both of which can enhance parasite transmission. Moreover, Plasmodium species may directly benefit for elevating temperatures, which provide stimulating conditions for parasite reproduction. To test for the link between climate change and malaria prevalence on a global scale for the first time, I used long‐term records on avian malaria, which is a key model for studying the dynamics of naturally occurring malarial infections. Following the variation in parasite prevalence in more than 3000 bird species over seven decades, I show that the infection rate by Plasmodium is strongly associated with temperature anomalies and has been augmented with accelerating tendency during the last 20 years. The impact of climate change on malaria prevalence varies across continents, with the strongest effects found for Europe and Africa. Migration habit did not predict susceptibility to the escalating parasite pressure by Plasmodium. Consequently, wild birds are at an increasing risk of malaria infection due to recent climate change, which can endanger both naïve bird populations and domesticated animals. The prevailing avian example may provide useful lessons for understanding the effect of climate change on malaria in humans.     

Climate change can alter competitive relationships between resident and migratory birds

Climate change could affect resource competition between resident and migratory bird species by changing the interval between their onsets of breeding or by altering their population densities. We studied interspecific nest-hole competition between resident great tits and migrant pied flycatchers in South-Western Finland over the past five decades (1953-2005). We found that appearance of fatal take-over trials, the cases where a pied flycatcher tried to take over a great tit nest but was killed by the tit, increased with a reduced interspecific laying date interval and with increasing densities of both tits and flycatchers. The probability of pied flycatchers taking over great tit nests increased with the density of pied flycatchers. Laying dates of the great tit and pied flycatcher are affected by the temperatures of different time periods, and divergent changes in these temperatures could consequently modify their competitive interactions. Densities are a result of reproductive success and survival, which can be affected by separate climatic factors in the resident great tit and trans-Saharan migrant pied flycatcher. On these bases we conclude that climate change has a great potential to alter the competitive balance between these two species.     

Age and size of European saltmarshes and the population genetic consequences for ground beetles

Enzyme and dispersal polymorphisms of the saltmarsh carabid beetles Pogonus chalceus and Dicheirotrichus gustavii were studied in European populations varying in size and in isolation in space and time. D. gustavii, a constantly fullwinged species, has a larger genetic diversity and a smaller genetic differentiation between populations than the wing-polymorphic P. chalceus. Clear relationships between population or site characteristics and genetic structure were not observed, except for the special position taken by some small populations in both species. The dispersal power of P. chalceus in small populations is larger than in large populations, suggesting that these populations are unstable and/or young. Small populations, however, do not always show a lower genetic diversity than large populations, as would be expected from genetic drift. Dispersal power in P. chalceus declines with increasing age of the saltmarsh, probably due to continuous emigration of winged individuals. Age and size of saltmarshes, although difficult to study independently, both appear to be important in determining the genetic structure of saltmarsh beetles. Maximum diversity in both parameters is therefore recommended as an optimal nature conservation strategy. Received: 22 September 1997 / Accepted: 29 December 1997     

Climate change affects populations of northern birds in boreal protected areas

Human land-use effects on species populations are minimized in protected areas and population changes can thus be more directly linked with changes in climate. In this study, bird population changes in 96 protected areas in Finland were compared using quantitative bird census data, between two time slices, 1981-1999 and 2000-2009, with the mean time span being 14 years. Bird species were categorized by distribution pattern and migratory strategy. Our results showed that northern bird species had declined by 21 per cent and southern species increased by 29 per cent in boreal protected areas during the study period, alongside a clear rise (0.7-0.8 °C) in mean temperatures. Distribution pattern was the main factor, with migratory strategy interacting in explaining population changes in boreal birds. Migration strategy interacted with distribution pattern so that, among northern birds, densities of both migratory and resident species declined, whereas among southern birds they both increased. The observed decline of northern species and increase in southern species are in line with the predictions of range shifts of these species groups under a warming climate, and suggest that the population dynamics of birds are already changing in natural boreal habitats in association with changing climate.     

Predicting landscape-genetic consequences of habitat loss, fragmentation and mobility for multiple species of woodland birds

Inference concerning the impact of habitat fragmentation on dispersal and gene flow is a key theme in landscape genetics. Recently, the ability of established approaches to identify reliably the differential effects of landscape structure (e.g. land-cover composition, remnant vegetation configuration and extent) on the mobility of organisms has been questioned. More explicit methods of predicting and testing for such effects must move beyond post hoc explanations for single landscapes and species. Here, we document a process for making a priori predictions, using existing spatial and ecological data and expert opinion, of the effects of landscape structure on genetic structure of multiple species across replicated landscape blocks. We compare the results of two common methods for estimating the influence of landscape structure on effective distance: least-cost path analysis and isolation-by-resistance. We present a series of alternative models of genetic connectivity in the study area, represented by different landscape resistance surfaces for calculating effective distance, and identify appropriate null models. The process is applied to ten species of sympatric woodland-dependant birds. For each species, we rank a priori the expectation of fit of genetic response to the models according to the expected response of birds to loss of structural connectivity and landscape-scale tree-cover. These rankings (our hypotheses) are presented for testing with empirical genetic data in a subsequent contribution. We propose that this replicated landscape, multi-species approach offers a robust method for identifying the likely effects of landscape fragmentation on dispersal.     

Species loss and gain in communities under future climate change: consequences for functional diversity

It is anticipated that anthropogenic climate change will lead to substantial reassembly within communities in coming decades as individual species shift their ranges to track optimal conditions for growth and survival. As species are lost and gained in communities, what are the consequences for functional trait diversity? Functional traits are the characteristics of species that affect individual performance and provide the vital link between biodiversity at the species level and ecosystem function. We investigated how projected changes in species richness in plant communities under climate change scenarios for the decade 2050 will affect the distribution and diversity of five functional traits. We aggregated range change projections made in Maxent for the decade 2050 across all species in the regional pool of littoral rainforest vines in eastern Australia (n = 163 species). The effect of richness changes on trait diversity was assessed in nine rainforest reserves along the east coast of Australia. Although richness was predicted to significantly decline across all communities, functional diversity remained stable, indicating a decoupling in response to climate change at these two different levels of biological organization. A high degree of redundancy in trait composition in communities may buffer against the loss of function in these plant communities. Scaling‐up our understanding of the impact of climate change from the species level to communities is a critical step towards developing conservation strategies aimed at preserving ecosystem function.     

Projected climate change effects on Rocky Mountain and Great Plains birds: generalities of biodiversity consequences

Climate change effects on biodiversity are already manifested, and yet no predictive knowledge characterizes the likely nature of these effects. Previous studies suggested an influence of topography on these effects, a possibility tested herein. Bird species with distributions restricted to montane (26 species) and Great Plains (19 species) regions of central and western North America were modeled, and climate change effects on their distributions compared: in general, plains species were more heavily influenced by climate change, with drastic area reductions (mode 35% of distributional area lost under assumption of no dispersal) and dramatic spatial movements (0–400 km shift of range centroid under assumption of no dispersal) of appropriate habitats. These results suggest an important generality regarding climate change effects on biodiversity, and provide useful guidelines for conservation planning.     

Climate envelope, life history traits and the resilience of birds facing global change

Few studies have examined how life history traits and the climate envelope influence the ability of species to respond to climate change and habitat degradation. In this study, we test whether 18 species-specific variables, related to the climate envelope, ecological envelope and life history, could predict recent population trends (over 17 years) of 71 common breeding bird species in France. Habitat specialists were declining at a much higher rate than generalists, a sign that habitat quality is decreasing globally. The lower the thermal maximum (temperature at the hot edge of the climate envelope), the more negative are the population trends and the less tolerant these species are climate warming, regardless of the thermal range over which these species occur. The life history trait 'the number of broods per year' was positively related to recent trends, suggesting that single-brooded species might be more sensitive to advances in food peak due to climate change, as it increases the risk of mistiming their single-breeding event. Annual fecundity explained long-term declines, as it is a good proxy for most other demographic rates, with shorter-lived species being more sensitive to global change: individuals of species with higher fecundity might have too short a life to learn to adapt to directional changes in their environment. Finally, there was evidence that natal dispersal was a predictor of recent trends, with species with high natal dispersal experiencing smaller population declines than species with low natal dispersal. This is expected if the higher the natal dispersal, the larger the ability to shift spatially when facing changes in local habitat or climate, in order to track optimal conditions and adapt to global change. Identifying decline-promoting factors allow us to infer mechanisms responsible for observed declines in wild bird populations facing global change, and by doing so allow for a more pre-emptive approach to conservation planning.     

Climate change causes rapid changes in the distribution and site abundance of birds in winter

Detecting coherent signals of climate change is best achieved by conducting expansive, long‐term studies. Here, using counts of waders (Charadrii) collected from ca. 3500 sites over 30 years and covering a major portion of western Europe, we present the largest‐scale study to show that faunal abundance is influenced by climate in winter. We demonstrate that the ‘weighted centroids’ of populations of seven species of wader occurring in internationally important numbers have undergone substantial shifts of up to 115 km, generally in a northeasterly direction. To our knowledge, this shift is greater than that recorded in any other study, but closer to what would be expected as a result of the spatial distribution of ecological zones. We establish that year‐to‐year changes in site abundance have been positively correlated with concurrent changes in temperature, but that this relationship is most marked towards the colder extremities of the birds' range, suggesting that shifts have occurred as a result of range expansion and that responses to climate change are temperature dependent. Many attempts to model the future impacts of climate change on the distribution of organisms, assume uniform responses or shifts throughout a species' range or with temperature, but our results suggest that this may not be a valid approach. We propose that, with warming temperatures, hitherto unsuitable sites in northeastern Europe will host increasingly important wader numbers, but that this may not be matched by declines elsewhere within the study area. The need to establish that such changes are occurring is accentuated by the statutory importance of this taxon in the designation of protected areas.     

Climate change and human colonization triggered habitat loss and fragmentation in Madagascar

The relative effect of past climate fluctuations and anthropogenic activities on current biome distribution is subject to increasing attention, notably in biodiversity hot spots. In Madagascar, where humans arrived in the last ~4 to 5,000 years, the exact causes of the demise of large vertebrates that cohabited with humans are yet unclear. The prevailing narrative holds that Madagascar was covered with forest before human arrival and that the expansion of grasslands was the result of human‐driven deforestation. However, recent studies have shown that vegetation and fauna structure substantially fluctuated during the Holocene. Here, we study the Holocene history of habitat fragmentation in the north of Madagascar using a population genetics approach. To do so, we infer the demographic history of two northern Madagascar neighbouring, congeneric and critically endangered forest dwelling lemur species—Propithecus tattersalli and Propithecus perrieri—using population genetic analyses. Our results highlight the necessity to consider population structure and changes in connectivity in demographic history inferences. We show that both species underwent demographic fluctuations which most likely occurred after the mid‐Holocene transition. While mid‐Holocene climate change probably triggered major demographic changes in the two lemur species range and connectivity, human settlements that expanded over the last four millennia in northern Madagascar likely played a role in the loss and fragmentation of the forest cover.     

Climate change is creating a mismatch between protected areas and suitable habitats for frogs and birds in Puerto Rico

Climate change is altering the spatial distribution of many species around the world. In response, we need to identify and protect suitable areas for a large proportion of the fauna so that they persist through time. This exercise must also evaluate the ability of existing protected areas to provide safe havens for species in the context of climate change. Here, we combined passive acoustic monitoring, semi-automatic species identification models, and species distribution models of 21 bird and frog species based on past (1980–1989), present (2005–2014), and future (2040–2060) climate scenarios to determine how species distributions relate to the current distribution of protected areas in Puerto Rico. Species detection/non-detection data were acquired across?~?700 sampling sites. We developed always-suitable maps that characterized suitable habitats in all three time periods for each species and overlaid these maps to identify regions with high species co-occurrence. These distributions were then compared with the distribution of existing protected areas. We show that Puerto Rico is projected to become dryer by 2040–2060, and precipitation in the warmest quarter was among the most important variables affecting bird and frog distributions. A large portion of always-suitable areas (ASA) is outside of protected areas (>?80%), and the percent of protected areas that overlaps with always-suitable areas is larger for bird (75%) than frog (39%) species. Our results indicate that present protected areas will not suffice to safeguard bird and frog species under climate change; however, the establishment of larger protected areas, buffer zones, and connectivity between protected areas may allow species to find suitable niches to withstand environmental changes.