Which function describes the species–area relationship best? A review and empirical evaluation

Aim The aims of this study are to resolve terminological confusion around different types of species–area relationships (SARs) and their delimitation from species sampling relationships (SSRs), to provide a comprehensive overview of models and analytical methods for SARs, to evaluate these theoretically and empirically, and to suggest a more consistent approach for the treatment of species–area data. Location Curonian Spit in north‐west Russia and archipelagos world‐wide. Methods First, I review various typologies for SARs and SSRs as well as mathematical models, fitting procedures and goodness‐of‐fit measures applied to SARs. This results in a list of 23 function types, which are applicable both for untransformed (S) and for log‐transformed (log S) species richness. Then, example data sets for nested plots in continuous vegetation (n = 14) and islands (n = 6) are fitted to a selection of 12 function types (linear, power, logarithmic, saturation, sigmoid) both for S and for log S. The suitability of these models is assessed with Akaike’s information criterion for S and log S, and with a newly proposed metric that addresses extrapolation capability. Results SARs, which provide species numbers for different areas and have no upper asymptote, must be distinguished from SSRs, which approach the species richness of one single area asymptotically. Among SARs, nested plots in continuous ecosystems, non‐nested plots in continuous ecosystems, and isolates can be distinguished. For the SARs of the empirical data sets, the normal and quadratic power functions as well as two of the sigmoid functions (Lomolino, cumulative beta‐P) generally performed well. The normal power function (fitted for S) was particularly suitable for predicting richness values over ten‐fold increases in area. Linear, logarithmic, convex saturation and logistic functions generally were inappropriate. However, the two sigmoid models produced unstable results with arbitrary parameter estimates, and the quadratic power function resulted in decreasing richness values for large areas. Main conclusions Based on theoretical considerations and empirical results, I suggest that the power law should be used to describe and compare any type of SAR while at the same time testing whether the exponent z changes with spatial scale. In addition, one should be aware that power‐law parameters are significantly influenced by methodology.     

Assessments of species’ vulnerability to climate change: from pseudo to science

Climate change vulnerability assessments (CCVAs) are important tools to plan for and mitigate potential impacts of climate change. However, CCVAs often lack scientific rigor, which can ultimately lead to poor conservation prioritization and associated ecological and economic costs. We discuss the need to improve comparability and consistency of CCVAs and either validate their findings or improve assessment of CCVA uncertainty and sensitivity to methodological assumptions.     

Which seed origin provides better tolerance to flooding and drought when restoring to face climate change?

Our goal was to establish the tolerance to flooding and drought of seedlings from a hydric gradient of different seed sources to provide recommendations for forest restoration in the face of climate change. We used Drimys winteri var. chilensis, a tree species that grows from extreme arid zones to wetlands along Chile, as the study subject. We expected that seedlings of xeric origin would perform better in drought conditions than populations from moist environments, and vice versa for flooding tolerance. We collected D. winteri seeds from xeric, mesic and wet environments. Seedlings at two development stages were submitted to an extreme flooding and drought treatment during 2 or 4 months in a common garden. After the flooding and drought assays finished, the number of surviving and damaged seedlings, lenticels and adventitious root presence, height, new leaves and specific leaf area, shoot/root ratio, water potential and/or chlorophyll fluorescence (Fv/Fm), were recorded. We found that flooding and drought affected almost all the parameters studied negatively. The xeric population seedlings, at both development stages studied, were the most tolerant to the drought and, unexpectedly, also to the flooding treatment. We recommend restoring with seedlings of xeric origin especially in arid areas where sudden flooding is frequent, as occurs in the Andes Mountains. In the face of climate change, we recommend carrying out common garden and field studies before advising which population origin should be used for restoration, since they do not always respond in accordance with expected patterns of local adaptation.     

Does daily climate variation have an effect on species’ elevational range size?

In their recent paper published in Science (2016, 351 , 1437–1439), Chan et al. analysed 137 montane gradients, concluding that they found a novel pattern—a negative relationship between mean elevational range size of species and daily temperature variation, which was claimed as empirical evidence for a novel macrophysiological principle (Gilchrist's hypothesis). This intriguing possibility was their key conceptual contribution. Unfortunately, as we show, the empirical evidence was flawed because of errors in the analyses and substantial sampling bias in the data. First, we re‐ran their analyses using their data, finding that their model should have been rejected. Second, we performed two additional re‐analyses of their data, addressing biases and pseudoreplication in different ways, both times again rejecting the evidence claimed to support Gilchrist's hypothesis. These results overturn the key empirical findings of Chan et al.'s study. Therefore, the “macrophysiological principle” should be regarded as currently remaining unsupported by empirical evidence.     

Hosts’ Responses to Parasitic Eggs: Which Cues Elicit Hosts’ Egg Discrimination?

Many hosts of the common cuckoo (Cuculus canorus) exhibit egg recognition, and reject parasitic eggs. How do hosts discriminate cuckoo eggs from their own? Hosts might be able to recognize their own eggs using the specific pigment pattern on the outer eggshell surface, which may serve as a cue for recognition. We tested if patterns of egg pigments (spottedness) contain this information by manipulating spot density of great reed warbler eggs (Acrocephalus arundinaceus). We also manipulated the colour of eggs when the original spot pattern remained the same. Spot density (approximately 15–75%) did not significantly affect rejection rate (8–20% rejection), but when spots fully covered the eggs, i.e. the eggshell was plain dark brown, rejection rate increased abruptly to 100%. A loglinear model revealed the significant influence of colour on rejection rates, although there was no interactive effect between spottedness and colour. Our results strongly support the differential use of egg markers in host’s egg discrimination, suggesting that spot density has limited importance compared to eggshell colour.     

Which practices co‐deliver food security,climate change mitigation and adaptation,and combat land degradation and desertification?

There is a clear need for transformative change in the land management and food production sectors to address the global land challenges of climate change mitigation, climate change adaptation, combatting land degradation and desertification, and delivering food security (referred to hereafter as “land challenges”). We assess the potential for 40 practices to address these land challenges and find that: Nine options deliver medium to large benefits for all four land challenges. A further two options have no global estimates for adaptation, but have medium to large benefits for all other land challenges. Five options have large mitigation potential (>3 Gt CO₂eq/year) without adverse impacts on the other land challenges. Five options have moderate mitigation potential, with no adverse impacts on the other land challenges. Sixteen practices have large adaptation potential (>25 million people benefit), without adverse side effects on other land challenges. Most practices can be applied without competing for available land. However, seven options could result in competition for land. A large number of practices do not require dedicated land, including several land management options, all value chain options, and all risk management options. Four options could greatly increase competition for land if applied at a large scale, though the impact is scale and context specific, highlighting the need for safeguards to ensure that expansion of land for mitigation does not impact natural systems and food security. A number of practices, such as increased food productivity, dietary change and reduced food loss and waste, can reduce demand for land conversion, thereby potentially freeing‐up land and creating opportunities for enhanced implementation of other practices, making them important components of portfolios of practices to address the combined land challenges.     

Altitudinally restricted communities of Schizophoran flies in Queensland’s Wet Tropics: vulnerability to climate change

The Australian Wet Tropics World Heritage Area (WTWHA) contains a number of highland vertebrates predicted to face extinction due to a warming climate, but little is known about risks to invertebrates, which are vital to ecosystem health. This study investigates the distribution and abundance patterns of the Dipteran sub-order Schizophora along an altitudinal transect in the Carbine Uplands of the WTWHA using Malaise traps. The season of peak abundance changed with altitude, with highland abundance peaking in October, and lowland abundance peaking in April. There was a high level of species turnover with altitude, and some evidence for distinct low-, mid-, and high-elevation assemblages, with the high-elevation assemblage containing the most restricted species. We would expect this high-elevation assemblage to be at risk of local extinction with 2–3° of warming, and the mid-elevation assemblage to be at risk with 4–5° warming. Future work should continue sampling to confirm patterns presented here and to monitor range shifts with climate change. A highland species—Helosciomyza ferruginea Hendel is suggested as a good indicator species for such monitoring.     

Mother’s curse and indirect genetic effects: Do males matter to mitochondrial genome evolution?

Maternal inheritance of mitochondrial DNA (mtDNA) was originally thought to prevent any response to selection on male phenotypic variation attributable to mtDNA, resulting in a male‐biased mtDNA mutation load (“mother's curse”). However, the theory underpinning this claim implicitly assumes that a male's mtDNA has no effect on the fitness of females he comes into contact with. If such “mitochondrially encoded indirect genetics effects” (mtIGEs) do in fact exist, and there is relatedness between the mitochondrial genomes of interacting males and females, male mtDNA‐encoded traits can undergo adaptation after all. We tested this possibility using strains of Drosophila melanogaster that differ in their mtDNA. Our experiments indicate that female fitness is influenced by the mtDNA carried by males that the females encounter, which could plausibly allow the mitochondrial genome to evolve via kin selection. We argue that mtIGEs are probably common, and that this might ameliorate or exacerbate mother's curse.     

Connecting soundscape to landscape: Which acoustic index best describes landscape configuration?

Soundscape assessment has been proposed as a remote ecological monitoring tool for measuring biodiversity, but few studies have examined how soundscape patterns vary with landscape configuration and condition. The goal of our study was to examine a suite of published acoustic indices to determine whether they provide comparable results relative to varying levels of landscape fragmentation and ecological condition in nineteen forest sites in eastern Australia. Our comparison of six acoustic indices according to time of day revealed that two indices, the acoustic complexity and the bioacoustic index, presented a similar pattern that was linked to avian song intensity, but was not related to landscape and biodiversity attributes. The diversity indices, acoustic entropy and acoustic diversity, and the normalized difference soundscape index revealed high nighttime sound, as well as a dawn and dusk chorus. These indices appear to be sensitive to nocturnal biodiversity which is abundant at night in warm, subtropical environments. We argue that there is need to better understand temporal partitioning of the soundscape by specific taxonomic groups, and this should involve integrated research on amphibians, insects and birds during a 24 h cycle. The three indices that best connected the soundscape with landscape characteristics, ecological condition and bird species richness were acoustic entropy, acoustic evenness and the normalized difference soundscape index. This study has demonstrated that remote soundscape assessment can be implemented as an ecological monitoring tool in fragmented Australian forest landscapes. However, further investigation should be dedicated to refining and/or combining existing acoustic indices and also to determine if these indices are appropriate in other landscapes and for other survey purposes.     

Multispecies comparisons of adaptability to climate change: A role for life‐history characteristics?

Phenological advancement allows individuals to adapt to climate change by timing life‐history events to the availability of key resources so that individual fitness is maximized. However, different trophic levels may respond to changes in their environment at different rates, potentially leading to a phenological mismatch. This may be especially apparent in the highly seasonal arctic environment that is experiencing the effects of climate change more so than any other region. During a 14‐year study near Utqia?vik (formerly Barrow), Alaska, we estimated phenological advancement in egg laying in relation to snowmelt for eight arctic‐breeding shorebirds and investigated potential linkages to species‐specific life‐history characteristics. We found that snowmelt advanced 0.8 days/year—six times faster than the prior 60‐year period. During this same time, six of the eight species exhibited phenological advancement in laying dates (varying among species from 0.1 to 0.9 days earlier per year), although no species appeared capable of keeping pace with advancing snowmelt. Phenological changes were likely the result of high phenotypic plasticity, as all species investigated in this study showed high interannual variability in lay dates. Commonality among species with similar response rates to timing of snowmelt suggests that nesting later and having an opportunistic settlement strategy may increase the adaptability of some species to changing climate conditions. Other life‐history characteristics, such as migration strategy, previous site experience, and mate fidelity did not influence the ability of individuals to advance laying dates. As a failure to advance egg laying is likely to result in greater phenological mismatch, our study provides an initial assessment of the relative risk of species to long‐term climatic changes.     

Predicting future distributions of mountain plants under climate change: does dispersal capacity matter?

Many studies have investigated the potential impacts of climate change on the distribution of plant species, but few have attempted to constrain projections through plant dispersal limitations. Instead, most studies published so far have simplified dispersal as either unlimited or null. However, depending on the dispersal capacity of a species, landscape fragmentation, and the rate of climatic change, these assumptions can lead to serious over- or underestimation of the future distribution of plant species.
To quantify the discrepancies between simulations accounting for dispersal or not, we carried out projections of future distribution over the 21st century for 287 mountain plant species in a study area of the western Swiss Alps. For each species, simulations were run for four dispersal scenarios (unlimited dispersal, no dispersal, realistic dispersal, and realistic dispersal with long-distance dispersal events) and under four climate change scenarios.
Although simulations accounting for realistic dispersal limitations did significantly differ from those considering dispersal as unlimited or null in terms of projected future distribution, the unlimited dispersal simplification did nevertheless provide good approximations for species extinctions under more moderate climate change scenarios. Overall, simulations accounting for dispersal limitations produced, for our mountainous study area, results that were significantly closer to unlimited dispersal than to no dispersal. Finally, analysis of the temporal pattern of species extinctions over the entire 21st century revealed that important species extinctions for our study area might not occur before the 2080–2100 period, due to the possibility of a large number of species shifting their distribution to higher elevation.     

Detrending climate data prior to climate–growth analyses in dendroecology: A common best practice?

Tree growth varies closely with high–frequency climate variability. Since the 1930s detrending climate data prior to comparing them with tree growth data has been shown to better capture tree growth sensitivity to climate. However, in a context of increasingly pronounced trends in climate, this practice remains surprisingly rare in dendroecology. In a review of Dendrochronologia over the 2018–2021 period, we found that less than 20 % of dendroecological studies detrended climate data prior to climate-growth analyses. With an illustrative study, we want to remind the dendroecology community that such a procedure is still, if not more than ever, rational and relevant. We investigated the effects of detrending climate data on climate–growth relationships across North America over the 1951–2000 period. We used a network of 2536 tree individual ring-width series from the Canadian and Western US forest inventories. We compared correlations between tree growth and seasonal climate data (Tmin, Tmax, Prec) both raw and detrended. Detrending approaches included a linear regression, 30-yr and 100-yr cubic smoothing splines. Our results indicate that on average the detrending of climate data increased climate–growth correlations. In addition, we observed that strong trends in climate data translated to higher variability in inferred correlations based on raw vs. detrended climate data. We provide further evidence that our results hold true for the entire spectrum of dendroecological studies using either mean site chronologies and correlations coefficients, or individual tree time series within a mixed-effects model framework where regression coefficients are used more commonly. We show that even without a change in correlation, regression coefficients can change a lot and we tend to underestimate the true climate impact on growth in case of climate variables containing trends. This study demonstrates that treating climate and tree-ring time series “like-for-like” is a necessary procedure to reduce false negatives and positives in dendroecological studies. Concluding, we recommend using the same detrending for climate and tree growth data when tree-ring time series are detrended with splines or similar frequency-based filters.     

Plant extinction risk under climate change: are forecast range shifts alone a good indicator of species vulnerability to global warming?

Models that couple habitat suitability with demographic processes offer a potentially improved approach for estimating spatial distributional shifts and extinction risk under climate change. Applying such an approach to five species of Australian plants with contrasting demographic traits, we show that: (i) predicted climate‐driven changes in range area are sensitive to the underlying habitat model, regardless of whether demographic traits and their interaction with habitat patch configuration are modeled explicitly; and (ii) caution should be exercised when using predicted changes in total habitat suitability or geographic extent to infer extinction risk, because the relationship between these metrics is often weak. Measures of extinction risk, which quantify threats to population persistence, are particularly sensitive to life‐history traits, such as recruitment response to fire, which explained approximately 60% of the deviance in expected minimum abundance. Dispersal dynamics and habitat patch structure have the strongest influence on the amount of movement of the trailing and leading edge of the range margin, explaining roughly 40% of modeled structural deviance. These results underscore the need to consider direct measures of extinction risk (population declines and other measures of stochastic viability), as well as measures of change in habitat area, when assessing climate change impacts on biodiversity. Furthermore, direct estimation of extinction risk incorporates important demographic and ecosystem processes, which potentially influence species’ vulnerability to extinction due to climate change.     

Nonlinear climate change and Andean feedbacks: an imminent turning point?

A 370 000‐year paleoecological record from Lake Titicaca provides a detailed record of past climate change in which interglacial periods are seen to have some elements of commonality, but also some key differences. We advance a conceptual feedback model to account for the observed changes that includes previously ignored lake effects. Today Lake Titicaca serves to warm the local environment by about 4–5 °C and also to increase rainfall. We observe that as water levels in the lake are drawn down due to warm, dry, interglacial conditions, there is a possible regional cooling as the lake effect on local microclimates diminishes. Positive feedback mechanisms promote drying until much of the lake basin is reduced to salt marsh. Consequently, the usual concept of upslope migration of species with warming would not be applicable in the Altiplano. If, as projected, the next century brings warmer and drier conditions than those of today, a tipping point appears to exist within ca. 1–2 °C of current temperatures, where the relatively benign agricultural conditions of the northern Altiplano would be replaced by inhospitable arid climates. Such a change would have profound implications for the citizens of the Bolivian capital, La Paz.     

Hybridization rate and climate change: are endangered species at risk?

Many species are altering their geographic range due to climate change creating new sympatric populations of otherwise allopatric populations. We investigated whether climate change will affect the distribution and thus the pattern of hybridization between two pairs of closely related damselfly species [Ischnura damula and I. demorsa, and I. denticollis and I. gemina (this, an endangered species)]. Thus, we estimated the strength of pre and postmating reproductive barriers between both pairs of species, and we predicted future potential distribution under four different Global Circulation Models and a realistic emissions scenario of climate change by using maximum entropy modelling technique. Our results showed that reproductive isolation (RI) is complete in I. damula × I. demorsa individuals: F₁ (first generation) hybrids are produced but do not reach sexual maturation. However, RI in I. denticollis × I. gemina hybrids is high but incomplete and unidirectional: only I. gemina females produced F₁ hybrids which mate with males and females of I. denticollis and between them producing BC₁ (backcrosses) and F₂ (second generation) viable hybrids. Maximum entropy models revealed a northern and westward shift and a general reduction of the potential geographic ranges. Based on the pattern of hybridization, for I. damula and I. demorsa there is a current threat as well as a rapid displacement and/or extinction of I. gemina by I. denticollis. However, the current pattern of extinction may not continue due to the contraction in ranges of the four species.     

Tree-ring growth and intra-annual density fluctuations of Pinus pinaster responses to climate: does size matter?

Dendroclimatology generally assumes that climate–growth relationships are age and size independent. However, there is evidence that climate response can be unstable across different age/size classes. In addition, the occurrence of some anatomical features, such as intra-annual density fluctuations (IADFs), is age dependent. The present study investigates whether the climate–growth responses and the occurrence of IADFs in an even-aged stand of Pinus pinaster Ait., growing under Mediterranean climate, are also size-dependent. We randomly selected 60 P. pinaster trees falling within two stem diameter classes: small (<27 cm) and large (>35 cm). Tree rings were crossdated, measured and IADFs identified according to their position within the ring. The residual chronologies of both size classes were strongly correlated, suggesting a common signal. In fact, similar growth–climate relationships were observed in large and small trees. The frequency of IADFs was higher in large than in small trees, suggesting that IADFs were more likely to occur in wider rings of fast-growing trees. In both size classes, most of the IADFs were found in latewood. Latewood IADFs were triggered by the combination of dry June, wet September, and warm December, whereas IADFs located at the end of earlywood were triggered by previous winter precipitation and favorable conditions before summer (high precipitation for large trees and lower temperature for small trees). Our results suggest that IADFs can be a mechanism used at the individual level for adaptation to drought in P. pinaster. The climatic signal of IADFs between earlywood and latewood was mediated by stem size suggesting that future tree-ring studies should include trees stratified by size to better estimate the sensitivity of IADFs to climate.