Macroecology and extinction risk correlates of frogs

Aim  Our aim was to test whether extinction risk of frog species could be predicted from their body size, fecundity or geographical range size. Because small geographical range size is a correlate of extinction risk in many taxa, we also tested hypotheses about correlates of range size in frogs.
Location  Global.
Methods  Using a large comparative data set ( n  = 527 species) compiled from the literature, we performed bivariate and multiple regressions through the origin of independent contrasts to test proposed macroecological patterns and correlates of extinction risk in frogs. We also created minimum adequate models to predict snout–vent length, clutch size, geographical range size and IUCN Red List status in frogs. Parallel non-phylogenetic analyses were also conducted. We verified the results of the phylogenetic analyses using gridded data accounting for spatial autocorrelation.
Results  The most threatened frog species tend to have small geographical ranges, although the relationship between range and extinction risk is not linear. In addition, tropical frogs with small clutches have the smallest ranges. Clutch size was strongly positively correlated with geographical range size ( r ² = 0.22) and body size ( r ² = 0.28).
Main conclusions  Our results suggest that body size and fecundity only affect extinction risk indirectly through their effect on geographical range size. Thus, although large frogs with small clutches tend to be endangered, there is no comparative evidence that this relationship is direct. If correct, this inference has consequences for conservation strategy: it would be inefficient to allocate conservation resources on the basis of low fecundity or large body size; instead it would be better to protect areas that contain many frog species with small geographical ranges.     

Diversity,extinction, and threat status in Lagomorphs

A quarter of all lagomorphs (pikas, rabbits, hares and jackrabbits) are threatened with extinction, including several genera that contain only one species. The number of species in a genus correlates with extinction risk in lagomorphs, but not in other mammal groups, and this is concerning because the non‐random extinction of small clades disproportionately threatens genetic diversity and phylogenetic history. Here, we use phylogenetic analyses to explore the properties of the lagomorph phylogeny and test if variation in evolution, biogeography and ecology between taxa explains current patterns of diversity and extinction risk. Threat status was not related to body size (and, by inference, its biological correlates), and there was no phylogenetic signal in extinction risk. We show that the lagomorph phylogeny has a similar clade‐size distribution to other mammals, and found that genus size was unrelated to present climate, topography, or geographic range size. Extinction risk was greater in areas of higher human population density and negatively correlated with anthropogenically modified habitat. Consistent with this, habitat generalists were less likely to be threatened. Our models did not predict threat status accurately for taxa that experience region‐specific threats. We suggest that pressure from human populations is so severe and widespread that it overrides ecological, biological, and geographic variation in extant lagomorphs.     

Predicting extinction risk in declining species

What biological attributes predispose species to the risk of extinction? There are many hypotheses but so far there has been no systematic analysis for discriminating between them. Using complete phylogenies of contemporary carnivores and primates, we present, to our knowledge, the first comparative test showing that high trophic level, low population density slow life history and, in particular, small geographical range size are all significantly and independently associated with a high extinction risk in declining species. These traits together explain nearly 50% of the total between-species variation in extinction risk. Much of the remaining variation can be accounted for by external anthropogenic factors that affect species irrespective of their biology.     

Determining variation in the success of New Zealand land birds

The New Zealand avifauna has changed considerably since the arrival of humans approximately 1000 years ago. The extinction of native species followed by their replacement with introduced exotic species is recognized as having greatly reduced the distinctiveness of the avifauna. Using phylogenetically independent methods, I tested whether life history and ecological correlates of extinction, introduction and persistence were significant across species and within taxa. Extinct land bird species were, on average, and independent of phylogenetic relatedness, larger-bodied than extant species. Introduction success was found to be significantly associated with increasing species body size, generation time and indices of human effort. In contrast, the geographical distribution of extant land birds was correlated with traits associated with high population growth rates (i.e. small, rapidly developing species, with high fecundity). My results suggest that selection pressures have changed significantly since the arrival of human colonists in New Zealand and that these changes have favoured different types of land birds, to the general detriment of endemic species. Comparative methods that explicitly examine historical changes are necessary to elucidate the changing roles of biological and extrinsic traits in the differential success of species in any assemblage.     

Minimum viable population sizes and global extinction risk are unrelated

Theoretical and empirical work has shown that once reduced in size and geographical range, species face a considerably elevated risk of extinction. We predict minimum viable population sizes (MVP) for 1198 species based on long-term time-series data and model-averaged population dynamics simulations. The median MVP estimate was 1377 individuals (90% probability of persistence over 100 years) but the overall distribution was wide and strongly positively skewed. Factors commonly cited as correlating with extinction risk failed to predict MVP but were able to predict successfully the probability of World Conservation Union Listing. MVPs were most strongly related to local environmental variation rather than a species' intrinsic ecological and life history attributes. Further, the large variation in MVP across species is unrelated to (or at least dwarfed by) the anthropogenic threats that drive the global biodiversity crisis by causing once-abundant species to decline.     

Geographic range size, life history and rates of diversification in Australian mammals

Abstract What causes species richness to vary among different groups of organisms? Two hypotheses are that large geographical ranges and fast life history either reduce extinction rates or raise speciation rates, elevating a clade's rate of diversification. Here we present a comparative analysis of these hypotheses using data on the phylogenetic relationships, geographical ranges and life history of the terrestrial mammal fauna of Australia. By comparing species richness patterns to null models, we show that species are distributed nonrandomly among genera. Using sister‐clade comparisons to control for clade age, we then find that faster diversification is significantly associated with larger geographical ranges and larger litters, but there is no evidence for an effect of body size or age at first breeding on diversification rates. We believe the most likely explanation for these patterns is that larger litters and geographical ranges increase diversification rates because they buffer species from extinction. We also discuss the possibility that positive effects of litter size and range size on diversification rates result from elevated speciation rates.     

Macroevolution of hoverflies (Diptera: Syrphidae): the effect of using higher‐level taxa in studies of biodiversity,and correlates of species richness

We test a near‐complete genus level phylogeny of hoverflies (Diptera: Syrphidae) for consistency with a null model of clade growth having uniform probabilities of speciation and extinction among contemporaneous species. The phylogeny is too unbalanced for this null model. Importantly, the degree of imbalance in the phylogeny depends on whether the phylogeny is analysed at the genus level or species level, suggesting that genera ought not to be used uncritically as surrogates for species in large‐scale evolutionary analyses. Tests for a range of morphological, life‐history and ecological correlates of diversity give equivocal results, but suggest that high species‐richness may be associated with sexual selection and diet breadth. We find no correlation between species‐richness and either body size or reproductive rate.     

Relationships between body size and geographical range size among Australian mammals: has human impact distorted macroecological patterns?

Extinction and artificial reduction in the size of geographical ranges of many species have occurred extensively across the globe because of human activities. In particular, Australian mammals have suffered heavily in the last two hundred years, with the highest number of reported cases of mammal extinctions anywhere. In the present study, we investigated the extent to which human impact has affected contemporary macroecological patterns in Australian terrestrial mammals. After examining patterns relating to body size and range size among the contemporary mammal fauna, we removed the effects of the last two hundred years of human impact by exploring patterns in the pre‐European assemblage. This permitted us to determine whether contemporary macroecological patterns are distortions of pre‐European patterns. In contrast to the expected pattern of a significant positive relationship between body size and range size, our results showed no significant association for the complete fauna in both cross‐species and phylogenetic analyses, even when data were corrected for species extinctions and range reductions. Analyses within families and among species with the same dietary strategy revealed three significant positive relationships (Macropodidae, Peramelidae, and herbivores) and one significant negative relationship (insectivores) within the contemporary assemblage that disappeared when the pre‐European assemblage was analysed. A positive relationship also emerged in the pre‐European assemblage for the Vombatidae that was not apparent in the contemporary fauna. Thus, correcting for human impact revealed important distortions among contemporary macroecological relationships that have been brought about by human‐induced range reduction and extinction. These findings not only provide further evidence that the Australian continent presents a unique and valuable opportunity with which to test the generality of macroecological patterns, but they also have important ramifications for the analysis and interpretation of contemporary macroecological datasets.     

Pattern and process in the geographical ranges of freshwater fishes

North American freshwater fishes were studied to determine whether they displayed the same relationships between log (geographical range size) and log (body size) and the same pattern of range shape as found among North American birds and mammals. The forces that produce these patterns were also investigated. The log (geographical range size) : log (body size) relationship was analysed for 121 North American freshwater fish species. Thirty‐two imperilled species were compared with 89 non‐imperilled species to determine if the overall relationship could result from differential extinction. Range geometries were analysed, within and among habitat guilds, to determine if general patterns could be detected. The log (geographical range size) : log (body size) pattern among freshwater fish species was triangular and qualitatively similar to that found for North American birds and mammals. The results suggest that below a minimum geographical range, the likelihood of extinction increases dramatically for freshwater fishes and that this minimum range size increases with body size. The pattern of fish species’ range shapes differs from that found for other North American vertebrate taxa because, on average, fish possess much smaller ranges than terrestrial species and most fish species’ geographical ranges extend further on a north–south axis than on an east–west axis. The log (geographical range size) : log (body size) pattern reveals that fish species’ geographical ranges are more constrained than those of terrestrial species. The triangular relationship may be caused by differential extinction of species with large bodies and small geographical ranges as well as higher speciation rates of small‐bodied fish. The restricted geographical ranges of freshwater fishes gives them much in common with terrestrial species on oceanic islands. Range shape patterns within habitat guilds reflect guild‐specific historical and current ecological forces. The overall pattern of range shapes emerges from the combination of ecologically different subunits.     

Using functional traits and phylogeny to understand local extinction risk in dragonflies and damselflies (Odonata)

Understanding the risk of local extinction of a species is vital in conservation biology, especially now when anthropogenic disturbances and global warming are severely changing natural habitats. Local extinction risk depends on species traits, such as its geographical range size, fresh body mass, dispersal ability, length of flying period, life history variation, and how specialized it is regarding its breeding habitat. We used a phylogenetic approach because closely related species are not independent observations in the statistical tests. Our field data contained the local extinction risk of 31 odonate (dragonflies and damselflies) species from Central Finland. Species relatedness (i.e., phylogenetic signal) did not affect local extinction risk, length of flying period, nor the geographical range size of a species. However, we found that closely related species were similar in hind wing length, length of larval period, and habitat of larvae. Both phylogenetically corrected (PGLS) and uncorrected (GLM) analysis indicated that the geographical range size of species was negatively related to local extinction risk. Contrary to expectations, habitat specialist species did not have higher local extinction rates than habitat generalist species nor was it affected by the relatedness of species. As predicted, species’ long larval period increased, and long wings decreased the local extinction risk when evolutionary relatedness was controlled. Our results suggest that a relatively narrow geographical range size is an accurate estimate for a local extinction risk of an odonate species, but the species with long life history and large habitat niche width of adults increased local extinction risk. Because the results were so similar between PGLS and GLM methods, it seems that using a phylogenetic approach does not improve predicting local extinctions.     

Species richness among birds: body size,life history,sexual selection or ecology?

Why do some avian families contain so many more species than other families? We use comparisons between sister taxa to test predictions arising from six explanations to this puzzle: that differences between families are due to chance, body size, life history, sexual selection, intrinsic ecological factors or extrinsic abiotic factors, respectively. In agreement with previous analyses, we find no support for the idea that differences in species richness are simply due to chance. However, contrary to most previous work, we also find no support for the hypotheses that high species richness is correlated with small body size and fast life history. Rather, high species diversity is strongly associated with pronounced plumage dichromatism, generalist feeding habits and good dispersal capabilities as well as large and fragmented geographical ranges. In addition, all of these relationships are robust to the removal of the two most speciose avian lineages, the Ciconiiformes and the Passeriformes. The supposed relationships between species richness and both body size and life history are, however, due to phylogenetic non-independence. Together with previous work showing that differences between avian lineages in extinction risk are associated with variation in body size and life history, these results indicate that extinction rates and speciation rates are not necessarily determined by the same factors. Hence, high extinction rates are not inevitably associated with low speciation rates. Extinction-prone lineages may, in fact, have a high rate of speciation. In such lineages a high proportion of ''vulnerable'' species would be a natural, ongoing phenomenon.     

Geographical distribution and extinction risk: lessons from Triassic–Jurassic marine benthic organisms

Aim To evaluate the influence of geographical distribution on the extinction risk of benthic marine invertebrates using data from the fossil record, both during times of background extinction and across a mass‐extinction episode. Total geographical range is contrasted with proxies of global abundance to assess the relationships between the two essential components of geographical distribution and extinction risk. Location A global occurrence data base of fossil benthic macro‐organisms from the Triassic and Jurassic periods was used for this study. Methods Geographical distributions and biodiversity dynamics were assessed for each genus (all taxa) or species (bivalves) based on a sample‐standardized data set and palaeogeographical reconstructions. Geographical ranges were measured by the maximum great circle distance of a taxon within a stratigraphic interval. Global abundance was assessed by the number of localities at which a taxon was recorded. Widespread and rare taxa were separated using median and percentile values of the frequency distributions of occurrences. Results The frequency distribution of geographical ranges is very similar to that for modern taxa. Although no significant correlation could be established between local abundance and geographical range, proxies of global abundance are strongly correlated with geographical range. Taxon longevities are correlated with both mean geographical range and mean global abundance, but range size appears to be more critical than abundance in determining extinction risk. These results are valid when geographical distribution is treated as a trait of taxa and when assessed for individual geological stages. Main conclusions Geographical distribution is a key predictor of extinction risk of Triassic and Jurassic benthic marine invertebrates. An important exception is in the end‐Triassic mass extinction, which equally affected geographically restricted and widespread genera, as well as common and rare genera. This suggests that global diversity crises may curtail the role of geographical distribution in determining extinction risk.     

Unravelling the structure of species extinction risk for predictive conservation science

Extinction risk varies across species and space owing to the combined and interactive effects of ecology/life history and geography. For predictive conservation science to be effective, large datasets and integrative models that quantify the relative importance of potential factors and separate rapidly changing from relatively static threat drivers are urgently required. Here, we integrate and map in space the relative and joint effects of key correlates of The International Union for Conservation of Nature-assessed extinction risk for 8700 living birds. Extinction risk varies significantly with species' broad-scale environmental niche, geographical range size, and life-history and ecological traits such as body size, developmental mode, primary diet and foraging height. Even at this broad scale, simple quantifications of past human encroachment across species' ranges emerge as key in predicting extinction risk, supporting the use of land-cover change projections for estimating future threat in an integrative setting. A final joint model explains much of the interspecific variation in extinction risk and provides a remarkably strong prediction of its observed global geography. Our approach unravels the species-level structure underlying geographical gradients in extinction risk and offers a means of disentangling static from changing components of current and future threat. This reconciliation of intrinsic and extrinsic, and of past and future extinction risk factors may offer a critical step towards a more continuous, forward-looking assessment of species' threat status based on geographically explicit environmental change projections, potentially advancing global predictive conservation science.     

Can intrinsic factors explain population declines in North American breeding shorebirds? A comparative analysis

Many shorebirds that breed in North America are declining. These trends reflect global patterns in shorebird populations. Here we ask what factors make some shorebird species more prone to decline than others. Specifically, we test the influence of migratory behaviour (route and distance), biogeography (population size and range), life history (body size, clutch size) and sexual selection (social mating system and testis size) on population trends in North American breeding shorebirds. Using phylogenetic comparative methods, we show that species that migrate across continental North America are more prone to decline than species that do not. Our finding that continental migrants are associated with population decline indicates that intrinsic factors may play an important role in predisposing a species to decline. Previous studies within the class Aves have failed to identify migration route as a correlate of decline or extinction risk. Two other intrinsic factors (oceanic migrants and threats on the non-breeding grounds) were also important in our overall models, although neither was significant alone. The moderate explanatory power of our variables indicates that other factors are also important for explaining shorebird declines. We suggest that contemporary threats, most notably habitat loss and degradation at migratory stopover sites, are likely to be important.     

Global gradients of avian longevity support the classic evolutionary theory of ageing

Senescence, the process of physiological deterioration associated with growing old, is a shared characteristic of a wide range of animals. Yet, lifespan varies dramatically among species. To explain this variation, the evolutionary theory of ageing has been proposed more than 50 yr ago. Although the theory has been tested experimentally and through comparative analyses, there remains debate whether its fundamental prediction is empirically supported. Here, we use a comprehensive database on avian life history traits to test the evolutionary theory of ageing at a global scale. We show that pronounced geographical gradients of maximum longevity exist, that they are predicted by measures of predator diversity and only partly depend on correlated life‐history traits. The results are consistent with species‐level analyses and can be replicated across bio‐geographical regions. Our analyses suggest that stochastic predation is an important driver of the evolution of lifespan, at least in birds.     

BIODIVERSITY RESEARCH: Nestedness for different reasons: the distributions of birds,lizards and small mammals on islands of an inundated lake

Aim We examined whether the community compositions of birds, lizards and small mammals were nested in a fragmented landscape in the Thousand Island Lake, China. We also assessed whether the mechanisms influencing nestedness differed among these taxonomic groups. Location Thousand Island Lake, China. Methods Presence/absence matrices were compiled for birds (42 islands) and lizards (42 islands) using line‐transect methods, and for small mammals (14 islands) using live‐trapping methods from 2006 to 2009. Nestedness was analysed using BINMATNEST, and statistical significance was assessed using the conservative null model 3. We used Spearman rank correlations and partial Spearman rank correlations to examine associations of nestedness and habitat variables (area, isolation, habitat diversity and plant richness) as well as life‐history traits (body size, habitat specificity, geographical range size and area requirement) related to species extinction and immigration tendencies. Results The community compositions of birds, lizards and small mammals were all significantly nested, but the causal factors underlying nestedness differed among taxonomic groups. For birds, island area, habitat specificity and area requirement were significantly correlated with nestedness after controlling for other independent variables. For lizards, habitat heterogeneity was the single best correlate of nestedness. For small mammals, island area, habitat heterogeneity and habitat specificity were significantly correlated with nestedness. The nested patterns of birds, lizards and small mammals were not attributable to passive sampling or selective colonization. Main conclusions The processes influencing nested patterns differed among taxonomic groups. Nestedness of bird assemblages was driven by selective extinction, and lizard assemblage was caused by habitat nestedness, while nestedness of small mammals resulted from both selective extinction and habitat nestedness. Therefore, we should take taxonomic differences into account when analysing nestedness to develop conservation guidelines and refrain from using single taxa as surrogates for others.     

Reductions in connectivity and habitat quality drive local extinctions in a plant diversity hotspot

It is well documented that habitat loss is a major cause of biodiversity decline. However, the roles of the different aspects of habitat loss in local extinctions are less understood. Anthropogenic destruction of an area of habitat causes immediate local extinction but subsequently three additional gradual drivers influence the likelihood of delayed extinction: decreased habitat patch size, lower connectivity and habitat deterioration. We investigated the role of these drivers in local extinctions of 82 declining species in a UK biodiversity hotspot. We combined a unique set of ≈ 7000 vegetation surveys and habitat maps from the 1930s with contemporary species’ occurrences. We extrapolated from these surveys to the whole 2500‐km² study area using habitat suitability surfaces. The strengths of drivers in explaining local extinctions over this 70 yr period were determined by contrasting connectivity, patch size and habitat quality loss for locations at which a species went extinct and those with persisting occurrences. Species’ occurrences declined on average by 60%, with half of local extinctions attributable to immediate habitat loss and half to the gradual processes causing delayed extinctions. On average, locations where a species persisted had a 73% higher contemporary connectivity than those suffering extinctions, but showed no differences in historical connectivity. Furthermore, locations with extinctions experienced a 37% greater decline in suitability associated with changes in habitat type. The strength of the drivers and the proportion of extinctions depended on the species’ habitat specialism, but were affected only minimally by life‐history characteristics. In conclusion, we identified a hierarchy of drivers influencing local extinction: with connectivity loss being the strongest, suitability change being moderately important, but changes in habitat patch size having only weak effects. We suggest conservation efforts could be most effective by strengthening connectivity along with reducing habitat deterioration, which would benefit a wide range of species.     

Drivers and interrelationships among multiple dimensions of rarity for freshwater fishes

Species can be rare or common in three different dimensions: geographic range size, habitat breadth, and local abundance. Understanding drivers of rarity are not only fundamentally interesting; it is also pertinent for their conservation. We addressed this challenge by analyzing the rarity of 291 native freshwater fishes occurring in ca 3500 independent stream reaches that span a broad environmental gradient across continental USA. Using phylogenetic regression and path analysis, we examined the concordance among the three rarity dimensions, and identified possible mechanisms by which species life‐history, habitat affinities, and biogeography drive variation in rarity. Weak double extinction jeopardies were driven by weakly positive correlations between habitat breadth and local abundance, and between habitat breadth and geographic range size. However, a triple extinction jeopardy was averted as local abundance and range size were not positively linked in our study. This is because large‐river and lacustrine habitat use mediated a trade‐off between local abundance and range size. Large rivers and lacustrine habitats represent important dispersal pathways and refugia that enabled fishes to acquire wide ranges; however, species using these habitats are less abundant overall because they are less adapted to small lotic channels, which comprise the majority of stream habitats in the US. Life‐history traits were key in governing the relationship between abundance and range size as large‐river and lacustrine habitat use were driven by body size, egg size, and parental care. Our analysis contributes novel insights into mechanisms that underlie multiple dimensions of rarity in freshwater fish and informs the prioritization of multiply rare species for conservation.     

Amphibians live longer at higher altitudes but not at higher latitudes

Why and how organisms differ in life‐history strategies across their range is a long‐standing topic of interest to evolutionary ecologists. Although many studies have addressed this issue for several life‐history traits, such as body size and clutch size, very few have been made for some others traits, including longevity. In the present study, we performed a comparative study aiming to develop general patterns of geographical variation in longevity of urodele and anuran amphibians using published information on demographic age derived from skeletochronology. We conducted within‐species meta‐analyses using datasets of two (ten urodele and 12 anuran species) and multiple (two urodele and nine anuran species) spatially‐separated populations and found that maturation, mean, and maximum age all increased with altitude but not with latitude in each sex of both amphibian groups. This geographical pattern held true across 33 urodele and 86 anuran species at common body sizes, independent of phylogeny. It is likely that metabolic rate, reproductive investment, and mortality risk, which are the key factors that affect longevity as suggested by ageing theory, vary systemically along altitudinal gradients but not along latitudinal gradients. The evolutionary causes behind these puzzling patterns deserve further investigation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 623–632.