A conceptual and statistical framework for adaptive radiations with a key role for diversity dependence

Abstract In this article we propose a new framework for studying adaptive radiations in the context of diversity-dependent diversification. Diversity dependence causes diversification to decelerate at the end of an adaptive radiation but also plays a key role in the initial pulse of diversification. In particular, key innovations (which in our definition include novel traits as well as new environments) may cause decoupling of the diversity-dependent dynamics of the innovative clade from the diversity-dependent dynamics of its ancestral clade. We present a likelihood-based inference method to test for decoupling of diversity dependence using molecular phylogenies. The method, which can handle incomplete phylogenies, identifies when the decoupling took place and which diversification parameters are affected. We illustrate our approach by applying it to the molecular phylogeny of the North American clade of the legume tribe Psoraleeae (47 extant species, of which 4 are missing). Two diversification rate shifts were previously identified for this clade; our analysis shows that the first, positive shift can be associated with decoupling of two Pediomelum subgenera from the other Psoraleeae lineages, while we argue that the second, negative shift can be attributed to speciation being protracted. The latter explanation yields nonzero extinction rates, in contrast to previous findings. Our framework offers a new perspective on macroevolution: new environments and novel traits (ecological opportunity) and diversity dependence (ecological limits) cannot be considered separately.     

Climate-influenced boreotropical survival and rampant introgressions explain the thriving of New World grapes in the north temperate zone

The north temperate region was characterized by a warm climate and a rich thermophilic flora before the Eocene, but early diversifications of the temperate biome under global climate change and biome shift remain uncertain. Moreover, it is becoming clear that hybridization/introgression is an important driving force of speciation in plant diversity. Here, we applied analyses from biogeography and phylogenetic networks to account for both introgression and incomplete lineage sorting based on geno...     

Evolutionary history of Gymnocarpos (Caryophyllaceae) in the arid regions from North Africa to Central Asia

Gymnocarpos has only about ten species distributed in the arid regions of Asia and Africa, but it exhibits a geographical disjunction between eastern Central Asia and western North Africa and Minor Asia. We sampled eight species of the genus and sequenced two chloroplast regions (rps16 and psbB–psbH), and the nuclear rDNA (ITS) to study the phylogeny and biogeography. The results of the phylogenetic analyses corroborated that Gymnocarpos is monophyletic, in the phylogenetic tree two well supported clades are recognized: clade 1 includes Gymnocarpos sclerocephalus and G. decandrus, mainly the North African group, whereas clade 2 comprises the remaining species, mainly in the Southern Arabian Peninsula. Molecular dating analysis revealed that the divergence age of Gymnocarpos was c. 31.33 Mya near the Eocene and Oligocene transition boundary, the initial diversification within Gymnocarpos dated to c. 6.69 Mya in the late Miocene, and the intraspecific diversification mostly occurred during the Quaternary climate oscillations. Ancestral area reconstruction suggested that the Southern Arabian Peninsula was the ancestral area for Gymnocarpos. Our conclusions revealed that the aridification since mid‐late Miocene significantly affected the diversification of the genus in these areas.     

Projected changes in climatic suitability for Kinosternon turtles by 2050 and 2070

Chelonians are expected to be negatively impacted by climate change due to limited vagility and temperature‐dependent sex determination. However, few studies have examined how freshwater turtle distributions may shift under different climate change scenarios. We used a maximum entropy approach to model the distribution of five widespread North American Kinosternon species (K. baurii, K. flavescens, K. hirtipes, K. sonoriense, and K. subrubrum) under four climate change scenarios. We found that areas with suitable climatic conditions for K. baurii and K. hirtipes are expected to decline substantially during the 21st century. In contrast, the area with suitable climate for K. sonoriense will remain essentially unchanged, while areas suitable for K. flavescens and K. subrubrum are expected to substantially increase. The centroid for the distribution of four of the five species shifted northward, while the centroid for K. sonoriense shifted slightly southward. Overall, centroids shifted at a median rate of 37.5 km per decade across all scenarios. Given the limited dispersal ability of turtles, it appears unlikely that range shifts will occur rapidly enough to keep pace with climate change during the 21st century. The ability of chelonians to modify behavioral and physiological responses in response to unfavorable conditions may allow turtles to persist for a time in areas that have become increasingly unsuitable, but this plasticity will likely only delay local extinctions.     

Upward ant distribution shift corresponds with minimum,not maximum,temperature tolerance

Rapid climate change may prompt species distribution shifts upward and poleward, but species movement in itself is not sufficient to establish climate causation. Other dynamics, such as disturbance history, may prompt species distribution shifts resembling those expected from rapid climate change. Links between species distributions, regional climate trends and physiological mechanism are needed to convincingly establish climate‐induced species shifts. We examine a 38‐year shift (1974–2012) in an elevation ecotone between two closely related ant species, Aphaenogaster picea and A. rudis. Even though A. picea and A. rudis are closely related with North American distributions that sometimes overlap, they also exhibit local‐ and regional‐scale differences in temperature requirements so that A. rudis is more southerly and inhabits lower elevations whereas A. picea is more northerly and inhabits high elevations. We find considerable movement by the warm‐habitat species upward in elevation between 1974 and 2012 with A. rudis, replacing the cold‐habitat species, A. picea, along the southern edge of the Appalachian Mountain chain in north Georgia, USA. Concomitant with the distribution shifts, regional mean and maximum temperatures remain steady (1974–2012), but minimum temperatures increase. We collect individuals from the study sites and subject them to thermal tolerance testing in a controlled setting and find that maximum and minimum temperature acclimatization occurs along the elevation gradient in both species, but A. rudis consistently becomes physiologically incapacitated at minimum and maximum temperatures 2 °C higher than A. picea. These results indicate that rising minimum temperatures allow A. rudis to move upward in elevation and displace A. picea. Given that Aphaenogaster ants are the dominant woodland seed dispersers in eastern deciduous forests, and that their thermal tolerances drive distinct differences in temperature‐cued synchrony with early blooming plants, these climate responses not only impact ant‐ant interactions, but might have wide implications for ant‐plant interactions.     

Among‐tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.     

Climate change and evolution of the New World pitviper genus Agkistrodon (Viperidae)

Aim We derived phylogenies, phylogeographies, and population demographies for two North American pitvipers, Agkistrodon contortrix (Linnaeus, 1766) and A. piscivorus (Lacépède, 1789) (Viperidae: Crotalinae), as a mechanism to evaluate the impact of rapid climatic change on these taxa. Location Midwestern and eastern North America. Methods We reconstructed maximum parsimony (MP) and maximum likelihood (ML) relationships based on 846 base pairs of mitochondrial DNA (mtDNA) ATPase 8 and ATPase 6 genes sequenced over 178 individuals. We quantified range expansions, demographic histories, divergence dates and potential size differences among clades since their last period of rapid expansion. We used the Shimodaira–Hasegawa (SH) test to compare our ML tree against three biogeographical hypotheses. Results A significant SH test supported diversification of A. contortrix from northeastern Mexico into midwestern–eastern North America, where its trajectory was sundered by two vicariant events. The first (c. 5.1 Ma) segregated clades at 3.1% sequence divergence (SD) along a continental east–west moisture gradient. The second (c. 1.4 Ma) segregated clades at 2.4% SD along the Mississippi River, coincident with the formation of the modern Ohio River as a major meltwater tributary. A single glacial refugium was detected within the Apalachicola region of southeastern North America. Significant support was also found for a hypothesis of trans‐Gulf rafting by the common ancestor of A. piscivorus from eastern Mexico (possibly the Yucatan Peninsula) to northern Florida. There, a Mid–Late Pliocene marine transgression separated it at 4.8% SD from mainland North America. Significant range expansions followed compressive glacial effects in three (of four) A. contortrix clades and in two (of three) A. piscivorus clades, with the Florida A. piscivorus clade exhibiting significant distributional stasis. Main conclusions Pliocene glaciations, rapidly developing western aridity, and Pleistocene glacial meltwaters seemingly led to the diversification of A. contortrix and A. piscivorus in North America. Both species were pushed southwards by Pleistocene climate change, with subsequent northward expansions uninhibited topographically. The subspecific taxonomy used for A. contortrix and A. piscivorus today, however, appear non‐representative. The monophyletic Florida subspecies of A. piscivorus may be a distinct species (at 4.8% SD), whereas two western subspecies of A. contortrix also appear to constitute a single distinct species, pending additional analyses. We conclude that both species of Agkistrodon can be used as suitable ectothermic models to gauge impacts of future climate change.     

Climate,physiological tolerance and sex‐biased dispersal shape genetic structure of Neotropical orchid bees

Understanding the impact of past climatic events on the demographic history of extant species is critical for predicting species' responses to future climate change. Palaeoclimatic instability is a major mechanism of lineage diversification in taxa with low dispersal and small geographical ranges in tropical ecosystems. However, the impact of these climatic events remains questionable for the diversification of species with high levels of gene flow and large geographical distributions. In this study, we investigate the impact of Pleistocene climate change on three Neotropical orchid bee species (Eulaema bombiformis, E. meriana and E. cingulata) with transcontinental distributions and different physiological tolerances. We first generated ecological niche models to identify species‐specific climatically stable areas during Pleistocene climatic oscillations. Using a combination of mitochondrial and nuclear markers, we inferred calibrated phylogenies and estimated historical demographic parameters to reconstruct the phylogeographical history of each species. Our results indicate species with narrower physiological tolerance experienced less suitable habitat during glaciations and currently exhibit strong population structure in the mitochondrial genome. However, nuclear markers with low and high mutation rates show lack of association with geography. These results combined with lower migration rate estimates from the mitochondrial than the nuclear genome suggest male‐biased dispersal. We conclude that despite large effective population sizes and capacity for long‐distance dispersal, climatic instability is an important mechanism of maternal lineage diversification in orchid bees. Thus, these Neotropical pollinators are susceptible to disruption of genetic connectivity in the event of large‐scale climatic changes.     

Exploring the tempo of species diversification in legumes

Whatever criteria are used to measure evolutionary success – species numbers, geographic range, ecological abundance, ecological and life history diversity, background diversification rates, or the presence of rapidly evolving clades – the legume family is one of the most successful lineages of flowering plants. Despite this, we still know rather little about the dynamics of lineage and species diversification across the family through the Cenozoic, or about the underlying drivers of diversification. There have been few attempts to estimate net species diversification rates or underlying speciation and extinction rates for legume clades, to test whether among-lineage variation in diversification rates deviates from null expectations, or to locate species diversification rate shifts on specific branches of the legume phylogenetic tree. In this study, time-calibrated phylogenetic trees for a set of species-rich legume clades – Calliandra, Indigofereae, Lupinus, Mimosa and Robinieae – and for the legume family as a whole, are used to explore how we might approach these questions. These clades are analysed using recently developed maximum likelihood and Bayesian methods to detect species diversification rate shifts and test for among-lineage variation in speciation, extinction and net diversification rates. Possible explanations for rate shifts in terms of extrinsic factors and/or intrinsic trait evolution are discussed. In addition, several methodological issues and limitations associated with these analyses are highlighted emphasizing the potential to improve our understanding of the evolutionary dynamics of legume diversification by using much more densely sampled phylogenetic trees that integrate information across broad taxonomic, geographical and temporal levels.     

Effects of changing climate on species diversification in tropical forest butterflies of the genus Cymothoe (Lepidoptera: Nymphalidae)

Extant clades may differ greatly in their species richness, suggesting differential rates of species diversification. Based on phylogenetic trees, it is possible to identify potential correlates of such differences. Here, we examine species diversification in a clade of 82 tropical African forest butterfly species (Cymothoe), together with its monotypic sister genus Harma. Our aim was to test whether the diversification of the Harma–Cymothoe clade correlates with end‐Miocene global cooling and desiccation, or with Pleistocene habitat range oscillations, both postulated to have led to habitat fragmentation. We first generated a species‐level phylogenetic tree for Harma and Cymothoe, calibrated within an absolute time scale, and then identified temporal and phylogenetic shifts in species diversification. Finally, we assessed correlations between species diversification and reconstructed global temperatures. Results show that, after the divergence of Harma and Cymothoe in the Miocene (15 Mya), net species diversification was low during the first 7 Myr. Coinciding with the onset of diversification of Cymothoe around 7.5 Mya, there was a sharp and significant increase in diversification rate, suggesting a rapid radiation, and correlating with a reconstructed period of global cooling and desiccation in the late Miocene, rather than with Pleistocene oscillations. Our estimated age of 4 Myr for a clade of montane species corresponds well with the uplift of the Eastern Arc Mountains where they occur. We conclude that forest fragmentation caused by changing climate in the late Miocene as well as the Eastern Arc Mountain uplift are both likely to have promoted species diversification in the Harma–Cymothoe clade. Cymothoe colonized Madagascar much later than most other insect lineages and, consequently, had less time available for diversification on the island. We consider the diversification of Cymothoe to be a special case compared with other butterfly clades studied so far, both in terms of its abrupt diversification rate increase and its recent occurrence (7 Myr). It is clear that larval host plant shift(s) cannot explain the difference in diversification between Cymothoe and Harma; however, such a shift(s) may have triggered differential diversification rates within Cymothoe. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, ●● , ●●–●●.     

Rapid diversification of Tragopogon and ecological associates in Eurasia

Tragopogon comprises approximately 150 described species distributed throughout Eurasia from Ireland and the UK to India and China with a few species in North Africa. Most of the species diversity is found in Eastern Europe to Western Asia. Previous phylogenetic analyses identified several major clades, generally corresponding to recognized taxonomic sections, although relationships both among these clades and among species within clades remain largely unresolved. These patterns are consistent with rapid diversification following the origin of Tragopogon, and this study addresses the timing and rate of diversification in Tragopogon. Using BEAST to simultaneously estimate a phylogeny and divergence times, we estimate the age of a major split and subsequent rapid divergence within Tragopogon to be ~2.6 Ma (and 1.7–5.4 Ma using various clock estimates). Based on the age estimates obtained with BEAST (HPD 1.7–5.4 Ma) for the origin of crown group Tragopogon and 200 estimated species (to accommodate a large number of cryptic species), the diversification rate of Tragopogon is approximately 0.84–2.71 species/Myr for the crown group, assuming low levels of extinction. This estimate is comparable in rate to a rapid Eurasian radiation in Dianthus (0.66–3.89 species/Myr), which occurs in the same or similar habitats. Using available data, we show that subclades of various plant taxa that occur in the same semi‐arid habitats of Eurasia also represent rapid radiations occurring during roughly the same window of time (1.7–5.4 Ma), suggesting similar causal events. However, not all species‐rich plant genera from the same habitats diverged at the same time, or at the same tempo. Radiations of several other clades in this same habitat (e.g. Campanula, Knautia, Scabiosa) occurred at earlier dates (45–4.28 Ma). Existing phylogenetic data and diversification estimates therefore indicate that, although some elements of these semi‐arid communities radiated during the Plio‐Pleistocene period, other clades sharing the same habitat appear to have diversified earlier.     

Co‐occurrence patterns of trees along macro‐climatic gradients and their potential influence on the present and future distribution of Fagus sylvatica L.

Aim During recent and future climate change, shifts in large‐scale species ranges are expected due to the hypothesized major role of climatic factors in regulating species distributions. The stress‐gradient hypothesis suggests that biotic interactions may act as major constraints on species distributions under more favourable growing conditions, while climatic constraints may dominate under unfavourable conditions. We tested this hypothesis for one focal tree species having three major competitors using broad‐scale environmental data. We evaluated the variation of species co‐occurrence patterns in climate space and estimated the influence of these patterns on the distribution of the focal species for current and projected future climates. Location Europe. Methods We used ICP Forest Level 1 data as well as climatic, topographic and edaphic variables. First, correlations between the relative abundance of European beech (Fagus sylvatica) and three major competitor species (Picea abies, Pinus sylvestris and Quercus robur) were analysed in environmental space, and then projected to geographic space. Second, a sensitivity analysis was performed using generalized additive models (GAM) to evaluate where and how much the predicted F. sylvatica distribution varied under current and future climates if potential competitor species were included or excluded. We evaluated if these areas coincide with current species co‐occurrence patterns. Results Correlation analyses supported the stress‐gradient hypothesis: towards favourable growing conditions of F. sylvatica, its abundance was strongly linked to the abundance of its competitors, while this link weakened towards unfavourable growing conditions, with stronger correlations in the south and at low elevations than in the north and at high elevations. The sensitivity analysis showed a potential spatial segregation of species with changing climate and a pronounced shift of zones where co‐occurrence patterns may play a major role. Main conclusions Our results demonstrate the importance of species co‐occurrence patterns for calibrating improved species distribution models for use in projections of climate effects. The correlation approach is able to localize European areas where inclusion of biotic predictors is effective. The climate‐induced spatial segregation of the major tree species could have ecological and economic consequences.     

Canary Grasses (Phalaris,Poaceae): biogeography,molecular dating and the role of floret structure in dispersal

Canary grasses (Phalaris, Poaceae) include 21 species, widely spread throughout the temperate and subtropical regions of the world with two centres of diversity: the Mediterranean Basin and western North America. The genus contains annual and perennial, endemic, cosmopolitan, wild, and invasive species with diploid, tetraploid and hexaploid cytotypes. As such, Phalaris presents an ideal platform to study diversification via historic hybridization and polyploidy events, and geographical dispersal in grasses. We present the first empirical phylogeographic study for Phalaris testing current, intuitive hypotheses on the centres of origin, historic dispersal events and diversification within a geological timeframe. Bayesian methods (beast , version 1.6.2) were used to establish divergence dates, and dispersal–vicariance analyses (rasp , version 2.1b) were implemented for ancestral node reconstructions. Our phylogeographic results indicate that the genus emerged during the Miocene epoch [20.6–8.4 Ma (million years ago)] in the Mediterranean basin followed by dispersal and vicariance events to Africa, Asia and the Americas. We propose that a diploid ancestor of P. arundinacea migrated to western North America via the Bering Strait, where further diversification emerged in the New World. It appears that polyploidy played a major role in the evolution of the genus in the Old World, while diversification in the New World followed a primarily diploid pathway. Dispersal to various parts of the Americas followed different routes. Fertile florets with hairy protruding sterile lemmas showed significant correlation with wider geographical distribution.     

Defining spatial and temporal patterns of phylogeographic structure in Madagascar's iguanid lizards (genus Oplurus)

Understanding the remarkably high species diversity and levels of endemism found among Madagascar’s flora and fauna has been the focus of many studies. One hypothesis that has received much attention proposes that Quaternary climate fluctuations spurred diversification. While spatial patterns of distribution and phylogenetic relationships can provide support for biogeographic predictions, temporal estimates of divergence are required to determine the fit of these geospatial patterns to climatic or biogeographic mechanisms. We use multilocus DNA sequence data to test whether divergence times among Malagasy iguanid lizards of the subfamily Oplurinae are compatible with a hypotheses of Pliocene–Pleistocene diversification. We estimate the oplurine species tree and associated divergence times under a relaxed‐clock model. In addition, we examine the phylogeographic structure and population divergence times within two sister species of Oplurus primarily distributed in the north‐west and south‐west of Madagascar (Oplurus cuvieri and Oplurus cyclurus, respectively). We find that divergence events among oplurine lineages occurred in the Oligocene and Miocene and are thus far older and incompatible with the hypothesis that recent climate fluctuations are related to current species diversity. However, the timing of intraspecific divergences and spatial patterns of population genetic structure within O. cuvieri and O. cyclurus suggest a role for both intrinsic barriers and recent climate fluctuations at population‐level divergences. Integrating information across spatial and temporal scales allows us to identify and better understand the mechanisms generating patterns diversity.     

Impacts of climate change on the global potential distribution of two notorious invasive crayfishes

1. Invasive alien species and climate change are two of the most serious global environmental threats. In particular, it is of great interest to understand how changing climates could impact the distribution of invaders that pose serious threats to ecosystems and human activities.
2. In this study, we developed ensemble species distribution models for predicting the current and future global distribution of the signal crayfish Pacifastacus leniusculus and the red swamp crayfish Procambarus clarkii, two of the most highly problematic invaders of freshwater ecosystems worldwide. We collected occurrence records of the species, from native and alien established ranges worldwide. These records in combination with averaged observations of current climatic conditions were used to calibrate a set of 10 distinct correlative models for estimating the climatic niche of each species. We next projected the estimated niches into the geographical space for the current climate conditions and for the 2050s and 2070s under representative concentration pathway 2.6 and 8.5 scenarios.
3. Our species distribution models had high predictive abilities and suggest that annual mean temperature is the main driver of the distribution of both species. Model predictions indicated that the two crayfish species have not fully occupied their suitable climates and will respond differently to future climate scenarios in different geographic regions. Suitable climate for P. leniusculus was predicted to shift poleward and to increase in extent in North America and Europe but decrease in Asia. Regions with suitable climate for P. clarkii are predicted to widen in Europe but contract in North America and Asia.
4. This study highlights that invasive species with different thermal preference are likely to respond differently to future climate changes. Our results provide important information for policy makers to design and implement anticipated measures for the prevention and control of these two problematic species.

     

Both temperature fluctuations and East Asian monsoons have driven plant diversification in the karst ecosystems from southern China

Karst ecosystems in southern China are species‐rich and have high levels of endemism, yet little is known regarding the evolutionary processes responsible for the origin and diversification of karst biodiversity. The genus Primulina (Gesneriaceae) comprises ca. 170 species endemic to southern China with high levels of ecological (edaphic) specialization, providing an exceptional model to study the plant diversification in karsts. We used molecular data from nine chloroplast and 11 nuclear regions and macroevolutionary analyses to assess the origin and cause of species diversification due to palaeoenvironmental changes and edaphic specialization in Primulina. We found that speciation was positively associated with changes in past temperatures and East Asian monsoons through the evolutionary history of Primulina. Climatic change around the mid‐Miocene triggered an early burst followed by a slowdown of diversification rate towards the present with the climate cooling. We detected different speciation rates among edaphic types, and transitions among soil types were infrequently and did not impact the overall speciation rate. Our findings suggest that both global temperature changes and East Asian monsoons have played crucial roles in floristic diversification within the karst ecosystems in southern China, such that speciation was higher when climate was warmer and wetter. This is the first study to directly demonstrate that past monsoon activity is positively correlated with speciation rate in East Asia. This case study could motivate further investigations to assess the impacts of past environmental changes on the origin and diversification of biodiversity in global karst ecosystems, most of which are under threat.     

Shifts in the climate space of temperate cyprinid fishes due to climate change are coupled with altered body sizes and growth rates

Predictions of species responses to climate change often focus on distribution shifts, although responses can also include shifts in body sizes and population demographics. Here, shifts in the distributional ranges (‘climate space’), body sizes (as maximum theoretical body sizes, L∞) and growth rates (as rate at which L∞ is reached, K) were predicted for five fishes of the Cyprinidae family in a temperate region over eight climate change projections. Great Britain was the model area, and the model species were Rutilus rutilus, Leuciscus leuciscus, Squalius cephalus, Gobio gobio and Abramis brama. Ensemble models predicted that the species' climate spaces would shift in all modelled projections, with the most drastic changes occurring under high emissions; all range centroids shifted in a north‐westerly direction. Predicted climate space expanded for R. rutilus and A. brama, contracted for S. cephalus, and for L. leuciscus and G. gobio, expanded under low‐emission scenarios but contracted under high emissions, suggesting the presence of some climate‐distribution thresholds. For R. rutilus, A. brama, S. cephalus and G. gobio, shifts in their climate space were coupled with predicted shifts to significantly smaller maximum body sizes and/or faster growth rates, aligning strongly to aspects of temperature‐body size theory. These predicted shifts in L∞ and K had considerable consequences for size‐at‐age per species, suggesting substantial alterations in population age structures and abundances. Thus, when predicting climate change outcomes for species, outputs that couple shifts in climate space with altered body sizes and growth rates provide considerable insights into the population and community consequences, especially for species that cannot easily track their thermal niches.     

Factors affecting the establishment and growth of annual legumes in semi-arid mediterranean grasslands

Legumes are an important component of mediterranean grasslands with a significant ecological and economic role. The aim of this study was to investigate the factors that affect their establishment and growth and how they survive in a highly variable and unpredictable environment. The research was carried out in a grassland characterised by a semi-arid mediterranean climate and located on a calcareous substrate at about 150 m a.s.l., in Macedonia, northern Greece. It was dominated by annual legumes such as Hippocrepis multisiliquosa, Medicago disciformis, Medicago minima, Onobrychis aequindentata, Trifolium angustifolium, Trifolium campestre and Trifolium scabrum. It was subjected to the following treatments for four consecutive years: prescribed burning, irrigation, digging, cutting, P fertilization and control. Total legume density was measured in late autumn and in the following spring each year, while total legume biomass was measured only in spring. Dominant legume species densities and biomasses were measured only in spring in the last 3 years. Also, monthly precipitation and air temperature were recorded in a nearby weather station. A great reduction of both legume density and biomass occurred at the third growing season due to adverse weather conditions. Among treatments, P fertilization affected the positively annual legume density and biomass. The other treatments such as burning, irrigation, digging and cutting influenced positively or negatively annual legume density and biomass depending on the climatic characteristics of the particular growing season involved. It is concluded that in semi-arid mediterranean grasslands with cold winters, weather conditions strongly interact with human disturbances in affecting establishment and growth of annual legumes.     

Explaining disparities in species richness between Mediterranean floristic regions: a case study in Gladiolus (Iridaceae)

Aim The causes of geographical variation in species richness in clades that do not follow the latitudinal diversity gradient have rarely been investigated. Here, we examine spatial asymmetries of diversity in Gladiolus (Iridaceae), a large genus (> 260 species) that is present in two mediterranean climate biomes: the Cape of southern Africa (106 species) and the Mediterranean Basin (7 species). Despite convergence of climatic conditions between the two regions, the species density of Gladiolus is over one order of magnitude higher in the Cape than in the Mediterranean Basin. We investigate whether the diversity disparities observed in the genus are better explained by recent colonization of species‐poor areas (temporal hypothesis) or by differential rates of diversification (evolutionary hypothesis). Location Africa, Madagascar and Eurasia Methods We employ a recently developed Bayesian method for the estimation of diversification rates and a biogeographical optimization approach within a phylogenetic framework. Results In Gladiolus, the ‘diversity anomaly’ between the two Mediterranean climate regions cannot be explained solely by the time available for speciation in the Cape, but is also due to locally reduced rates of diversification in the Mediterranean Basin. Furthermore, high overall diversity in southern Africa stems from an ancient origin in the Cape allied with high rates of diversification in the summer‐rainfall region of the subcontinent. Main conclusions Both evolutionary and temporal hypotheses must be taken into account in order to explain the diversity anomaly between the Mediterranean Basin and the Cape. Our results suggest that regions at comparable latitudes and/or with similar climate may not converge in diversity levels due to heterogeneity of diversification rates and contrasting biogeographical histories.