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1.

The Chelonian lineage has been exposed to several climate change events along its evolutionary history, but the rapid contemporary change in climate patterns has the potential to erode turtle populations. This study focuses on (1) evaluating the climatically suitable area available for 15 species of mud turtles of the genus Kinosternon, and on (2) assessing whether or not these species retain their ancestral climate niche. Occurrence data was collected for these species and, using the Maxent algorithm and WorldClim bioclimatic variables, suitable present and future climatic niche areas were modeled. In addition, we also carried out climatic niche similarity analyses between pairs of species to evaluate whether these conserve their climatic niche. Our models suggest that most species of Kinosternon will lose a high proportion of their suitable habitat in the future. Most mud turtle species seem to conserve their climatic niche, suggesting the prevalence of niche conservatism in the group. Our results indicate that several mud turtle species could be at severe risk of disappearing over the next few decades due to the loss of climatically suitable areas and of the conservation of their climatic niches.

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2.
Macroecological analyses often test hypotheses at the global scale, or among more closely related species in a single region (e.g. continent). Here, we test several hypotheses about climatic niche widths among relatively closely related species that occur across multiple continents, and compare patterns within and across continents to see if they differ. We focus on the lizard genus Varanus (monitor lizards), which occurs in diverse environments in Africa, Asia, and Australia. We address three main questions. 1) How do climatic niche breadths of species on a given niche axis change based on the position of species along that niche axis? (E.g. are species that occur in more extreme environments more narrowly specialized for those conditions?) 2) Are there trade‐offs in niche breadths on temperature and precipitation axes among species, or are niche widths on different axes positively related? 3) Is variation in niche breadths among species explained primarily by within‐locality seasonal variation, or by differences in climatic conditions among localities across the species range? We generate a new time‐calibrated phylogeny for Varanus and test these hypotheses within and between continents using climatic data and phylogenetic methods. Our results show that patterns on each continent often parallel each other and global patterns. However, in many other cases, the strength of relationships can change dramatically among closely related species on different continents. Overall, we found that: 1) species in warmer environments have narrower temperature niche breadths, but there is no relationship between precipitation niche breadth and niche position; 2) temperature and precipitation niche breadths tend to be positively related among species, rather than showing trade‐offs; and 3) within‐locality seasonal variation explains most variation in climatic niche breadths. Some of these results are concordant with previous studies (in amphibians and North American lizards), and might represent general macroecological patterns.  相似文献   

3.
Ecological requirements and environmental conditions can influence diversification across temporal and spatial scales. Understanding the role of ecological niche evolution under phylogenetic contexts provides insights on speciation mechanisms and possible responses to future climatic change. Large‐scale phyloclimatic studies on the megadiverse Neotropics, where biomes with contrasting vegetation types occur in narrow contact, are rare. We integrate ecological and biogeographic data with phylogenetic comparative methods, to investigate the relative roles of biogeographic events and niche divergence and conservatism on the diversification of the lizard genus Kentropyx Spix, 1825 (Squamata: Teiidae), distributed in South American rainforests and savannas. Using five molecular markers, we estimated a dated species tree, which recovered three clades coincident with previously proposed species groups diverging during the mid‐Miocene. Biogeography reconstruction indicates a role of successive dispersal events from an ancestral range in the Brazilian Shield and western Amazonia. Ancestral reconstruction of climatic tolerances and niche overlap metrics indicates a trend of conservatism during the diversification of groups from the Amazon Basin and Guiana Shield, and a strong signal of niche divergence in the Brazilian Shield savannas. Our results suggest that climatic‐driven divergence at dynamic forest‐savanna borders might have resulted in adaptation to new environmental niches, promoting habitat shifts and shaping speciation patterns of Neotropical lizards. Dispersal and ecological divergence could have a more important role in Neotropical diversification than previously thought.  相似文献   

4.
Climate may play important roles in speciation, such as causing the range fragmentation that underlies allopatric speciation (through niche conservatism) or driving divergence of parapatric populations along climatic gradients (through niche divergence). Here, we developed new methods to test the frequency of climate niche conservatism and divergence in speciation, and applied it to species pairs of squamate reptiles (lizards and snakes). We used a large‐scale phylogeny to identify 242 sister species pairs for analysis. From these, we selected all terrestrial allopatric pairs with sufficient occurrence records (= 49 pairs) and inferred whether each originated via climatic niche conservatism or climatic niche divergence. Among the 242 pairs, allopatric pairs were most common (41.3%), rather than parapatric (19.4%), partially sympatric (17.7%), or fully sympatric species pairs (21.5%). Among the 49 selected allopatric pairs, most appeared to have originated via climatic niche divergence (61–76%, depending on the details of the methods). Surprisingly, we found greater climatic niche divergence between allopatric sister species than between parapatric pairs, even after correcting for geographic distance. We also found that niche divergence did not increase with time, further implicating niche divergence in driving lineage splitting. Overall, our results suggest that climatic niche divergence may often play an important role in allopatric speciation, and the methodology developed here can be used to address the generality of these findings in other organisms.  相似文献   

5.
Understanding biodiversity patterns on islands has long been a central aim in ecology and conservation biology. Island‐specific biogeographical processes play substantial roles in the formation of endemic biota. Here, we examined how climate niche conservatism and geohistorical factors are interactively associated with in situ diversification of Tertiary relict flora in the east Asian continental islands. We generated two novel datasets for species distribution and phylogeny that included all of the known vascular plant species in Japan (5575). Then we tested phylogenetic signal of climatic tolerance, in terms of absolute minimum temperature and water balance, and explored environmental predictors of phylogenetic structure (evolutionary derivedness and clustering) of species assemblages. Although phylogenetic signal of climatic tolerance was significant across the phylogeny of most species, the strength of climatic niche conservatism differed among ferns, gymnosperms, angiosperm trees, and angiosperm herbs. For angiosperm trees, cold temperatures acted as environmental filters that generated phylogenetic derivedness/clustering of species assemblages. For fern and angiosperm herb species, however, phylogenetic properties were not associated with climatic harshness. These contrasting patterns among groups reflected climate niche evolution in vascular plants with different growth forms and traits; for example, diversification of angiosperm trees (but not fern and herb) occurred in response to historical climatic cooling. More importantly, geographical constraints contributed to evolutionary radiation that resulted from isolation by distance from the continent or by elevation. Quaternary climate change was also associated with clade‐specific radiation in refugial habitats. The degree to which geographical, geological, and palaeoclimatic variables explain the phylogenetic structure underscores the importance of isolation‐ and habitat‐stability‐related geohistorical processes in driving in situ diversification despite climatic niche conservatism. We propose that the highly endemic flora of the east Asian islands resulted from the interplay of idiosyncratic regional factors, and ecological and evolutionary processes, such as climate niche assembly and adaptive/nonadaptive radiation.  相似文献   

6.
Spatial responses of species to past climate change depend on both intrinsic traits (climatic niche breadth, dispersal rates) and the scale of climatic fluctuations across the landscape. New capabilities in generating and analysing population genomic data, along with spatial modelling, have unleashed our capacity to infer how past climate changes have shaped populations, and by extension, complex communities. Combining these approaches, we uncover lineage diversity across four codistributed lizards from the Australian Monsoonal Tropics and explore how varying climatic tolerances interact with regional climate history to generate common vs. disparate responses to late Pleistocene change. We find more divergent spatial structuring and temporal demographic responses in the drier Kimberley region compared to the more mesic and consistently suitable Top End. We hypothesize that, in general, the effects of species’ traits on sensitivity to climate fluctuation will be more evident in climatically marginal regions. If true, this points to the need in climatically marginal areas to craft more species‐(or trait)‐specific strategies for persistence under future climate change.  相似文献   

7.
The progressive expansion of the Australian arid zone during the last 20 Ma appears to have spurred the diversification of several families of plants, vertebrates and invertebrates, yet such taxonomic groups appear to show limited niche radiation. Here, we test whether speciation is associated with niche conservatism (constraints on ecological divergence) or niche divergence in a tribe of marsupial mice (Sminthopsini; 23 taxa) that includes the most speciose genus of living dasyurids, the sminthopsins. To that end, we integrated phylogenetic data with ecological niche modelling, to enable us to reconstruct the evolution of climatic suitability within Sminthopsini. Niche overlap among species was low‐moderate (but generally higher than expected given environmental background similarity), and the degree of phylogenetic clustering increased with aridity. Climatic niche reconstruction illustrates that there has been little apparent evolution of climatic tolerance within clades. Accordingly, climatic disparity tends to be accumulated among clades, suggesting considerable niche conservatism. Our results also indicate that evolution of climatic tolerances has been heterogeneous across different dimensions of climate (temperature vs. precipitation) and across phylogenetic clusters (Sminthopsis murina group vs. other groups). Although some results point to the existence of shifts in climatic niches during the speciation of sminthopsins, our study provides evidence for substantial phylogenetic niche conservatism in the group. We conclude that niche diversification had a low impact on the speciation of this tribe of small, but highly mobile marsupials.  相似文献   

8.
Studies on niche evolution allow us to establish how species niches have changed over time and to identify how long‐term evolutionary processes have led to present‐day species distributions. Here, we investigate the patterns of climatic niche evolution in Tynanthus (Bignonieae, Bignoniaceae), a genus of narrowly distributed species. We test the hypothesis that niche conservatism has played an important role in the history of this group of Neotropical lianas. We perform univariate and multivariate comparisons between climatic niches of species and associated environmental data with information on phylogenetic relationships. We encountered considerable divergence in niches among species, indicating that niche conservatism in climatic variables does not seem to have played a key role in the history of the genus. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 95–109.  相似文献   

9.
In this study, we explore the interplay of population demography with the evolution of ecological niches during or after speciation in Hordeum. While large populations maintain a high level of standing genetic diversity, gene flow and recombination buffers against fast alterations in ecological adaptation. Small populations harbour lower allele diversity but can more easily shift to new niches if they initially survive under changed conditions. Thus, large populations should be more conservative regarding niche changes in comparison to small populations. We used environmental niche modelling together with phylogenetic, phylogeographic and population genetic analyses to infer the correlation of population demography with changes in ecological niche dimensions in 12 diploid Hordeum species from the New World, forming four monophyletic groups. Our analyses found both shifts and conservatism in distinct niche dimensions within and among clades. Speciation due to vicariance resulted in three species with no pronounced climate niche differences, while species originating due to long‐distance dispersals or otherwise encountering genetic bottlenecks mostly revealed climate niche shifts. Niche convergence among clades indicates a niche‐filling pattern during the last 2 million years in South American Hordeum. We provide evidence that species, which did not encounter population reductions mainly showed ecoclimatic niche conservatism, while major niche shifts occurred in species which have undergone population bottlenecks. Our data allow the conclusion that population demography influences adaptation and niche shifts or conservatism in South American Hordeum species.  相似文献   

10.
Rising temperatures are predicted to melt all perennial ice cover in the Arctic by the end of this century, thus opening up suitable habitat for temperate and subarctic species. Canopy‐forming seaweeds provide an ideal system to predict the potential impact of climate‐change on rocky‐shore ecosystems, given their direct dependence on temperature and their key role in the ecological system. Our primary objective was to predict the climate‐change induced range‐shift of Fucus distichus, the dominant canopy‐forming macroalga in the Arctic and subarctic rocky intertidal. More specifically, we asked: which Arctic/subarctic and cold‐temperate shores of the northern hemisphere will display the greatest distributional change of Fdistichus and how will this affect niche overlap with seaweeds from temperate regions? We used the program MAXENT to develop correlative ecological niche models with dominant range‐limiting factors and 169 occurrence records. Using three climate‐change scenarios, we projected habitat suitability of Fdistichus – and its niche overlap with three dominant temperate macroalgae – until year 2200. Maximum sea surface temperature was identified as the most important factor in limiting the fundamental niche of Fdistichus. Rising temperatures were predicted to have low impact on the species' southern distribution limits, but to shift its northern distribution limits poleward into the high Arctic. In cold‐temperate to subarctic regions, new areas of niche overlap were predicted between Fdistichus and intertidal macroalgae immigrating from the south. While climate‐change threatens intertidal seaweeds in warm‐temperate regions, seaweed meadows will likely flourish in the Arctic intertidal. Although this enriches biodiversity and opens up new seaweed‐harvesting grounds, it will also trigger unpredictable changes in the structure and functioning of the Arctic intertidal ecosystem.  相似文献   

11.
Niche divergence among closely related lineages can be informative on the ecological and evolutionary processes involved in differentiation, particularly in the case of cryptic species complexes. Here we compared phylogenetic relationships and niche similarity between pairs of lineages included in the Podarcis hispanicus complex to examine patterns of niche divergence and its role in the organization of current diversity patterns, as allopatric, parapatric, and sympatric lineages occur in the Western Mediterranean Basin. First, we used ecological niche models to characterize the realized climatic niche of each Podarcis hispanicus complex lineage based on topographic and climatic variables, to identify important variables, and to test for niche conservatism or divergence between pairs of lineages. Variables related to precipitation generally exhibited the highest contribution to niche models, highlighting the importance of rainfall levels in shaping distributions of Podarcis wall lizards. We found that most forms have significant differences in realized climatic niches that do not follow the pattern of mitochondrial divergence. These results lend support to the hypothesis that genetic divergence across Podarcis hispanicus complex most likely occurred in allopatric conditions, mostly with significant niche divergence. Competition after secondary contact is also suggested by the common occurrence of niche overlap between lineages that exhibit strictly parapatric distribution. The almost continuous distribution of Podarcis lizards in the study area appears to be a result of a combination of complementary suitable niches and competition, which seem two important mechanisms limiting geographic distributions and restricting the existence of extensive contact zones.  相似文献   

12.
Phenotypic plasticity can contribute to the process of adaptive radiation by facilitating population persistence in novel environments. West Indian Anolis lizards provide a classic example of an adaptive radiation, in which divergence has occurred along two primary ecological axes: structural microhabitat and climate. Adaptive plasticity in limb morphology is hypothesized to have facilitated divergence along the structural niche axis in Anolis, but very little work has explored plasticity in physiological traits. Here, we experimentally ask whether Puerto Rican Anolis cristatellus from mesic and xeric habitats differ in desiccation rates, and whether these lizards exhibit an acclimation response to changes in relative humidity. We first present microclimatic data collected at lizard perch sites that demonstrate that abiotic conditions experienced by lizards differ between mesic and xeric habitat types. In Experiment 1, we measured desiccation rates of lizards from both habitats maintained under identical laboratory conditions. This experiment demonstrated that desiccation rates differ between populations; xeric lizards lose water more slowly than mesic lizards. In Experiment 2, lizards from each habitat were either maintained under the conditions of Experiment 1, or under extremely low relative humidity. Desiccation rates did not differ between lizards from the same habitat maintained under different treatments and xeric lizards maintained lower desiccation rates than mesic lizards within each treatment. Our results demonstrate that A. cristatellus does not exhibit an acclimation response to abrupt changes of hydric conditions, and suggest that tropical Anolis lizards might be unable to exhibit physiological plasticity in desiccation rates in response to varying climatic conditions.  相似文献   

13.
To generate realistic projections of species’ responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species’s climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.  相似文献   

14.
Although climatic niche conservatism has been assumed by a large number of studies focused on climatic niche evolution, there are examples of climatic niche diversification and adaptation to changing climates. In this article, we reconstruct a climatic niche of scaly tree ferns (Cyatheaceae) using a rigorous analytical procedure which combines climatic niche modelling with reconstruction of continuous characters given a phylogenetic hypothesis. To estimate the limits to climatic niches of species, we used climate envelope modelling and ordination. Ancestral climatic niches of species were reconstructed by maximum likelihood and least‐squares analyses. We observed a trend towards niche conservatism with occasional events of niche transformations in scaly tree ferns. We discuss the implications of our study with respect to the potential and limitations for applications of niche modelling to evolutionary studies. We suggest that future studies of evolution of climatic niches could be considerably improved by employing approaches enabling reconstruction of continuous response to climatic gradients. Further progress may also be achieved by exploring models of character evolution other than the Brownian motion model. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 1–19.  相似文献   

15.
Cold‐adapted taxa are experiencing severe range shifts due to climate change and are expected to suffer a significant reduction of their climatically suitable habitats in the next few decades. However, it has been proposed that taxa with sufficient standing genetic and ecologic diversity will better withstand climate change. These taxa are typically more broadly distributed in geographic and ecological niche space, therefore they are likely to endure higher levels of populations loss than more restricted, less diverse taxa before the effects of those losses impact their overall diversity and resilience. Here, we explore the potential relationship between intraspecific genetic and ecological diversity and future resilience, using the cold‐adapted plant Primula farinosa. We employ high‐throughput sequencing to assess the genomic diversity of phylogeographic lineages in P. farinosa. Additionally, we use current climatic variables to define niche breadth and niche differentiation across lineages. Finally, we calibrate species distribution models (SDMs) and project the climatic preferences of each lineage on future climate to predict lineage‐specific shifts in climatically suitable habitats. Our study predicts relative persistence of future suitable habitats for the most genetically and ecologically diverse lineages of the cold‐adapted P. farinosa, but significant reduction of them for two out of its four lineages. While we do not provide specific experiments aimed at identifying the causal links between genetic diversity and resilience to climate change, our results indicate that greater genetic diversity and wider ecological breadth may buffer species responses to rapid climatic changes. This study further highlights the importance of integrating knowledge of intraspecific diversity for predicting species fate in response to climate change.  相似文献   

16.
Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.  相似文献   

17.
With rates of climate change exceeding the rate at which many species are able to shift their range or adapt, it is important to understand how future changes are likely to affect biodiversity at all levels of organisation. Understanding past responses and extent of niche conservatism in climatic tolerance can help predict future consequences. We use an integrated approach to determine the genetic consequences of past and future climate changes on a bat species, Plecotus austriacus. Glacial refugia predicted by palaeo‐modelling match those identified from analyses of extant genetic diversity and model‐based inference of demographic history. Former refugial populations currently contain disproportionately high genetic diversity, but niche conservatism, shifts in suitable areas and barriers to migration mean that these hotspots of genetic diversity are under threat from future climate change. Evidence of population decline despite recent northward migration highlights the need to conserve leading‐edge populations for spearheading future range shifts.  相似文献   

18.

Background

The impact of global climate change on plant distribution, speciation and extinction is of current concern. Examining species climatic preferences via bioclimatic niche modelling is a key tool to study this impact. There is an established link between bioclimatic niche models and phylogenetic diversification. A next step is to examine future distribution predictions from a phylogenetic perspective. We present such a study using Cyclamen (Myrsinaceae), a group which demonstrates morphological and phenological adaptations to its seasonal Mediterranean-type climate. How will the predicted climate change affect future distribution of this popular genus of garden plants?

Results

We demonstrate phylogenetic structure for some climatic characteristics, and show that most Cyclamen have distinct climatic niches, with the exception of several wide-ranging, geographically expansive, species. We reconstruct climate preferences for hypothetical ancestral Cyclamen. The ancestral Cyclamen lineage has a preference for the seasonal Mediterranean climate characteristic of dry summers and wet winters. Future bioclimatic niches, based on BIOCLIM and Maxent models, are examined with reference to a future climate scenario for the 2050s. Over the next 50 years we predict a northward shift in the area of climatic suitability, with many areas of current distribution becoming climatically unsuitable. The area of climatic suitability for every Cyclamen species is predicted to decrease. For many species, there may be no areas with a suitable climate regardless of dispersal ability, these species are considered to be at high risk of extinction. This risk is examined from a phylogenetic perspective.

Conclusion

Examining bioclimatic niches from a phylogenetic perspective permits novel interpretations of these models. In particular, reconstruction of ancestral niches can provide testable hypothesis about the historical development of lineages. In the future we can expect a northwards shift in climatic suitability for the genus Cyclamen. If this proves to be the case then dispersal is the best chance of survival, which seems highly unlikely for ant-dispersed Cyclamen. Human-assisted establishment of Cyclamen species well outside their native ranges offers hope and could provide the only means of dispersal to potentially suitable future environments. Even without human intervention the phylogenetic perspective demonstrates that major lineages could survive climate change even if many species are lost.  相似文献   

19.
Thick‐billed Parrots (Rhynchopsitta pachyrhyncha) and Maroon‐fronted Parrots (Rhynchopsitta terrisi) are the only parrots in Mexico found in high‐elevation coniferous forests. Both species are critically endangered due to logging, and climate change is expected to further reduce their available habitat. Our objectives were to assess the present and future availability of a suitable habitat for these parrots using ecological niche models. Future climatic scenarios were estimated by overlaying the present distributions of these parrots on maps of projected biome distributions generated using a North American vegetation model. Our climatic scenarios revealed that the distribution of key habitats for both parrots will likely be affected as the climate becomes more suitable for xeric biomes. The climate associated with coniferous forests in the current range of Maroon‐fronted Parrots is predicted to disappear by 2090, and the climate associated with the key coniferous habitats of Thick‐billed Parrots may contract. However, our results also indicate that suitable climatic conditions will prevail for the high‐elevation coniferous biomes where Thick‐billed Parrots nest. The degree to which both species of parrots will be able to adapt to the new scenarios is uncertain. Some of their life history traits may allow them to respond with a combination of adaptive and spatial responses to climatic change and, in addition, suitable climatic conditions will prevail in some portions of their ranges. Actions needed to ensure the conservation of these parrots include strict control of logging and integration of rapid response teams for fire management within the potential foraging ranges of nesting pairs. A landscape with a greater proportion of restored forests would also aid in the recovery of current populations of Thick‐billed and Maroon‐fronted parrots and facilitate their responses to climate change.  相似文献   

20.
Climate adaptation has major consequences in the evolution and ecology of all living organisms. Though phytophagous insects are an important component of Earth's biodiversity, there are few studies investigating the evolution of their climatic preferences. This lack of research is probably because their evolutionary ecology is thought to be primarily driven by their interactions with their host plants. Here, we use a robust phylogenetic framework and species‐level distribution data for the conifer‐feeding aphid genus Cinara to investigate the role of climatic adaptation in the diversity and distribution patterns of these host‐specialized insects. Insect climate niches were reconstructed at a macroevolutionary scale, highlighting that climate niche tolerance is evolutionarily labile, with closely related species exhibiting strong climatic disparities. This result may suggest repeated climate niche differentiation during the evolutionary diversification of Cinara. Alternatively, it may merely reflect the use of host plants that occur in disparate climatic zones, and thus, in reality the aphid species' fundamental climate niches may actually be similar but broad. Comparisons of the aphids' current climate niches with those of their hosts show that most Cinara species occupy the full range of the climatic tolerance exhibited by their set of host plants, corroborating the hypothesis that the observed disparity in Cinara species' climate niches can simply mirror that of their hosts. However, 29% of the studied species only occupy a subset of their hosts' climatic zone, suggesting that some aphid species do indeed have their own climatic limitations. Our results suggest that in host‐specialized phytophagous insects, host associations cannot always adequately describe insect niches and abiotic factors must be taken into account.  相似文献   

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