保护演化生物学——保护生物学的新分支
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摘要
离开演化谈保护,往往难窥其道.保护生物学建立之初旨在阐释受人类活动或其他因素干扰下的物种、种群、群落和生态系统的保护问题,在其发展过程中其逐渐从生态、行为等宏观层面深入到生理、遗传、基因组、适应性演化等分子机制层面.特别是近年来,越来越多的研究聚焦到物种演化历史及其成因、适应性演化机制与演化潜力等方面,这些研究超越了传统保护生物学分支学科如保护生态学和行为学等的研究内容.将演化生物学原理引入保护生物学研究则能更好地揭示问题的本质,因而越来越受到保护生物学家的重视.因此,我们提出"保护演化生物学"这一新分支学科,以强化演化生物学原理和方法在解决物种保护问题中的应用.保护演化生物学是将演化生物学原理和方法整合进保护生物学研究中,旨在从演化的视角探讨物种的过去、现在与未来,揭示物种如何适应和响应环境变化以维持长期生存的机制,阐明物种濒危过程与演化潜力,以期为制定前瞻性的物种保护策略提供科学依据.本文从保护演化生物学的产生、发展、研究内容、研究方法及研究意义等方面进行了简要介绍,以推动保护演化生物学分支学科的发展.
"Nothing in Biology Makes Sense Except in the Light of Evolution".Conservation biology was established to address the processes that endangered biodiversity in the form of species,populations,communities,and ecosystems.Traditionally conservation biologists were mainly concerned with aspects related to ecology,demography,behavior,and physiology.As human impacts on earth's biota intensify and diversify across scales,the focus of the field has expanded to address other aspects from ecology to molecular mechanisms such as genetic diversity and adaptive evolution.More and more studies are now addressing how biodiversity responds to evolving threats whose impacts on biodiversity span years to decades and centuries.These studies go beyond the traditional fields in conservation biology such as conservation ecology and behavior.Integrating evolutionary principles into conservation biology science and practice can reveal the nature of the problem,which has attracted more and more attention from conservation biologists.Hereby,we propose a new branch of conservation biology:the conservation evolutionary biology in order to strengthen the application of evolutionary principles in solving the conservation issues of species.Conservation evolutionary biology integrates the principles and methods of evolutionary biology into conservation biology research.It aims to explore the past,present and future of species status from the perspective of evolution.Likewise,it aims to reveal how species adapt to and respond to environmental changes to ensure long-term persistence,and to clarify the process of species endangerment and evolutionary potential.A major goal is to provide scientific basis for formulating forward-looking strategies for species conservation.Here,the origin,development,research contents,research methods and significance of conservation evolutionary biology are briefly introduced to stimulate further development of the field.
"Nothing in Biology Makes Sense Except in the Light of Evolution".Conservation biology was established to address the processes that endangered biodiversity in the form of species,populations,communities,and ecosystems.Traditionally conservation biologists were mainly concerned with aspects related to ecology,demography,behavior,and physiology.As human impacts on earth's biota intensify and diversify across scales,the focus of the field has expanded to address other aspects from ecology to molecular mechanisms such as genetic diversity and adaptive evolution.More and more studies are now addressing how biodiversity responds to evolving threats whose impacts on biodiversity span years to decades and centuries.These studies go beyond the traditional fields in conservation biology such as conservation ecology and behavior.Integrating evolutionary principles into conservation biology science and practice can reveal the nature of the problem,which has attracted more and more attention from conservation biologists.Hereby,we propose a new branch of conservation biology:the conservation evolutionary biology in order to strengthen the application of evolutionary principles in solving the conservation issues of species.Conservation evolutionary biology integrates the principles and methods of evolutionary biology into conservation biology research.It aims to explore the past,present and future of species status from the perspective of evolution.Likewise,it aims to reveal how species adapt to and respond to environmental changes to ensure long-term persistence,and to clarify the process of species endangerment and evolutionary potential.A major goal is to provide scientific basis for formulating forward-looking strategies for species conservation.Here,the origin,development,research contents,research methods and significance of conservation evolutionary biology are briefly introduced to stimulate further development of the field.
引文
1 Dobzhansky T.Nothing in biology makes sense except in the light of evolution.Am Biol Teacher,1973,35:125-129
2 Wei F W,Nie Y G,Miao H X,et al.Advancements of the researches on biodiversity loss mechanisms(in Chinese).Chin Sci Bull(Chin Ver),2014,59:430-437[魏辅文,聂永刚,苗海霞,等.生物多样性丧失机制研究进展.科学通报,2014,59:430-437]
3 Wei F W.Evolutionary conservation biology of giant pandas(in Chinese).Sci Sin Vitae,2018,48:1048-1053[魏辅文.大熊猫演化保护生物学研究.中国科学:生命科学,2018,48:1048-1053]
4 Ferriere R,Dieckmann U,Couver D.Evolutionary Conservation Biology.New York:Cambridge University Press,2004
5 Zhao S,Zheng P,Dong S,et al.Whole-genome sequencing of giant pandas provides insights into demographic history and local adaptation.Nat Genet,2013,45:67-71
6 Zhou X,Guang X,Sun D,et al.Population genomics of finless porpoises reveal an incipient cetacean species adapted to freshwater.Nat Commun,2018,9:1276
7 Bay R A,Harrigan R J,Underwood V L,et al.Genomic signals of selection predict climate-driven population declines in a migratory bird.Science,2018,359:83-86
8 Kimura M.Evolutionary rate at the molecular level.Nature,1968,217:624-626
9 Levine J M,Hille Ris Lambers J.The importance of niches for the maintenance of species diversity.Nature,2009,461:254-257
10 Pavlogiannis A,Tkadlec J,Chatterjee K,et al.Construction of arbitrarily strong amplifiers of natural selection using evolutionary graph theory.Commun Biol,2018,1:71
11 Zhou X,Sun F,Xu S,et al.Baiji genomes reveal low genetic variability and new insights into secondary aquatic adaptations.Nat Commun,2013,4:2708
12 Zhou X,Wang B,Pan Q,et al.Whole-genome sequencing of the snub-nosed monkey provides insights into folivory and evolutionary history.Nat Genet,2014,46:1303-1310
13 Li H,Durbin R.Inference of human population history from individual whole-genome sequences.Nature,2011,475:493-496
14 Zhu L,Wu Q,Dai J,et al.Evidence of cellulose metabolism by the giant panda gut microbiome.Proc Natl Acad Sci USA,2011,108:17714-17719
15 Wei F,Wu Q,Hu Y,et al.Conservation metagenomics:a new branch of conservation biology.Sci China Life Sci,2019,62:168-178
16 Zhang Z,Xu D,Wang L,et al.Convergent evolution of rumen microbiomes in high-altitude mammals.Curr Biol,2016,26:1873-1879
17 Shao C,Bao B,Xie Z,et al.The genome and transcriptome of Japanese flounder provide insights into flatfish asymmetry.Nat Genet,2017,49:119-124
18 Girod C,Vitalis R,Leblois R,et al.Inferring population decline and expansion from microsatellite data:A simulation-based evaluation of the Msvar method.Genetics,2011,188:165-179
19 Elmer K R,Reggio C,Wirth T,et al.Pleistocene desiccation in East Africa bottlenecked but did not extirpate the adaptive radiation of Lake Victoria haplochromine cichlid fishes.Proc Natl Acad Sci USA,2009,106:13404-13409
20 Fahrig L.Ecological responses to habitat fragmentation Per Se.Annu Rev Ecol Evol Syst,2017,48:1-23
21 Franks S J,Kane N C,O’Hara N B,et al.Rapid genome-wide evolution in Brassica rapa populations following drought revealed by sequencing of ancestral and descendant gene pools.Mol Ecol,2016,25:3622-3631
22 Lindbladh M,Oswald W W,Foster D R,et al.A late-glacial transition from Picea glauca to Picea mariana in southern New England.Quat res,2007,67:502-508
23 Cruzan M B,Templeton A R.Paleoecology and coalescence:Phylogeographic analysis of hypotheses from the fossil record.Trends Ecol Evol,2000,15:491-496
24 Guichoux E,Lagache L,Wagner S,et al.Current trends in microsatellite genotyping.Mol Ecol Res,2011,11:591-611
25 Schiffels S,Durbin R.Inferring human population size and separation history from multiple genome sequences.Nat Genet,2014,46:919-925
26 Hewitt G.The genetic legacy of the Quaternary ice ages.Nature,2000,405:907-913
27 Hewitt G M.Genetic consequences of climatic oscillations in the Quaternary.Philos Trans R Soc London Ser B-Biol Sci,2004,359:183-195
28 Konnert M,Bergmann F.The geographical distribution of genetic variation of silver fir(Abies alba,Pinaceae)in relation to its migration history.Pl Syst Evol,1995,196:19-30
29 Liu Z,Ren B,Wu R,et al.The effect of landscape features on population genetic structure in Yunnan snub-nosed monkeys(Rhinopithecus bieti)implies an anthropogenic genetic discontinuity.Mol Ecol,2009,18:3831-3846
30 Ko A M,Zhang Y,Yang M A,et al.Mitochondrial genome of a 22,000-year-old giant panda from southern China reveals a new panda lineage.Curr Biol,2018,28:R693-R694
31 Li Y,Liu Z,Shi P,et al.The hearing gene Prestin unites echolocating bats and whales.Curr Biol,2010,20:R55-R56
32 Zhao H,Yang J R,Xu H,et al.Pseudogenization of the umami taste receptor gene Tas1r1 in the giant panda coincided with its dietary switch to bamboo.Mol Biol Evol,2010,27:2669-2673
33 Hu Y,Wu Q,Ma S,et al.Comparative genomics reveals convergent evolution between the bamboo-eating giant and red pandas.Proc Natl Acad Sci USA,2017,114:1081-1086
34 Shan L,Wu Q,Wang L,et al.Lineage-specific evolution of bitter taste receptor genes in the giant and red pandas implies dietary adaptation.Integrat Zool,2018,13:152-159
35 Zhu K,Zhou X,Xu S,et al.The loss of taste genes in cetaceans.BMC Evol Biol,2014,14:218
36 Liu Y,Cotton J A,Shen B,et al.Convergent sequence evolution between echolocating bats and dolphins.Curr Biol,2010,20:R53-R54
37 Shen Y Y,Liang L,Li G S,et al.Parallel evolution of auditory genes for echolocation in bats and toothed whales.PLo S Genet,2012,8:e1002788
38 Wei F W.A new perspective on animal adaptive evolution:Animal gut microbiome.Sci Sin Vitae,2016,46:1-3[魏辅文.动物适应性进化研究的新视角:动物肠道微生物组.中国科学:生命科学,2016,46:1-3]
39 Gaynor K M,Hojnowski C E,Carter N H,et al.The influence of human disturbance on wildlife nocturnality.Science,2018,360:1232-1235
40 Fritz S A,Bininda-Emonds O R P,Purvis A.Geographical variation in predictors of mammalian extinction risk:Big is bad,but only in the tropics.Ecol Lett,2009,12:538-549
41 Krauss J,Bommarco R,Guardiola M,et al.Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels.Ecol Lett,2010,13:597-605
42 Sih A,Ferrari M C O,Harris D J.Evolution and behavioural responses to human-induced rapid environmental change.Evolary Appl,2011,4:367-387
43 Barelli C,Albanese D,Donati C,et al.Habitat fragmentation is associated to gut microbiota diversity of an endangered primate:implications for conservation.Sci Rep,2015,5:14862
44 Allendorf F W,England P R,Luikart G,et al.Genetic effects of harvest on wild animal populations.Trends Ecol Evol,2008,23:327-337
45 J?rgensen C,Enberg K,Dunlop E S,et al.Ecology:Managing evolving fish stocks.Science,2007,318:1247-1248
46 Diaz Pauli B,Sih A.Behavioural responses to human-induced change:Why fishing should not be ignored.Evol Appl,2017,10:231-240
47 Andres S,Ribeyre F,Tourencq J N,et al.Interspecific comparison of cadmium and zinc contamination in the organs of four fish species along a polymetallic pollution gradient(Lot River,France).Sci Total Environ,2000,248:11-25
48 Mc Comb B C,Curtis L,Chambers C L,et al.Acute toxic hazard evaluations of glyphosate herbicide on terrestrial vertebrates of the oregon coast range.Environ Sci Pollut Res,2008,15:266-272
49 H?lker F,Wolter C,Perkin E K,et al.Light pollution as a biodiversity threat.Trends Ecol Evol,2010,25:681-682
50 Tuxbury S M,Salmon M.Competitive interactions between artificial lighting and natural cues during seafinding by hatchling marine turtles.Biol Conserv,2005,121:311-316
51 Clarke C A,Mani G S,Wynne G.Evolution in reverse:clean air and the peppered moth.Biol J Linnean Soc,1985,26:189-199
52 Van't Hof A E,Campagne P,Rigden D J,et al.The industrial melanism mutation in British peppered moths is a transposable element.Nature,2016,534:102-105
53 Altizer S,Harvell D,Friedle E.Rapid evolutionary dynamics and disease threats to biodiversity.Trends Ecol Evol,2003,18:589-596
54 Gozlan R E,St-Hilaire S,Feist S W,et al.Disease threat to European fish.Nature,2005,435:1046
55 Pavlovich S S,Lovett S P,Koroleva G,et al.The Egyptian rousette genome reveals unexpected features of bat antiviral immunity.Cell,2018,173:1098-1110.e18
56 Pinsky M L,Worm B,Fogarty M J,et al.Marine taxa track local climate velocities.Science,2013,341:1239-1242
57 Urban M C.Accelerating extinction risk from climate change.Science,2015,348:571-573
58 Scheffers B R,De Meester L,Bridge T C L,et al.The broad footprint of climate change from genes to biomes to people.Science,2016,354:aaf7671
59 Iknayan K J,Beissinger S R.Collapse of a desert bird community over the past century driven by climate change.Proc Natl Acad Sci USA,2018,115:8597-8602
60 Bellard C,Bertelsmeier C,Leadley P,et al.Impacts of climate change on the future of biodiversity.Ecol Lett,2012,15:365-377
61 Phillips S J,Dudík M,Elith J,et al.Sample selection bias and presence-only distribution models:Implications for background and pseudoabsence data.Ecol Appl,2009,19:181-197
62 Chen I C,Hill J K,Ohlemüller R,et al.Rapid range shifts of species associated with high levels of climate warming.Science,2011,333:1024-1026
63 Rinawati F,Stein K,Lindner A.Climate change impacts on biodiversity-The setting of a lingering global crisis.Diversity,2013,5:114-123
64 Karell P,Ahola K,Karstinen T,et al.Climate change drives microevolution in a wild bird.Nat Commun,2011,2:208
65 Frankham R,Ballou J,Briscoe D.Introduction To Conservation Genetics.1st ed.Cambridge:Cambridge University Press,2002
66 Kekkonen J.Evolutionary and Conservation Biology of the Finnish House Sparrow.Dissertation for Doctoral Degree.Helsinki:University of Helsinki,2011.8-9
67 Hartl D L,Clark A G.Principles of Population Genetics,4th ed.Los Angeles:Sinauer Associates,2007
68 Maynard Smith J.Evolutionary Genetics.2nd.Oxford:Oxford University Press,2000
69 Barrett R D H,Schluter D.Adaptation from standing genetic variation.Trends Ecol Evol,2008,23:38-44
70 IPBES.Unedited advance summaries for policymakers of the four regional assessments and the thematic assessment on land degradation and restoration(https://www.ipbes.net/outcomes),2018
2 Wei F W,Nie Y G,Miao H X,et al.Advancements of the researches on biodiversity loss mechanisms(in Chinese).Chin Sci Bull(Chin Ver),2014,59:430-437[魏辅文,聂永刚,苗海霞,等.生物多样性丧失机制研究进展.科学通报,2014,59:430-437]
3 Wei F W.Evolutionary conservation biology of giant pandas(in Chinese).Sci Sin Vitae,2018,48:1048-1053[魏辅文.大熊猫演化保护生物学研究.中国科学:生命科学,2018,48:1048-1053]
4 Ferriere R,Dieckmann U,Couver D.Evolutionary Conservation Biology.New York:Cambridge University Press,2004
5 Zhao S,Zheng P,Dong S,et al.Whole-genome sequencing of giant pandas provides insights into demographic history and local adaptation.Nat Genet,2013,45:67-71
6 Zhou X,Guang X,Sun D,et al.Population genomics of finless porpoises reveal an incipient cetacean species adapted to freshwater.Nat Commun,2018,9:1276
7 Bay R A,Harrigan R J,Underwood V L,et al.Genomic signals of selection predict climate-driven population declines in a migratory bird.Science,2018,359:83-86
8 Kimura M.Evolutionary rate at the molecular level.Nature,1968,217:624-626
9 Levine J M,Hille Ris Lambers J.The importance of niches for the maintenance of species diversity.Nature,2009,461:254-257
10 Pavlogiannis A,Tkadlec J,Chatterjee K,et al.Construction of arbitrarily strong amplifiers of natural selection using evolutionary graph theory.Commun Biol,2018,1:71
11 Zhou X,Sun F,Xu S,et al.Baiji genomes reveal low genetic variability and new insights into secondary aquatic adaptations.Nat Commun,2013,4:2708
12 Zhou X,Wang B,Pan Q,et al.Whole-genome sequencing of the snub-nosed monkey provides insights into folivory and evolutionary history.Nat Genet,2014,46:1303-1310
13 Li H,Durbin R.Inference of human population history from individual whole-genome sequences.Nature,2011,475:493-496
14 Zhu L,Wu Q,Dai J,et al.Evidence of cellulose metabolism by the giant panda gut microbiome.Proc Natl Acad Sci USA,2011,108:17714-17719
15 Wei F,Wu Q,Hu Y,et al.Conservation metagenomics:a new branch of conservation biology.Sci China Life Sci,2019,62:168-178
16 Zhang Z,Xu D,Wang L,et al.Convergent evolution of rumen microbiomes in high-altitude mammals.Curr Biol,2016,26:1873-1879
17 Shao C,Bao B,Xie Z,et al.The genome and transcriptome of Japanese flounder provide insights into flatfish asymmetry.Nat Genet,2017,49:119-124
18 Girod C,Vitalis R,Leblois R,et al.Inferring population decline and expansion from microsatellite data:A simulation-based evaluation of the Msvar method.Genetics,2011,188:165-179
19 Elmer K R,Reggio C,Wirth T,et al.Pleistocene desiccation in East Africa bottlenecked but did not extirpate the adaptive radiation of Lake Victoria haplochromine cichlid fishes.Proc Natl Acad Sci USA,2009,106:13404-13409
20 Fahrig L.Ecological responses to habitat fragmentation Per Se.Annu Rev Ecol Evol Syst,2017,48:1-23
21 Franks S J,Kane N C,O’Hara N B,et al.Rapid genome-wide evolution in Brassica rapa populations following drought revealed by sequencing of ancestral and descendant gene pools.Mol Ecol,2016,25:3622-3631
22 Lindbladh M,Oswald W W,Foster D R,et al.A late-glacial transition from Picea glauca to Picea mariana in southern New England.Quat res,2007,67:502-508
23 Cruzan M B,Templeton A R.Paleoecology and coalescence:Phylogeographic analysis of hypotheses from the fossil record.Trends Ecol Evol,2000,15:491-496
24 Guichoux E,Lagache L,Wagner S,et al.Current trends in microsatellite genotyping.Mol Ecol Res,2011,11:591-611
25 Schiffels S,Durbin R.Inferring human population size and separation history from multiple genome sequences.Nat Genet,2014,46:919-925
26 Hewitt G.The genetic legacy of the Quaternary ice ages.Nature,2000,405:907-913
27 Hewitt G M.Genetic consequences of climatic oscillations in the Quaternary.Philos Trans R Soc London Ser B-Biol Sci,2004,359:183-195
28 Konnert M,Bergmann F.The geographical distribution of genetic variation of silver fir(Abies alba,Pinaceae)in relation to its migration history.Pl Syst Evol,1995,196:19-30
29 Liu Z,Ren B,Wu R,et al.The effect of landscape features on population genetic structure in Yunnan snub-nosed monkeys(Rhinopithecus bieti)implies an anthropogenic genetic discontinuity.Mol Ecol,2009,18:3831-3846
30 Ko A M,Zhang Y,Yang M A,et al.Mitochondrial genome of a 22,000-year-old giant panda from southern China reveals a new panda lineage.Curr Biol,2018,28:R693-R694
31 Li Y,Liu Z,Shi P,et al.The hearing gene Prestin unites echolocating bats and whales.Curr Biol,2010,20:R55-R56
32 Zhao H,Yang J R,Xu H,et al.Pseudogenization of the umami taste receptor gene Tas1r1 in the giant panda coincided with its dietary switch to bamboo.Mol Biol Evol,2010,27:2669-2673
33 Hu Y,Wu Q,Ma S,et al.Comparative genomics reveals convergent evolution between the bamboo-eating giant and red pandas.Proc Natl Acad Sci USA,2017,114:1081-1086
34 Shan L,Wu Q,Wang L,et al.Lineage-specific evolution of bitter taste receptor genes in the giant and red pandas implies dietary adaptation.Integrat Zool,2018,13:152-159
35 Zhu K,Zhou X,Xu S,et al.The loss of taste genes in cetaceans.BMC Evol Biol,2014,14:218
36 Liu Y,Cotton J A,Shen B,et al.Convergent sequence evolution between echolocating bats and dolphins.Curr Biol,2010,20:R53-R54
37 Shen Y Y,Liang L,Li G S,et al.Parallel evolution of auditory genes for echolocation in bats and toothed whales.PLo S Genet,2012,8:e1002788
38 Wei F W.A new perspective on animal adaptive evolution:Animal gut microbiome.Sci Sin Vitae,2016,46:1-3[魏辅文.动物适应性进化研究的新视角:动物肠道微生物组.中国科学:生命科学,2016,46:1-3]
39 Gaynor K M,Hojnowski C E,Carter N H,et al.The influence of human disturbance on wildlife nocturnality.Science,2018,360:1232-1235
40 Fritz S A,Bininda-Emonds O R P,Purvis A.Geographical variation in predictors of mammalian extinction risk:Big is bad,but only in the tropics.Ecol Lett,2009,12:538-549
41 Krauss J,Bommarco R,Guardiola M,et al.Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels.Ecol Lett,2010,13:597-605
42 Sih A,Ferrari M C O,Harris D J.Evolution and behavioural responses to human-induced rapid environmental change.Evolary Appl,2011,4:367-387
43 Barelli C,Albanese D,Donati C,et al.Habitat fragmentation is associated to gut microbiota diversity of an endangered primate:implications for conservation.Sci Rep,2015,5:14862
44 Allendorf F W,England P R,Luikart G,et al.Genetic effects of harvest on wild animal populations.Trends Ecol Evol,2008,23:327-337
45 J?rgensen C,Enberg K,Dunlop E S,et al.Ecology:Managing evolving fish stocks.Science,2007,318:1247-1248
46 Diaz Pauli B,Sih A.Behavioural responses to human-induced change:Why fishing should not be ignored.Evol Appl,2017,10:231-240
47 Andres S,Ribeyre F,Tourencq J N,et al.Interspecific comparison of cadmium and zinc contamination in the organs of four fish species along a polymetallic pollution gradient(Lot River,France).Sci Total Environ,2000,248:11-25
48 Mc Comb B C,Curtis L,Chambers C L,et al.Acute toxic hazard evaluations of glyphosate herbicide on terrestrial vertebrates of the oregon coast range.Environ Sci Pollut Res,2008,15:266-272
49 H?lker F,Wolter C,Perkin E K,et al.Light pollution as a biodiversity threat.Trends Ecol Evol,2010,25:681-682
50 Tuxbury S M,Salmon M.Competitive interactions between artificial lighting and natural cues during seafinding by hatchling marine turtles.Biol Conserv,2005,121:311-316
51 Clarke C A,Mani G S,Wynne G.Evolution in reverse:clean air and the peppered moth.Biol J Linnean Soc,1985,26:189-199
52 Van't Hof A E,Campagne P,Rigden D J,et al.The industrial melanism mutation in British peppered moths is a transposable element.Nature,2016,534:102-105
53 Altizer S,Harvell D,Friedle E.Rapid evolutionary dynamics and disease threats to biodiversity.Trends Ecol Evol,2003,18:589-596
54 Gozlan R E,St-Hilaire S,Feist S W,et al.Disease threat to European fish.Nature,2005,435:1046
55 Pavlovich S S,Lovett S P,Koroleva G,et al.The Egyptian rousette genome reveals unexpected features of bat antiviral immunity.Cell,2018,173:1098-1110.e18
56 Pinsky M L,Worm B,Fogarty M J,et al.Marine taxa track local climate velocities.Science,2013,341:1239-1242
57 Urban M C.Accelerating extinction risk from climate change.Science,2015,348:571-573
58 Scheffers B R,De Meester L,Bridge T C L,et al.The broad footprint of climate change from genes to biomes to people.Science,2016,354:aaf7671
59 Iknayan K J,Beissinger S R.Collapse of a desert bird community over the past century driven by climate change.Proc Natl Acad Sci USA,2018,115:8597-8602
60 Bellard C,Bertelsmeier C,Leadley P,et al.Impacts of climate change on the future of biodiversity.Ecol Lett,2012,15:365-377
61 Phillips S J,Dudík M,Elith J,et al.Sample selection bias and presence-only distribution models:Implications for background and pseudoabsence data.Ecol Appl,2009,19:181-197
62 Chen I C,Hill J K,Ohlemüller R,et al.Rapid range shifts of species associated with high levels of climate warming.Science,2011,333:1024-1026
63 Rinawati F,Stein K,Lindner A.Climate change impacts on biodiversity-The setting of a lingering global crisis.Diversity,2013,5:114-123
64 Karell P,Ahola K,Karstinen T,et al.Climate change drives microevolution in a wild bird.Nat Commun,2011,2:208
65 Frankham R,Ballou J,Briscoe D.Introduction To Conservation Genetics.1st ed.Cambridge:Cambridge University Press,2002
66 Kekkonen J.Evolutionary and Conservation Biology of the Finnish House Sparrow.Dissertation for Doctoral Degree.Helsinki:University of Helsinki,2011.8-9
67 Hartl D L,Clark A G.Principles of Population Genetics,4th ed.Los Angeles:Sinauer Associates,2007
68 Maynard Smith J.Evolutionary Genetics.2nd.Oxford:Oxford University Press,2000
69 Barrett R D H,Schluter D.Adaptation from standing genetic variation.Trends Ecol Evol,2008,23:38-44
70 IPBES.Unedited advance summaries for policymakers of the four regional assessments and the thematic assessment on land degradation and restoration(https://www.ipbes.net/outcomes),2018