生境破碎化对动植物遗传多样性的影响研究进展
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摘要
生境破碎化是生物多样性降低的主要原因之一,此前研究多集中在生境破碎化对物种和生态系统多样性的影响.综述了生境破碎化的成因、度量和理论基础,以及对动植物遗传多样性的影响机理和生态学效应.生境破碎化主要通过增加种群间隔离程度和降低有效种群大小等途径限制种群间基因交流,提高遗传漂变频率,从而影响动植物遗传多样性;但是影响效应取决于植物繁殖策略和动物迁移能力,以及传粉方式、种群大小、生命周期、生境的景观特性等因素;而且生境破碎化对动植物遗传多样性的影响大多存在滞后效应.由此可见,生境破碎化对动植物遗传多样性的影响因素多且复杂,而且不易及时发现,从而忽略生境破碎化对遗传多样性的负面影响;未来应加强生境破碎化对动植物遗传多样性的影响研究,特别是在更长时间尺度上的生境破碎化研究.(表1参73)
Habitat fragmentation(HF) is a key factor in the decrease in biological diversity. Previous studies focused on species and ecosystem diversity, whereas the effects of HF on genetic diversity and its mechanism remains unclear. Here we reviewed the causes, measurements, and theoretical foundations of HF, and the ecological consequences and effects of HF on genetic diversity of plants and animals. Habitat fragmentation affects genetic diversity through increasing isolation distance among populations and decreasing effective population size, with the alteration of gene flow frequency among populations and genetic drift within populations. However, the HF effects depend on the reproductive strategy of plants and the mobility of animals, which interact with other factors, such as mating patterns, population size, life cycle, landscape attributes, and so on. For example, HF significantly affects animals with low mobility, such as small mammals and amphibians, whereas animals with high mobility, such as birds, are less sensitive to HF. Trees can buffer the effects of the genetic drift because of their longevity, but grasses and shrubs with short life cycles cannot. Moreover, there is often a time-delayed effect of HF on genetic diversity. These results indicate that the mechanism of HF on genetic diversity is complicated, and its negative effects are likely to be ignored because of the time-delayed phenomenon. Hence, it is important to study the effects of HF on genetic diversity,especially at a longer time scale.
Habitat fragmentation(HF) is a key factor in the decrease in biological diversity. Previous studies focused on species and ecosystem diversity, whereas the effects of HF on genetic diversity and its mechanism remains unclear. Here we reviewed the causes, measurements, and theoretical foundations of HF, and the ecological consequences and effects of HF on genetic diversity of plants and animals. Habitat fragmentation affects genetic diversity through increasing isolation distance among populations and decreasing effective population size, with the alteration of gene flow frequency among populations and genetic drift within populations. However, the HF effects depend on the reproductive strategy of plants and the mobility of animals, which interact with other factors, such as mating patterns, population size, life cycle, landscape attributes, and so on. For example, HF significantly affects animals with low mobility, such as small mammals and amphibians, whereas animals with high mobility, such as birds, are less sensitive to HF. Trees can buffer the effects of the genetic drift because of their longevity, but grasses and shrubs with short life cycles cannot. Moreover, there is often a time-delayed effect of HF on genetic diversity. These results indicate that the mechanism of HF on genetic diversity is complicated, and its negative effects are likely to be ignored because of the time-delayed phenomenon. Hence, it is important to study the effects of HF on genetic diversity,especially at a longer time scale.
引文
1 Hammond RA,Hudson MD.Environmental management of UK golf courses for biodiversity-attitudes and actions[J].Landscape Urban Plan,2007,83:127-136
2武晶,刘志民.生境破碎化对生物多样性的影响研究综述[J].生态学杂志,2014,33(7):1946-1952[Wu J,Liu ZM.Effect of habitat fragmentation on biodiversity[J].Chin J Ecol,2014,33(7):1946-1952]
3马克平,钱迎倩.生物多样性保护及其研究进展[J].应用与环境生物学报,1998,4(1):95-99[Ma KP,Qian YQ.Biodiversity conservation and it.s research progress[J].Chin J Appl Environ Biol,1998,4(1):95-99]
4武正军,李义明.生境破碎化对动物种群存活的影响[J].生态学报,2003,23(11):2424-2435[Wu ZJ,Li YM.Effects of habitat fragmentation on survival of animal population[J].Acta Ecol Sin,2003,23(11):2424-2435]
5 Myers N,Mittermeier RA,Mittermeier CG,Fonseca GABD,Kent J.Biodiversity hotspots for conservation priorities[J].Nature,2000,403(6772):853-858
6 Wright SJ,Mullerlandau HC.The future of tropical forest species[J].Biotropica,2006,38(3):287-301
7 Wade TG,Riitters KH,Wickham JD,Jones KB.Distribution and causes of global forest fragmentation[J].Conserv Ecol,2003,7(2):1850-1851
8 Haddad NM,Brudvig LA,Clobert J,Davies KF,Gonzalez A,Holt RD,Lovejoy TE,Sexton JO,Austin MP,Collins CD.Habitat fragmentation and its lasting impact on Earth's ecosystems[J].Sci Adv,2015,1(2):e1500052
9 Laurance WF,Camargo JLC,Luizao RCC,Laurance SG,Pimm SL,Bruna EM,Stouffer PC,Bruce WG,Benitezmalvido J,Vasconcelos HL.The fate of Amazonian forest fragments:a 32-year investigation[J].Biol Conserv,2011,144(1):56-67
10 Simpson J,Weiner E.The Oxford English Dictionary Vol X[M].Oxford:Oxford University Press,1989
11 Fahrig L.Rethinking patch size and isolation effects:the habitat amount hy pothesis[J].J Biogeogr,2013,40(9):1649-1663
12 Fahrig L.Eff'ects of habitat fragmentation on biodiversity[J].Annu Rev Ecol Ecol,2003,34(1):487-515
13 Young A,Boyle T,Brown T.The population genetic consequences of habitat fragmentation for plants[J].Trends Ecol Evol,1996,11(10):413-418
14 Xu WH,Wang XZ,Ouyang ZY,Zhang JD,Li ZQ,Xiao Y,Zheng H.Conservation of giant panda habuitat in South Minshan,China,after the May 2008 earthquake[J].Front Ecol Environ,2009,7(7):353-358
15 Iwamura T,Fuller RA.Migratory connectivity magnifies the consequences of habitat loss from sea-level rise for shorebird populations[J].Proc R Soc Lond B Biol,2013,280(1761):20130325
16 Laurance WF,Clements GR,Sloan S,O'Connell CS,Mueller ND,Goosem M,Venter O,Edwards DP,Phalan B,Balmford A.A global strategy for road building[J].Nature,2014,513(7517):229-232
17 Laurance WF,Goosem M,Laurance SG.Impacts of roads and linear clearings on tropical forests[J].Trends Ecol Evol,2009,24(12):659-669
18 Finer M,Jenkins CN.Proliferation of hydroelectric dams in the andean amazon and implications for andes-amazon connectivity[J].P LoS ONE,2012,7(4):e35126
19谭珊珊,胡广,邵德钰,胡仁勇,徐高福,于明坚.千岛湖库区种子植物区系研究[J].广西植物,2010,30(6):770-775[Tan SS,Hu G,Shao DJ,Hu RY,Xu GF,Yu MJ.Floristic analysis of seed plants in the Thousand Island Lake region[J].Guihaia,2010,30(6):770-775]
20 Brunet J,Valtinat K,Mayr ML,Felton A,Lindbladh M,Bruun HH.Understory succession in post-agricultural oak forests:habitat fragmentation affects forest specialists and generalists differently[J].For Ecol Manage,2011,262(9):1863-1871
21 Laurance WF.Theory meets reality:How habitat fragmentation research has transcended island biogeographic theory[J].Biol Conserv,2008,141(7):1731-1744
22吴倩倩,梁宗锁,刘佳佳,于明坚,胡广.中国生境片段化对生物多样性影响研究进展[J].生态学杂志,2017,36(9):2605-2614[Wu QQ,Liang ZS,Liu JJ,Yu MJ,Hu G.Effects of habitat fragmentation on biodiversity in China[J].Chin J Ecol,2017,36(9):2605-2614]
23 Newman BJ,Ladd P,Brundrett M,Dixon KW.Effects of habitat fragmentation on plant reproductive success and population viability at the landscape and habitat scale[J].Biol Conserv,2013,159:16-23
24 Suarez-Esteban A,Fahrig L,Delibes M,Fedriani JM.Can anthropogenic linear gaps increase plant abundance and diversity?[J].La ndscape Ecol,2016,31(4):721-729
25 Parejo-Farnes C,Robledo-Arnuncio JJ,Albaladejo RG,Rubio-Perez E,Aparicio A.Effects of habitat fragmentation on parental correlations in the seed rain of a bird-dispersed species[J].Tree Genet Genome,2017,13(1):17
26 Hermansen TD,Minchinton TE,Ayre DJ.Habitat fragmentation leads to reduced pollinator visitation,fruit production and recruitment in urban mangrove forests[J].Oecologia,2017,185(2):221-231
27 Soons MB,Messelink JH,Jongejans E,Heil GW.Habitat fragmentation reduces grassland connectivity for both short-distance and long-distance wind-dispersed forbs[J].J Ecol,2005,93(6):1214-1225
28 Zhang XM,Gao LM,Moeller M,Li DZ.Molecular evidence for fragmentation among populations of Taxus wallichiana var.mairei,a highly endangered conifer in China[J].Can J For Res,2009,39(4):755-764
29 Browne L,Karubian J.Habitat loss and fragmentation reduce effective gene flow by disrupting seed dispersal in a neotropical palm[J].Mol Ecol,2018,27:3055-3069
30 Broeck AV,Tobias C,Gunter K,Maurice H,Olivier H,Joachim M.Dispersal constraints for the conservation of the grassland herb Thymus pulegioides L.in a highly fragmented agricultural landscape[J].Conserv Genet,2015,16(4):765-776
31 Bijlsma R,Loeschcke V.Genetic erosion impedes adaptive responses to stressful environments[J].Evol Appl,2012,5(2):117-129
32 Lienert J.Habitat fragmentation effects on fitness of plant populations一a review[J].J Nat Conserv,2004,12(1):53-72
33 Luquet E,Garner TWJ,Lena JP,Bruel C,Joly P,Lengagne T,Grolet O,Plenet S.Genetic erosion in wild populations makes resistance to a pathogen more costly[J].Evolution,2012,66(6):1942-1952
34 Vandepitte K,Jacquemyn H,Roldan-Ruiz I,Honnay O.Landscape genetics of the self-compatible forest herb Geum urbanum:effects of habitat age,fragmentation and local environ ment[J].Mol Ecol,2007,16(19):4171-4179
35 Rusterholz HP,Baur B.Delayed response in a plant-pollinator system to experimental grassland fragmentation[J].Oecologia,2010,163(1):141-152
36 Heinken T,Weber E.Consequences of habitat fragmentation for plant species:Do we know enough?[J].Perspect Plant Ecol,2013,15(4):205-216
37 Wiberg RAW,Scobie AR,Ahara S,Ennos RA,Cottrell J.The genetic consequences of long term habitat fragmentation on a self-incompatibleclonal plant,Linnaea borealis L[J].Biol Conser v,2016,20 1:405-413
38 Breed MF,Ottewell KM,Gardner MG,Marklund MHK,Dormott EE,Lowe AJ.Mating patterns and pollinator mobility are critical traits in forest fragmentation genetics[J].Heredity,2015,115(2):108-114
39 Battocletti A.The population and ecological genetic effects of habitat fragmentation[D].Washington DC:Georgetown Vniversity,2017
40 Zhang X,Shi MM,Shen DW,Chen XY.Habitat loss other than fragmentation per se decreased nuclear and chloroplast genetic diversity in a monoecious tree[J].PLoS ONE,2012,7(6):e39146
41 Lowe AJ,Boshier D,Ward M,Bacles CF,Navarro C.Genetic resource impacts of habitat loss and degradation;reconciling empirical evidence and predicted theory for neotropical trees[J].Heredity,2005,95(4):255-273
42 Holderegger R,Giulio MD.The genetic effects of roads:a review of empirical evidence[J].Basic Appl Ecol,2010,11(6):522-531
43 Rivera-Ortiz FA,Aguilar R,Arizmendi MDC,Quesada-Avendano M,Oyama K.Habitat fragmentation and genetic variability of tetrapod populations[J].Anim Conserv,2015,18(3):249-258
44 Marsh DM,Page RB,Hanlon TJ,Corritone R,Little EC,Seifert DE,Cabe PR.Effects of roads on patterns of genetic differentiation in redbacked salamanders,Plethodon cinereus[J].Conserv Genet,2008,9(3):603-613
45 Buchmann CM,Schurr FM,Ran N,Jeltsch F.Habitat loss and fragmentation affecting mammal and bird communities—The role of interspecific competition and individual space use[J].Ecol Inform,2013,14:90-98
46 Canales-Delgadillo J,Scott-Morales L,Korb J.The influence of habitat fragmentation on genetic diversity of a rare bird species that commonly faces environmental fluctuations[J].J Avian Biol,2012,43(2):168-176
47 Dixo M,Metzger JP,Morgante JS,Zamudio KR.Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest[J].Biol Conserv,2009,142(8):1560-1569
48 Garridogarduno T,Tellezvaldes O,Manel S,Vazquezdominguez E.Role of habitat heterogeneity and landscape connectivity in shaping gene flow and spatial population structure of a dominant rodent species in a tropical dry forest[J].J Zool,2016,298(4):293-302
49 Moniem HEMA,Schemerhorn BJ,Dewoody JA,Holland JD.Landscape genetics of a pollinator longhorn beetle[Typocerus velutinus(Olivier)]on a continuous habitat surface[J].Mol Ecol,2016,25(20):5015-5028
50 Schuler MS,Chase JM,Knight TM.Habitat size modulates the influence of heterogeneity on species richness patterns in a model zooplankton community[J].Ecology,2017,98(6):1651-1659
51 Jangjoo M,Matter SF,Roland J,Keyghobadi N.Connectivity rescues genetic diversity after a demographic bottleneck in a butterfly population network[J].PNAS,2016,113(39):10914-10919
52 Munoz PT,Torres FP,Megias AG.Effects of roads on insects:a review[J].Biodiv Conser v,2015,24(3):659-682
53 Crawford JA,Peterman WE,Kuhns AR,Eggert LS.Altered functional connectivity and genetic diversity of a threatened salamander in an agroecosystem[J].La ndscape Ecol,2016,31(10):2231-2244
54 Heinenkay JL,Noel HG,Layman CA,Langerhans RB.Human-caused habitat fragmentation can drive rapid divergence of male genitalia[J].Evol Appl,2014,7(10):1252-1267
55 Vandergast AG,Wood DA,Thompson AR,Fisher M,Barrows CW,Grant TJ.Drifting to oblivion?Rapid genetic differentiation in an endangered lizard following habitat fragmentation and drought[J].Divers Distrib,2016,22(3):344-357
56 Kuussaari M,Bommarco R,Heikkinen RK,Helm A,Krauss J,Lindborg R,Ockinger E,Partel M,Pino J,Roda F.Extinction debt:a challenge for biodiversity conservation[J].Trends Ecol Evol,2009,24(10):564-571
57 Essl F,Dullinger S,Rabitsch W,Hulme PE,Pysek P,Wilson JRL,Richardson DM.Delayed biodiversity change:no time to waste[J].Trends Ecol Evol,2015,30(7):375-378
58 Garcia MB,Dahlgren JP,Ehrlen J.No evidence of senescence in a300-year-old mountain herb[J].J Ecol,2011,99(6):1424-1430
59 Mergeay J,Vanoverbeke J,Verschuren D,De Meester L.Extinction,recolonization,and dispersal through time in a planktonic crustacean[J].Ecology,2007,88(12):3032-3043
60 Lee AM,Saether B,Engen S.Demographic stochasticity,allee effects,and extinction:the influence of mating system and sex ratio[J].Am Na t.2011,177(3):301-313
61 Hanski I,Ovaskainen O.Extinction debt at extinction threshold[J].Conserv Biol,2002,16(3):666-673
62 Otsu C,Lijima H,Nagaike T,Hoshino Y.Evidence of extinction debt through the survival and colonization of each species in semi-natural grasslands[J].J Veg Sci,2017,28(3):464-474
63 Vranckx G,Jacquemyn H,Muys B,Honnay O.Meta-analysis of susceptibility of woody plants to loss of genetic diversity through habitat fragmentation[J].Conserv Biol,2012,26(2):228-237
64 Otto R,Garzonmachado V,Arco MD,Fernandezlugo S,De Nascimentc L,Oromi P,Baez M,Ibanez M,Alonso MR,Fernandezpalacios JM.Unpaid extinction debts for endemic plants and invertebrates as a legacy of habitat loss on oceanic islands[J].Divers Distrib,2017,23(9):1031-1041
65 Mona S,Arenas M,Excoffier L.Genetic consequences of habitat fragmentation during a range expansion[J].Heredity,2014,112(3):291-299
66 Krauss J,Bommarco R,Guardiola M,Heikkinen RK,Helm A.Kuussaari M,Lindborg R,Ockinger E,Partel M,Pino J,Poyry J.Raatikainen KM,Sang A,Stefanescu C,Teder T,Zobel M,SteffanDewenter I.Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels[J].Ecol Lett,2010,13(5):597-605
67 Bommarco R,Lindborg R,Marini L,Ockinger E.Extinction debt for plants and flower-visiting insects in landscapes with contrasting land use history[J].Divers Distrib,2014,20(5):591-599
68 Lindborg R,Eriksson O.Historical landscape connectivity affects present plant species diversity[J].Ecology,2004,85(7):1840-1845
69 Vellend M,Verheyen K,Jacquemyn H,Kolb A,Van Calster H,Peterken G,Hermy M.Extinction debt of forest plants persists for more than a century following habitat fragmentation[J].Ecology,2006,87(3):542-548
70 Saar L,Takkis K,Partel M,Helm A.Which plant traits predict species loss in calcareous grasslands with extinction debt[J].Divers Distrib.2012,18(8):808-817
71 Alofs KM,Gonzalez AV,Fowler NL.Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales[J].Plant Ecol,2014,215(10):1139-1151
72 Ortego J,Bonal R,Munoz A.Genetic consequences of habital fragmentation in long-lived tree species:the case of the Mediterranean holm oak(Quercus ilex L.)[J].J Hered,2010,101(6):717-726
73 Eva G,Kevin M,Van Karine D,Hilde D,Xiang L,Konj ev D,Frederik H.Loss of genetic diversity and increased genetic structuring in response to forest area reduction in a ground dwelling insect:a case study of the flightless carabid beetle Carabus problematicus(Coleoptera,Carabidae)[J].Insect Conserv Diver,2013,6(4):473-482
2武晶,刘志民.生境破碎化对生物多样性的影响研究综述[J].生态学杂志,2014,33(7):1946-1952[Wu J,Liu ZM.Effect of habitat fragmentation on biodiversity[J].Chin J Ecol,2014,33(7):1946-1952]
3马克平,钱迎倩.生物多样性保护及其研究进展[J].应用与环境生物学报,1998,4(1):95-99[Ma KP,Qian YQ.Biodiversity conservation and it.s research progress[J].Chin J Appl Environ Biol,1998,4(1):95-99]
4武正军,李义明.生境破碎化对动物种群存活的影响[J].生态学报,2003,23(11):2424-2435[Wu ZJ,Li YM.Effects of habitat fragmentation on survival of animal population[J].Acta Ecol Sin,2003,23(11):2424-2435]
5 Myers N,Mittermeier RA,Mittermeier CG,Fonseca GABD,Kent J.Biodiversity hotspots for conservation priorities[J].Nature,2000,403(6772):853-858
6 Wright SJ,Mullerlandau HC.The future of tropical forest species[J].Biotropica,2006,38(3):287-301
7 Wade TG,Riitters KH,Wickham JD,Jones KB.Distribution and causes of global forest fragmentation[J].Conserv Ecol,2003,7(2):1850-1851
8 Haddad NM,Brudvig LA,Clobert J,Davies KF,Gonzalez A,Holt RD,Lovejoy TE,Sexton JO,Austin MP,Collins CD.Habitat fragmentation and its lasting impact on Earth's ecosystems[J].Sci Adv,2015,1(2):e1500052
9 Laurance WF,Camargo JLC,Luizao RCC,Laurance SG,Pimm SL,Bruna EM,Stouffer PC,Bruce WG,Benitezmalvido J,Vasconcelos HL.The fate of Amazonian forest fragments:a 32-year investigation[J].Biol Conserv,2011,144(1):56-67
10 Simpson J,Weiner E.The Oxford English Dictionary Vol X[M].Oxford:Oxford University Press,1989
11 Fahrig L.Rethinking patch size and isolation effects:the habitat amount hy pothesis[J].J Biogeogr,2013,40(9):1649-1663
12 Fahrig L.Eff'ects of habitat fragmentation on biodiversity[J].Annu Rev Ecol Ecol,2003,34(1):487-515
13 Young A,Boyle T,Brown T.The population genetic consequences of habitat fragmentation for plants[J].Trends Ecol Evol,1996,11(10):413-418
14 Xu WH,Wang XZ,Ouyang ZY,Zhang JD,Li ZQ,Xiao Y,Zheng H.Conservation of giant panda habuitat in South Minshan,China,after the May 2008 earthquake[J].Front Ecol Environ,2009,7(7):353-358
15 Iwamura T,Fuller RA.Migratory connectivity magnifies the consequences of habitat loss from sea-level rise for shorebird populations[J].Proc R Soc Lond B Biol,2013,280(1761):20130325
16 Laurance WF,Clements GR,Sloan S,O'Connell CS,Mueller ND,Goosem M,Venter O,Edwards DP,Phalan B,Balmford A.A global strategy for road building[J].Nature,2014,513(7517):229-232
17 Laurance WF,Goosem M,Laurance SG.Impacts of roads and linear clearings on tropical forests[J].Trends Ecol Evol,2009,24(12):659-669
18 Finer M,Jenkins CN.Proliferation of hydroelectric dams in the andean amazon and implications for andes-amazon connectivity[J].P LoS ONE,2012,7(4):e35126
19谭珊珊,胡广,邵德钰,胡仁勇,徐高福,于明坚.千岛湖库区种子植物区系研究[J].广西植物,2010,30(6):770-775[Tan SS,Hu G,Shao DJ,Hu RY,Xu GF,Yu MJ.Floristic analysis of seed plants in the Thousand Island Lake region[J].Guihaia,2010,30(6):770-775]
20 Brunet J,Valtinat K,Mayr ML,Felton A,Lindbladh M,Bruun HH.Understory succession in post-agricultural oak forests:habitat fragmentation affects forest specialists and generalists differently[J].For Ecol Manage,2011,262(9):1863-1871
21 Laurance WF.Theory meets reality:How habitat fragmentation research has transcended island biogeographic theory[J].Biol Conserv,2008,141(7):1731-1744
22吴倩倩,梁宗锁,刘佳佳,于明坚,胡广.中国生境片段化对生物多样性影响研究进展[J].生态学杂志,2017,36(9):2605-2614[Wu QQ,Liang ZS,Liu JJ,Yu MJ,Hu G.Effects of habitat fragmentation on biodiversity in China[J].Chin J Ecol,2017,36(9):2605-2614]
23 Newman BJ,Ladd P,Brundrett M,Dixon KW.Effects of habitat fragmentation on plant reproductive success and population viability at the landscape and habitat scale[J].Biol Conserv,2013,159:16-23
24 Suarez-Esteban A,Fahrig L,Delibes M,Fedriani JM.Can anthropogenic linear gaps increase plant abundance and diversity?[J].La ndscape Ecol,2016,31(4):721-729
25 Parejo-Farnes C,Robledo-Arnuncio JJ,Albaladejo RG,Rubio-Perez E,Aparicio A.Effects of habitat fragmentation on parental correlations in the seed rain of a bird-dispersed species[J].Tree Genet Genome,2017,13(1):17
26 Hermansen TD,Minchinton TE,Ayre DJ.Habitat fragmentation leads to reduced pollinator visitation,fruit production and recruitment in urban mangrove forests[J].Oecologia,2017,185(2):221-231
27 Soons MB,Messelink JH,Jongejans E,Heil GW.Habitat fragmentation reduces grassland connectivity for both short-distance and long-distance wind-dispersed forbs[J].J Ecol,2005,93(6):1214-1225
28 Zhang XM,Gao LM,Moeller M,Li DZ.Molecular evidence for fragmentation among populations of Taxus wallichiana var.mairei,a highly endangered conifer in China[J].Can J For Res,2009,39(4):755-764
29 Browne L,Karubian J.Habitat loss and fragmentation reduce effective gene flow by disrupting seed dispersal in a neotropical palm[J].Mol Ecol,2018,27:3055-3069
30 Broeck AV,Tobias C,Gunter K,Maurice H,Olivier H,Joachim M.Dispersal constraints for the conservation of the grassland herb Thymus pulegioides L.in a highly fragmented agricultural landscape[J].Conserv Genet,2015,16(4):765-776
31 Bijlsma R,Loeschcke V.Genetic erosion impedes adaptive responses to stressful environments[J].Evol Appl,2012,5(2):117-129
32 Lienert J.Habitat fragmentation effects on fitness of plant populations一a review[J].J Nat Conserv,2004,12(1):53-72
33 Luquet E,Garner TWJ,Lena JP,Bruel C,Joly P,Lengagne T,Grolet O,Plenet S.Genetic erosion in wild populations makes resistance to a pathogen more costly[J].Evolution,2012,66(6):1942-1952
34 Vandepitte K,Jacquemyn H,Roldan-Ruiz I,Honnay O.Landscape genetics of the self-compatible forest herb Geum urbanum:effects of habitat age,fragmentation and local environ ment[J].Mol Ecol,2007,16(19):4171-4179
35 Rusterholz HP,Baur B.Delayed response in a plant-pollinator system to experimental grassland fragmentation[J].Oecologia,2010,163(1):141-152
36 Heinken T,Weber E.Consequences of habitat fragmentation for plant species:Do we know enough?[J].Perspect Plant Ecol,2013,15(4):205-216
37 Wiberg RAW,Scobie AR,Ahara S,Ennos RA,Cottrell J.The genetic consequences of long term habitat fragmentation on a self-incompatibleclonal plant,Linnaea borealis L[J].Biol Conser v,2016,20 1:405-413
38 Breed MF,Ottewell KM,Gardner MG,Marklund MHK,Dormott EE,Lowe AJ.Mating patterns and pollinator mobility are critical traits in forest fragmentation genetics[J].Heredity,2015,115(2):108-114
39 Battocletti A.The population and ecological genetic effects of habitat fragmentation[D].Washington DC:Georgetown Vniversity,2017
40 Zhang X,Shi MM,Shen DW,Chen XY.Habitat loss other than fragmentation per se decreased nuclear and chloroplast genetic diversity in a monoecious tree[J].PLoS ONE,2012,7(6):e39146
41 Lowe AJ,Boshier D,Ward M,Bacles CF,Navarro C.Genetic resource impacts of habitat loss and degradation;reconciling empirical evidence and predicted theory for neotropical trees[J].Heredity,2005,95(4):255-273
42 Holderegger R,Giulio MD.The genetic effects of roads:a review of empirical evidence[J].Basic Appl Ecol,2010,11(6):522-531
43 Rivera-Ortiz FA,Aguilar R,Arizmendi MDC,Quesada-Avendano M,Oyama K.Habitat fragmentation and genetic variability of tetrapod populations[J].Anim Conserv,2015,18(3):249-258
44 Marsh DM,Page RB,Hanlon TJ,Corritone R,Little EC,Seifert DE,Cabe PR.Effects of roads on patterns of genetic differentiation in redbacked salamanders,Plethodon cinereus[J].Conserv Genet,2008,9(3):603-613
45 Buchmann CM,Schurr FM,Ran N,Jeltsch F.Habitat loss and fragmentation affecting mammal and bird communities—The role of interspecific competition and individual space use[J].Ecol Inform,2013,14:90-98
46 Canales-Delgadillo J,Scott-Morales L,Korb J.The influence of habitat fragmentation on genetic diversity of a rare bird species that commonly faces environmental fluctuations[J].J Avian Biol,2012,43(2):168-176
47 Dixo M,Metzger JP,Morgante JS,Zamudio KR.Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest[J].Biol Conserv,2009,142(8):1560-1569
48 Garridogarduno T,Tellezvaldes O,Manel S,Vazquezdominguez E.Role of habitat heterogeneity and landscape connectivity in shaping gene flow and spatial population structure of a dominant rodent species in a tropical dry forest[J].J Zool,2016,298(4):293-302
49 Moniem HEMA,Schemerhorn BJ,Dewoody JA,Holland JD.Landscape genetics of a pollinator longhorn beetle[Typocerus velutinus(Olivier)]on a continuous habitat surface[J].Mol Ecol,2016,25(20):5015-5028
50 Schuler MS,Chase JM,Knight TM.Habitat size modulates the influence of heterogeneity on species richness patterns in a model zooplankton community[J].Ecology,2017,98(6):1651-1659
51 Jangjoo M,Matter SF,Roland J,Keyghobadi N.Connectivity rescues genetic diversity after a demographic bottleneck in a butterfly population network[J].PNAS,2016,113(39):10914-10919
52 Munoz PT,Torres FP,Megias AG.Effects of roads on insects:a review[J].Biodiv Conser v,2015,24(3):659-682
53 Crawford JA,Peterman WE,Kuhns AR,Eggert LS.Altered functional connectivity and genetic diversity of a threatened salamander in an agroecosystem[J].La ndscape Ecol,2016,31(10):2231-2244
54 Heinenkay JL,Noel HG,Layman CA,Langerhans RB.Human-caused habitat fragmentation can drive rapid divergence of male genitalia[J].Evol Appl,2014,7(10):1252-1267
55 Vandergast AG,Wood DA,Thompson AR,Fisher M,Barrows CW,Grant TJ.Drifting to oblivion?Rapid genetic differentiation in an endangered lizard following habitat fragmentation and drought[J].Divers Distrib,2016,22(3):344-357
56 Kuussaari M,Bommarco R,Heikkinen RK,Helm A,Krauss J,Lindborg R,Ockinger E,Partel M,Pino J,Roda F.Extinction debt:a challenge for biodiversity conservation[J].Trends Ecol Evol,2009,24(10):564-571
57 Essl F,Dullinger S,Rabitsch W,Hulme PE,Pysek P,Wilson JRL,Richardson DM.Delayed biodiversity change:no time to waste[J].Trends Ecol Evol,2015,30(7):375-378
58 Garcia MB,Dahlgren JP,Ehrlen J.No evidence of senescence in a300-year-old mountain herb[J].J Ecol,2011,99(6):1424-1430
59 Mergeay J,Vanoverbeke J,Verschuren D,De Meester L.Extinction,recolonization,and dispersal through time in a planktonic crustacean[J].Ecology,2007,88(12):3032-3043
60 Lee AM,Saether B,Engen S.Demographic stochasticity,allee effects,and extinction:the influence of mating system and sex ratio[J].Am Na t.2011,177(3):301-313
61 Hanski I,Ovaskainen O.Extinction debt at extinction threshold[J].Conserv Biol,2002,16(3):666-673
62 Otsu C,Lijima H,Nagaike T,Hoshino Y.Evidence of extinction debt through the survival and colonization of each species in semi-natural grasslands[J].J Veg Sci,2017,28(3):464-474
63 Vranckx G,Jacquemyn H,Muys B,Honnay O.Meta-analysis of susceptibility of woody plants to loss of genetic diversity through habitat fragmentation[J].Conserv Biol,2012,26(2):228-237
64 Otto R,Garzonmachado V,Arco MD,Fernandezlugo S,De Nascimentc L,Oromi P,Baez M,Ibanez M,Alonso MR,Fernandezpalacios JM.Unpaid extinction debts for endemic plants and invertebrates as a legacy of habitat loss on oceanic islands[J].Divers Distrib,2017,23(9):1031-1041
65 Mona S,Arenas M,Excoffier L.Genetic consequences of habitat fragmentation during a range expansion[J].Heredity,2014,112(3):291-299
66 Krauss J,Bommarco R,Guardiola M,Heikkinen RK,Helm A.Kuussaari M,Lindborg R,Ockinger E,Partel M,Pino J,Poyry J.Raatikainen KM,Sang A,Stefanescu C,Teder T,Zobel M,SteffanDewenter I.Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels[J].Ecol Lett,2010,13(5):597-605
67 Bommarco R,Lindborg R,Marini L,Ockinger E.Extinction debt for plants and flower-visiting insects in landscapes with contrasting land use history[J].Divers Distrib,2014,20(5):591-599
68 Lindborg R,Eriksson O.Historical landscape connectivity affects present plant species diversity[J].Ecology,2004,85(7):1840-1845
69 Vellend M,Verheyen K,Jacquemyn H,Kolb A,Van Calster H,Peterken G,Hermy M.Extinction debt of forest plants persists for more than a century following habitat fragmentation[J].Ecology,2006,87(3):542-548
70 Saar L,Takkis K,Partel M,Helm A.Which plant traits predict species loss in calcareous grasslands with extinction debt[J].Divers Distrib.2012,18(8):808-817
71 Alofs KM,Gonzalez AV,Fowler NL.Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales[J].Plant Ecol,2014,215(10):1139-1151
72 Ortego J,Bonal R,Munoz A.Genetic consequences of habital fragmentation in long-lived tree species:the case of the Mediterranean holm oak(Quercus ilex L.)[J].J Hered,2010,101(6):717-726
73 Eva G,Kevin M,Van Karine D,Hilde D,Xiang L,Konj ev D,Frederik H.Loss of genetic diversity and increased genetic structuring in response to forest area reduction in a ground dwelling insect:a case study of the flightless carabid beetle Carabus problematicus(Coleoptera,Carabidae)[J].Insect Conserv Diver,2013,6(4):473-482