21. Lect_21_Community_Biodiv_1 . ( ) ( ). . . . . : . . . ,

: ,

R.H. MacArthur and E.O. Wilson

The theory of island biogeography

1967

Princeton University Press

Robert MacArthur (1930-1972)Edward O. Wilson (b. 1929)

.

. , dispersal assembled niche assembled

:

. (J. Diamond)

, , ( , )

- ( , )

-

, ( 50 ):

(-) - 316 269 823 - 1186 ( 52 ) 1123 ( 25 !!!)

: Wills . Harms K.E., Condit R. et al. 2006. Nonrandom processes maintain diversity in tropical forests // Science. Vol. 311. P. 527 - 531

, , , .

( - ) ,

. , , . -

(), . www.ecuador-travel.net

. 1. A . ( 2 ), ( ) . , , , , ( ). 20 20 . B , , 20 20 ( observed quadrat) , ( null expectation), . , , , , . , , . . Science ( 2 ), ( ) . , , , ( ).

( Y) ( ). Peter B. Adler, Janneke HilleRisLambers and Jonathan M. Levine. A niche for neutrality // Ecology Letters. 2007. V. 10 (2). P. 95104 .. // . 2007. N 11. . 29-37. http://rogov.zwz.ru/Macroevolution/ghilarov_neutral.pdf

, : Peter B. Adler, Janneke HilleRisLambers and Jonathan M. Levine. A niche for neutrality // Ecology Letters. 2007. V. 10 (2). P. 95104

?

?

- , . , .

:

, ( ) . ()

2010 . Science Nature , . : . 25.08.10 Elementy.ru: http://elementy.ru/news/431384

Liza S. Comita, Helene C. Muller-Landau, Salomn Aguilar, Stephen P. Hubbell

Asymmetric density dependence shapes species abundances in a tropical tree community // Science. 2010. V. 329. P. 330332

50 , , 5 ( 1 ), ( 20 )

1 1 2, . 30 . , 180

, - 180 , ,

, ,

Liza S. Comita et al. 2010 // Science. V. 329. P. 330332 . ,

( ),

Scott A. Mangan, Stefan A. Schnitzer, Edward A. Herre, Keenan M. L. Mack, Mariana C. Valencia, Evelyn I. Sanchez, James D. Bever

Negative plantsoil feedback predicts tree-species relative abundance in a tropical forest // Nature. 2010. V. 466. P. 752755

, . -

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BIODIVERSITY

Wenzel Peter, 1745 1829. Adamo ed Eva nel Paradiso Terrestre Pinacotecca Vatican

, biodiversity

Trends in Ecology and Evolution, 1996, Vol. 11 (7): 304-306

( !) , 20000

Luehea seemanniiTerry L. Erwin

Terry L. Erwin

Terry Erwin (1982): () Luehea seemannii 1200 , - 163 50 000 8 150 000

40% . 20

, 30

- , 4-5

, , . Novotny V., Basset Y., Miller S., et al. // Nature. 2002. V. 416. P. 841-844

51 , -13 (Ficus) 4 Psychotria. 50 . , 935 , , (),

:

32.9 1.7 ,

26.1 2.4 (),

20.6 0.9

( ), :

7.9 , 13.3 2.9

() ! , 1, 2, 3, 4 ( 14) (A) (B). Lep-L Lepidoptera (, ) Hym-L (, ), Col-L (; ), Col-A - ; , Ort-A

Errata! 3- 0.01

1

3.2

3.52

4.04

4.36

4.56

5.04

4.4

3.72

0.7

lg

(: Fenchel, 1993)

1

0.1 3.2

1 3.52

0.01 4.04

0.1 4.36

1 4.56

1 5.04

10 4.4

1 3.72

10 0.7

1

0

0

0

0

0

0

0

0

0

lg

(: Fenchel, 1993)

2

&A

&P

3

&A

&P

4

&A

&P

5

&A

&P

6

&A

&P

7

&A

&P

8

&A

&P

9

&A

&P

10

&A

&P

11

&A

&P

12

&A

&P

13

&A

&P

14

&A

&P

15

&A

&P

16

&A

&P

1

4.2

5.35

5.8

5.1

4.05

3.85

2.9

2.3

0.8

lg

(: May, 1988)

1

0.3 - 1 4.2

1 - 3.2 5.35

3.2 -10 5.8

1- 3.2 5.1

3.2-10 4.05

10-32 3.85

32-100 2.9

1 -3.2 2.3

3.2 - 10 0.8

1

lg

(: May, 1988)

2

&A

&P

3

&A

&P

4

&A

&P

5

&A

&P

6

&A

&P

7

&A

&P

8

&A

&P

9

&A

&P

10

&A

&P

11

&A

&P

12

&A

&P

13

&A

&P

14

&A

&P

15

&A

&P

16

&A

&P

.. : ? : ?

1

400003000

VERTEBRATA

CILIATA

1

12

400003000

15001500

5100

1

00

VERTEBRATA

CILIATA

2

00

VERTEBRATA

CILIATA

3

00

VERTEBRATA

CILIATA

100 2

2

15001500

VERTEBRATA

CILIATA

1

12

400003000

15001500

5100

1

00

VERTEBRATA

CILIATA

2

00

VERTEBRATA

CILIATA

3

00

VERTEBRATA

CILIATA

100 2

3

5100

VERTEBRATA

CILIATA

100 2

1

12

400003000

15001500

5100

1

00

VERTEBRATA

CILIATA

2

00

VERTEBRATA

CILIATA

3

00

VERTEBRATA

CILIATA

100 2

: ? : ? ..

1

5100

15001200

400003000

VERTEBRATES

CILIATES

1

AREASvSc

5.00E+14400003000

1.00E+1115001500

1.00E+045100

1.00E+00

1.00E+045100

North Europe1.00E+1115001200

The World5.00E+14400003000

1

VERTEBRATES

CILIATES

2

3

..

1

1.8751.333333333313.09523809521222.7272727273

0.6250.1254.76190476191015.4090909091

0.0250.05833333331.19047619054.44.0909090909

0.0093750.03333333330.47619047621.83.4090909091

0.0050.09166666670.26190476191.22.7272727273

0.0006254.73333333330.1190476190.613.6363636364

0.021250.61.23809523814.967.5454545455

0.31251.33333333332.380952381813.6363636364

0.0531252.55.95238095242.6815.6363636364

0.1368750.6517.53.244.8181818182

0.481250.82510.85714285717.2413.1818181818

1.093750.06666666670.76190476195.845.6818181818

1.6156250.33333333331.19047619051.642.8181818182

0.16750.11.071428571444.9545454545

0.076250.43333333330.14285714292.67.6363636364

0.083750.45833333330.23809523813.46.3636363636

0.691250.18333333330.26190476193.45.8636363636

0.5850.250.30952380953.42.9090909091

0.203750.09166666670.26190476193.481.7727272727

0.413750.4250.3809523811.9210.4545454545

0.443750.5255.595238095212.721.1818181818

0.93750.98333333332.7142857143104.1818181818

0.1506250.81666666673.97619047622.5909090909

0.0150.74166666672.85714285712.2727272727

0.38750.351.57142857148.9545454545

0.12250.50.78571428572.5363636364

0.0506250.48.9090909091

0.1931250.541666666714.2727272727

0.0343750.258333333321.6363636364

0.00250.3252.8636363636

0.0068750.4538.3181818182

0.018752.716666666711

0.03750.2608333333

0.093751.9083333333

0.1093752.6916666667

0.068750.575

0.0918750.5583333333

0.1656250.0866666667

0.011250.9416666667

0.03750.9583333333

0.043750.4833333333

0.0618750.775

0.07375

0.04375

0.045625

0.021875

0.015

0.088125

0.0725

0.096875

0.136875

0.051875

0.083125

0.075625

0.0925

0.05

0.084375

0.035625

0.034375

0.074375

0.00875

0.254375

0.039375

0.026875

0.003125

0.0075

0.625

(2)

(%)

Ris43

. 43. , (S'),

N ( ). ) ;

) ; ) ; ) ; ) ;

) . .

(D - ).

Ris43

1.8815653696

1.9059848046

1.9237933832

1.9201794939

2.0156709149

2.0474435232

1.9860330529

2.0579051114

1.9480033716

1.9901586178

1.991150562

1.8839869146

2.0472080092

1.8994948937

2.1516763219

2.0369567504

2.0597609955

1.9993218301

1.9856421329

2.0938205643

2.233727836

lg N

lg S'

Ris44

0.6478276701

0.6001467785

0.5859624158

0.8000257795

0.6330944486

0.729278014

0.8143965023

0.7100546897

0.9358337329

1.1805453655

0.6833582124

0.7603299246

0.9554304934

1.1186224164

0.5631818378

0.6277389923

0.7105407514

0.9156838075

0.9633926852

0.6076849134

0.7575853461

0.9675348724

0.6391845897

0.7835192526

0.9629257528

1.0992725051

0.8853926852

0.8859657882

0.9828758012

0.939080662

1.0773833601

1.102973348

1.1513861365

lg N

lg S'

Ris 45

0.8281916795

1.1126650254

1.0349616679

1.1146694218

1.2055702739

1.0890271321

1.1890136242

1.2718291549

1.0955354695

1.1953072803

1.2367890994

1.2844307338

1.137690199

1.2039850386

1.2654031354

1.1607058903

1.2307809503

1.3336222236

1.2904565454

1.3828043062

1.3457671266

1.5169708448

1.7445719233

lg N

lg S'

Ris46

1.3853285766

1.3747132391

1.4849571888

1.4620663497

1.4982873435

1.2307607526

1.3994279936

1.3991280087

1.4448854992

1.4690191789

1.3222865871

1.4010180201

1.5116600582

1.5494540058

1.5708521505

1.5428230807

1.6384216852

1.5824651957

1.6888696277

1.7169221765

lg N

lg S'

Ris47

0.70667769070.88500826630.7066776907

0.8928525251.01536491160.892852525

0.78718063931.10288238450.7871806393

0.87254726231.14455268560.8725472623

0.9555138680.99886048990.955513868

0.90574761731.11128168210.9057476173

0.78246542681.3237244140.7824654268

0.87472300121.32748178520.8747230012

0.76685392921.05470188130.7668539292

0.85949915031.13688863750.8594991503

0.99564400281.17657201360.9956440028

1.10893268461.21321251381.1089326846

0.77301205750.7730120575

0.84764567580.8476456758

0.94910331660.9491033166

1.0002558391.000255839

1.05842870431.0584287043

0.96094363910.9609436391

1.06788332481.0678833248

0.96179589690.9617958969

1.0791812461.079181246

1.13830269821.1383026982

1.18326984371.1832698437

0.82577027410.8257702741

0.87147146150.8714714615

0.95635835860.9563583586

1.01767188571.0176718857

1.14261062231.1426106223

0.96469060640.9646906064

1.04657774591.0465777459

1.09822951481.0982295148

0.77441301070.7744130107

0.90410584890.9041058489

1.0185146531.018514653

0.97603414870.9760341487

1.05199975411.0519997541

1.14208711711.1420871171

1.10389743841.1038974384

0.8850082663

1.0153649116

1.1028823845

1.1445526856

0.9988604899

1.1112816821

1.323724414

1.3274817852

1.0547018813

1.1368886375

1.1765720136

1.2132125138

D30 m

lg N

lg S'

1.42829066051.68643594941.4282906605

1.5072977541.5866433391.507297754

1.57574001561.61640301141.5757400156

1.56733317861.66443484151.5673331786

1.69129400081.73848197161.6912940008

1.57441290691.70853086821.5744129069

1.49009202121.77664668811.4900920212

1.723196081.68546233331.72319608

1.75727586961.58237846111.7572758696

1.61438068871.63467296291.6143806887

1.6646466631.67306935381.664646663

1.71060695271.69900508791.7106069527

1.63505999131.74945769961.6350599913

1.50167872991.64011556531.5016787299

1.5682061141.67867662811.568206114

1.71210202461.71619298931.7121020246

1.55509444861.71841086471.5550944486

1.60745502321.75743497261.6074550232

1.64345267651.89817649931.6434526765

1.70757017611.61936421571.7075701761

1.75587485571.80999136711.7558748557

1.6854909751.6864359494

1.74813778411.586643339

1.76292924311.6164030114

1.78707809241.6644348415

1.82177173431.7384819716

2.09870193271.7085308682

2.01163994131.7766466881

1.6854623333

1.5823784611

1.6346729629

1.6730693538

1.6990050879

1.7494576996

1.6401155653

1.6786766281

1.7161929893

1.7184108647

1.7574349726

1.8981764993

1.6193642157

1.8099913671

1.685490975

1.7481377841

1.7629292431

1.7870780924

1.8217717343

2.0987019327

2.0116399413

D1 m

lg N

lg S'

. 44. , (S''),

(D - ).

) ; ) ; ) ;

) ; ) ; ) .

1.8175653696

1.8419848046

1.8281005841

1.8137485782

1.865155917

1.8847261691

1.9268700173

1.8745921577

1.9055724558

1.8814264979

1.902290691

1.8695579593

1.8977920314

1.8986969379

2.0020165345

1.9617278788

2.0247364406

1.922746993

1.956027975

1.9817566887

2.064845798

Distance, m

-1.2218487496

1.1924366607

-0.920818754

-0.920818754

1.217788144

1.1695820248

-0.7447274949

-0.7447274949

-0.6575773192

1.1515434727

1.2287733507

1.2848663333

-0.5228787453

1.1515434727

1.1958054143

1.1950876951

1.211109298

1.2205636266

1.2131119139

1.2479065241

1.2509004748

1.2528262278

1.2405976741

1.1910135318

1.17951902

1.2309485778

1.2685237449

1.3178538799

1.2509004748

1.2755919151

1.2700479848

1.2775975736

1.3500947125

1.4073078175

1.2777400479

1.2965099541

1.2722059188

1.2912119392

1.3574877695

1.4306487157

1.4512515102

1.4609411222

1.4406349337

1.4861414474

1.5682017241

1.5762727564

1.533517568

1.5719190291

1.6067213323

1.6826438862

Distance (m)

1.4363440395

1.4313637642

1.5097400156

1.5013331786

1.4800077366

1.4375575584

1.4471580313

1.5218438056

1.4620022366

1.5344200094

1.5377253043

1.5216069527

1.4337077169

1.4243310317

1.4438977364

1.53061773

1.4611730899

1.4665729852

1.4556099592

1.5392179017

1.4936012226

1.5218438056

1.5454936712

1.5282926643

1.548854258

1.5138620674

1.5559235952

1.5301180565

1.5538701896

1.5313910923

1.5367249147

1.5281606263

1.5071356811

1.5106276136

1.5783112489

1.5424413959

1.4709184365

1.5988035917

1.543906341

1.6219104416

1.512888963

1.6550569394

1.5950757192

1.5638011697

1.6436967744

1.6381730888

1.6416864973

1.7093801392

1.7837377813

Distance, m

0.4683473304

0.4578818967

0.443697534

0.5126644398

0.4708060498

0.4827012164

0.4563965023

0.4748758789

0.5320781246

0.5810781559

0.5348049318

0.5624710915

0.561860059

0.6407635833

0.4926285572

0.4728973129

0.5063935195

0.5158249744

0.7392219363

0.5236196858

0.5399261672

0.5541640923

0.6615906384

0.67233135

0.7053318015

0.7080846025

0.8235103351

0.8612988959

0.9582089089

0.9019492338

0.9559635331

1.0187808105

1.0447822023

Distance, m

1.4885818651

1.4928249449

1.6030688946

1.7140368769

1.8531865085

1.5947607526

1.5907220298

1.8264522553

1.8240685672

1.7581530304

1.6914928181

1.8084360543

1.7674016014

1.9092822127

1.9306803574

1.8732582956

2.0723922123

2.0395133983

2.1123041051

2.0209144453

Distance, m

0.8569372253

1.0219016629

0.877462315

0.9056939397

0.9285620128

0.8966082997

0.9285620128

0.8486215409

0.9196732869

0.9514123187

0.9248613588

0.9044702142

0.9653518871

0.9636139478

0.9569992655

0.9964046204

0.9984469014

0.9652226167

0.9900897174

0.9479176302

1.0473965432

1.2012125138

1.1236575401

Distance, m

. 45.

( ).

1.5991.3850.6890.98251.63851.8380

1.5991.4350.6890.98251.71851.8383

1.8191.6851.2891.28252.13852.488

()

()

( )

. 46.

.

) ; ) %% .

3000160550300500

100015200250339

4075011090

154204575

811113060

1568515300

347252124166

500160100200300

8530025067344

2197873581106

77099456181290

1750832146125

258540504162

2681245100109

12252665168

134551085140

1106221185129

93630138564

32611118739

662511648230

7106323531826

150011811425092

2419816757

248912050

6204266197

196603355.8

8148196

30965314

5531476

43963

1154843

30326242

6031.3

150229

175323

11069

14767

26510.4

18113

60115

7058

9993

118

70

73

35

24

141

116

155

219

83

133

121

148

80

135

57

55

119

14

407

63

43

5

12

1000

(2)

1.87513.0952380952121.333333333322.7272727273

0.6254.7619047619100.12515.4090909091

0.0251.19047619054.40.05833333334.0909090909

0.0093750.47619047621.80.03333333333.4090909091

0.0050.26190476191.20.09166666672.7272727273

0.0006250.1190476190.64.733333333313.6363636364

0.021251.23809523814.960.67.5454545455

0.31252.38095238181.333333333313.6363636364

0.0531255.95238095242.682.515.6363636364

0.13687517.53.240.654.8181818182

0.4812510.85714285717.240.82513.1818181818

1.093750.76190476195.840.06666666675.6818181818

1.6156251.19047619051.640.33333333332.8181818182

0.16751.071428571440.14.9545454545

0.076250.14285714292.60.43333333337.6363636364

0.083750.23809523813.40.45833333336.3636363636

0.691250.26190476193.40.18333333335.8636363636

0.5850.30952380953.40.252.9090909091

0.203750.26190476193.480.09166666671.7727272727

0.413750.3809523811.920.42510.4545454545

0.443755.595238095212.720.5251.1818181818

0.93752.7142857143100.98333333334.1818181818

0.1506253.97619047620.81666666672.5909090909

0.0152.85714285710.74166666672.2727272727

0.38751.57142857140.358.9545454545

0.12250.78571428570.52.5363636364

0.0506250.48.9090909091

0.1931250.541666666714.2727272727

0.0343750.258333333321.6363636364

0.00250.3252.8636363636

0.0068750.4538.3181818182

0.018752.716666666711

0.03750.2608333333

0.093751.9083333333

0.1093752.6916666667

0.068750.575

0.0918750.5583333333

0.1656250.0866666667

0.011250.9416666667

0.03750.9583333333

0.043750.4833333333

0.0618750.775

0.07375

0.04375

0.045625

0.021875

0.015

0.088125

0.0725

0.096875

0.136875

0.051875

0.083125

0.075625

0.0925

0.05

0.084375

0.035625

0.034375

0.074375

0.00875

0.254375

0.039375

0.026875

0.003125

0.0075

0.625

(2)

(%)

. 47. "-"

.

0.14798290660.0090427090.009042709

0.10658692680.00502877190.0050287719

0.07476097320.00622140990.0062214099

0.06577658430.00502478230.0050247823

0.11536249850.01255384690.0125538469

Diatomea

Ciliata

Nematoda

Polychaeta

All Macrofauna

,

, Z

1.07893803820.0090427090.009042709

1.30942172560.00502877190.0050287719

1.66555818780.00622140990.0062214099

1.83267955890.00502478230.0050247823

0.84518104610.01255384690.0125538469

Diatomea

Ciliata

Nematoda

Polychaeta

All Macrofauna

,

, a

... , , . , . ( ) . .

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