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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
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?
- , . , .
:
, ( ) . ()
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
, . -
6 , , ( ). , , - ,
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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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