814:
732:
dataset is appropriate for the questions at hand. In practice, the interest is usually in the species diversity of areas so large that not all individuals in them can be observed and identified to species, but a sample of the relevant individuals has to be obtained. Extrapolation from the sample to the underlying population of interest is not straightforward, because the species diversity of the available sample generally gives an underestimation of the species diversity in the entire population. Applying different
786:
800:
774:), the species diversity of the resulting set can be expected to be higher than if all individuals are drawn from a similar environment. Increasing the area sampled increases observed species diversity both because more individuals get included in the sample and because large areas are environmentally more heterogeneous than small areas.
752:
In general, sets with many individuals can be expected to have higher species diversity than sets with fewer individuals. When species diversity values are compared among sets, sampling efforts need to be standardised in an appropriate way for the comparisons to yield ecologically meaningful results.
731:
Depending on the purposes of quantifying species diversity, the data set used for the calculations can be obtained in different ways. Although species diversity can be calculated for any data-set where individuals have been identified to species, meaningful ecological interpretations require that the
736:
will lead to different sets of individuals being observed for the same area of interest, and the species diversity of each set may be different. When a new individual is added to a dataset, it may introduce a species that was not yet represented. How much this increases species diversity depends on
710:
When interpreted in ecological terms, each one of these indices corresponds to a different thing, and their values are therefore not directly comparable. Species richness quantifies the actual rather than effective number of species. The
Shannon index equals
529:
719:, q = 2, and quantifies the probability that two individuals taken at random from the dataset (with replacement of the first individual before taking the second) represent the same species. The Gini-Simpson index equals 1 - 1/
183:
346:
769:
The observed species diversity is affected not only by the number of individuals but also by the heterogeneity of the sample. If individuals are drawn from different environmental conditions (or different
53:. Species richness is a simple count of species. Taxonomic or phylogenetic diversity is the genetic relationship between different groups of species. Species evenness quantifies how equal the
761:
and the number of species only represented by one or a few individuals can be used to help in estimating how representative the available sample is of the population from which it was drawn.
645:
value. In many real datasets, the least abundant species is represented by a single individual, and then the effective number of species would equal the number of individuals in the dataset.
931:
Tuomisto, H. (2010) A diversity of beta diversities: straightening up a concept gone awry. Part 1. Defining beta diversity as a function of alpha and gamma diversity. Ecography, 33, 2-22.
715:), that is, q approaching 1, and in practice quantifies the uncertainty in the species identity of an individual that is taken at random from the dataset. The Simpson index equals 1/
648:
The same equation can be used to calculate the diversity in relation to any classification, not only species. If the individuals are classified into genera or functional types,
591:
increases the weight given to the most abundant species, and fewer equally abundant species are hence needed to reach mean proportional abundance. Consequently, large values of
1025:
Tuomisto, H. (2010) A diversity of beta diversities: straightening up a concept gone awry. Part 2. Quantifying beta diversity and related phenomena. Ecography, 33, 23-45.
416:
1065:
Studies in the
Numerical Analysis of Complex Rain-Forest Communities: II. The Problem of Species-Sampling. Journal of Ecology, Vol. 55, No. 2, Jul., 1967, pp. 525-538
1041:
Colwell, R. K. and
Coddington, J. A. (1994) Estimating terrestrial biodiversity through extrapolation. Philosophical Transactions: Biological Sciences, 345, 101-118.
673:
643:
581:
392:
237:
1097:(2005) Species richness estimation. Pages 7909-7916 in N. Balakrishnan, C. B. Read, and B. Vidakovic, eds. Encyclopedia of Statistical Sciences. New York, Wiley.
1085:
Good, I. J. and
Toulmin, G. H. (1956) The number of new species, and the increase in population coverage, when a sample is increased. Biometrika, 43, 45-63.
79:
245:
1905:
1158:
41:(a dataset). The effective number of species refers to the number of equally abundant species needed to obtain the same mean proportional
2049:
2119:
970:
Xu, S., Böttcher, L., and Chou, T. (2020). Diversity in biology: definitions, quantification and models. Physical
Biology, 17, 031001.
880:
45:
as that observed in the dataset of interest (where all species may not be equally abundant). Meanings of species diversity may include
1691:
1656:
614:
are not used, because then the effective number of species (diversity) would exceed the actual number of species (richness). As
2129:
1857:
2725:
2134:
2322:
954:
Tuomisto, H. 2010. A consistent terminology for quantifying species diversity? Yes, it does exist. Oecologia 4: 853–860.
17:
2720:
2075:
1898:
1739:
1151:
1411:
239:. The proportional abundances themselves are used as weights. The equation is often written in the equivalent form:
1872:
2154:
1867:
1734:
1446:
2539:
2412:
2585:
2184:
2139:
1891:
1144:
890:
1671:
2017:
535:
1016:
Jost, L. (2007) Partitioning diversity into independent alpha and beta components. Ecology, 88, 2427–2439.
2710:
2374:
1265:
915:
Hill, M. O. (1973) Diversity and evenness: a unifying notation and its consequences. Ecology, 54, 427–432
848:
208:
is the total number of species (species richness) in the dataset, and the proportional abundance of the
2439:
2159:
1646:
1363:
1260:
885:
2620:
2232:
2124:
1982:
1967:
1962:
1641:
1353:
745:= 0, each new actual species causes species diversity to increase by one effective species, but when
1136:
723:
and quantifies the probability that the two randomly taken individuals represent different species.
2610:
2605:
2575:
2379:
1842:
1724:
1883:
1514:
2454:
2317:
2227:
2095:
1977:
1947:
1804:
1769:
1489:
1456:
1431:
1051:
758:
754:
524:{\displaystyle \lim _{q\rightarrow 1}{}^{q}\!D=\exp \left(-\sum _{i=1}^{S}p_{i}\ln p_{i}\right),}
599:
for the same dataset. If all species are equally abundant in the dataset, changing the value of
2600:
2544:
2479:
2342:
2277:
2212:
1774:
1562:
1270:
1250:
189:
70:
1119:. William and Flora Hewlett Foundation, the Maxfield Foundation, and the Connexions Consortium
2504:
2449:
2312:
2297:
2080:
2037:
2027:
2022:
1779:
1759:
1615:
1605:
1547:
1542:
1378:
1230:
855:
819:
2630:
2595:
2590:
2514:
2509:
2464:
2362:
2332:
2327:
2179:
2042:
2032:
1577:
1416:
1205:
651:
621:
559:
370:
215:
8:
2680:
2655:
2519:
2489:
2434:
2347:
2237:
2222:
2169:
2002:
1937:
1819:
1749:
1681:
1280:
749:
is large, adding a rare species to a dataset has little effect on its species diversity.
733:
54:
38:
2691:
2640:
2635:
2444:
2407:
2149:
2105:
2070:
1927:
1852:
1754:
1686:
1676:
1610:
1557:
1368:
1313:
1275:
1200:
1069:
986:
Krebs, C. J. (1999) Ecological
Methodology. Second edition. Addison-Wesley, California.
860:
791:
2580:
2549:
2337:
1972:
1837:
1814:
1552:
1328:
1240:
1225:
1210:
1190:
1030:
936:
875:
193:
42:
995:
Magurran, A. E. (2004) Measuring biological diversity. Blackwell
Publishing, Oxford.
2715:
2534:
2397:
2389:
2307:
2189:
2174:
2110:
2090:
2007:
1997:
1992:
1957:
1789:
1729:
1600:
1401:
1343:
1255:
1215:
1050:
1026:
971:
955:
932:
696:
197:
66:
50:
46:
707:, and the complement of the Simpson index (also known as the Gini-Simpson index).
2670:
2529:
2499:
2494:
2484:
2417:
2402:
2282:
2262:
2144:
2012:
1918:
1809:
1719:
1661:
1245:
1171:
870:
843:
828:
692:
545:
2650:
2474:
2427:
2357:
2352:
2247:
2114:
1987:
1794:
1784:
1764:
1567:
1532:
1471:
1348:
1303:
1195:
1130:
1059:
975:
865:
833:
805:
959:
2704:
2675:
1651:
1625:
1582:
1572:
1527:
1494:
1386:
1220:
1175:
785:
704:
700:
360:
178:{\displaystyle {}^{q}\!D={1 \over {\sqrt{\sum _{i=1}^{S}p_{i}p_{i}^{q-1}}}}}
2660:
2645:
2302:
2272:
2217:
2100:
2065:
1942:
1441:
1055:
838:
341:{\displaystyle {}^{q}\!D=\left({\sum _{i=1}^{S}p_{i}^{q}}\right)^{1/(1-q)}}
618:
approaches negative infinity, the generalized mean approaches the minimum
1952:
1499:
1461:
1436:
1426:
1391:
1338:
1318:
1110:
1112:
2665:
2242:
2207:
1847:
1799:
1744:
1714:
1620:
1537:
1481:
1358:
1308:
1073:
1063:
69:
of species proportional abundances in the dataset, and then taking the
2570:
2524:
2252:
1696:
1666:
1466:
1421:
1396:
1333:
1323:
1298:
1290:
1235:
1166:
1094:
757:
can be used to bring samples of different sizes to a common footing.
65:
Species diversity in a dataset can be calculated by first taking the
2625:
2554:
2085:
1913:
1592:
1504:
1451:
1406:
683:
equals genus diversity or functional type diversity, respectively.
553:
2615:
2422:
2292:
2287:
1914:
1862:
1522:
1167:
771:
34:
691:
Often researchers have used the values given by one or more
1007:
Jost, L. (2006) Entropy and diversity. Oikos, 113, 363–375
595:
lead to smaller species diversity than small values of
398:
is equal to the number of species or species richness,
654:
624:
603:
has no effect, but species diversity at any value of
562:
419:
373:
248:
218:
82:
781:
695:
to quantify species diversity. Such indices include
27:
Number of distinct species in a biological community
1111:Harrison, Ian; Laverty, Melina; Sterling, Eleanor.
1068:, Journal of Ecology, British Ecological Society,
667:
637:
575:
523:
386:
340:
231:
177:
587:modifies species weighting, such that increasing
445:
258:
92:
2702:
421:
950:
948:
946:
944:
927:
925:
923:
921:
196:in the dataset as calculated with the weighted
556:, the generalized mean approaches the maximum
49:, taxonomic or phylogenetic diversity, and/or
1899:
1152:
675:represents the proportional abundance of the
941:
918:
406:= 1 is undefined, except that the limit as
60:
2120:Latitudinal gradients in species diversity
1906:
1892:
1159:
1145:
1003:
1001:
911:
909:
907:
881:Latitudinal gradients in species diversity
726:
2018:Predator–prey (Lotka–Volterra) equations
1657:Tritrophic interactions in plant defense
1131:Creative Commons 1.0 Attribution Generic
394:values cancel out, with the result that
2050:Random generalized Lotka–Volterra model
998:
904:
14:
2703:
1858:Herbivore adaptations to plant defense
1887:
1140:
964:
1873:Predator avoidance in schooling fish
686:
2323:Intermediate disturbance hypothesis
24:
2076:Ecological effects of biodiversity
25:
2737:
1412:Generalist and specialist species
1104:
679:th genus or functional type, and
2135:Occupancy–abundance relationship
1031:10.1111/j.1600-0587.2009.06148.x
937:10.1111/j.1600-0587.2009.05880.x
812:
798:
784:
534:which is the exponential of the
359:= 0 corresponds to the weighted
37:that are represented in a given
2155:Relative abundance distribution
1868:Plant defense against herbivory
1735:Competitive exclusion principle
1447:Mesopredator release hypothesis
1088:
1079:
1044:
1035:
355:determines which mean is used.
1740:Consumer–resource interactions
1019:
1010:
989:
980:
428:
410:approaches 1 is well defined:
333:
321:
13:
1:
2586:Biological data visualization
2413:Environmental niche modelling
2140:Population viability analysis
2071:Density-dependent inhibition
7:
2726:Measurement of biodiversity
2540:Liebig's law of the minimum
2375:Resource selection function
1266:Metabolic theory of ecology
849:Measurement of biodiversity
777:
33:is the number of different
10:
2742:
2440:Niche apportionment models
2160:Relative species abundance
1364:Primary nutritional groups
1261:List of feeding behaviours
886:Relative species abundance
73:of this. The equation is:
2721:Environmental terminology
2689:
2621:Ecosystem based fisheries
2563:
2463:
2388:
2261:
2233:Interspecific competition
2198:
2125:Minimum viable population
2058:
1983:Maximum sustainable yield
1968:Intraspecific competition
1963:Effective population size
1926:
1843:Anti-predator adaptations
1828:
1707:
1634:
1591:
1513:
1480:
1377:
1354:Photosynthetic efficiency
1289:
1183:
960:10.1007/s00442-010-1812-0
891:Species–area relationship
764:
607:equals species richness.
192:equals mean proportional
2611:Ecological stoichiometry
2576:Alternative stable state
976:10.1088/1478-3975/ab6754
897:
759:Species discovery curves
61:Calculation of diversity
2455:Ontogenetic niche shift
2318:Ideal free distribution
2228:Ecological facilitation
1978:Malthusian growth model
1948:Consumer-resource model
1805:Paradox of the plankton
1770:Energy systems language
1490:Chemoorganoheterotrophy
1457:Optimal foraging theory
1432:Heterotrophic nutrition
1062:; Lance, G. N. (1969),
727:Sampling considerations
544:= 2 corresponds to the
2601:Ecological forecasting
2545:Marginal value theorem
2343:Landscape epidemiology
2278:Cross-boundary subsidy
2213:Biological interaction
1563:Microbial intelligence
1251:Green world hypothesis
669:
639:
577:
525:
486:
388:
342:
291:
233:
204:- 1. In the equation,
179:
127:
2606:Ecological humanities
2505:Ecological energetics
2450:Niche differentiation
2313:Habitat fragmentation
2081:Ecological extinction
2028:Small population size
1780:Feed conversion ratio
1760:Ecological succession
1692:San Francisco Estuary
1606:Ecological efficiency
1548:Microbial cooperation
856:Biotic homogenization
820:Earth sciences portal
670:
668:{\displaystyle p_{i}}
640:
638:{\displaystyle p_{i}}
578:
576:{\displaystyle p_{i}}
526:
466:
389:
387:{\displaystyle p_{i}}
343:
271:
234:
232:{\displaystyle p_{i}}
180:
107:
2631:Evolutionary ecology
2596:Ecological footprint
2591:Ecological economics
2515:Ecological threshold
2510:Ecological indicator
2380:Source–sink dynamics
2333:Land change modeling
2328:Insular biogeography
2180:Species distribution
1919:Modelling ecosystems
1578:Microbial metabolism
1417:Intraguild predation
1206:Biogeochemical cycle
1172:Modelling ecosystems
1117:Connexions (cnx.org)
652:
622:
583:value. In practice,
560:
417:
371:
246:
216:
80:
57:of the species are.
2681:Theoretical ecology
2656:Natural environment
2520:Ecosystem diversity
2490:Ecological collapse
2480:Bateman's principle
2435:Limiting similarity
2348:Landscape limnology
2170:Species homogeneity
2008:Population modeling
2003:Population dynamics
1820:Trophic state index
1113:"Species Diversity"
610:Negative values of
306:
158:
18:Taxonomic diversity
2711:Ecological metrics
2692:Outline of ecology
2641:Industrial ecology
2636:Functional ecology
2500:Ecological deficit
2445:Niche construction
2408:Ecosystem engineer
2185:Species–area curve
2106:Introduced species
1921:: Other components
1853:Deimatic behaviour
1755:Ecological network
1687:North Pacific Gyre
1672:hydrothermal vents
1611:Ecological pyramid
1558:Microbial food web
1369:Primary production
1314:Foundation species
861:Coexistence theory
792:Environment portal
755:Resampling methods
665:
635:
573:
521:
435:
384:
338:
292:
229:
175:
138:
2698:
2697:
2581:Balance of nature
2338:Landscape ecology
2223:Community ecology
2165:Species diversity
2101:Indicator species
2096:Gradient analysis
1973:Logistic function
1881:
1880:
1838:Animal coloration
1815:Trophic mutualism
1553:Microbial ecology
1344:Photoheterotrophs
1329:Myco-heterotrophy
1241:Ecosystem ecology
1226:Carrying capacity
1191:Abiotic component
876:Genetic diversity
693:diversity indices
687:Diversity indices
420:
194:species abundance
173:
171:
43:species abundance
31:Species diversity
16:(Redirected from
2733:
2398:Ecological niche
2370:selection theory
2190:Umbrella species
2175:Species richness
2111:Invasive species
2091:Flagship species
1998:Population cycle
1993:Overexploitation
1958:Ecological yield
1908:
1901:
1894:
1885:
1884:
1790:Mesotrophic soil
1730:Climax community
1662:Marine food webs
1601:Biomagnification
1402:Chemoorganotroph
1256:Keystone species
1216:Biotic component
1161:
1154:
1147:
1138:
1137:
1129:(Licensed under
1128:
1126:
1124:
1098:
1092:
1086:
1083:
1077:
1076:
1048:
1042:
1039:
1033:
1023:
1017:
1014:
1008:
1005:
996:
993:
987:
984:
978:
968:
962:
952:
939:
929:
916:
913:
822:
817:
816:
815:
808:
803:
802:
801:
794:
789:
788:
734:sampling methods
697:species richness
674:
672:
671:
666:
664:
663:
644:
642:
641:
636:
634:
633:
582:
580:
579:
574:
572:
571:
530:
528:
527:
522:
517:
513:
512:
511:
496:
495:
485:
480:
444:
443:
438:
434:
393:
391:
390:
385:
383:
382:
347:
345:
344:
339:
337:
336:
320:
311:
307:
305:
300:
290:
285:
257:
256:
251:
238:
236:
235:
230:
228:
227:
198:generalized mean
184:
182:
181:
176:
174:
172:
170:
159:
157:
146:
137:
136:
126:
121:
105:
100:
91:
90:
85:
67:weighted average
51:species evenness
47:species richness
21:
2741:
2740:
2736:
2735:
2734:
2732:
2731:
2730:
2701:
2700:
2699:
2694:
2685:
2671:Systems ecology
2559:
2530:Extinction debt
2495:Ecological debt
2485:Bioluminescence
2466:
2459:
2428:marine habitats
2403:Ecological trap
2384:
2264:
2257:
2200:
2194:
2150:Rapoport's rule
2145:Priority effect
2086:Endemic species
2054:
2013:Population size
1929:
1922:
1912:
1882:
1877:
1830:
1824:
1810:Trophic cascade
1720:Bioaccumulation
1703:
1630:
1587:
1509:
1476:
1373:
1285:
1246:Ecosystem model
1179:
1165:
1122:
1120:
1107:
1102:
1101:
1093:
1089:
1084:
1080:
1060:Williams, W. T.
1049:
1045:
1040:
1036:
1024:
1020:
1015:
1011:
1006:
999:
994:
990:
985:
981:
969:
965:
953:
942:
930:
919:
914:
905:
900:
895:
871:Gamma diversity
844:Diversity index
829:Alpha diversity
818:
813:
811:
804:
799:
797:
790:
783:
780:
767:
729:
689:
659:
655:
653:
650:
649:
629:
625:
623:
620:
619:
567:
563:
561:
558:
557:
546:arithmetic mean
536:Shannon entropy
507:
503:
491:
487:
481:
470:
462:
458:
439:
437:
436:
424:
418:
415:
414:
378:
374:
372:
369:
368:
316:
312:
301:
296:
286:
275:
270:
266:
265:
252:
250:
249:
247:
244:
243:
223:
219:
217:
214:
213:
160:
147:
142:
132:
128:
122:
111:
106:
104:
99:
86:
84:
83:
81:
78:
77:
63:
28:
23:
22:
15:
12:
11:
5:
2739:
2729:
2728:
2723:
2718:
2713:
2696:
2695:
2690:
2687:
2686:
2684:
2683:
2678:
2673:
2668:
2663:
2658:
2653:
2651:Microecosystem
2648:
2643:
2638:
2633:
2628:
2623:
2618:
2613:
2608:
2603:
2598:
2593:
2588:
2583:
2578:
2573:
2567:
2565:
2561:
2560:
2558:
2557:
2552:
2550:Thorson's rule
2547:
2542:
2537:
2532:
2527:
2522:
2517:
2512:
2507:
2502:
2497:
2492:
2487:
2482:
2477:
2475:Assembly rules
2471:
2469:
2461:
2460:
2458:
2457:
2452:
2447:
2442:
2437:
2432:
2431:
2430:
2420:
2415:
2410:
2405:
2400:
2394:
2392:
2386:
2385:
2383:
2382:
2377:
2372:
2360:
2358:Patch dynamics
2355:
2353:Metapopulation
2350:
2345:
2340:
2335:
2330:
2325:
2320:
2315:
2310:
2305:
2300:
2295:
2290:
2285:
2280:
2275:
2269:
2267:
2259:
2258:
2256:
2255:
2250:
2248:Storage effect
2245:
2240:
2235:
2230:
2225:
2220:
2215:
2210:
2204:
2202:
2196:
2195:
2193:
2192:
2187:
2182:
2177:
2172:
2167:
2162:
2157:
2152:
2147:
2142:
2137:
2132:
2130:Neutral theory
2127:
2122:
2117:
2115:Native species
2108:
2103:
2098:
2093:
2088:
2083:
2078:
2073:
2068:
2062:
2060:
2056:
2055:
2053:
2052:
2047:
2046:
2045:
2040:
2030:
2025:
2020:
2015:
2010:
2005:
2000:
1995:
1990:
1988:Overpopulation
1985:
1980:
1975:
1970:
1965:
1960:
1955:
1950:
1945:
1940:
1934:
1932:
1924:
1923:
1911:
1910:
1903:
1896:
1888:
1879:
1878:
1876:
1875:
1870:
1865:
1860:
1855:
1850:
1845:
1840:
1834:
1832:
1826:
1825:
1823:
1822:
1817:
1812:
1807:
1802:
1797:
1795:Nutrient cycle
1792:
1787:
1785:Feeding frenzy
1782:
1777:
1772:
1767:
1765:Energy quality
1762:
1757:
1752:
1747:
1742:
1737:
1732:
1727:
1725:Cascade effect
1722:
1717:
1711:
1709:
1705:
1704:
1702:
1701:
1700:
1699:
1694:
1689:
1684:
1679:
1674:
1669:
1659:
1654:
1649:
1644:
1638:
1636:
1632:
1631:
1629:
1628:
1623:
1618:
1613:
1608:
1603:
1597:
1595:
1589:
1588:
1586:
1585:
1580:
1575:
1570:
1568:Microbial loop
1565:
1560:
1555:
1550:
1545:
1540:
1535:
1533:Lithoautotroph
1530:
1525:
1519:
1517:
1515:Microorganisms
1511:
1510:
1508:
1507:
1502:
1497:
1492:
1486:
1484:
1478:
1477:
1475:
1474:
1472:Prey switching
1469:
1464:
1459:
1454:
1449:
1444:
1439:
1434:
1429:
1424:
1419:
1414:
1409:
1404:
1399:
1394:
1389:
1383:
1381:
1375:
1374:
1372:
1371:
1366:
1361:
1356:
1351:
1349:Photosynthesis
1346:
1341:
1336:
1331:
1326:
1321:
1316:
1311:
1306:
1304:Chemosynthesis
1301:
1295:
1293:
1287:
1286:
1284:
1283:
1278:
1273:
1268:
1263:
1258:
1253:
1248:
1243:
1238:
1233:
1228:
1223:
1218:
1213:
1208:
1203:
1198:
1196:Abiotic stress
1193:
1187:
1185:
1181:
1180:
1164:
1163:
1156:
1149:
1141:
1135:
1134:
1106:
1105:External links
1103:
1100:
1099:
1087:
1078:
1043:
1034:
1018:
1009:
997:
988:
979:
963:
940:
917:
902:
901:
899:
896:
894:
893:
888:
883:
878:
873:
868:
866:Dark diversity
863:
858:
853:
852:
851:
846:
836:
834:Beta diversity
831:
825:
824:
823:
809:
806:Ecology portal
795:
779:
776:
766:
763:
728:
725:
688:
685:
662:
658:
632:
628:
570:
566:
532:
531:
520:
516:
510:
506:
502:
499:
494:
490:
484:
479:
476:
473:
469:
465:
461:
457:
454:
451:
448:
442:
433:
430:
427:
423:
381:
377:
349:
348:
335:
332:
329:
326:
323:
319:
315:
310:
304:
299:
295:
289:
284:
281:
278:
274:
269:
264:
261:
255:
226:
222:
212:th species is
200:with exponent
186:
185:
169:
166:
163:
156:
153:
150:
145:
141:
135:
131:
125:
120:
117:
114:
110:
103:
98:
95:
89:
62:
59:
26:
9:
6:
4:
3:
2:
2738:
2727:
2724:
2722:
2719:
2717:
2714:
2712:
2709:
2708:
2706:
2693:
2688:
2682:
2679:
2677:
2676:Urban ecology
2674:
2672:
2669:
2667:
2664:
2662:
2659:
2657:
2654:
2652:
2649:
2647:
2644:
2642:
2639:
2637:
2634:
2632:
2629:
2627:
2624:
2622:
2619:
2617:
2614:
2612:
2609:
2607:
2604:
2602:
2599:
2597:
2594:
2592:
2589:
2587:
2584:
2582:
2579:
2577:
2574:
2572:
2569:
2568:
2566:
2562:
2556:
2553:
2551:
2548:
2546:
2543:
2541:
2538:
2536:
2535:Kleiber's law
2533:
2531:
2528:
2526:
2523:
2521:
2518:
2516:
2513:
2511:
2508:
2506:
2503:
2501:
2498:
2496:
2493:
2491:
2488:
2486:
2483:
2481:
2478:
2476:
2473:
2472:
2470:
2468:
2462:
2456:
2453:
2451:
2448:
2446:
2443:
2441:
2438:
2436:
2433:
2429:
2426:
2425:
2424:
2421:
2419:
2416:
2414:
2411:
2409:
2406:
2404:
2401:
2399:
2396:
2395:
2393:
2391:
2387:
2381:
2378:
2376:
2373:
2371:
2369:
2365:
2361:
2359:
2356:
2354:
2351:
2349:
2346:
2344:
2341:
2339:
2336:
2334:
2331:
2329:
2326:
2324:
2321:
2319:
2316:
2314:
2311:
2309:
2308:Foster's rule
2306:
2304:
2301:
2299:
2296:
2294:
2291:
2289:
2286:
2284:
2281:
2279:
2276:
2274:
2271:
2270:
2268:
2266:
2260:
2254:
2251:
2249:
2246:
2244:
2241:
2239:
2236:
2234:
2231:
2229:
2226:
2224:
2221:
2219:
2216:
2214:
2211:
2209:
2206:
2205:
2203:
2197:
2191:
2188:
2186:
2183:
2181:
2178:
2176:
2173:
2171:
2168:
2166:
2163:
2161:
2158:
2156:
2153:
2151:
2148:
2146:
2143:
2141:
2138:
2136:
2133:
2131:
2128:
2126:
2123:
2121:
2118:
2116:
2112:
2109:
2107:
2104:
2102:
2099:
2097:
2094:
2092:
2089:
2087:
2084:
2082:
2079:
2077:
2074:
2072:
2069:
2067:
2064:
2063:
2061:
2057:
2051:
2048:
2044:
2041:
2039:
2036:
2035:
2034:
2031:
2029:
2026:
2024:
2021:
2019:
2016:
2014:
2011:
2009:
2006:
2004:
2001:
1999:
1996:
1994:
1991:
1989:
1986:
1984:
1981:
1979:
1976:
1974:
1971:
1969:
1966:
1964:
1961:
1959:
1956:
1954:
1951:
1949:
1946:
1944:
1941:
1939:
1936:
1935:
1933:
1931:
1925:
1920:
1916:
1909:
1904:
1902:
1897:
1895:
1890:
1889:
1886:
1874:
1871:
1869:
1866:
1864:
1861:
1859:
1856:
1854:
1851:
1849:
1846:
1844:
1841:
1839:
1836:
1835:
1833:
1827:
1821:
1818:
1816:
1813:
1811:
1808:
1806:
1803:
1801:
1798:
1796:
1793:
1791:
1788:
1786:
1783:
1781:
1778:
1776:
1773:
1771:
1768:
1766:
1763:
1761:
1758:
1756:
1753:
1751:
1748:
1746:
1743:
1741:
1738:
1736:
1733:
1731:
1728:
1726:
1723:
1721:
1718:
1716:
1713:
1712:
1710:
1706:
1698:
1695:
1693:
1690:
1688:
1685:
1683:
1680:
1678:
1675:
1673:
1670:
1668:
1665:
1664:
1663:
1660:
1658:
1655:
1653:
1650:
1648:
1645:
1643:
1640:
1639:
1637:
1633:
1627:
1626:Trophic level
1624:
1622:
1619:
1617:
1614:
1612:
1609:
1607:
1604:
1602:
1599:
1598:
1596:
1594:
1590:
1584:
1583:Phage ecology
1581:
1579:
1576:
1574:
1573:Microbial mat
1571:
1569:
1566:
1564:
1561:
1559:
1556:
1554:
1551:
1549:
1546:
1544:
1541:
1539:
1536:
1534:
1531:
1529:
1528:Bacteriophage
1526:
1524:
1521:
1520:
1518:
1516:
1512:
1506:
1503:
1501:
1498:
1496:
1495:Decomposition
1493:
1491:
1488:
1487:
1485:
1483:
1479:
1473:
1470:
1468:
1465:
1463:
1460:
1458:
1455:
1453:
1450:
1448:
1445:
1443:
1442:Mesopredators
1440:
1438:
1435:
1433:
1430:
1428:
1425:
1423:
1420:
1418:
1415:
1413:
1410:
1408:
1405:
1403:
1400:
1398:
1395:
1393:
1390:
1388:
1387:Apex predator
1385:
1384:
1382:
1380:
1376:
1370:
1367:
1365:
1362:
1360:
1357:
1355:
1352:
1350:
1347:
1345:
1342:
1340:
1337:
1335:
1332:
1330:
1327:
1325:
1322:
1320:
1317:
1315:
1312:
1310:
1307:
1305:
1302:
1300:
1297:
1296:
1294:
1292:
1288:
1282:
1279:
1277:
1274:
1272:
1269:
1267:
1264:
1262:
1259:
1257:
1254:
1252:
1249:
1247:
1244:
1242:
1239:
1237:
1234:
1232:
1229:
1227:
1224:
1222:
1221:Biotic stress
1219:
1217:
1214:
1212:
1209:
1207:
1204:
1202:
1199:
1197:
1194:
1192:
1189:
1188:
1186:
1182:
1177:
1173:
1169:
1162:
1157:
1155:
1150:
1148:
1143:
1142:
1139:
1132:
1118:
1114:
1109:
1108:
1096:
1091:
1082:
1075:
1071:
1067:
1066:
1061:
1057:
1056:Tracey, J. G.
1053:
1047:
1038:
1032:
1028:
1022:
1013:
1004:
1002:
992:
983:
977:
973:
967:
961:
957:
951:
949:
947:
945:
938:
934:
928:
926:
924:
922:
912:
910:
908:
903:
892:
889:
887:
884:
882:
879:
877:
874:
872:
869:
867:
864:
862:
859:
857:
854:
850:
847:
845:
842:
841:
840:
837:
835:
832:
830:
827:
826:
821:
810:
807:
796:
793:
787:
782:
775:
773:
762:
760:
756:
750:
748:
744:
740:
737:the value of
735:
724:
722:
718:
714:
708:
706:
705:Simpson index
702:
701:Shannon index
698:
694:
684:
682:
678:
660:
656:
646:
630:
626:
617:
613:
608:
606:
602:
598:
594:
590:
586:
568:
564:
555:
551:
547:
543:
539:
537:
518:
514:
508:
504:
500:
497:
492:
488:
482:
477:
474:
471:
467:
463:
459:
455:
452:
449:
446:
440:
431:
425:
413:
412:
411:
409:
405:
401:
397:
379:
375:
366:
363:, which is 1/
362:
361:harmonic mean
358:
354:
351:The value of
330:
327:
324:
317:
313:
308:
302:
297:
293:
287:
282:
279:
276:
272:
267:
262:
259:
253:
242:
241:
240:
224:
220:
211:
207:
203:
199:
195:
191:
167:
164:
161:
154:
151:
148:
143:
139:
133:
129:
123:
118:
115:
112:
108:
101:
96:
93:
87:
76:
75:
74:
72:
68:
58:
56:
52:
48:
44:
40:
36:
32:
19:
2661:Regime shift
2646:Macroecology
2367:
2363:
2303:Edge effects
2273:Biogeography
2218:Commensalism
2164:
2066:Biodiversity
1943:Allee effect
1682:kelp forests
1635:Example webs
1500:Detritivores
1339:Organotrophs
1319:Kinetotrophs
1271:Productivity
1121:. Retrieved
1116:
1090:
1081:
1064:
1046:
1037:
1021:
1012:
991:
982:
966:
839:Biodiversity
768:
751:
746:
742:
738:
730:
720:
716:
712:
709:
690:
680:
676:
647:
615:
611:
609:
604:
600:
596:
592:
588:
584:
549:
541:
540:
533:
407:
403:
399:
395:
367:because the
364:
356:
352:
350:
209:
205:
201:
187:
64:
30:
29:
2298:Disturbance
2201:interaction
2023:Recruitment
1953:Depensation
1745:Copiotrophs
1616:Energy flow
1538:Lithotrophy
1482:Decomposers
1462:Planktivore
1437:Insectivore
1427:Heterotroph
1392:Bacterivore
1359:Phototrophs
1309:Chemotrophs
1281:Restoration
1231:Competition
1052:Webb, L. J.
552:approaches
190:denominator
2705:Categories
2666:Sexecology
2243:Parasitism
2208:Antibiosis
2043:Resistance
2038:Resilience
1928:Population
1848:Camouflage
1800:Oligotroph
1715:Ascendency
1677:intertidal
1667:cold seeps
1621:Food chain
1422:Herbivores
1397:Carnivores
1324:Mixotrophs
1299:Autotrophs
1178:components
1123:1 February
55:abundances
2571:Allometry
2525:Emergence
2253:Symbiosis
2238:Mutualism
2033:Stability
1938:Abundance
1750:Dominance
1708:Processes
1697:tide pool
1593:Food webs
1467:Predation
1452:Omnivores
1379:Consumers
1334:Mycotroph
1291:Producers
1236:Ecosystem
1201:Behaviour
501:
468:∑
464:−
456:
429:→
328:−
273:∑
165:−
152:−
109:∑
39:community
2626:Endolith
2555:Xerosere
2467:networks
2283:Ecocline
1829:Defense,
1505:Detritus
1407:Foraging
1276:Resource
1095:Chao, A.
778:See also
772:habitats
554:infinity
2716:Habitat
2616:Ecopath
2423:Habitat
2293:Ecotype
2288:Ecotone
2265:ecology
2263:Spatial
2199:Species
2059:Species
1930:ecology
1915:Ecology
1863:Mimicry
1831:counter
1775:f-ratio
1523:Archaea
1211:Biomass
1184:General
1176:Trophic
1168:Ecology
1074:2257891
741:: when
71:inverse
35:species
1647:Rivers
1543:Marine
1072:
765:Trends
703:, the
699:, the
2564:Other
2465:Other
2418:Guild
2390:Niche
1642:Lakes
1070:JSTOR
898:Notes
548:. As
1652:Soil
1125:2011
711:log(
188:The
1027:doi
972:doi
956:doi
933:doi
453:exp
422:lim
2707::
2113:/
1917::
1174::
1170::
1133:).
1115:.
1058:;
1054:;
1000:^
943:^
920:^
906:^
538:.
498:ln
402:.
2368:K
2366:/
2364:r
1907:e
1900:t
1893:v
1160:e
1153:t
1146:v
1127:.
1029::
974::
958::
935::
747:q
743:q
739:q
721:D
717:D
713:D
681:D
677:i
661:i
657:p
631:i
627:p
616:q
612:q
605:q
601:q
597:q
593:q
589:q
585:q
569:i
565:p
550:q
542:q
519:,
515:)
509:i
505:p
493:i
489:p
483:S
478:1
475:=
472:i
460:(
450:=
447:D
441:q
432:1
426:q
408:q
404:q
400:S
396:D
380:i
376:p
365:S
357:q
353:q
334:)
331:q
325:1
322:(
318:/
314:1
309:)
303:q
298:i
294:p
288:S
283:1
280:=
277:i
268:(
263:=
260:D
254:q
225:i
221:p
210:i
206:S
202:q
168:1
162:q
155:1
149:q
144:i
140:p
134:i
130:p
124:S
119:1
116:=
113:i
102:1
97:=
94:D
88:q
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
