803:
721:
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
775:
789:
763:), 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.
741:
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.
720:
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
725:
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
699:
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
518:
708:, 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/
172:
335:
758:
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
42:. 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
750:
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.
634:
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.
920:
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.
704:), 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/
637:
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,
580:
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
1014:
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.
405:
1054:
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
1030:
Colwell, R. K. and
Coddington, J. A. (1994) Estimating terrestrial biodiversity through extrapolation. Philosophical Transactions: Biological Sciences, 345, 101-118.
662:
632:
570:
381:
226:
1086:(2005) Species richness estimation. Pages 7909-7916 in N. Balakrishnan, C. B. Read, and B. Vidakovic, eds. Encyclopedia of Statistical Sciences. New York, Wiley.
1074:
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.
68:
234:
1894:
1147:
30:(a dataset). The effective number of species refers to the number of equally abundant species needed to obtain the same mean proportional
2038:
2108:
959:
Xu, S., Böttcher, L., and Chou, T. (2020). Diversity in biology: definitions, quantification and models. Physical
Biology, 17, 031001.
869:
34:
as that observed in the dataset of interest (where all species may not be equally abundant). Meanings of species diversity may include
1680:
1645:
603:
are not used, because then the effective number of species (diversity) would exceed the actual number of species (richness). As
2118:
1846:
2714:
2123:
2311:
943:
Tuomisto, H. 2010. A consistent terminology for quantifying species diversity? Yes, it does exist. Oecologia 4: 853–860.
2709:
2064:
1887:
1728:
1140:
1400:
228:. The proportional abundances themselves are used as weights. The equation is often written in the equivalent form:
1861:
2143:
1856:
1723:
1435:
2528:
2401:
2574:
2173:
2128:
1880:
1133:
879:
1660:
2006:
524:
1005:
Jost, L. (2007) Partitioning diversity into independent alpha and beta components. Ecology, 88, 2427–2439.
2699:
2363:
1254:
904:
Hill, M. O. (1973) Diversity and evenness: a unifying notation and its consequences. Ecology, 54, 427–432
837:
197:
is the total number of species (species richness) in the dataset, and the proportional abundance of the
2428:
2148:
1635:
1352:
1249:
874:
2609:
2221:
2113:
1971:
1956:
1951:
1630:
1342:
734:= 0, each new actual species causes species diversity to increase by one effective species, but when
1125:
712:
and quantifies the probability that the two randomly taken individuals represent different species.
2599:
2594:
2564:
2368:
1831:
1713:
1872:
1503:
2443:
2306:
2216:
2084:
1966:
1936:
1793:
1758:
1478:
1445:
1420:
1040:
747:
743:
513:{\displaystyle \lim _{q\rightarrow 1}{}^{q}\!D=\exp \left(-\sum _{i=1}^{S}p_{i}\ln p_{i}\right),}
588:
for the same dataset. If all species are equally abundant in the dataset, changing the value of
2589:
2533:
2468:
2331:
2266:
2201:
1763:
1551:
1259:
1239:
178:
59:
1108:. William and Flora Hewlett Foundation, the Maxfield Foundation, and the Connexions Consortium
2493:
2438:
2301:
2286:
2069:
2026:
2016:
2011:
1768:
1748:
1604:
1594:
1536:
1531:
1367:
1219:
844:
808:
2619:
2584:
2579:
2503:
2498:
2453:
2351:
2321:
2316:
2168:
2031:
2021:
1566:
1405:
1194:
640:
610:
548:
359:
204:
8:
2669:
2644:
2508:
2478:
2423:
2336:
2226:
2211:
2158:
1991:
1926:
1808:
1738:
1670:
1269:
738:
is large, adding a rare species to a dataset has little effect on its species diversity.
722:
43:
27:
2680:
2629:
2624:
2433:
2396:
2138:
2094:
2059:
1916:
1841:
1743:
1675:
1665:
1599:
1546:
1357:
1302:
1264:
1189:
1058:
975:
Krebs, C. J. (1999) Ecological
Methodology. Second edition. Addison-Wesley, California.
849:
780:
2569:
2538:
2326:
1961:
1826:
1803:
1541:
1317:
1229:
1214:
1199:
1179:
1019:
925:
864:
182:
31:
984:
Magurran, A. E. (2004) Measuring biological diversity. Blackwell
Publishing, Oxford.
2704:
2523:
2386:
2378:
2296:
2178:
2163:
2099:
2079:
1996:
1986:
1981:
1946:
1778:
1718:
1589:
1390:
1332:
1244:
1204:
1039:
1015:
960:
944:
921:
685:
186:
55:
39:
35:
696:, and the complement of the Simpson index (also known as the Gini-Simpson index).
2659:
2518:
2488:
2483:
2473:
2406:
2391:
2271:
2251:
2133:
2001:
1907:
1798:
1708:
1650:
1234:
1160:
859:
832:
817:
681:
534:
2639:
2463:
2416:
2346:
2341:
2236:
2103:
1976:
1783:
1773:
1753:
1556:
1521:
1460:
1337:
1292:
1184:
1119:
1048:
964:
854:
822:
794:
948:
2693:
2664:
1640:
1614:
1571:
1561:
1516:
1483:
1375:
1209:
1164:
774:
693:
689:
349:
167:{\displaystyle {}^{q}\!D={1 \over {\sqrt{\sum _{i=1}^{S}p_{i}p_{i}^{q-1}}}}}
2649:
2634:
2291:
2261:
2206:
2089:
2054:
1931:
1430:
1044:
827:
330:{\displaystyle {}^{q}\!D=\left({\sum _{i=1}^{S}p_{i}^{q}}\right)^{1/(1-q)}}
607:
approaches negative infinity, the generalized mean approaches the minimum
1941:
1488:
1450:
1425:
1415:
1380:
1327:
1307:
1099:
1101:
2654:
2231:
2196:
1836:
1788:
1733:
1703:
1609:
1526:
1470:
1347:
1297:
1062:
1052:
58:
of species proportional abundances in the dataset, and then taking the
2559:
2513:
2241:
1685:
1655:
1455:
1410:
1385:
1322:
1312:
1287:
1279:
1224:
1155:
1083:
746:
can be used to bring samples of different sizes to a common footing.
54:
Species diversity in a dataset can be calculated by first taking the
2614:
2543:
2074:
1902:
1581:
1493:
1440:
1395:
672:
equals genus diversity or functional type diversity, respectively.
542:
2604:
2411:
2281:
2276:
1903:
1851:
1511:
1156:
760:
23:
680:
Often researchers have used the values given by one or more
996:
Jost, L. (2006) Entropy and diversity. Oikos, 113, 363–375
584:
lead to smaller species diversity than small values of
387:
is equal to the number of species or species richness,
643:
613:
592:
has no effect, but species diversity at any value of
551:
408:
362:
237:
207:
71:
770:
684:
to quantify species diversity. Such indices include
16:
Number of distinct species in a biological community
1100:Harrison, Ian; Laverty, Melina; Sterling, Eleanor.
1057:, Journal of Ecology, British Ecological Society,
656:
626:
564:
512:
375:
329:
220:
166:
576:modifies species weighting, such that increasing
434:
247:
81:
2691:
410:
939:
937:
935:
933:
916:
914:
912:
910:
185:in the dataset as calculated with the weighted
545:, the generalized mean approaches the maximum
38:, taxonomic or phylogenetic diversity, and/or
1888:
1141:
664:represents the proportional abundance of the
930:
907:
395:= 1 is undefined, except that the limit as
49:
2109:Latitudinal gradients in species diversity
1895:
1881:
1148:
1134:
992:
990:
900:
898:
896:
870:Latitudinal gradients in species diversity
715:
2007:Predator–prey (Lotka–Volterra) equations
1646:Tritrophic interactions in plant defense
1120:Creative Commons 1.0 Attribution Generic
383:values cancel out, with the result that
2039:Random generalized Lotka–Volterra model
987:
893:
2692:
1847:Herbivore adaptations to plant defense
1876:
1129:
953:
1862:Predator avoidance in schooling fish
675:
2312:Intermediate disturbance hypothesis
13:
2065:Ecological effects of biodiversity
14:
2726:
1401:Generalist and specialist species
1093:
668:th genus or functional type, and
2124:Occupancy–abundance relationship
1020:10.1111/j.1600-0587.2009.06148.x
926:10.1111/j.1600-0587.2009.05880.x
801:
787:
773:
523:which is the exponential of the
348:= 0 corresponds to the weighted
26:that are represented in a given
2144:Relative abundance distribution
1857:Plant defense against herbivory
1724:Competitive exclusion principle
1436:Mesopredator release hypothesis
1077:
1068:
1033:
1024:
344:determines which mean is used.
1729:Consumer–resource interactions
1008:
999:
978:
969:
417:
399:approaches 1 is well defined:
322:
310:
1:
2575:Biological data visualization
2402:Environmental niche modelling
2129:Population viability analysis
2060:Density-dependent inhibition
7:
2715:Measurement of biodiversity
2529:Liebig's law of the minimum
2364:Resource selection function
1255:Metabolic theory of ecology
838:Measurement of biodiversity
766:
22:is the number of different
10:
2731:
2429:Niche apportionment models
2149:Relative species abundance
1353:Primary nutritional groups
1250:List of feeding behaviours
875:Relative species abundance
62:of this. The equation is:
2710:Environmental terminology
2678:
2610:Ecosystem based fisheries
2552:
2452:
2377:
2250:
2222:Interspecific competition
2187:
2114:Minimum viable population
2047:
1972:Maximum sustainable yield
1957:Intraspecific competition
1952:Effective population size
1915:
1832:Anti-predator adaptations
1817:
1696:
1623:
1580:
1502:
1469:
1366:
1343:Photosynthetic efficiency
1278:
1172:
949:10.1007/s00442-010-1812-0
880:Species–area relationship
753:
596:equals species richness.
181:equals mean proportional
2600:Ecological stoichiometry
2565:Alternative stable state
965:10.1088/1478-3975/ab6754
886:
748:Species discovery curves
50:Calculation of diversity
2444:Ontogenetic niche shift
2307:Ideal free distribution
2217:Ecological facilitation
1967:Malthusian growth model
1937:Consumer-resource model
1794:Paradox of the plankton
1759:Energy systems language
1479:Chemoorganoheterotrophy
1446:Optimal foraging theory
1421:Heterotrophic nutrition
1051:; Lance, G. N. (1969),
716:Sampling considerations
533:= 2 corresponds to the
2590:Ecological forecasting
2534:Marginal value theorem
2332:Landscape epidemiology
2267:Cross-boundary subsidy
2202:Biological interaction
1552:Microbial intelligence
1240:Green world hypothesis
658:
628:
566:
514:
475:
377:
331:
280:
222:
193:- 1. In the equation,
168:
116:
2595:Ecological humanities
2494:Ecological energetics
2439:Niche differentiation
2302:Habitat fragmentation
2070:Ecological extinction
2017:Small population size
1769:Feed conversion ratio
1749:Ecological succession
1681:San Francisco Estuary
1595:Ecological efficiency
1537:Microbial cooperation
845:Biotic homogenization
809:Earth sciences portal
659:
657:{\displaystyle p_{i}}
629:
627:{\displaystyle p_{i}}
567:
565:{\displaystyle p_{i}}
515:
455:
378:
376:{\displaystyle p_{i}}
332:
260:
223:
221:{\displaystyle p_{i}}
169:
96:
2620:Evolutionary ecology
2585:Ecological footprint
2580:Ecological economics
2504:Ecological threshold
2499:Ecological indicator
2369:Source–sink dynamics
2322:Land change modeling
2317:Insular biogeography
2169:Species distribution
1908:Modelling ecosystems
1567:Microbial metabolism
1406:Intraguild predation
1195:Biogeochemical cycle
1161:Modelling ecosystems
1106:Connexions (cnx.org)
641:
611:
572:value. In practice,
549:
406:
360:
235:
205:
69:
46:of the species are.
2670:Theoretical ecology
2645:Natural environment
2509:Ecosystem diversity
2479:Ecological collapse
2469:Bateman's principle
2424:Limiting similarity
2337:Landscape limnology
2159:Species homogeneity
1997:Population modeling
1992:Population dynamics
1809:Trophic state index
1102:"Species Diversity"
599:Negative values of
295:
147:
2700:Ecological metrics
2681:Outline of ecology
2630:Industrial ecology
2625:Functional ecology
2489:Ecological deficit
2434:Niche construction
2397:Ecosystem engineer
2174:Species–area curve
2095:Introduced species
1910:: Other components
1842:Deimatic behaviour
1744:Ecological network
1676:North Pacific Gyre
1661:hydrothermal vents
1600:Ecological pyramid
1547:Microbial food web
1358:Primary production
1303:Foundation species
850:Coexistence theory
781:Environment portal
744:Resampling methods
654:
624:
562:
510:
424:
373:
327:
281:
218:
164:
127:
2687:
2686:
2570:Balance of nature
2327:Landscape ecology
2212:Community ecology
2154:Species diversity
2090:Indicator species
2085:Gradient analysis
1962:Logistic function
1870:
1869:
1827:Animal coloration
1804:Trophic mutualism
1542:Microbial ecology
1333:Photoheterotrophs
1318:Myco-heterotrophy
1230:Ecosystem ecology
1215:Carrying capacity
1180:Abiotic component
865:Genetic diversity
682:diversity indices
676:Diversity indices
409:
183:species abundance
162:
160:
32:species abundance
20:Species diversity
2722:
2387:Ecological niche
2359:selection theory
2179:Umbrella species
2164:Species richness
2100:Invasive species
2080:Flagship species
1987:Population cycle
1982:Overexploitation
1947:Ecological yield
1897:
1890:
1883:
1874:
1873:
1779:Mesotrophic soil
1719:Climax community
1651:Marine food webs
1590:Biomagnification
1391:Chemoorganotroph
1245:Keystone species
1205:Biotic component
1150:
1143:
1136:
1127:
1126:
1118:(Licensed under
1117:
1115:
1113:
1087:
1081:
1075:
1072:
1066:
1065:
1037:
1031:
1028:
1022:
1012:
1006:
1003:
997:
994:
985:
982:
976:
973:
967:
957:
951:
941:
928:
918:
905:
902:
811:
806:
805:
804:
797:
792:
791:
790:
783:
778:
777:
723:sampling methods
686:species richness
663:
661:
660:
655:
653:
652:
633:
631:
630:
625:
623:
622:
571:
569:
568:
563:
561:
560:
519:
517:
516:
511:
506:
502:
501:
500:
485:
484:
474:
469:
433:
432:
427:
423:
382:
380:
379:
374:
372:
371:
336:
334:
333:
328:
326:
325:
309:
300:
296:
294:
289:
279:
274:
246:
245:
240:
227:
225:
224:
219:
217:
216:
187:generalized mean
173:
171:
170:
165:
163:
161:
159:
148:
146:
135:
126:
125:
115:
110:
94:
89:
80:
79:
74:
56:weighted average
40:species evenness
36:species richness
2730:
2729:
2725:
2724:
2723:
2721:
2720:
2719:
2690:
2689:
2688:
2683:
2674:
2660:Systems ecology
2548:
2519:Extinction debt
2484:Ecological debt
2474:Bioluminescence
2455:
2448:
2417:marine habitats
2392:Ecological trap
2373:
2253:
2246:
2189:
2183:
2139:Rapoport's rule
2134:Priority effect
2075:Endemic species
2043:
2002:Population size
1918:
1911:
1901:
1871:
1866:
1819:
1813:
1799:Trophic cascade
1709:Bioaccumulation
1692:
1619:
1576:
1498:
1465:
1362:
1274:
1235:Ecosystem model
1168:
1154:
1111:
1109:
1096:
1091:
1090:
1082:
1078:
1073:
1069:
1049:Williams, W. T.
1038:
1034:
1029:
1025:
1013:
1009:
1004:
1000:
995:
988:
983:
979:
974:
970:
958:
954:
942:
931:
919:
908:
903:
894:
889:
884:
860:Gamma diversity
833:Diversity index
818:Alpha diversity
807:
802:
800:
793:
788:
786:
779:
772:
769:
756:
718:
678:
648:
644:
642:
639:
638:
618:
614:
612:
609:
608:
556:
552:
550:
547:
546:
535:arithmetic mean
525:Shannon entropy
496:
492:
480:
476:
470:
459:
451:
447:
428:
426:
425:
413:
407:
404:
403:
367:
363:
361:
358:
357:
305:
301:
290:
285:
275:
264:
259:
255:
254:
241:
239:
238:
236:
233:
232:
212:
208:
206:
203:
202:
149:
136:
131:
121:
117:
111:
100:
95:
93:
88:
75:
73:
72:
70:
67:
66:
52:
17:
12:
11:
5:
2728:
2718:
2717:
2712:
2707:
2702:
2685:
2684:
2679:
2676:
2675:
2673:
2672:
2667:
2662:
2657:
2652:
2647:
2642:
2640:Microecosystem
2637:
2632:
2627:
2622:
2617:
2612:
2607:
2602:
2597:
2592:
2587:
2582:
2577:
2572:
2567:
2562:
2556:
2554:
2550:
2549:
2547:
2546:
2541:
2539:Thorson's rule
2536:
2531:
2526:
2521:
2516:
2511:
2506:
2501:
2496:
2491:
2486:
2481:
2476:
2471:
2466:
2464:Assembly rules
2460:
2458:
2450:
2449:
2447:
2446:
2441:
2436:
2431:
2426:
2421:
2420:
2419:
2409:
2404:
2399:
2394:
2389:
2383:
2381:
2375:
2374:
2372:
2371:
2366:
2361:
2349:
2347:Patch dynamics
2344:
2342:Metapopulation
2339:
2334:
2329:
2324:
2319:
2314:
2309:
2304:
2299:
2294:
2289:
2284:
2279:
2274:
2269:
2264:
2258:
2256:
2248:
2247:
2245:
2244:
2239:
2237:Storage effect
2234:
2229:
2224:
2219:
2214:
2209:
2204:
2199:
2193:
2191:
2185:
2184:
2182:
2181:
2176:
2171:
2166:
2161:
2156:
2151:
2146:
2141:
2136:
2131:
2126:
2121:
2119:Neutral theory
2116:
2111:
2106:
2104:Native species
2097:
2092:
2087:
2082:
2077:
2072:
2067:
2062:
2057:
2051:
2049:
2045:
2044:
2042:
2041:
2036:
2035:
2034:
2029:
2019:
2014:
2009:
2004:
1999:
1994:
1989:
1984:
1979:
1977:Overpopulation
1974:
1969:
1964:
1959:
1954:
1949:
1944:
1939:
1934:
1929:
1923:
1921:
1913:
1912:
1900:
1899:
1892:
1885:
1877:
1868:
1867:
1865:
1864:
1859:
1854:
1849:
1844:
1839:
1834:
1829:
1823:
1821:
1815:
1814:
1812:
1811:
1806:
1801:
1796:
1791:
1786:
1784:Nutrient cycle
1781:
1776:
1774:Feeding frenzy
1771:
1766:
1761:
1756:
1754:Energy quality
1751:
1746:
1741:
1736:
1731:
1726:
1721:
1716:
1714:Cascade effect
1711:
1706:
1700:
1698:
1694:
1693:
1691:
1690:
1689:
1688:
1683:
1678:
1673:
1668:
1663:
1658:
1648:
1643:
1638:
1633:
1627:
1625:
1621:
1620:
1618:
1617:
1612:
1607:
1602:
1597:
1592:
1586:
1584:
1578:
1577:
1575:
1574:
1569:
1564:
1559:
1557:Microbial loop
1554:
1549:
1544:
1539:
1534:
1529:
1524:
1522:Lithoautotroph
1519:
1514:
1508:
1506:
1504:Microorganisms
1500:
1499:
1497:
1496:
1491:
1486:
1481:
1475:
1473:
1467:
1466:
1464:
1463:
1461:Prey switching
1458:
1453:
1448:
1443:
1438:
1433:
1428:
1423:
1418:
1413:
1408:
1403:
1398:
1393:
1388:
1383:
1378:
1372:
1370:
1364:
1363:
1361:
1360:
1355:
1350:
1345:
1340:
1338:Photosynthesis
1335:
1330:
1325:
1320:
1315:
1310:
1305:
1300:
1295:
1293:Chemosynthesis
1290:
1284:
1282:
1276:
1275:
1273:
1272:
1267:
1262:
1257:
1252:
1247:
1242:
1237:
1232:
1227:
1222:
1217:
1212:
1207:
1202:
1197:
1192:
1187:
1185:Abiotic stress
1182:
1176:
1174:
1170:
1169:
1153:
1152:
1145:
1138:
1130:
1124:
1123:
1095:
1094:External links
1092:
1089:
1088:
1076:
1067:
1032:
1023:
1007:
998:
986:
977:
968:
952:
929:
906:
891:
890:
888:
885:
883:
882:
877:
872:
867:
862:
857:
855:Dark diversity
852:
847:
842:
841:
840:
835:
825:
823:Beta diversity
820:
814:
813:
812:
798:
795:Ecology portal
784:
768:
765:
755:
752:
717:
714:
677:
674:
651:
647:
621:
617:
559:
555:
521:
520:
509:
505:
499:
495:
491:
488:
483:
479:
473:
468:
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458:
454:
450:
446:
443:
440:
437:
431:
422:
419:
416:
412:
370:
366:
338:
337:
324:
321:
318:
315:
312:
308:
304:
299:
293:
288:
284:
278:
273:
270:
267:
263:
258:
253:
250:
244:
215:
211:
201:th species is
189:with exponent
175:
174:
158:
155:
152:
145:
142:
139:
134:
130:
124:
120:
114:
109:
106:
103:
99:
92:
87:
84:
78:
51:
48:
15:
9:
6:
4:
3:
2:
2727:
2716:
2713:
2711:
2708:
2706:
2703:
2701:
2698:
2697:
2695:
2682:
2677:
2671:
2668:
2666:
2665:Urban ecology
2663:
2661:
2658:
2656:
2653:
2651:
2648:
2646:
2643:
2641:
2638:
2636:
2633:
2631:
2628:
2626:
2623:
2621:
2618:
2616:
2613:
2611:
2608:
2606:
2603:
2601:
2598:
2596:
2593:
2591:
2588:
2586:
2583:
2581:
2578:
2576:
2573:
2571:
2568:
2566:
2563:
2561:
2558:
2557:
2555:
2551:
2545:
2542:
2540:
2537:
2535:
2532:
2530:
2527:
2525:
2524:Kleiber's law
2522:
2520:
2517:
2515:
2512:
2510:
2507:
2505:
2502:
2500:
2497:
2495:
2492:
2490:
2487:
2485:
2482:
2480:
2477:
2475:
2472:
2470:
2467:
2465:
2462:
2461:
2459:
2457:
2451:
2445:
2442:
2440:
2437:
2435:
2432:
2430:
2427:
2425:
2422:
2418:
2415:
2414:
2413:
2410:
2408:
2405:
2403:
2400:
2398:
2395:
2393:
2390:
2388:
2385:
2384:
2382:
2380:
2376:
2370:
2367:
2365:
2362:
2360:
2358:
2354:
2350:
2348:
2345:
2343:
2340:
2338:
2335:
2333:
2330:
2328:
2325:
2323:
2320:
2318:
2315:
2313:
2310:
2308:
2305:
2303:
2300:
2298:
2297:Foster's rule
2295:
2293:
2290:
2288:
2285:
2283:
2280:
2278:
2275:
2273:
2270:
2268:
2265:
2263:
2260:
2259:
2257:
2255:
2249:
2243:
2240:
2238:
2235:
2233:
2230:
2228:
2225:
2223:
2220:
2218:
2215:
2213:
2210:
2208:
2205:
2203:
2200:
2198:
2195:
2194:
2192:
2186:
2180:
2177:
2175:
2172:
2170:
2167:
2165:
2162:
2160:
2157:
2155:
2152:
2150:
2147:
2145:
2142:
2140:
2137:
2135:
2132:
2130:
2127:
2125:
2122:
2120:
2117:
2115:
2112:
2110:
2107:
2105:
2101:
2098:
2096:
2093:
2091:
2088:
2086:
2083:
2081:
2078:
2076:
2073:
2071:
2068:
2066:
2063:
2061:
2058:
2056:
2053:
2052:
2050:
2046:
2040:
2037:
2033:
2030:
2028:
2025:
2024:
2023:
2020:
2018:
2015:
2013:
2010:
2008:
2005:
2003:
2000:
1998:
1995:
1993:
1990:
1988:
1985:
1983:
1980:
1978:
1975:
1973:
1970:
1968:
1965:
1963:
1960:
1958:
1955:
1953:
1950:
1948:
1945:
1943:
1940:
1938:
1935:
1933:
1930:
1928:
1925:
1924:
1922:
1920:
1914:
1909:
1905:
1898:
1893:
1891:
1886:
1884:
1879:
1878:
1875:
1863:
1860:
1858:
1855:
1853:
1850:
1848:
1845:
1843:
1840:
1838:
1835:
1833:
1830:
1828:
1825:
1824:
1822:
1816:
1810:
1807:
1805:
1802:
1800:
1797:
1795:
1792:
1790:
1787:
1785:
1782:
1780:
1777:
1775:
1772:
1770:
1767:
1765:
1762:
1760:
1757:
1755:
1752:
1750:
1747:
1745:
1742:
1740:
1737:
1735:
1732:
1730:
1727:
1725:
1722:
1720:
1717:
1715:
1712:
1710:
1707:
1705:
1702:
1701:
1699:
1695:
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1684:
1682:
1679:
1677:
1674:
1672:
1669:
1667:
1664:
1662:
1659:
1657:
1654:
1653:
1652:
1649:
1647:
1644:
1642:
1639:
1637:
1634:
1632:
1629:
1628:
1626:
1622:
1616:
1615:Trophic level
1613:
1611:
1608:
1606:
1603:
1601:
1598:
1596:
1593:
1591:
1588:
1587:
1585:
1583:
1579:
1573:
1572:Phage ecology
1570:
1568:
1565:
1563:
1562:Microbial mat
1560:
1558:
1555:
1553:
1550:
1548:
1545:
1543:
1540:
1538:
1535:
1533:
1530:
1528:
1525:
1523:
1520:
1518:
1517:Bacteriophage
1515:
1513:
1510:
1509:
1507:
1505:
1501:
1495:
1492:
1490:
1487:
1485:
1484:Decomposition
1482:
1480:
1477:
1476:
1474:
1472:
1468:
1462:
1459:
1457:
1454:
1452:
1449:
1447:
1444:
1442:
1439:
1437:
1434:
1432:
1431:Mesopredators
1429:
1427:
1424:
1422:
1419:
1417:
1414:
1412:
1409:
1407:
1404:
1402:
1399:
1397:
1394:
1392:
1389:
1387:
1384:
1382:
1379:
1377:
1376:Apex predator
1374:
1373:
1371:
1369:
1365:
1359:
1356:
1354:
1351:
1349:
1346:
1344:
1341:
1339:
1336:
1334:
1331:
1329:
1326:
1324:
1321:
1319:
1316:
1314:
1311:
1309:
1306:
1304:
1301:
1299:
1296:
1294:
1291:
1289:
1286:
1285:
1283:
1281:
1277:
1271:
1268:
1266:
1263:
1261:
1258:
1256:
1253:
1251:
1248:
1246:
1243:
1241:
1238:
1236:
1233:
1231:
1228:
1226:
1223:
1221:
1218:
1216:
1213:
1211:
1210:Biotic stress
1208:
1206:
1203:
1201:
1198:
1196:
1193:
1191:
1188:
1186:
1183:
1181:
1178:
1177:
1175:
1171:
1166:
1162:
1158:
1151:
1146:
1144:
1139:
1137:
1132:
1131:
1128:
1121:
1107:
1103:
1098:
1097:
1085:
1080:
1071:
1064:
1060:
1056:
1055:
1050:
1046:
1045:Tracey, J. G.
1042:
1036:
1027:
1021:
1017:
1011:
1002:
993:
991:
981:
972:
966:
962:
956:
950:
946:
940:
938:
936:
934:
927:
923:
917:
915:
913:
911:
901:
899:
897:
892:
881:
878:
876:
873:
871:
868:
866:
863:
861:
858:
856:
853:
851:
848:
846:
843:
839:
836:
834:
831:
830:
829:
826:
824:
821:
819:
816:
815:
810:
799:
796:
785:
782:
776:
771:
764:
762:
751:
749:
745:
739:
737:
733:
729:
726:the value of
724:
713:
711:
707:
703:
697:
695:
694:Simpson index
691:
690:Shannon index
687:
683:
673:
671:
667:
649:
645:
635:
619:
615:
606:
602:
597:
595:
591:
587:
583:
579:
575:
557:
553:
544:
540:
536:
532:
528:
526:
507:
503:
497:
493:
489:
486:
481:
477:
471:
466:
463:
460:
456:
452:
448:
444:
441:
438:
435:
429:
420:
414:
402:
401:
400:
398:
394:
390:
386:
368:
364:
355:
352:, which is 1/
351:
350:harmonic mean
347:
343:
340:The value of
319:
316:
313:
306:
302:
297:
291:
286:
282:
276:
271:
268:
265:
261:
256:
251:
248:
242:
231:
230:
229:
213:
209:
200:
196:
192:
188:
184:
180:
156:
153:
150:
143:
140:
137:
132:
128:
122:
118:
112:
107:
104:
101:
97:
90:
85:
82:
76:
65:
64:
63:
61:
57:
47:
45:
41:
37:
33:
29:
25:
21:
2650:Regime shift
2635:Macroecology
2356:
2352:
2292:Edge effects
2262:Biogeography
2207:Commensalism
2153:
2055:Biodiversity
1932:Allee effect
1671:kelp forests
1624:Example webs
1489:Detritivores
1328:Organotrophs
1308:Kinetotrophs
1260:Productivity
1110:. Retrieved
1105:
1079:
1070:
1053:
1035:
1026:
1010:
1001:
980:
971:
955:
828:Biodiversity
757:
740:
735:
731:
727:
719:
709:
705:
701:
698:
679:
669:
665:
636:
604:
600:
598:
593:
589:
585:
581:
577:
573:
538:
530:
529:
522:
396:
392:
388:
384:
356:because the
353:
345:
341:
339:
198:
194:
190:
176:
53:
19:
18:
2287:Disturbance
2190:interaction
2012:Recruitment
1942:Depensation
1734:Copiotrophs
1605:Energy flow
1527:Lithotrophy
1471:Decomposers
1451:Planktivore
1426:Insectivore
1416:Heterotroph
1381:Bacterivore
1348:Phototrophs
1298:Chemotrophs
1270:Restoration
1220:Competition
1041:Webb, L. J.
541:approaches
179:denominator
2694:Categories
2655:Sexecology
2232:Parasitism
2197:Antibiosis
2032:Resistance
2027:Resilience
1917:Population
1837:Camouflage
1789:Oligotroph
1704:Ascendency
1666:intertidal
1656:cold seeps
1610:Food chain
1411:Herbivores
1386:Carnivores
1313:Mixotrophs
1288:Autotrophs
1167:components
1112:1 February
44:abundances
2560:Allometry
2514:Emergence
2242:Symbiosis
2227:Mutualism
2022:Stability
1927:Abundance
1739:Dominance
1697:Processes
1686:tide pool
1582:Food webs
1456:Predation
1441:Omnivores
1368:Consumers
1323:Mycotroph
1280:Producers
1225:Ecosystem
1190:Behaviour
490:
457:∑
453:−
445:
418:→
317:−
262:∑
154:−
141:−
98:∑
28:community
2615:Endolith
2544:Xerosere
2456:networks
2272:Ecocline
1818:Defense,
1494:Detritus
1396:Foraging
1265:Resource
1084:Chao, A.
767:See also
761:habitats
543:infinity
2705:Habitat
2605:Ecopath
2412:Habitat
2282:Ecotype
2277:Ecotone
2254:ecology
2252:Spatial
2188:Species
2048:Species
1919:ecology
1904:Ecology
1852:Mimicry
1820:counter
1764:f-ratio
1512:Archaea
1200:Biomass
1173:General
1165:Trophic
1157:Ecology
1063:2257891
730:: when
60:inverse
24:species
1636:Rivers
1532:Marine
1061:
754:Trends
692:, the
688:, the
2553:Other
2454:Other
2407:Guild
2379:Niche
1631:Lakes
1059:JSTOR
887:Notes
537:. As
1641:Soil
1114:2011
700:log(
177:The
1016:doi
961:doi
945:doi
922:doi
442:exp
411:lim
2696::
2102:/
1906::
1163::
1159::
1122:).
1104:.
1047:;
1043:;
989:^
932:^
909:^
895:^
527:.
487:ln
391:.
2357:K
2355:/
2353:r
1896:e
1889:t
1882:v
1149:e
1142:t
1135:v
1116:.
1018::
963::
947::
924::
736:q
732:q
728:q
710:D
706:D
702:D
670:D
666:i
650:i
646:p
620:i
616:p
605:q
601:q
594:q
590:q
586:q
582:q
578:q
574:q
558:i
554:p
539:q
531:q
508:,
504:)
498:i
494:p
482:i
478:p
472:S
467:1
464:=
461:i
449:(
439:=
436:D
430:q
421:1
415:q
397:q
393:q
389:S
385:D
369:i
365:p
354:S
346:q
342:q
323:)
320:q
314:1
311:(
307:/
303:1
298:)
292:q
287:i
283:p
277:S
272:1
269:=
266:i
257:(
252:=
249:D
243:q
214:i
210:p
199:i
195:S
191:q
157:1
151:q
144:1
138:q
133:i
129:p
123:i
119:p
113:S
108:1
105:=
102:i
91:1
86:=
83:D
77:q
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
