Biology:Beta diversity
In ecology, beta diversity (β-diversity or true beta diversity) is the ratio between regional and local species diversity. The term was introduced by R. H. Whittaker[1] together with the terms alpha diversity (α-diversity) and gamma diversity (γ-diversity). The idea was that the total species diversity in a landscape (γ) is determined by two different things: the mean species diversity at the local level (α) and the differentiation among local sites (β). Other formulations for beta diversity include "absolute species turnover", "Whittaker's species turnover" and "proportional species turnover".[citation needed]
Whittaker proposed several ways of quantifying differentiation, and subsequent generations of ecologists have invented more. As a result, there are now many defined types of beta diversity.[2][3] Some use beta diversity to refer to any of several indices related to compositional heterogeneity.[4][5][6] Confusion is avoided by using distinct names for other formulations.[2][3][7][8][9][10]
Beta diversity as a measure of species turnover overemphasizes the role of rare species as the difference in species composition between two sites or communities is likely reflecting the presence and absence of some rare species in the assemblages. Beta diversity can also be a measure of nestedness, which occurs when species assemblages in species-poor sites are a subset of the assemblages in more species-rich sites.[11] Moreover, pairwise beta diversity are inadequate in building all biodiversity partitions (some partitions in a Venn diagram of 3 or more sites cannot be expressed by alpha and beta diversity). Consequently, some macroecological and community patterns cannot be fully expressed by alpha and beta diversity. Due to these two reasons, a new way of measuring species turnover, coined Zeta diversity (ζ-diversity),[12] has been proposed and used to connect all existing incidence-based biodiversity patterns.
Types
Whittaker beta diversity
Gamma diversity and alpha diversity can be calculated directly from species inventory data.[2][13] The simplest of Whittaker's original definitions of beta diversity is
β = γ/α
Here gamma diversity is the total species diversity of a landscape and alpha diversity is the mean species diversity per site. Because the limits among local sites and landscapes are diffuse and to some degree subjective, it has been proposed that gamma diversity can be quantified for any inventory dataset and that alpha and beta diversity can be quantified whenever the dataset is divided into subunits. Then gamma diversity is the total species diversity in the dataset and alpha diversity the mean species diversity per subunit. Beta diversity quantifies how many subunits there would be if the total species diversity of the dataset and the mean species diversity per subunit remained the same, but the subunits shared no species.[2][7]
Absolute species turnover
Some researchers have preferred to partition gamma diversity into additive rather than multiplicative components.[14][15] Then the beta component of diversity becomes
βA = γ - α
This quantifies how much more species diversity the entire dataset contains than an average subunit within the dataset. This can also be interpreted as the total amount of species turnover among the subunits in the dataset.[2]
When there are two subunits, and presence-absence data are used, this can be calculated with the following equation:
[math]\displaystyle{ \beta_A=(S_1-c)+(S_2-c) }[/math][16]
where, S1= the total number of species recorded in the first community, S2= the total number of species recorded in the second community, and c= the number of species common to both communities.
Whittaker's species turnover
If absolute species turnover is divided by alpha diversity, a measure is obtained that quantifies how many times the species composition changes completely among the subunits of the dataset. This measure was proposed by Whittaker,[17] so it has been called Whittaker's species turnover.[2] It is calculated as
βW = (γ - α)/α = γ/α - 1
When there are two subunits, and presence-absence data are used, this equals the one-complement of the Sørensen similarity index.[2][18]
Proportional species turnover
If absolute species turnover is divided by gamma diversity, a measure is obtained that quantifies what proportion of the species diversity in the dataset is not contained in an average subunit.[2] It is calculated as
βP = (γ - α)/γ = 1 - α/γ
When there are two subunits, and presence-absence data are used, this measure as ranged to the interval [0, 1] equals the one-complement of the Jaccard similarity index.[2]
β-diversity patterns
Although understanding the change in species composition from local to regional scales (β-diversity) is a central theme in ecology and biogeography, studies often reached different conclusions as to the fundamental patterns in β-diversity. For example, niche compression hypothesis predicted higher β-diversity at lower latitudes.[20][19][21][22][23] Studies comparing natural local sites with human-modified local sites are no different. Kitching et al.[24] sampled moths in primary and logged forests of Danum valley, Borneo to show that β-diversity in primary forests was higher than logged forests. Contrastingly, Berry et al.[25] sampled trees in the same study area to show that β-diversity in logged forests was higher than primary forests. The results of these two studies were completely different from the results of a recent quantitative synthesis,[26] which showed that β-diversity in primary forests were similar to β-diversity in all types of human-modified local sites (secondary forests, plantations, pasture and urban). Therefore, there is a clear lack of consensus on β-diversity patterns among studies. Sreekar et al.[19] suggested that most of these inconsistencies were due to the differences in grain size and/or spatial extent among studies. They showed that spatial scale changes the relationship between β-diversity and latitude.
Diversity partitioning in the geologic past
Major diversification events in the geologic past were associated with shifts in the relative contributions of alpha- and beta-diversity (diversity partitioning). Examples include the Cambrian explosion,[27] the great Ordovician biodiversification event,[28] and the recoveries from the end-Permian[29] and end-Triassic[30] mass extinction events. Empirical data from these case studies confirm theoretical predictions that an increasing number of species will increase beta-diversity relative to alpha diversity because of the effects from interspecific competition; yet, alpha diversity may increase again once options for increasing geographic turnover are exhausted.[31]
See also
- Alpha diversity
- Biotic homogenization
- Bray–Curtis dissimilarity
- Dark diversity
- Diversity index
- Gamma diversity
- Global biodiversity
- Jaccard distance
- Measurement of biodiversity
- Zeta diversity
References
- ↑ "Vegetation of the Siskiyou Mountains, Oregon and California.". Ecological Monographs 30 (3): 279–338. 1960. doi:10.2307/1943563. Bibcode: 1960EcoM...30..279W.
- ↑ 2.0 2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 "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. 2010. doi:10.1111/j.1600-0587.2009.05880.x. Bibcode: 2010Ecogr..33....2T.
- ↑ 3.0 3.1 "A diversity of beta diversities: straightening up a concept gone awry. Part 2. Quantifying beta diversity and related phenomena.". Ecography 33 (1): 23–45. 2010. doi:10.1111/j.1600-0587.2009.06148.x. Bibcode: 2010Ecogr..33...23T.
- ↑ "Measuring beta diversity for presence–absence data.". Journal of Animal Ecology 72 (3): 367–382. 2003. doi:10.1046/j.1365-2656.2003.00710.x. Bibcode: 2003JAnEc..72..367K.
- ↑ "Navigating the multiple meanings of β diversity: a roadmap for the practicing ecologist". Ecology Letters 14 (1): 19–28. January 2011. doi:10.1111/j.1461-0248.2010.01552.x. PMID 21070562. Bibcode: 2011EcolL..14...19A.
- ↑ "Commentary: Do we have a consistent terminology for species diversity? The fallacy of true diversity". Oecologia 167 (4): 885–8; discussion 903–11. December 2011. doi:10.1007/s00442-011-2124-8. PMID 21947497. Bibcode: 2011Oecol.167..885G.
- ↑ 7.0 7.1 Tuomisto, H. 2010. A consistent terminology for quantifying species diversity? Yes, it does exist. Oecologia 4: 853–860. doi:10.1007/s00442-010-1812-0
- ↑ "Commentary: do we have a consistent terminology for species diversity? We are on the way". Oecologia 167 (4): 893–902; discussion 903–11. December 2011. doi:10.1007/s00442-011-2126-6. PMID 21938639. Bibcode: 2011Oecol.167..893J.
- ↑ "Commentary: Do we have a consistent terminology for species diversity? Back to basics and toward a unifying framework". Oecologia 167 (4): 889–92; discussion 903–11. December 2011. doi:10.1007/s00442-011-2125-7. PMID 21965142. Bibcode: 2011Oecol.167..889M.
- ↑ "Commentary: do we have a consistent terminology for species diversity? Yes, if we choose to use it.". Oecologia 167 (4): 903–911. 2011. doi:10.1007/s00442-011-2128-4. Bibcode: 2011Oecol.167..903T.
- ↑ "Partitioning the turnover and nestedness components of beta diversity" (in en). Global Ecology and Biogeography 19 (1): 134–143. January 2010. doi:10.1111/j.1466-8238.2009.00490.x. ISSN 1466-8238. Bibcode: 2010GloEB..19..134B.
- ↑ "Zeta diversity as a concept and metric that unifies incidence-based biodiversity patterns.". American Naturalist 184 (5): 684–694. 2014. doi:10.1086/678125. PMID 25325751.
- ↑ "Phylogenetic diversity measures based on Hill numbers". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 365 (1558): 3599–609. November 2010. doi:10.1098/rstb.2010.0272. PMID 20980309.
- ↑ "Statistics and partitioning of species diversity, and similarity among multiple communities.". Oikos 1 (1): 5–13. May 1996. doi:10.2307/3545743. Bibcode: 1996Oikos..76....5L.
- ↑ "The additive partitioning of species diversity: recent revival of an old idea.". Oikos 99 (1): 3–9. October 2002. doi:10.1034/j.1600-0706.2002.990101.x. Bibcode: 2002Oikos..99....3V.
- ↑ Albert, James S.; Reis, Roberto E. (8 March 2011). Historical Biogeography of Neotropical Freshwater Fishes. University of California Press. p. 308. ISBN 978-0-520-26868-5. https://books.google.com/books?id=_Suu7a-ERdMC&pg=PA308. Retrieved 28 June 2011.
- ↑ "Evolution and measurement of species diversity.". Taxon 21 (2–3): 213–251. 1972. doi:10.2307/1218190.
- ↑ "A method of establishing groups of equal amplitude in plant sociology based on similarity of species content, and its application to analyses of the vegetation on Danish commons.". Kongelige Danske Videnskabernes Selskabs Biologiske Skrifter 5: 1–34. 1948.
- ↑ 19.0 19.1 19.2 "Spatial scale changes the relationship between beta diversity, species richness and latitude". Royal Society Open Science 5 (9): 181168. September 2018. doi:10.1098/rsos.181168. PMID 30839691. Bibcode: 2018RSOS....581168S.
- ↑ "Patterns of species diversity.". Biological Reviews 40 (4): 510–33. November 1965. doi:10.1111/j.1469-185X.1965.tb00815.x.
- ↑ "Are there latitudinal gradients in species turnover?". Global Ecology and Biogeography 12 (6): 483–98. November 2003. doi:10.1046/j.1466-822X.2003.00056.x. Bibcode: 2003GloEB..12..483K.
- ↑ "Disentangling the drivers of β diversity along latitudinal and elevational gradients". Science (New York, N.Y.) 333 (6050): 1755–8. September 2011. doi:10.1126/science.1208584. PMID 21940897. Bibcode: 2011Sci...333.1755K.
- ↑ "Drivers of β‐diversity along latitudinal gradients revisited.". Global Ecology and Biogeography 22 (6): 659–70. June 2013. doi:10.1111/geb.12020. Bibcode: 2013GloEB..22..659Q.
- ↑ "Distance‐driven species turnover in Bornean rainforests: homogeneity and heterogeneity in primary and post‐logging forests.". Ecography 36 (6): 675–82. June 2013. doi:10.1111/j.1600-0587.2012.00023.x.
- ↑ "Impacts of selective logging on tree diversity across a rainforest landscape: the importance of spatial scale.". Landscape Ecology 23 (8): 915–29. October 2008. doi:10.1007/s10980-008-9248-1.
- ↑ "Global patterns of terrestrial assemblage turnover within and among land uses.". Ecography 39 (12): 1151–63. December 2016. doi:10.1111/ecog.01932. Bibcode: 2016Ecogr..39.1151N.
- ↑ "Diversity partitioning during the Cambrian radiation". Proceedings of the National Academy of Sciences of the United States of America 112 (15): 4702–6. April 2015. doi:10.1073/pnas.1424985112. PMID 25825755. Bibcode: 2015PNAS..112.4702N.
- ↑ "Alpha, beta, or gamma: where does all the diversity go?". Paleobiology 14 (3): 221–34. 1988. doi:10.1017/s0094837300011969. PMID 11542147. Bibcode: 1988Pbio...14..221S.
- ↑ "Diversity partitioning in Permian and Early Triassic benthic ecosystems of the Western USA: a comparison" (in en). Historical Biology 29 (7): 918–930. 2017-10-03. doi:10.1080/08912963.2016.1263626. Bibcode: 2017HBio...29..918H.
- ↑ "Diversity partitioning in Jurassic level-bottom communities" (in en). Historical Biology 28 (6): 849–857. 2016. doi:10.1080/08912963.2015.1051482. Bibcode: 2016HBio...28..849H.
- ↑ Hautmann, Michael (2014). "Diversification and diversity partitioning" (in en). Paleobiology 40 (2): 162–176. doi:10.1666/13041.
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