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Origins & Maintenance of Diversity A Hierarchy of Processes

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1 Origins & Maintenance of Diversity A Hierarchy of Processes
Please do not use the images in these PowerPoint slides without permission.

2 Origins & Maintenance of Diversity A Hierarchy of Processes
Processes operate on a variety of temporal & spatial scales to influence diversity The levels are nested Unique (chance) events may occur at any level Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Figure 30.2 from Ricklefs & Schluter (1993)

3 Origins & Maintenance of Diversity A Hierarchy of Processes
Processes operate on a variety of temporal & spatial scales to influence diversity The levels are nested Unique (chance) events may occur at any level Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Figure 30.2 from Ricklefs & Schluter (1993)

4 Origins & Maintenance of Diversity A Hierarchy of Processes
Processes operate on a variety of temporal & spatial scales to influence diversity The levels are nested Unique (chance) events may occur at any level Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Figure 30.2 from Ricklefs & Schluter (1993)

5 Origins & Maintenance of Diversity A Hierarchy of Processes
Processes operate on a variety of temporal & spatial scales to influence diversity The levels are nested Unique (chance) events may occur at any level Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Figure 30.2 from Ricklefs & Schluter (1993)

6 Origins & Maintenance of Diversity A Hierarchy of Processes
Processes operate on a variety of temporal & spatial scales to influence diversity The levels are nested Unique (chance) events may occur at any level Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Figure 30.2 from Ricklefs & Schluter (1993)

7 Origins & Maintenance of Diversity A Hierarchy of Processes
Processes operate on a variety of temporal & spatial scales to influence diversity The levels are nested Unique (chance) events may occur at any level Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Figure 30.2 from Ricklefs & Schluter (1993)

8 Origins & Maintenance of Diversity A Hierarchy of Processes
Biotal interchange Stochastic extinction Regional diversity Local diversity Species production Habitat selection Competitive exclusion Mass extinction Predatory exclusion Please do not use the images in these PowerPoint slides without permission. This is similar to some of the flow diagrams presented in the first lecture on “What is Community Ecology?” R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. A major challenge confronting advances to a research program that investigates this multi-scale set of processes is the reconstruction of historical events So, community ecologists have added phylogenetic & biogeographic analyses (e.g., phylogeography) to their toolkits… Redrawn from figure 30.1, Ricklefs & Schluter (1993)

9 Origins & Maintenance of Diversity Phylogenetic Perspectives
Please do not use the images in these PowerPoint slides without permission.

10 Origins & Maintenance of Diversity Phylogenetic Perspectives
Losos (1996) “If species adapted rapidly and without constraint, and if any lineage could occur in any community, then we would expect differences in community structure to be indicative of environmental differences. Because lineages differ in their evolutionary potential and are geographically restricted, however, comparisons of community structure must take account of communities’ histories…” “…differences in the structure of communities may result as much from the differences in the lineages that occur in different communities as from differences in environmental conditions” Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: “Phylogenetic information about the constituent lineages in a community can allow lineage effects to be factored out, thus allowing an assessment of environmental determinants of community structure”

11 Origins & Maintenance of Diversity Phylogenetic Perspectives
Ricklefs & Schluter (1993) An example from passerine birds compared between Panama and Illinois: Why do different areas vary in species richness of particular clades? “The approximately twofold difference in average age of the taxa matches the twofold difference in diversity, although age and diversity are not well correlated among individual clades.” Please do not use the images in these PowerPoint slides without permission. R. E. Ricklefs & D. Schluter, eds Species Diversity in Ecological Communities. University of Chicago Press, Chicago, IL, USA. Table 30.2 from Ricklefs & Schluter (1993)

12 Origins & Maintenance of Diversity Phylogenetic Perspectives
Losos (1990, 1996) An example from Anolis lizards inhabiting Caribbean islands: Where & under what community conditions did patterns of niche-use originate? Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77:

13 Origins & Maintenance of Diversity Phylogenetic Perspectives
27 islands of the Lesser Antilles contain either one or two species of Anolis lizards; on 9 of the 10 two-species islands, the species differ considerably in size (one large & one small), whereas on 16 of 17 one-species islands, the species are intermediate in size Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: Figure from Losos (1996)

14 Origins & Maintenance of Diversity Phylogenetic Perspectives
Hypotheses: (1) “Size adjustment (character displacement): Species of similar size colonize an island & evolve in opposite directions in situ to minimize resource competition” Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: From Losos (1990)

15 Origins & Maintenance of Diversity Phylogenetic Perspectives
Size adjustment Large 2 Large 2 Large 2 Small 2 Small 2 Small 2 Int. 1 Int. 1 Int. 1 Int. 1 (Int. 1) (Int. 1) (Int. 1) (Int. 1) (Int. 1) (Int. 1) (Int. 1) (Int. 1) Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Presumably the first three large species reading from the left at the branch tips are individually paired with the next three small species. In the case of size adjustment (character displacement), each species evolves to a new size upon colonizing an island with another species. (Int. 1) Increase in size Decrease in size Switch to 2-species island Redrawn from Losos (1990)

16 Origins & Maintenance of Diversity Phylogenetic Perspectives
Hypotheses: (1) “Size adjustment (character displacement): Species of similar size colonize an island & evolve in opposite directions in situ to minimize resource competition” (2) “Size assortment: Competitive exclusion or other processes prevent similar-sized species from colonizing the same island; only species that are already dissimilar in size can successfully colonize and coexist” Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: From Losos (1990)

17 Origins & Maintenance of Diversity Phylogenetic Perspectives
Size assortment Large 2 Large 2 Large 2 Small 2 Small 2 Small 2 Int. 1 Int. 1 Int. 1 Int. 1 (Int. 1) (Large 1) (Small 1) (Int. 1) (Large 1) (Small 1) (Int. 1) (Int. 1) Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Presumably the first three large species reading from the left at the branch tips are individually paired with the next three small species. In the case of size assortment, a species only colonizes an island in which its previously evolved size allows it to colonize with another species present. (Int. 1) Increase in size Decrease in size Switch to 2-species island Redrawn from Losos (1990)

18 Origins & Maintenance of Diversity
Phylogenetic Perspectives Size assortment & size adjustment Large 2 Large 2 Large 2 Small 2 Small 2 Small 2 Int. 1 Int. 1 Int. 1 Int. 1 (Int. 1) (Large 2) (Small 2) (Int. 1) (Large 2) (Small 2) (Int. 1) (Int. 1) Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Presumably the first three large species reading from the left at the branch tips are individually paired with the next three small species. In the case of size adjustment & assortment, “size adjustment occurs when two similar-sized species first co-occur; subsequent successful colonization of additional islands by descendents of both forms constitutes size assortment.” As drawn, it could be either a case of sympatric speciation and character divergence, followed by colonization of additional islands of large and small morphs, or colonization of a single island by two intermediate-sized ancestors that then diverge on their shared island and subsequently colonize other islands. (Int. 1) Increase in size Decrease in size Switch to 2-species island Redrawn from Losos (1990)

19 Origins & Maintenance of Diversity
Phylogenetic Perspectives No evidence for size assortment or size adjustment Large 2 Large 2 Int. 2 Small 2 Small 1 Small 1 Int. 1 Int. 2 Small 1 Int. 1 (Int. 1) (Large 2) (Small 1) (Int. 1) (Int. 2) (Small 1) (Int. 1) (Int. 1) Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Both the size pattern at the tips and the directionality of changes do not clearly suggest either size assortment nor size adjustment. (Int. 1) Increase in size Decrease in size Switch to 2-species island Redrawn from Losos (1990)

20 Origins & Maintenance of Diversity Phylogenetic Perspectives
“In the northern Lesser Antilles, large & small size appear to have evolved simultaneously when two lineages came into sympatry… exactly the prediction of a hypothesis of character displacement…” Dominica Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: Figure from Losos (1996); quote from Losos (1990)

21 Origins & Maintenance of Diversity Phylogenetic Perspectives
“By contrast, in the southern Lesser Antilles, evolutionary change in body size appears to have been unrelated to whether a species occurred in sympatry with congeners… Consequently, the existence of size patterns must have resulted from a process of ecological sorting in which only dissimilar-sized species can successfully colonize and coexist on the same island…” Dominica Martinique Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: Figure from Losos (1996); quote from Losos (1990)

22 Origins & Maintenance of Diversity Phylogenetic Perspectives
“Thus, the relative importance of co-evolutionary processes in determining community structure differs between the northern and southern Lesser Antilles, a result that is only apparent when these lizards are studied in a historical [phylogenetic] context” St. Maarten St. Eustatius Dominica Martinique Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B A phylogenetic analysis of character displacement in Caribbean Anolis lizards. Evolution 44: Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: St. Eustatius & St. Maarten figure prominently in Pacala & Roughgarden (1982) Science – 2 species on St. Eustatius have greater degree of resource partitioning and less competition compared to 2 species on St. Maarten. Figure from Losos (1996); quote from Losos (1990)

23 Origins & Maintenance of Diversity Phylogenetic Perspectives
McPeek (1995) An example from damselflies inhabiting freshwater lakes: Where and under what community conditions did patterns of niche-use originate? Please do not use the images in these PowerPoint slides without permission. McPeek, Mark A Morphological evolution mediated by behavior in the damselflies of two communities. Evolution 49:

24 Origins & Maintenance of Diversity Phylogenetic Perspectives
E. geminatum E. antennatum E. aspersum E. traviatum E. signatum E. vesperum E. boreale E. hageni 0.0 +0.1 +2.0 0.0 -0.5 0.0 +1.5 -0.3 +0.4 0.0 +0.9 0.0 0.0 0.0 Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: McPeek, Mark A Morphological evolution mediated by behavior in the damselflies of two communities. Evolution 49: Numbers represent the amount of evolutionary change in limb length between ancestor & descendant, as deduced using parsimony methods for reconstruction of ancestral character states Redrawn from Losos (1996), based on McPeek (1995)

25 Origins & Maintenance of Diversity Phylogenetic Perspectives
E. geminatum E. antennatum E. aspersum E. traviatum E. signatum E. vesperum E. boreale E. hageni 0.0 +0.1 +2.0 0.0 -0.5 0.0 +1.5 -0.3 +0.4 0.0 +0.9 0.0 0.0 0.0 Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: McPeek, Mark A Morphological evolution mediated by behavior in the damselflies of two communities. Evolution 49: The common ancestor inhabited lakes occupied by fish, where its best strategy to avoid predation was to hide Redrawn from Losos (1996), based on McPeek (1995)

26 Origins & Maintenance of Diversity Phylogenetic Perspectives
E. geminatum E. antennatum E. aspersum E. traviatum E. signatum E. vesperum E. boreale E. hageni 0.0 +0.1 +2.0 0.0 -0.5 0.0 +1.5 -0.3 +0.4 0.0 +0.9 0.0 0.0 0.0 Please do not use the images in these PowerPoint slides without permission. Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: McPeek, Mark A Morphological evolution mediated by behavior in the damselflies of two communities. Evolution 49: Bars represent transitions to fishless lakes, where a damselfly’s best strategy to avoid predation by dragonflies is to swim or run away Redrawn from Losos (1996), based on McPeek (1995)

27 Origins & Maintenance of Diversity Phylogenetic Perspectives
Cadle & Greene (1993) An example from Neotropical snake assemblages: For areas that differ in the species richness of particular clades, do contrasting characteristics of those clades contribute to emergent community-level properties? Please do not use the images in these PowerPoint slides without permission. Cadle, John E. & Harry W. Green Phylogenetic patterns, biogeography, and the ecological structure of Neotropical snake assemblages. Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL.

28 Origins & Maintenance of Diversity Phylogenetic Perspectives
Cadle & Greene (1993) Observations: Three main lineages of Neotropical snakes within the family Colubridae “Goo-eaters” Please do not use the images in these PowerPoint slides without permission. Cadle, John E. & Harry W. Green Phylogenetic patterns, biogeography, and the ecological structure of Neotropical snake assemblages. Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL.

29 Origins & Maintenance of Diversity Phylogenetic Perspectives
Cadle & Greene (1993) Observations: Three main lineages of Neotropical snakes within the family Colubridae The xenodontine lineages originated & diversified relatively isolated from one another, even though the current distributions overlap substantially Principal zones of radiation Colubrines radiated relatively recently worldwide Please do not use the images in these PowerPoint slides without permission. Cadle, John E. & Harry W. Green Phylogenetic patterns, biogeography, and the ecological structure of Neotropical snake assemblages. Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL.

30 Origins & Maintenance of Diversity Phylogenetic Perspectives
Cadle & Greene (1993) Observations: Three main lineages of Neotropical snakes within the family Colubridae The xenodontine lineages originated & diversified relatively isolated from one another, even though the current distributions overlap substantially The lineages differ in mean body size, diet, and etc. Please do not use the images in these PowerPoint slides without permission. Cadle, John E. & Harry W. Green Phylogenetic patterns, biogeography, and the ecological structure of Neotropical snake assemblages. Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL.

31 Origins & Maintenance of Diversity Phylogenetic Perspectives
Cadle & Greene (1993) An example from Neotropical snake assemblages: For areas that differ in the species richness of particular clades, do contrasting characteristics of those clades contribute to emergent community-level properties? Observation: Sites in Central & South America differ in body size distribution, diet & etc. of their snakes, but those differences are largely due to differences in the dominant clades occupying those sites Please do not use the images in these PowerPoint slides without permission. Cadle, John E. & Harry W. Green Phylogenetic patterns, biogeography, and the ecological structure of Neotropical snake assemblages. Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL. For example, the differential history of colonization of different sites by snakes of the three lineages is largely responsible for among-site differences in overall size distributions of their snake faunas, as opposed to evolutionary and ecological processes that maintain a particular size distribution in each site. Conclusion: The overall size distribution, diet & etc. of a community bears a direct relationship to the clade composition (i.e., history matters)

32 Origins & Maintenance of Diversity Phylogenetic Perspectives
Cadle & Greene (1993) An example from Neotropical snake assemblages: For areas that differ in the species richness of particular clades, do contrasting characteristics of those clades contribute to emergent community-level properties? Suggested a reinterpretation of patterns (e.g., no “goo-eaters”) from Brazilian caatinga… In contrast to Vitt and Vangilder (1983), “we do not need to ‘explain’ the absence of invertebrate-eating snakes [in the caatinga community] in terms of present-day ecological factors if historical events resulted in the absence of appropriate lineages from the community” Please do not use the images in these PowerPoint slides without permission. Cadle, John E. & Harry W. Green Phylogenetic patterns, biogeography, and the ecological structure of Neotropical snake assemblages. Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL. Vitt, L. J. & L. D. Vangilder Ecology of a snake community in northeastern Brazil. Amphibia-Reptilia 4:

33 Origins & Maintenance of Diversity Phylogenetic Perspectives
Losos (1996), after Brooks & McLennan (1991, 1993) “Four components contributing to community structure”: 1. Species that interact in the same way that their ancestors did in other communities 2. Species that colonized a community but did not lead to ecological shifts in residents (new associations of lineages, but ancestral ecologies were maintained) Please do not use the images in these PowerPoint slides without permission. All of these possibilities would require historical information for support or refutation. Brooks, Daniel R. & Deborah A. McLennan Phylogeny, Ecology, and Behavior. Chicago U. Press, Chicago, IL, USA. Brooks, Daniel R. & Deborah A. McLennan Pp in Robert E. Ricklefs & Dolph Schulter, eds., Species Diversity in Ecological Communities: Historical and Geographical Perspectives. The University of Chicago Press, Chicago, IL. Losos, Jonathan B Phylogenetic perspectives on community ecology. Ecology 77: 3. Species that arose in situ and evolved different ecological requirements 4. Species that colonized a community and interacted with species already present, leading to ecological shifts in the colonizer, the resident, or both

34 Origins & Maintenance of Diversity Phylogenetic Perspectives
Webb et al. (2002) “The differences among species that co-occur in an ecological community are the result of modifications to a common ancestor that the species all ultimately share” Phylogenetic relationships provide “a new dimension of information… with which to make sense of these differences among species” Differences among species are due to divergence (either in sympatry or allopatry), which may have occurred recently due to changes in the focal species, or in the distant past due to changes in the ancestors of the focal species Similarities among species are either due to shared ancestry, or convergence (either in sympatry or allopatry) Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O., David D. Ackerly, Mark A. McPeek & Micheal J. Donoghue Phylogenies and community ecology. Annual Review of Ecology and Systematics 33:

35 Origins & Maintenance of Diversity Phylogenetic Perspectives
Webb (2000) An example from rainforest trees: Is the distribution of species among habitats (or samples) in a community nonrandom with respect to phylogeny? “The demonstration of nonrandom spatial association of species with habitats is a necessary but not sufficient criterion in demonstrating that habitat partitioning is important in enabling many species to coexist…” “However, to be able to show that co-occurring species in different habitats are more ecologically similar than expected by chance would support the case that species are partitioning habitat according to their autecology” Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156: “On the other hand, if species in local communities were less ecologically similar than expected by chance, this would suggest that negative neighborhood interactions [e.g., competition] were causing increased mortality among ecologically similar species…”

36 Origins & Maintenance of Diversity Phylogenetic Perspectives
Webb (2000) Specific question: Are the tree species in 0.16-ha plots more or less related than expected if such communities were formed from a random sampling of available species in the larger area (150 ha)? “…because of the conservatism of many species traits in the evolution of a lineage, we expect, in general, a positive relationship between a measure of the phylogenetic relatedness of two species and a measure of their overall life-history and ecological similarity…” Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156:

37 Origins & Maintenance of Diversity Phylogenetic Perspectives
Webb (2000) What about character displacement? “Under certain circumstances, where new niches are encountered (e.g., on islands) or where competition with an ecologically similar species is strong and predictable, particular ecological traits of an organism may change adaptively over time… leading to a breakdown of the correlation between phylogenetic relatedness and ecological similarity for some traits… However, even in these organisms we would still expect that a measure of ‘overall’ ecological similarity, taking many traits into account, would be correlated with phylogenetic relatedness” Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156:

38 Origins & Maintenance of Diversity Phylogenetic Perspectives
Webb (2000) Data: 28 tree plots of 40 x 40 m (0.16 ha) scattered throughout 150 ha of lowland tropical forest, Indonesia How to efficiently estimate relatedness? Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156:

39 Origins & Maintenance of Diversity Phylogenetic Perspectives
Webb (2000) Data: 28 tree plots of 40 x 40 m (0.16 ha) scattered throughout 150 ha of lowland tropical forest, Indonesia How to efficiently estimate relatedness? Created “supertrees” from published phylogenies and then counted nodes separating terminal taxa Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156:

40 Mean pairwise nodal dist. = (1+2+4+2+4+3)/6 = 2.66
Greatest possible mean pairwise nodal distance for a community of 4 taxa (given this phylogeny) = 3.66 nodes (for A, B, E, F) Phylogeny A B C D E F Community 1: A, B, C, D Community 2: A, B, E, F Nodal distances: Nodal distances: A B C D A B E F A 1 2 4 A 1 5 5 B 2 4 B 5 5 C 3 E 1 Please do not use the images in these PowerPoint slides without permission. The smaller the mean pairwise nodal distance, the larger the NRI. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156: Mean pairwise nodal dist. = ( )/6 = 2.66 Mean pairwise nodal dist. = ( )/6 = 3.66 Net Relatedness Index = 1 - (2.66/3.66) = 0.273 Net Relatedness Index = 1 - (3.66/3.66) = 0.0 From Webb (2000)

41 Mean nearest nodal dist. = (1+1+2+3)/4 = 1.75
Greatest possible mean nearest nodal distance for a community of 4 taxa (given this phylogeny) = 2.00 nodes (for A, C, D, F) Phylogeny A B C D E F Community 1: A, B, C, D Community 2: A, B, E, F Nodal distances: Nodal distances: A B C D A B E F A 1 2 4 A 1 5 5 B 2 4 B 5 5 C 3 E 1 Please do not use the images in these PowerPoint slides without permission. The smaller the mean nearest nodal distance, the larger the NTI. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156: Mean nearest nodal dist. = ( )/4 = 1.75 Mean nearest nodal dist. = ( )/4 = 1.0 Nearest Taxa Index = 1 - (1.75/2.0) = 0.125 Nearest Taxa Index = 1 - (1.0/2.0) = 0.5 From Webb (2000)

42 Origins & Maintenance of Diversity Phylogenetic Perspectives
Methods: Webb (2000) created 1000 sets of plots with randomized species membership, following two simple rules: (1) each species occurred in the same total number of plots as observed; (2) each plot contained the same total number of species as observed Randomized plots represent expected distributions of species if membership in plots occurs at random with respect to the overall species pool of 324 observed species throughout the 150-ha area Compared the observed Net Relatedness Index (NRI) and the observed Nearest Taxa Index (NTI) to the appropriate distributions of expected values Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156: Results: The mean NRI did not differ significantly from the expectation of the null hypothesis, but the mean NTI was significantly greater than expected

43 Origins & Maintenance of Diversity Phylogenetic Perspectives
Conclusion: Webb (2000) concluded that species in the 0.16-ha plots were more likely to be found with species separated by fewer nodes than expected by chance (e.g., congeners) We can therefore reject the null hypothesis that species are assembled into local communities at random; there is evidence that species occur with closely related species more than we expect by chance Environmental filtering? Primarily abiotic? Biotic? Trait-based? Please do not use the images in these PowerPoint slides without permission. Webb, Campbell O Exploring phylogenetic structure of ecological communities: An example for rain forest trees. The American Naturalist 156:

44 Phylogenetic & Trait-based Community Structure
Lorelei E. Patrick Please do not use the images in these PowerPoint slides without permission. See Lori Patrick’s presentation from Fall 2009! “Figure 3. Environmental filters, interspecific competition, trait lability and the structure of communities. Species composition within and among communities can be influenced by environmental filtering, interspecific interactions and the potential for evolutionary change of traits associated with niche occupancy. (a) Both phylogenetic and phenotypic clustering in three communities (squares) consistent with the existence of both the conserved evolution of traits associated with niche occupancy (represented by circle size and color) and strong environmental filtering. Such a pattern is observed in the Gigasporaceae and Glomeraceae lineages of Arbuscular mycorrhizal fungi on Plantago lanceolata [81]. (b) Both phylogenetic and phenotypic overdispersion consistent with conserved evolution of traits associated with niche occupancy, and greater importance for species interactions over environmental filtering in determining species composition. Such a pattern is inferred for wood warblers in North America [9]. (c) Phylogenetic clustering and phenotypic overdispersion consistent with evolutionary change in traits associated with niche occupancy and adaptive radiation as observed among Anolis lizards on different Caribbean islands [53]. (d) Phylogenetic overdispersion and phenotypic clustering consistent with evolutionary change in traits associated with niche occupancy and strong environmental filtering, as observed in Floridian oak communities [8]. Modified with permission from Ref. [8].” Emerson, Brent C. & Rosemary G. Gillespie Phylogenetic analysis of community assembly and structure over space and time. Trends in Ecology & Evolution 23: From Emerson & Gillespie (2008)

45 Phylogenetic Niche Conservatism (PNC)
“The tendency of species to retain ancestral traits” (Wiens & Graham 2005) Please do not use the images in these PowerPoint slides without permission. From Lori Patrick’s Ph.D. defense – April 2014 – PatrickLori_2014_PhDDefenseLSU.pptx Emerson, Brent C. & Rosemary G. Gillespie Phylogenetic analysis of community assembly and structure over space and time. Trends in Ecology & Evolution 23:

46 Habitat Filtering versus Competition
Community Phenotypes Habitat filtering: Only phenotypes capable of surviving environmental conditions can persist in community Competition: Only one species of each phenotype can persist in community Please do not use the images in these PowerPoint slides without permission. From Lori Patrick’s Ph.D. defense – April 2014 – PatrickLori_2014_PhDDefenseLSU.pptx

47 Phylogenetic Community Structure (PCS) with PNC
Habitat filtering Competition Please do not use the images in these PowerPoint slides without permission. From Lori Patrick’s Ph.D. defense – April 2014 – PatrickLori_2014_PhDDefenseLSU.pptx PNC PNC Phylogenetic & phenotypic clustering Phylogenetic & phenotypic over-dispersion

48 PCS with Convergent Evolution
Habitat filtering Competition Please do not use the images in these PowerPoint slides without permission. From Lori Patrick’s Ph.D. defense – April 2014 – PatrickLori_2014_PhDDefenseLSU.pptx Emerson, Brent C. & Rosemary G. Gillespie Phylogenetic analysis of community assembly and structure over space and time. Trends in Ecology & Evolution 23: Conver- gence Conver- gence Phylogenetic clustering & phenotypic over-dispersion Phylogenetic over-dispersion & phenotypic clustering


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