Species Diversity in Communities Please do not use the images in these PowerPoint slides without permission. A landscape contains a variety of communities: e.g., alpine meadow, alpine tundra, forest, etc… Wikipedia “Tundra” page; accessed 3-XI-2014 [By John Holm from Leadville, CO, USA (tundra on Spaulding Ridge Uploaded by Hike395) [CC-BY-2.0 (http://creativecommons.org/licenses/by/2.0)], via Wikimedia Commons] Photo from Wikimedia Commons

Community Assembly Ecological Filters Whereas the figure is drawn as if these processes occur sequentially, in reality they occur more-or-less simultaneously or in any order Please do not use the images in these PowerPoint slides without permission. Note that the biotic filter could either be the presence or the absence of a species that could influence local-community membership. In other words, in general we would consider the presence of a dominant competitor to be a biotic filter if the competitor excludes (or reduces the chances of establishment by) subordinate competitors. In contrast, the absence of a keystone predator could be considered a biotic filter, since in the absence of the keystone predator some species might be excluded from gaining membership in the local community (or have reduced chances of establishment). Bowman, Hacker & Cain (2017), Fig. 19.4

Co-occurrence  Coexistence Species “able to persist indefinitely together are deemed to ‘coexist’…” “If some mechanism promotes the coexistence of two or more species, each species must be able to increase when it is rare and the others are at their typical abundances; this invasibility criterion is fundamental evidence for species coexistence regardless of the mechanism.” “…some subset of the co-occurring species are either slowly being driven extinct by others in the assemblage… stochastically [drifting] to extinction via neutral dynamics… or maintained in a local area by dispersal from other areas (i.e., sink populations…)…” Please do not use the images in these PowerPoint slides without permission. Siepielski, Adam M. & Mark A. McPeek. 2010. On the evidence for species coexistence: a critique of the coexistence program. Ecology 91:3153-3164. Quotes from Siepielski & McPeek (2010) Ecology

Coexistence – Resource Partitioning Species-packing mechanisms that illustrate “resource-partitioning” or “niche-differentiation” explanations for differences in diversity between two sites Please do not use the images in these PowerPoint slides without permission. Ricklefs (1987) used a “balls in a box” metaphor, whereas Roughgarden (2009) used a “books on a bookshelf” metaphor to illustrate these species-packing ideas. Remsen, Jr., J. V. 1990. Community ecology of Neotropical kingfishers. Univ. Calif. Publ. Zool. 124:1-116. Figure from Remsen (1991) Univ. Calif. Publ. Zool.

Coexistence – Resource Partitioning Species-packing mechanisms that illustrate “resource-partitioning” or “niche-differentiation” explanations for differences in diversity between two sites Please do not use the images in these PowerPoint slides without permission. Ricklefs (1987) used a “balls in a box” metaphor, whereas Roughgarden (2009) used a “books on a bookshelf” metaphor to illustrate these species-packing ideas. Remsen, Jr., J. V. 1990. Community ecology of Neotropical kingfishers. Univ. Calif. Publ. Zool. 124:1-116. Figure from Remsen (1991) Univ. Calif. Publ. Zool.

Coexistence – Resource Partitioning Species-packing mechanisms that illustrate “resource-partitioning” or “niche-differentiation” explanations for differences in diversity between two sites Please do not use the images in these PowerPoint slides without permission. Ricklefs (1987) used a “balls in a box” metaphor, whereas Roughgarden (2009) used a “books on a bookshelf” metaphor to illustrate these species-packing ideas. Remsen, Jr., J. V. 1990. Community ecology of Neotropical kingfishers. Univ. Calif. Publ. Zool. 124:1-116. Figure from Remsen (1991) Univ. Calif. Publ. Zool.

Coexistence – Spatial Variability Spatial variability can foster coexistence Please do not use the images in these PowerPoint slides without permission. Robertson et al. (1988)… Bowman, Hacker & Cain (2017), Fig. 19.11; after Robertson et al. (1988)

Coexistence – Spatial Variability Spatial variability can foster coexistence Please do not use the images in these PowerPoint slides without permission. R. H. MacArthur (1958)… Bowman, Hacker & Cain (2017), Fig. 19.9; after MacArthur (1958)

Coexistence – Spatial Variability Spatial variability can foster coexistence Please do not use the images in these PowerPoint slides without permission. Bowman, Hacker & Cain (2017), Fig. 19.10; after MacArthur & MacArthur (1961)

Coexistence – Temporal Variability Temporal variability can foster coexistence G. E. Hutchinson’s (1961) “Paradox of the Plankton” i.e., that many more species of plankton apparently coexist in lakes than there are limiting nutrients; concluded that plankton rarely achieve equilibrium owing to ever-changing environmental conditions Please do not use the images in these PowerPoint slides without permission. Hutchinson, G. E. 1961. The paradox of the plankton. The American Naturalist 95:137-145. Note that Hutchinson’s explanation for the co-occurrence of so many species of plankton is that environmental conditions continually change and do not allow any species to be best favored by the environment and to outcompete the others. Bowman, Hacker & Cain (2017), Fig. 19.13

Coexistence – Temporal Variability Temporal variability can foster coexistence Storage Effect “…models in which stable coexistence results from environmental fluctuations are models of temporal niches: species are not distinguished by the resources they use but by when they are most actively using them…” (Chesson 2000) Please do not use the images in these PowerPoint slides without permission. Chesson, Peter. 2000. Mechanisms of maintenance of species diversity. Annual Review of Ecology and Systematics 31:343-366. Chesson, P. L. & R. R. Warner. 1981. Environmental variability promotes coexistence in lottery competitive systems. The American Naturalist 117:923-943. Quote from Chesson (2000) Ann. Rev. Ecol. Syst.

Coexistence – Intermediate Disturbance Hypothesis Disturbance can foster coexistence Please do not use the images in these PowerPoint slides without permission. Connell, J. H. 1978. Diversity in tropical rain forests and coral reefs. Science 199:1302-1310. Bowman, Hacker & Cain (2017), Fig. 19.14, after Connell (1978) Science

Coexistence – Intermediate Disturbance Hypothesis Disturbance can foster coexistence Please do not use the images in these PowerPoint slides without permission. Sousa, Wayne P. 1979. Disturbance in marine intertidal boulder fields: the nonequilibrium maintenance of species diversity. Ecology 60:1225-1239. Bowman, Hacker & Cain (2017), Fig. 19.15, after Sousa (1979) Ecology

Coexistence – Dynamic Equilibrium Model Disturbance can foster coexistence Please do not use the images in these PowerPoint slides without permission. Huston, Michael. 1979. A general hypothesis of species diversity. The American Naturalist 113:81-101. Bowman, Hacker & Cain (2017), Fig. 19.12, after Huston (1979) The American Naturalist

Coexistence – Keystone Predation Enemies can foster coexistence “The removal of Pisaster has resulted in a pronounced decrease in diversity… from a 15 to an eight-species system” Please do not use the images in these PowerPoint slides without permission. Paine, Robert T. 1966. Food web complexity and species diversity. The American Naturalist 100:65-75. Quote from Paine (1966) The American Naturalist; photo of Pisaster consuming a Mytilus from Wikimedia Commons; photo of Paine from http://naturalhistoriesproject.org/conversations/anemone-like

Coexistence – Janzen-Connell Model Enemies can foster coexistence Please do not use the images in these PowerPoint slides without permission. Connell, J. H. 1971. On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. Pp. 298-313 in P. J. den Boer & G. R. Gradwell, eds. Dynamics of Populations. Centre for Agricultural Publishing and Documentation, Wageningen, The Netherlands. Janzen, D. H. 1970. Herbivores and the number of tree species in tropical forests. The American Naturalist 104:501-528. Adapted from Janzen (1970) The American Naturalist

Coexistence – Facilitators Positive interactions can foster coexistence Please do not use the images in these PowerPoint slides without permission. Hacker, S. D. & S. D. Gaines. 1997. Some implications of direct positive interactions for community species diversity. Ecology 78:1990-2003. Bowman, Hacker & Cain (2017), Fig. 19.17, after Hacker & Gaines (1997) Ecology

Coexistence – Facilitators Positive interactions can foster coexistence Please do not use the images in these PowerPoint slides without permission. Hacker, S. D. & S. D. Gaines. 1997. Some implications of direct positive interactions for community species diversity. Ecology 78:1990-2003. Bowman, Hacker & Cain (2017), Fig. 19.18, after Hacker & Gaines (1997) Ecology

Coexistence – Facilitators Positive interactions can foster coexistence Please do not use the images in these PowerPoint slides without permission. Wikipedia “Bluestreak cleaner wrasse” page; accessed 10-III-2015 ["Cleaner wrasse with a client" by Alexander Vasenin - Own work. Licensed under CC BY-SA 3.0 via Wikimedia Commons - http://commons.wikimedia.org/wiki/File:Cleaner_wrasse_with_a_client.JPG#mediaviewer/File:Cleaner_wrasse_with_a_client.JPG] Grutter, Alexandra S., Jan Maree Murphy & J. Howard Choat. 2003. Cleaner fish drives local fish diversity on coral reefs. Current Biology 13:64-67. “Figure 1. Mean (s.e.) Visiting Client Fish Numbers per Reef at Two Sites Counted by Remote Video and by a Snorkeler (A) The number of visiting species on reefs with (closed symbols) and without (open symbols) cleaner fish at Casuarina Beach (circles) and Lagoon (squares) sites. (B) The log10 (x+1) abundance of visiting client fish per reef. Symbols are as in (A). Counts in (A) and (B) were pooled across different times of the day.” “reefs with (closed symbols) and without (open symbols) cleaner fish at Casuarina Beach (circles) and Lagoon (squares) sites” Figure from Grutter et al. (2003) Current Biology; photo of cleaner wrasse & client from Wikimedia Commons

Coexistence – Relative Importance of Factors A combination of predators and disturbance/stress can foster coexistence Please do not use the images in these PowerPoint slides without permission. Menge, Bruce A. & John P. Sutherland. 1976. Species diversity gradients: synthesis of the roles of predation, competition, and temporal heterogeneity. The American Naturalist 110:351-369. Menge, Bruce A. & John P. Sutherland. 1987. Community regulation: variation in disturbance, competition, and predation in relation to environmental stress and recruitment. The American Naturalist 130:730-757. Bowman, Hacker & Cain (2017), Fig. 19.19, after Menge & Sutherland (1987) The American Naturalist

Long-term Co-occurrence: Lottery and Neutral Models Experimental removal and numbers of replacements by 3 species of fishes Please do not use the images in these PowerPoint slides without permission. Sale, P. F. 1979. Recruitment, loss and coexistence in a guild of territorial coral reef fishes. Oecologia 42:159-177. Bowman, Hacker & Cain (2017), Fig. 19.20, after Sale (1979) Oecologia

Long-term Co-occurrence: Lottery and Neutral Models Steve Hubbell What if all individuals have the same per capita influence? Please do not use the images in these PowerPoint slides without permission. Neutral Theory and Ecological Drift Photo of S. P. Hubbell from UCLA

Please do not use the images in these PowerPoint slides without permission. Vellend, Mark. 2010. Conceptual synthesis in community ecology. Quarterly Review of Biology 85:183-206. Figure 2. Frequency Dynamics of Two Species under Drift and Selection. Dynamics are shown for two species, A and B, with non-overlapping generations in 10 simulated communities of constant size, J, with (A) no fitness differential and J 500, (B) no fitness differential and J 50, (C) a 5% fitness advantage to species A and J 500, and (D) a 5% fitness advantage to species A and J 50. Figure from Vellend (2010)

Diversity-Ecosystem Function (e.g., Productivity) Relationships Please do not use the images in these PowerPoint slides without permission. Bowman, Hacker & Cain (2017), Fig. 19.21A

Diversity-Ecosystem Function (e.g., Productivity) Relationships Drought resistance (measured as biomass change) in plots that varied in pre-drought species richness Separate experiment in which plots were constructed with varying numbers of species and cover after 2 yr was measured Please do not use the images in these PowerPoint slides without permission. Tilman, David, D. Wedin & J. Knops. 1996. Productivity and sustainability influences by biodiversity in grassland ecosystems. Nature 379:718-720.  Tilman, David & J. A. Downing. 1994. Biodiversity and stability in grasslands. Nature 367:363-365. Bowman, Hacker & Cain (2017), Fig. 19.21B after Tilman & Downing (1994), Fig. 19.21C after Tilman et al. (1996)

Diversity-Ecosystem Function Relationships Please do not use the images in these PowerPoint slides without permission. Bowman, Hacker & Cain (2017), Fig. 19.22, after Naeem et al. (1995) Phil. Trans. Roy. Soc.