1. Species Diversity in Communities
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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 ( via Wikimedia Commons]
Photo from Wikimedia Commons

2. Community Assembly – Ecological Filters
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Cain, Bowman & Hacker (2014), Fig. 19.4

3. 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…)…”
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Siepielski, Adam M. & Mark A. McPeek On the evidence for species coexistence: a critique of the coexistence program. Ecology 91:
Quotes from Siepielski & McPeek (2010) Ecology

4. Coexistence – Resource Partitioning Species-packing mechanisms
that illustrate
“resource-partitioning” or “niche-differentiation” explanations
for differences in diversity between two sites
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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 Community ecology of Neotropical kingfishers. Univ. Calif. Publ. Zool. 124:1-116.
Figure from Remsen (1991) Univ. Calif. Publ. Zool.

5. 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 Community ecology of Neotropical kingfishers. Univ. Calif. Publ. Zool. 124:1-116.
Figure from Remsen (1991) Univ. Calif. Publ. Zool.

6. 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 Community ecology of Neotropical kingfishers. Univ. Calif. Publ. Zool. 124:1-116.
Figure from Remsen (1991) Univ. Calif. Publ. Zool.

7. 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
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Wikipedia “Plankton” page; accessed 10-III-2015
[“Plankton collage" by Kils at the German language Wikipedia. Licensed under CC BY-SA 3.0 via Wikimedia Commons -
Hutchinson, G. E The paradox of the plankton. The American Naturalist 95:
Photomontage of plankton from Wikimedia Commons

8. 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)
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Chesson, Peter Mechanisms of maintenance of species diversity. Annual Review of Ecology and Systematics 31:
Chesson, P. L. & R. R. Warner Environmental variability promotes coexistence in lottery competitive systems. The American Naturalist 117:
Quote from Chesson (2000) Ann. Rev. Ecol. Syst.

9. Coexistence – Intermediate Disturbance Hypothesis
Disturbance can foster coexistence
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Connell, J. H Diversity in tropical rain forests and coral reefs. Science 199:
Cain, Bowman & Hacker (2014), Fig , after Connell (1978) Science

10. Coexistence – Intermediate Disturbance Hypothesis
Disturbance can foster coexistence
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Sousa, Wayne P Disturbance in marine intertidal boulder fields: the nonequilibrium maintenance of species diversity. Ecology 60:
Cain, Bowman & Hacker (2014), Fig , after Sousa (1979) Ecology

11. Coexistence – Dynamic Equilibrium Model
Disturbance can foster coexistence
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Huston, Michael A general hypothesis of species diversity. The American Naturalist 113:
Cain, Bowman & Hacker (2014), Fig , after Huston (1979) The American Naturalist

12. 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”
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Paine, Robert T 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

13. Coexistence – Janzen-Connell Model Enemies can foster coexistence
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Connell, J. H On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees. Pp in P. J. den Boer & G. R. Gradwell, eds. Dynamics of Populations. Centre for Agricultural Publishing and Documentation, Wageningen, The Netherlands.
Janzen, D. H Herbivores and the number of tree species in tropical forests. The American Naturalist 104:
Adapted from Janzen (1970) The American Naturalist

14. Coexistence – Facilitators
Positive interactions can foster coexistence
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Hacker, S. D. & S. D. Gaines Some implications of direct positive interactions for community species diversity. Ecology 78:
Cain, Bowman & Hacker (2014), Fig , after Hacker & Gaines (1997) Ecology

15. Coexistence – Facilitators
Positive interactions can foster coexistence
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Hacker, S. D. & S. D. Gaines Some implications of direct positive interactions for community species diversity. Ecology 78:
Cain, Bowman & Hacker (2014), Fig , after Hacker & Gaines (1997) Ecology

16. Coexistence – Facilitators
Positive interactions can foster coexistence
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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 -
Grutter, Alexandra S., Jan Maree Murphy & J. Howard Choat 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

17. Coexistence – Facilitators
A combination of predators and disturbance/stress
can foster coexistence
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Menge, Bruce A. & John P. Sutherland Species diversity gradients: synthesis of the roles of predation, competition, and temporal heterogeneity. The American Naturalist 110:
Menge, Bruce A. & John P. Sutherland Community regulation: variation in disturbance, competition, and predation in relation to environmental stress and recruitment. The American Naturalist 130:
Cain, Bowman & Hacker (2014), Fig , after Menge & Sutherland (1987) The American Naturalist

18. Long-term co-occurrence: Lottery and Neutral Models
Experimental removal and numbers of replacements by 3 species of fishes
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Sale, P. F Recruitment, loss and coexistence in a guild of territorial coral reef fishes. Oecologia 42:
Cain, Bowman & Hacker (2014), Fig , after Sale (1979) Oecologia

19. 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
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Tilman, David, D. Wedin & J. Knops Productivity and sustainability influences by biodiversity in grassland ecosystems. Nature 379:  
Tilman, David & J. A. Downing Biodiversity and stability in grasslands. Nature 367:
Cain, Bowman & Hacker (2014), Fig A after Tilman & Downing (1994), Fig B after Tilman et al. (1996)

20. Diversity-Ecosystem Function
Relationships
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Peterson, G., C. R. Allen & C. S. Holling Ecological resilience, biodiversity, and scale. Ecosystems 1:6-8.
Cain, Bowman & Hacker (2014), Fig , after Peterson et al. (1998) Ecosystems