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Gary G. Mittelbach PDF Print E-mail



cover of Community Ecology textbook

For information on this book see bottom of webpage

Professor of Zoology/Intergrative Biology                                       

Ph.D. Michigan State University, 1980

W. K. Kellogg Biological Station                    
Michigan State University
Hickory Corners, MI 49060
Phone: (269) 671-2216
Fax: (269) 671-2104
Email: Mittelbach@kbs.msu.edu

Research Interests

As an ecologist, I am interested in the evolution and maintenance of biodiversity (the variety of life), and in particular, what determines species diversity at different spatial scales. At the local scale, the number and type of species found in a community depends on biotic and abiotic interactions, species sorting, and dispersal of colonists from a regional species pool. At broad spatial scales (regions, continents), we need to consider the factors that drive rates of diversification (speciation and extinction), as well as the dispersal of species between regions. Studying biodiversity at these different spatial scales requires different tools and different approaches. Local communities often lend themselves to experimental manipulation and my current research in this area includes experimental studies in freshwater communities (working with fish and aquatic invertebrates), and a long-term, collaborative study on the effects of resource heterogeneity on species diversity in a terrestrial grassland. To study the causes of biodiversity patterns at broad spatial scales, I work collaboratively with a group of ecologists, evolutionary biologists, and paleontologists, focusing particularly on the evolution of the latitudinal gradient in biodiversity. Below are brief sketches of these research areas and references to recently published papers.


Biodiversity in aquatic communities (Most of this research is done in collaboration with my graduate students, who work with a variety of organisms in different aquatic habitats).

Mittelbach, G.G., E. A. Garcia, and Y. Taniguchi. 2006. Fish reintroductions reveal smooth transitions between lake community states. Ecology 87:312-318. Whether communities respond smoothly or discontinuously to changing environmental conditions has important consequences for the preservation and restoration of ecosystems. In this seventeen year study of a Michigan lake, we show that manipulating the fish community from high plantivore density to low planktivore density back to high planktivore density results in dramatic, but smooth transitions between ecosystem states.

Wojdak, J.M. and G. G. Mittelbach (2007). Consequences of niche overlap for ecosystem functioning: an experimental test of pond grazers. Ecology 88:2072-2083. We outline an experimental approach for testing how niche complimentarity contributes to positive biodiversity effects on ecosystem functioning and apply this approach in an experimental study with pond grazers (gastropods).

Garcia, E.A. and G. G. Mittelbach (2008). Regional coexistence and local dominance in Chaoborus: species sorting along a predation gradient. Ecology 89:1703-1713. By experimentally manipulating fish density in small ponds, we were able to show that variation in predation pressure, in combination with species traits, determines the distribution and local dominance of Chaoborus (phantom midge) species across a landscape of lakes and ponds.

Mittelbach, G.G., N.G. Ballew, and M.K.Kjelvik. (2014). Fish behavioral types and their ecological consequences. Can. J. Fish. Aquat. Sci. 71:927-944. In this review and perspective, we discuss the factors that lead to the formation and maintenance of behavioral ("personality") types in fish populations, as well as the importance of these behavioral types to fisheries management, conservation, and basic ecology.


Consequences of resource heterogeneity, productivity, and clonality to species diversity in a low-productivity grassland (in collaboration by Drs. Katherine Gross and Heather Reynolds, plus students and postdocs).

Coexistence theory predicts that greater heterogeneity of resources or other fitness-constraining environmental factors will promote species diversity, yet this classic mechanism of coexistence has rarely been tested in manipulative field experiments. We have conducted two large-scale field experiments manipulating soil resource heterogeneity, in combination with species introductions (by seed), and the presence/absence of clonal species, to test this hypothesis in a low productivity grassland in Michigan. Our results show that the large foraging area of clonal plants limits the positive impact of resource heterogeneity on species coexistence, and that the growth response of tall, clonal species to fertilzation has a particularly strong negative effect on plant species richness.

Gross, K.L., G.G. Mittelbach, and H.L. Reynolds. 2005. Grassland invasibility and diversity: responses to nutrients, seed input, and disturbance. Ecology 86:476-486.

Reynolds, H.L., G.G. Mittelbach, T.L. Darcy-Hall, G.R. Houseman, and K.L. Gross. 2007. No effect of varying soil resource heterogeneity on plant species richness in a low fertility grassland. Journal of Ecology 95:723-733.

Golubski, A.J., K.L. Gross, and G.G. Mittelbach. 2008. Competition among plant species that interact with their environment at different spatial scales. Proceedings of the Royal Society B 275:1897-1906.

Golubski, A.J., K.L. Gross, and G.G. Mittelbach. 2010. Recycling-mediated facilitation and coexistence based on plant size. American Naturalist 176:588-600.

Eilts, J.A., G.G. Mittelbach, H.L. Reynolds, and K.L. Gross. 2011. Resource heterogeneity, soil fertility, and species diversity: effects of clonal species on plant communities. American Naturalist 177:574-588.

Dickson, T.L., G.G. Mittelbach, H.L. Reynolds, and K.L. Gross. 2014. Height and clonality traits determine plant community responses to fertilization. Ecology 95:2443-2452.


Patterns in Species Diversity The dramatic increase in biodiversity as one moves from the poles to the tropics, generally known as the Latitudinal Diversity Gradient, “..is the major, unexplained pattern in natural history…, one that mocks our ignorance” (Robert Ricklefs). My interest in the latitudinal diversity gradient stems from a general interest in understanding how climate and evolutionary history affect broad scale patterns in biodiversity, and most of my work in this area is the result of collaborations developed from three working groups supported by the National Center for Ecological Analysis and Synthesis in Santa Barbara, CA. Listed below are some representative publications.

Hawkins, B.A., R. Field, H.V. Cornell, et al. 2003. Energy, water, and broad-scale geographic patterns of species richness. Ecology 84:3105-3117

Currie, D.J., G.G. Mittelbach, H.V. Cornell, et al. 2004. Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness. Ecology Letters 7:1121-1134.

Mittelbach, G.G., D.W. Schemske, H.V. Cornell, et al. 2007. Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography. Ecology Letters 10:315-331.

Schemske, D.W., G.G. Mittelbach, H.V. Cornell, J.M. Sobel, and K. Roy. 2009. Latitudinal variation in the strength of biotic interactions: evidence and consequences for the biodiversity gradient. Annual Review of Ecology, Evolution, and Systematics 40:245-269.

Mittelbach, G.G. and D.W. Schemske. 2015. Ecological and evolutionary perspectives on commmunity assembly. Trends in Ecology and Evolution (in press).


cover of Community Ecology textbook



Published in 2012 and available from the publisher, Sinauer Associates, as well as from Amazon, Barnes and Noble, and elsewhere.  For more information, including a full table of contents, a downloadable sample chapter on Biodiversity and Ecosystem Functioning, and instructions for ordering or requesting an examination copy, please see the Sinauer web site at:


About the book: Community ecology has undergone a transformation in recent years, from a discipline largely focused on processes occurring within a local area, to a discipline encompassing a much richer domain of study, including the linkages between communities separated in space (metacommunity dynamics), niche and neutral theory, the interplay between ecology and evolution (eco-evolutionary dynamics), and the influence of historical and regional processes in shaping patterns of biodiversity. To fully understand these new developments, however, students need a strong foundation in the study of species interactions and how these interactions are assembled into food webs and other ecological networks.

This book, written for graduate students, advanced undergraduates, and practicing ecologists, presents a broad, up-to-date coverage of ecological concepts at an advanced level, including both the ‘new’ and ‘traditional’ aspects of community ecology. It is divided into five sections: 1) The Big Picture: Patterns, Causes, and Consequences of Biodiversity, 2) The Nitty-Gritty: Species Interactions in Simple Modules, 3) Putting the Pieces Together: Food Webs and Ecological Networks, 4) Spatial Ecology: Metapopulations and Metacommunities, and 5) Species in Changing Environments: Ecology and Evolution. Applied aspects of community ecology (e.g., resource harvesting, invasive species, community restoration) are treated throughout the book as natural extensions of basic theoretical and empirical work. Theoretical concepts are developed using simple equations and there is an emphasis on the graphical presentation of ideas. Each chapter includes a summary.



1 Community Ecology's Roots 1

PART I The Big Picture: Patterns, Causes and Consequences of Biodiversity

2 Patterns of Biological Diversity 13

3 Biodiversity and Ecosystem Functioning 41

PART II The Nitty-Gritty: Species Interactions in Simple Modules

4 Population Growth and Density-Dependence 65

5 The Fundamentals of Predator-Prey Interactions 83

6 Selective Predators and Responsive Prey 103

7 Interspecific Competition: Simple Theory 125

8 Competition in Nature: Empirical Patterns and Tests of Theory 149

9 Beneficial Interactions in Communities: Mutualism and Facilitation 175

PART III Putting the Pieces Together: Food Webs and Ecological Networks

10 Species Interactions in Ecological Networks 197

11 Food Chains and Food Webs: Controlling Factors and Cascading Effects 223

PART IV Spatial Ecology: Metapopulations and Metacommunities

12 Patchy Environments, Metapopulations, and Fugitive Species 251

13 Metacommunities and the Neutral Theory 267

PART V Species in Changing Environments: Ecology and Evolution

14 Species Coexistence in Variable Environments 289

15 Evolutionary Community Ecology 317

16 Some Concluding Remarks and a Look Ahead 339



I will provide a list of all errata as they are found.  Please e-mail me at mittelbach@kbs.msu.edu if you find a mistake.

Chapter 2: typos

page 16, measures the difference in species composition

page 17, fourth line from top: "probably" should be "probability"

page 29, stronger and have a greater effect on species dynamics

page 31, fourth line from bottom: "descriptor of" should be "descriptor"

page 31, very productive environments

Chapter 3: typos

page 54 heading: "Invasiblity" should be "Invasibility"

page 62, It is still unknown what fraction of species could be lost

Chapter 4: typo

page 84, insight in the processes that govern the outcome of species interactions

Chapter 5:  multiple places where equations describing the rate of change of the predator population are incorrectly written as dN/dt instead of dP/dt.

page 88, equations 5.6 and 5.7 should read dP/dt not dN/dt

page 89 (Isocline analyis), second line of equations should read dP/dt not dN/dt

page 92, equation 5.10 should read dP/dt not dN/dt

page 95, the two growth rates equations for the predator population should read dP/dt not dN/dt

page 84, "the govern" should be "that govern"

Chapter 6:

page 105, citation to (Mittelbach 2001) in Figure 6.1 legend is incorrect.  Should be (Mittelbach 2002), which is missing from the Literature Cited.  Missing citation: Mittelbach, G.G. 2002. Fish foraging and habitat choice: a theoretical perspective. Pages 251-266 in P.J.B. Hart and J.D. Reynolds, editors. Handbook of Fish Biology and Fisheries. Volume 1, Fish Biology. Blackwell, Oxford, UK.

page 116, grazing algae from the stream bed and growing through a series of instars

page 117, sixth line from the top: "of the how the" should be "of how the"

page 117, Table 6.2. remove "(no predation)" from beneath "Natural mayfly population"

page 122, line 14: "need" should be "needed"

Chapter 7:

page 132, forefront, in part because they provide.....

page 135, Figure 7.5A.  Point labeled R* should be R* with subscript 2

page 143, In the second paragraph, under equation 7.7, the sentence should read "You should recognize Equation 7.7..."  The next paragraph should read "Like the L-V model, Equation 7.7..."  And then, "Equation 7.8 differes from the standard..."

page 144, Almost at the end of the page: should refer to Figure 10.13C

page 144, Near the top of the page, reference to Equation 7.11 should be Equation 7.10

Chaper 8:

page 161, coastal British Columbia

page 169, The idea that interspecific competition may affect the...

page 169, The goal of disentangling the factors that determine species

page 188 but not in small anemones

page 193 In the next chapter, we will examine the properties

Chapter 10:

Page 204, Table 10.1.  The formula for the "dynamic index" is incorrect as printed.  The correct version should be:  (ln(N/D))/Yt.

Page 212, Figure 10.13. For "indirect mutualism", the plus and minus signs for the interactions between P1 and B1, and betwen P2 and B2 should be reversed. For "habitat facilitation", the arrow pointing from P2 to B should be reversed.

Chapter 11:

page 234, may allow for the coexistence of N1 and N2

page 247, last paragraph before Summary: "In the next chapter..." should read "In Chapter 14..."

Chapter 12:

page 252, Equation 12.2 dP/dt=

page 253, Need ")" on third to last line: "....where D=1-m/c), the species..."

page 261, eight to last line: should be just Figure 12.7, not 12.7A, because there is no B

pag 264,  figure caption for Figure 12.9: Cordia nodosa is not an acacia.  Change to "...living symibotically on a single plant species (Cordia nodosa)

Chapter 13:

page 273,  period missing at the end of the last sentence of the first paragraph

page 277, Figure 13.8 caption, "Mapped species abundance distributions (SADs)..." should read, "Mapped distributions..."

page 278, last sentence on page, "...by the immigration to a ..." should read, "...by the immigration of species to a"

Chapter 14:

page 293, Figure 14.1(4), the green square containing species 1 should be changed to species 2

page 308, Fig. 14.8 legend bottom, if environmental conditions return to a point below c1

I thank Saara DeWalt and Bob Holt for bringing many of these errors to my attention.

Last Updated on Tuesday, 10 March 2015 18:21