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        Volume 88, Number 1, January 2007

Cover Photo: The consequences of climate change for ecosystem structure and function remain largely unknown. A 20-year study of a high-elevation riparian system in Arizona shows substantial changes in both structure and function related to climate change. Seven species of birds were found to initiate earlier breeding associated with an increase in spring temperature across years. Climate had much larger consequences for these species by affecting trophic levels below (plants) and above (predators) them. The Orange-crowned Warbler (Vermivora celata ) depends on maple for nesting and feeding. It has shown a major decline in abundance over the 20 years, associated with the decline in maple, but also associated with decreasing summer precipitation and increasing nest predation rates.

See the Photo Gallery for other photographs associated with this article by Thomas E. Martin.

Visit the Photo Gallery for more photographs submitted by our scientific journal authors.

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Table of Contents
(click on a title to view that section)

Governing Board

Request for an Editor: New Web Sites

Society Notices
Results of Elections for ESA Offices
Call for Nominations: ESA Honorary Member Award
Student Awards for Excellence in Ecology
Call for Student Award Judges

Other Notices
Vegetation Classification Draft Standard

Resolution of Respect: Frank B. Golley

Letter to the Editor of Ecology and Ecological Monographs
Reply from the Editor-in-Chief

Society Actions
Minutes of the 5–6 August Governing Board Meeting
Minutes of the 11 August Governing Board Meeting

PHOTO GALLERY: Images from upcoming articles in our scientific journals

Littorine Snails Compete by Altering a Shared Resource. J. Long, R. S. Hamilton, and J. L. Mitchell

Trophic Changes in a High-elevation Riparian System. T. E. Martin

Thermal Stress in the Rocky Intertidal Zone. B. Helmuth, B. R. Broitman, C. A. Blanchette, S. Gilman, P. Halpin, C. D. G. Harley, M. J. O’Donnell, G. E. Hoffman, B. Menge, and D. Strickland

Mangrove Forest Structure. W. P. Sousa, P. G. Kennedy, B. J. Mitchell, and B. M Ordóñez L.


Understanding Food Chains and Food Webs. F. N. Egerton

Advocacy and Integrity. S. Sorooshian

A History of the Ecological Sciences, Part 23. Linnaeus and the Economy of Nature. F. N. Egerton

Click here for a full complement of this series by Frank N. Egerton


Public Affairs Perspective
Rapid Response Team Fall Newsletter
ESA Statement on Global Climate Change

Ecology 101
Preparing for the Field Season. C. Wong

The Paper Trail
Essay on William S. Cooper’s “The Fundamentals of Vegetational Change.” S. T. A. Pickett

Reports of Symposia at the ESA Annual Meeting
Upstart View of Restoration Icons. J. B. Zedler, D. A. Falk, and D. J. Larkin
Urban Food Webs: Errata: Revised text and figure. P. Warren et al.

Society Section and Chapter News
Canada Chapter Newsletter
Southeastern Chapter Newsletter

Meeting Calendar
Urban Wildlife Management National Conference

Instructions for Contributors

is published quarterly by the
Ecological Society of America, 1707 H Street, N.W., Suite 400, Washington, DC 20006.
It is available online only, free of charge, at
Issues published prior to January 2004 are available through

Bulletin Editor-in-Chief E. A. Johnson

Bulletin of the Ecological Society of America, 1707 H Street, NW, Washington DC 20006
Phone (403) 220-7635, Fax (403) 289-9311,
E-mail: [email protected]

Associate Editor
David A. Gooding

ESA Publications Office,
127 W. State Street, Suite 301,
Ithaca, NY 14850-5427
E-mail: [email protected]

Production Editor
Regina Przygocki
ESA Publications Office,
127 W. State Street, Suite 301,
Ithaca, NY 14850-5427
E-mail: [email protected]

Section Editor, Ecology 101
H. Ornes
College of Sciences, SB310A, Southern Utah University
Cedar City, UT 84720 E-mail: [email protected]

Section Editor, Public Affairs Perspective
N. Lymn
Director for Public Affairs, ESA Headquarters,
1707 H Street, NW, Suite 400,
Washington, DC 20036 E-mail: [email protected]

Section Editors,
Emerging Technologies
D. W. Inouye
Department of Biology,
University of Maryland
College Park, MD 20742
E-mail: [email protected]
and S. Scheiner
Div. of Environmental Biology
Natl. Science Foundation
4201 Wilson Blvd.
Arlington, VA 22230
E-mail: [email protected]

Section Editors,
Ecological Education: K–12

S. Barker

Dept. of Secondary Education
350 Education South,
University of Alberta
Edmonton, Alberta
T6G 2G5 Canada
E-mail: [email protected]
and C. W. Anderson
319A Erickson Hall, Michigan State University
East Lansing, MI 48824 USA.
E-mail: [email protected]

The Ecological Society of America

Alan Covich, Institute of Ecology, University of Georgia, Athens, GA 30602
Norm Christensen, Nicholas School of the Environment and Earth Sciences, Duke University, Durham, NC 27708
Nancy B. Grimm, School of Life Sciences, Arizona State University, Tempe, AZ 85287-4501
Vice President for Science:
Gus R. Shaver, The Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA 02543
Vice President for Finance:
William J. Parton, Natural Resource Ecology Laboratory, Colorado State University, Ft. Collins, CO 80523-1499
Vice President for Public Affairs:
Richard V. Pouyat, 3315 Hudson St., Baltimore, MD 21224
Vice President for Education and Human Resources:
Margaret D. Lowman, Biology and Environmental Studies, New College of Florida, Sarasota, FL 34243-2109
David W. Inouye, Department of Biology, University of Maryland, College Park, MD 20742-4415
Dennis Ojima, Natural Resource Ecology Laboratory, Colorado State University, Ft. Collins, CO 80523-1499
Jayne Belnap, USGS Cayonlands Field Station, Southwest Biological Science Center, Moab, UT 84532
Juan J. Armesto, Departmento de Biologia, Facultad de Ciencias, Universidad de Chile, Santiago, Chile


The Ecological Society of America was founded in 1915 for the purpose of unifying the sciences of ecology, stimulating research in all aspects of the discipline, encouraging communication among ecologists, and promoting the responsible application of ecological data and principles to the solution of environmental problems. Ecology is the scientific discipline that is concerned with the relationships between organisms and their past, present, and future environments. These relationships include physiological responses of individuals, structure and dynamics of populations, interactions among species, organization of biological communities, and processing of energy and matter in ecosystems.

Membership is open to persons who are interested in the advancement of ecology or its applications, and to those who are engaged in any aspect of the study of organisms in relation to environment. The classes of membership and their annual dues for 2007 are as follows:
Regular member: Income level Dues
  <$40,000 $50.00
  $40,000—60,000 $75.00
  >$60,000 $95.00
Student member:
Emeritus member:   Free
Life member:
Contact Member and Subscriber Services (see below)  

Subscriptions to the journals are not included in the dues.
Special membership rates are available for individuals in developing countries. Contact Member and Subscriber services (address below) for details.

The Society publishes a bulletin, three print journals, and an electronic data archive. The Bulletin of the Ecological Society of America, issued quarterly, contains announcements of meetings of the Society and related organizations, programs, awards, articles, and items of current interest to members. The journal Ecology, issued monthly, publishes essays and articles that report and interpret the results of original scientific research in basic and applied ecology. Ecological Monographs is a quarterly journal for longer ecological research articles. Ecological Applications, published six times per year, contains ecological research and discussion papers that have specific relevance to environmental management and policy. Frontiers in Ecology and the Environment, with 10 issues each year, focuses on current ecological issues and environmental challenges: it is international in scope and interdisciplinary in approach. Ecological Archives is published on the Internet at ‹http://esapubs.org/Archive› and contains supplemental material to ESA journal articles and data papers.
No responsibility for the views expressed by the authors in ESA publications is assumed by the editors or the publisher, the Ecological Society of America.
Subscriptions for 2007 are available to ESA members as follows:
Regular Student
Ecology $65.00 $50.00
ulletin of the Ecological Society of America Free to members
cological Monographs $30.00 $25.00

Ecological Applications $50.00 $40.00
Frontiers in Ecology Free to members
Ecological Archives

Application blanks for membership may be obtained from the Ecological Society of America, Member and Subscriber Services, 1707 H Street, NW, Suite 400, Washington, DC 20006, to which all correspondence concerning membership should be addressed. Checks accompanying membership applications should be made payable to the Ecological Society of America.
For additional information on the Society and its publications, visit ESA's home page on the World Wide Web http://esa.org›.

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New Web Sites

I would like to add a new Department to the Bulletin of the Ecological Society of America featuring web sites that would be of interest to the widest number of ecologists.

The Editor of this Department will submit for each Bulletin several web sites and a brief (one paragraph or less) commentary on each site. If you would like to be considered for this position, please send me a CV and an example of how you think the Department could be set up.

E. A. Johnson

E-mail: [email protected]

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Society Notices

ESA 2007 Election Results

The following members have been elected to serve as Society officers, Governing Board members, and committee members. Their terms begin at the end of the 2007 Annual Meeting in San Jose.

(President-Elect August 2007–August 2008, President August 2008–August 2009,
Past-President August 2009–August 2010)
Alison (Sunny) Powers

Vice President for Science
(August 2007–August 2010)
Robert B. Jackson

Secretary (August 2007–August 2010)
David W. Inouye


(August 2007–August 2009)
Ann P. Kinzig

Board of Professional Certification
(January 2007–December 2009)
David B. Breshears
Carmen R. Cid
Steven N. Handel
Diane Wickland

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Call for Nominations: ESA Honorary Member Award

Honorary Membership in the Ecological Society is given to a distinguished ecologist who has made exceptional contributions to ecology and whose principal residence and site of ecological research are outside of North America. The Honorary Member receives a lifetime membership in the Ecological Society of America, lifetime subscriptions to all the ESA journals, and funding to attend the ESA Annual Meeting to receive a plaque at the awards ceremony. Recent honorees include Henri Decamps, John Robert Lewis, Norman Owen-Smith, Madhav Gadgil, and Carlos Herrara.

To submit a nomination, please e-mail a one-page letter of nomination and the nominee’s CV to the Committee Chair, Sandy Tartowski, at ‹[email protected]› by 9 February 2006.



Murray F. Buell Award and E. Lucy Braun Award

Murray F. Buell had a long and distinguished record of service and accomplishment in the Ecological Society of America. Among other things, he ascribed great importance to the participation of students in meetings and to excellence in the presentation of papers. To honor his selfless dedication to the younger generation of ecologists, the Murray F. Buell Award for Excellence in Ecology is given to a student for the outstanding oral paper presented at the ESA Annual Meeting.

E. Lucy Braun, an eminent plant ecologist and one of the charter members of the Society, studied and mapped the deciduous forest regions of eastern North America and described them in her classic book, The Deciduous Forests of Eastern North America. To honor her, the E. Lucy Braun Award for Excellence in Ecology is given to a student for the outstanding poster presentation at the ESA Annual Meeting.

A candidate for these awards must be an undergraduate, a graduate student, or a recent doctorate not more than 9 months past graduation at the time of the meeting. The paper or poster must be presented as part of the program sponsored by the Ecological Society of America, but the student need not be an ESA member. To be eligible for these awards the student must be the sole or senior author of the oral paper (Note: symposium talks are ineligible) or poster. Papers and posters will be judged on the significance of ideas, creativity, quality of methodology, validity of conclusions drawn from results, and clarity of presentation. While all students are encouraged to participate, winning papers and posters typically describe fully completed projects. The students selected for these awards will be announced in the ESA Bulletin following the Annual Meeting. A certificate and a check for $500 will be presented to each recipient at the next ESA Annual Meeting.

If you wish to be considered for either of these awards at the 2007 Annual Meeting, you must send the following to the Chair of the Student Awards Subcommittee: (1) the application form below, (2) a copy of your abstract, and (3) a 250-word or less description of why/how the research presented will advance the field of ecology. Because of the large number of applications for the Buell and Braun awards in recent years, applicants may be pre-screened prior to the meeting, based on the quality of the abstract and this description of the significance of their research. The application form, abstract, and research justification must be sent by mail, fax, or e-mail (e-mail is preferred; send e-mail to [email protected]) to the Chair of the Student Awards Subcommittee: Dr. Anita L. Davelos Baines, Dept. of Biology, The University of Texas-Pan American, 1201 W. University Drive, Edinburg, TX 78541-2999 USA. If you have questions, write, call (956) 380-8732, fax (956) 381-3657, or e-mail: [email protected]. You will be provided with suggestions for enhancing a paper or poster. The deadline for submission of form and abstract is 1 March 2006; applications sent after 1 March 2006 will not be considered. This submission is in addition to the regular abstract submission. Buell/Braun participants who fail to notify the B/B Chair by 1 May of withdrawal from the meeting will be ineligible, barring exceptional circumstances, for consideration in the future. Electronic versions of the Application Form are available on the ESA web site, or you can send an e-mail to [email protected] and request that an electronic version be sent to you as an attachment.

Application Form for Buell or Braun Award

Name _______________________________________________________________________________________

Current Mailing Address _____________________________________________________________________________

Current Telephone _________________________________________________________________________________

E-mail __________________________________________________________________________________________

College/University Affiliation ___________________________________________________________________________

Title of Presentation _________________________________________________________________________________

Presentation: Paper (Buell Award) ______ Poster (Braun Award) _______

At the time of presentation I will be (check one):
______an undergraduate student ______a graduate student______a recent doctorate not more than 9 months past graduation

I will be the sole ____ /senior ____ author (check one) of the paper/poster.

Signed (electronic signatures are OK)

Please attach a copy of your abstract and 250-word or less description of why/how the research presented will advance the field of ecology.

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Murray F. Buell Award
E. Lucy Braun Award

Judges are needed to evaluate candidates for the Murray F. Buell Award for the outstanding oral presentation by a student and the E. Lucy Braun Award for the outstanding poster presentation by a student at the Annual ESA Meeting at Memphis, Tennessee in 2006. We need to provide each candidate with at least four judges competent in the specific subject of the presentation. Each judge is asked to evaluate 3–5 papers and/or posters. Current graduate students are not eligible to judge. This is a great way to become involved in an important ESA activity. We desperately need your help!

Please complete and send this form by mail, fax, or e-mail to the Chair of the Student Awards Subcommittee:Dr. Anita L. Davelos Baines, Dept. of Biology, The University of Texas–Pan American, 1201 W. University Drive, Edinburg, TX 78541-2999 USA. (956) 380-8732, fax (956) 381-3657, or e-mail: [email protected].

If you have judged in the past several years, this information is on file. If you do not have to update your information, simply send me an e-mail message, “Yes, I can judge this year.”

Name ______________________________________________________________________________________________
Current mailing address _______________________________________________________________________________
June/July mailing address _____________________________________________________________________________
Current telephone Summer telephone ____________________________________________________________________
E-mail Fax __________________________________________________________________________________________
Year M.S. received Year Ph.D received ______________________________________

Areas of expertise (check all that apply):
— Discipline Research approach (please rank) Organisms
— Botany Population ecology Vertebrates
— Zoology Community ecology Types:______________________________________________________________________
— Microbiology Ecosystem ecology Invertebrates
— Applied ecology Types:__________________________________________________________________________________
— Habitat Physiological ecology Plants
— Soil Behavioral ecology Types:____________________________________________________________________________
— Terrestrial Paleoecology Fungi
— Freshwater Theoretical ecology Microbes
— Marine Evolutionary ecology Types:_________________________________________________________________________

Provide a few key words or phrases that describe your interests and expertise: _________________________



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Other Notices

Vegetation Classification Panel and the Federal Geographic Data Committee Vegetation Subcommittee

The Vegetation Classification Panel of the Ecological Society of America has been working with the Federal Geographic Data Committee Vegetation Subcommittee for several years to revise the established standards for vegetation classification, and add a mechanism for inclusion of the finest levels of the classification hierarchy (Associations and Alliances). A new draft federal standard (to replace and augment the 1997 standard) is now being circulated to the federal agencies for comment. Your comments would also be appreciated. I will compile any comments I receive and forward them to FGDC.

I attach an executive summary by Panel member Don Faber-Langendoen.

The full draft standard may be seen at ‹http://www.bio.unc.edu/faculty/peet/lab/temp/FGDCDraftStd_Aug2006.pdf

The key things to note are:

1) The upper levels of the hierarchy have been greatly modified.

2) Cultural vegetation is pulled out at the highest level.

3) The whole classification is now shifting to a dynamic content standard (a first for FGDC), where types can be added, deleted, or changed through a peer-review process without going through the entire FGDC agency approval process. ESA is likely to participate in that peer review, though the standard does not include the implementation plan.

Robert Peet
University of North Carolina
Chapel Hill, NC 27599-3275 USA
(919) 368-4971
E-mail: [email protected]


Vegetation Subcommittee

Federal Geographic Data Committee

SUMMMARY – NatureServe version

31 August 2006


The United States Federal Geographic Data Committee (hereafter called the FGDC) is tasked to develop geospatial data standards that will enable sharing of spatial data among producers and users. In 1997, the FGDC Vegetation Subcommittee developed a National Vegetation Classification (NVC) Standard (FGDC-STD-005-1997). This document is a proposed revision of the 1997 standard and, if approved, will replace that document.
Prior to formal submission, the document is being sent for review to federal agency and non-agency partners. The standards were developed in consultation with guidelines provided by the Ecological Society of America’s Vegetation Classification Panel and from the subcommittee’s Hierarchy Revisions Working Group.

The standard requires that vegetation types be defined and characterized using appropriate data. New vegetation types will be defined and previously defined types will be refined as data continue to be collected, analyzed, and correlated over time. This process is referred to as successive refinement (or successive approximation). Managing the vegetation classification (content standard) dynamically as the classification process is implemented will allow development of the national vegetation classification system (i.e. data classification standard) to proceed efficiently.

The overall purpose of this National Vegetation Classification Standard (hereafter referred to as the “Standard”) is to support the development and use of a consistent national vegetation classification (hereafter referred to as the “NVC”) in order to produce uniform statistics about vegetation resources across the nation, based on vegetation data gathered at local, regional, or national levels. This will facilitate cooperation on vegetation management issues that transcend jurisdictional boundaries. It is therefore important that, as agencies map or inventory vegetation, they collect enough data to translate it for national reporting, aggregation, and comparisons. The ability to crosswalk other vegetation classifications and map legends to the NVC will facilitate the compilation of regional and national summaries.


This Standard establishes national procedures for classifying existing vegetation for the United States and its Trust Territories that shall be used by Federal agencies to share vegetation information and report national statistics. Existing vegetation is the plant cover, or floristic composition and vegetation structure, documented to occur at a specific location and time.


Each Federal agency is free to develop vegetation classification systems that meet their own information and business needs. The ecological characteristics of such local vegetation types can help guide the design of map legends (sets of map units) to address varying land management issues at multiple spatial scales. The NVC is expected to provide the common link to compare and relate these various map legends to each other and facilitate information sharing between federal agencies and other organizations.


The structure of the revised NVC hierarchy is a substantial revision of the 1997 hierarchy. The revised hierarchy addresses the following issues, among others:

  • uses vegetation criteria to define all types, de-emphasizing abiotic criteria (e.g., eliminating direct use of hydrologic regimes in wetland formations, and using vegetation growth forms instead),
  • provides a clear “up-front” distinction between natural and cultural vegetation wherever these can be observed from broad growth form patterns (rather than combining natural and cultural vegetation initially and separating them at lower levels),
  • for natural vegetation, defines the upper levels based on broad growth form patterns that reflect ecological relationships (rather than detailed structural criteria, which are more appropriate lower down);
  • provides a new set of middle-level natural units that bridge the large conceptual gap between alliance and formation,
  • integrates the physiognomic and floristic hierarchy levels based on ecologic vegetation patterns, rather than developing the physiognomic and floristic levels independently and then forcing them into a hierarchy,
  • provides detailed standards for plot data collection, type description and classification, data management and peer review of natural vegetation, and
  • for cultural vegetation, provides an independent set of levels that addresses the particular needs of cultural vegetation.

Examples of how the NVC can be linked to various land cover classification schemes are also provided.


Table 1. Major categories used to organize the hierarchy, and the set of Level 1 units proposed to replace the current Level 1 units.



Level 1 – Formation Class



Forest and Woodland (Mesomorphic Tree Vegetation)



Shrubland and Grassland (Mesomorphic Shrub and Herb Vegetation)


Semi-Desert Vegetation (Xeromorphic Vegetation)


Polar and High Montane Vegetation (Cryomorphic Vegetation)



Aquatic Vegetation (Hydromorphic Vegetation)



Nonvascular and Sparse Vascular Vegetation (Lithomorphic Vegetation)



Agricultural Vegetation


Developed Vegetation


Not included in the NVC.

Table 2. Revised hierarchy for natural vegetation with an example.

Revised Hierarchy for Natural Vegetation


Upper Levels

1 – Formation Class

Scientific Name: Mesomorphic Shrub and Herb Vegetation

Colloquial Name: Shrubland and Grassland

2 – Formation Subclass

Scientific Name: Temperate and Boreal Shrub and Herb Vegetation

Colloquial Name: Temperate and Boreal Shrubland & Grassland

3 - Formation

Scientific Name: Temperate Shrub and Herb Vegetation

Colloquial Name: Temperate Shrubland & Grassland

Mid Levels

4 – Division

Scientific Name:Andropogon – Stipa – Bouteloua Grassland & Shrubland Division

Colloquial Name: North American Great Plains Grassland & Shrubland

5 – Macrogroup

Scientific Name:Andropogon gerardii – Schizachyrium scoparium – Sorghastrum nutans Grassland & Shrubland Macrogroup

Colloquial Name: Great Plains Tall Grassland & Shrubland

6 - Group

Scientific Name:Andropogon gerardii – Sporobolus heterolepis Grassland Group

Colloquial Name: Great Plains Tallgrass Mesic Grassland

Lower Levels

7 – Alliance

Scientific Name:Andropogon gerardii – (Clamagrostis Canadensis – Panicum virgatum) Herbaceous Alliance

Colloquial Name: Wet-mesic Tallgrass Prairie

8 – Association

Scientific Name:Andropogon gerardii – Panicum virgatum – Helianthus grosseserratus Herbaceous Vegetation

Colloquial Name: Central Wet-mesic Tallgrass Prairie

Table 3. Revised hierarchy for cultural vegetation with examples (from National Resources Inventory).

Revised Hierarchy for Natural Vegetation




Level 1 – Cultural Class

Agricultural Vegetation

Agricultural Vegetation

Level 2 – Cultural Subclass

Herbaceous Agricultural Vegetation

Woody Agricultural Vegetation

Level 3 – Cultural Formation [NRI]

Cultivated Crop

Woody Horticultural Crop

Level 4 – Cultural Subformation

Row Crop



Level 3 – Cultural Group [optional]

Temperate and Tropical Row Cro p

Temperate and Tropical Orchard

Level 4 – Cultural Subgroup (NRI)




Level 7 – Cultural Type (NRI)

Sweet Corn


Level 8 – Cultural Subtype [optional]


Data sources

Describes how to collect field plot data for vegetation classification purposes. Also provides guidelines for using literature and other data sources in addition to plot data.

Classification and description

Decribes how to prepare vegetation plot data or literature-based data, how to analyze and interpret the data, build a description of a type, including nomenclature.

Peer review of proposed vegetation types

Describes how to submit a vegetation type for peer review to an FGDC sponsored Peer Review team.

Data management and dissemination

Describes the databases, meta data standards, and web sites needed to maintain vegetation classification plot data and NVC type descriptions.


At this time the critical components for describing and classifying cultural vegetation are not well-developed. It is suggested that, for Agricultural Vegetation, the National Resource Inventory classification be a starting point for the NVC standard, and that where desirable, the standards developed for natural vegetation can be applied.


Based on comments received on this document (due 6 October), the FGDC vegetation subcommittee will revise the standards document and submit it for official approval to the FGDC Standards Working Group (SWG). The SWG is charged with conducting a public review of the proposed revised standard, and, provided comments and corrections are incorporated to the satisfaction of the SWG, would move to adopt the standard.

At the same time, the FGDC vegetation subcommittee with partners to solicit funds to maintain the standard, to develop an implementation standard, and to work with agencies to discuss how crosswalking to the standard can be accomplished.

Appendix B. Relation of USNVC to Land Cover Classifications

Table B.1. Comparison of FAO LCCS Land Cover Types (based on structural domains) and National Land Cover Database (NLCD) types with that of NVC Level 1 (see Di Gregorio and Jansen 1996, USGS 2001).





LCCS Major Land Cover Type with Structural Domain

NLCD (* indicates applies to Alaskan tundra only).

NVC Level 1



TERRESTRIAL: A12. Natural and Semi-Natural Terrestrial Vegetation

Forest & Woodland

Thicket & Shrubland


Sparse Vegetation



• Shrubland

– Dwarf Shrub*

– Shrub/Scrub


– Grassland/ Herbaceous

– Sedge Herbaceous*

• Non-Vascular

– Lichens*

– Moss* •


Forested Wetland

Scrub/Shrub Wetland

Emergent Herb Wetland

Aquatic Bed

Forest & Woodland

Shrubland & Grassland


Polar & High Montane Vegetation

Aquatic Vegetation

Nonvascular & Sparse Vascular Vegetation

WETLAND/AQUATIC:A24. Natural and Semi-Natural Aquatic or Regularly Flooded Vegetation

Forest & Woodland

Closed Shrubs & Open Shrubs


Sparse Vegetation




A11. Cultivated and Managed Terrestrial Areas

• Agriculture

Tree Crops

Shrub Crops

Herbaceous Crops

• Developed

Managed Lands

- parks (woody)

- parkland (scattered woody)

- lawns (herb)

• Agriculture

- Cultivated Crops (woody)

- Cultivated Crops (herb)

- Pasture/Hay

• Developed

– Developed, Open Space

• Agricultural Vegetation

- Woody Ag. Vegetation

Woody Horticultural Crops

Other Woody Ag. Vegetation

- Herbaceous Ag. Vegetation

Cultivated Crop


Other Herbaceous Ag Vegetation

• Developed Vegetation

- Herbaceous and Woody Developed Vegetation


A23. Cultivated Aquatic or Regularly Flooded Areas

- Aquatic Or Regularly Flooded Graminoid Crops

- Aquatic Or Regularly Flooded Non-Graminoid Crops





B16. Bare Areas

Consolidated Areas

Unconsolidated Areas

• Barren

– Rock/Sand/Clay

– Unconsolidated Shore**

FAO (informative)


B28. Natural Waterbodies, Snow and Ice

Natural Waterbodies



• W ater

– Open Water

– Perennial Ice/Snow

FAO (informative)



B15. Artificial Surfaces and Associated Areas

Built-Up Areas (Developed)

Non Built-Up Areas (Waste)

• Developed

– Low Intensity

– Medium Intensity

– High Intensity

(FAO informative)


B27. Artificial Surfaces and Associated Areas

Artificial Waterbodies

Artificial Snow

Artificial Ice

• Water

– Open Water

– Perennial Ice/Snow

FAO (informative)

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Resolution of Respect

Frank B. Golley (1930–2006)

Frank Benjamin Golley III was born on 24 September 1930 in Chicago, Illinois, to Frank Benjamin and Katharine Baird Golley, and spent his early years there. In January 1958 he took a position as assistant professor of zoology at the University of North Carolina. However, he was convinced by Eugene Odum to come to the University of Georgia in September of that year to work with him on Atomic Energy Commission (AEC) contract research at the recently established site of the Savannah River Plant in South Carolina. For Golley, this was the end of an academic odyssey, beginning with a B.S. in agriculture from Purdue University, followed by a M.S. in wildlife management from Washington State University, and a Ph.D. in zoology from Michigan State University. Golley embraced the ecosystem ecology of the Odum brothers, and this was the start of a long and productive collaboration that involved wide-ranging ecological research and the development of the ecology program at the University of Georgia.

Much of the research carried out for the AEC at the Savannah River Plant was from the very start basic in character. The philosophy behind this decision, which the AEC endorsed, was that in order to be prepared to deal with undefined future environmental problems with radionuclides, it was first necessary to understand the basic principles of ecosystem science and to develop a cadre of experienced scientists upon whom the AEC could call when needed (J. Wolfe, personal communication). Golley was insistent that the research be independent, and that it be published in the open literature without prior review by the plant management. Although this raised security issues and created friction with managers, Golley stood his ground. It proved to be the best policy for all parties, because it gave the work credibility. With expansion of the AEC contract and a move into larger quarters on the Savannah River site in 1961, Golley became the first Director of the Savannah River Radiation Ecology Laboratory (“radiation” was later officially dropped from the name), where he remained until 1962. He then moved to the Athens campus in 1967 to become Executive Director of the newly established Institute of Ecology. Shortly thereafter, the University received matching funds from NSF for construction of a building for the Institute. Golley took a strong interest in the design of that building, creating an aesthetically appealing structure that organically fostered interaction of the scientists. He especially wanted to involve graduate students and make them feel that they were an important, valued part of the Institute. It is typical of his vision for the Institute that each graduate student’s cubicle had a glass wall looking out on an attractive courtyard.

In 1979 Golley took leave from the University to serve as Director of the Division of Environmental Biology at the National Science Foundation. After returning to Athens, he served as director of the Institute of Ecology from 1984 to 1987. Throughout his life, his was a multifarious career of administration, research, and teaching. As the first Director of SREL and again as the first Executive Director of the Institute, Golley took up the challenge of creating a viable, successful organization from rather meager beginnings. The success of both organizations owes as much to Golley’s dedicated leadership and vision as to Odum’s initiative.

Golley served as President of ESA, the International Association for Ecology, and the International Society of Tropical Ecology. He served on the Advisory Council of the Smithsonian Institution, the National Institute of Ecology, and the Organization for Tropical Studies. He served on the editorial board of the journal Ethics and the Environment, and was the founder and first editor of the journal Landscape Ecology. He was a member of the committee to establish the State Botanical Garden of Georgia, and served as its interim director. In the course of his career, Golley was author or editor of 17 books and more than 170 technical articles and book chapters spanning ecology, biology, social sciences, the philosophy of science, and history.

Although originally trained in mammalogy, Golley’s interests and professional activities spanned much of ecology, even the history of ecology, and went beyond ecology as such to include environmental ethics. This range of professional interests was backed up by a remarkable fund of information. He could discourse extemporaneously on any of those subjects. In practice, his research spanned much of the world. In Puerto Rico, he helped assess the impact of a copper mine. In Greece, he assisted with planning a school of natural resources. He had cooperative research projects in Panama, Poland, Costa Rica, and Venezuela, and projects and a lecture series in Japan on city greenness and environmental conservation. Fluent in Spanish, he helped to coordinate and teach a postgraduate course on rural planning and the environment at the Instituto Agronomico Mediterraneo de Zaragoza over a period of 30 years. After retirement in 2000, he continued to be a constant presence and influence at the Institute of Ecology, remaining enthusiastically involved in graduate-level teaching.

Beyond his professional interests, Golley followed in the footsteps of one of his boat-building ancestors and crafted a small sailboat. He was an avid gardener and a skilled leatherworker. He enjoyed playing the guitar and dulcimer, and had recently taken up icon painting. His thirst for learning went far beyond ecology and his role in it. In conversation, he would often casually display detailed knowledge of some unusual subject, such as 20th century Japanese printmaking. Last year, he enrolled in a university class to learn more about one of his favorite authors, Vladimir Nabokov.

He is survived by his wife, Priscilla McKinzie Golley, to whom he was married for 53 years; his son, Frank Benjamin Golley IV and wife, Ruth of Decatur Georgia; his daughter Dr. Priscilla Sue Golley and husband Milton Carlson of Sacramento, California; his daughter, Kathleen Annette Anderson of Athens, Georgia; and four grandchildren, Stuart B. Golley and Peter M. Golley of Decatur, and Zachary R. Anderson and Ashley A. Anderson of Athens.

L. R. Pomeroy, David C. Coleman, and D. A. Crossley, Jr.
Institute of Ecology
University of Georgia
Athens, Georgia

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Rejecting Editorial Rejections: a Critique to Avoid Real Revisions
in Submitted Papers

The divulgation of ecological studies is essential to improve our knowledge of the natural systems. The adequate spread of this information to the scientific community minimizes the duplication of research efforts, gives reason to sustain or discard hypotheses, and stimulates the generation of new ideas. To do this, researchers often try to publish their studies in specialized journals. However, competition for space in high-ranking journals is severe, and these journals require an objective way to accept the best works.

Traditionally, the quality of a manuscript is evaluated through a “peer-review” procedure. A submitted manuscript is sent to several specialists in the same field, and the editor makes a decision based on the reviewers’ comments and his/her own opinion. The review of a manuscript by several colleagues reduces subjectivity and improves the quality of suggestions. Overall, in a good peer-review process everybody wins: authors, referees, and the journal. Authors and reviewers learn something about the topic of the work, and if the paper is accepted for publication, the published manuscript is a better version than the previously submitted one. However, this fruitful practice is now decaying.

More and more ecological journals are shortening this process, and rejecting some papers based only on the opinion of one person: the subject-editor. The main justification is that this saves authors from wasting time waiting for the revisions that will surely be negative, allowing them to quickly send the rejected paper to another, obviously, lower-ranked journal. Ironically, the argument is given in terms of benefits for authors. I am convinced that this explanation is irrelevant and unconstructive for science progress.

First, saving time by not waiting for colleagues’ revisions to re-send the same manuscript elsewhere as soon as possible is not a genuine advantage for anybody. As an author, I am not in such a hurry. I want to gain knowledge from nature and not to collect published papers as quickly as possible. Hence I prefer to wait, learn from the reviewer’s comments, and if eventually the manuscript is rejected, to send a better version to another journal. As a reader, I wish to read the better—not the worst—version of a paper. It is clear that a real revision of a manuscript implies a profit for the author and the readers: the author improves in knowledge, and the scientific community reads a better paper. Moreover, the reason why specialized journals reject papers without real revisions (i.e., saving time for authors) is now slight. The use of the Internet currently reduces the review process to 4–6 weeks, surely a sufficiently short period for all but perhaps the fastest-moving fields of biology (certainly fast enough for ecology). Moreover, this wait could be even shorter if reviewers and editors did a more efficient job.

Second, a decision made by only one person is inevitably biased by the strengths and weaknesses of this person, and will often work against the spirit of discussion essential for the progress of science. It is difficult to understand how the subject-editor can prejudge the opinion of other experts about the quality of a paper. Although the subject-editor is often an authority on the general topic of the study, he/she will seldom be a specialist on the specific subject of the submitted manuscript. Normally, this fact generates a bias against publishing scientific novelty (Nature Publishing Group 2003). Furthermore, the opinion of one person, independently of his/her expertise, is always subjective. This is the reason why in the traditional peer review process of submission, papers are sent to 2–3 reviewers and not to only one. A revision performed only by the subject editor is like a study designed without real replications (e.g., n = 1); it weakens the inference that can be made from it (e.g., about the quality of the manuscript).

In sum, the practice of rejecting submitted papers without multiple reviews weakens the spirit of forum that is crucial for progress in science. Neither authors nor reviewers (nor journals) learn from evading reviews, and the only advantage, saving time for the author, is trivial and unhelpful. It is unquestionable that journals must establish quality standards and need to have acceptance criteria. Perhaps it is time to rethink which type of manuscript evaluation is more constructive: a monarchical criterion supported in the opinion of only one person, or a parliamentary criterion supported in several contrasting opinions.


I thank W. Eberhard for his comments. This note is dedicated to the editors of several ecological journals that, ironically, rejected it without revision.

Literature cited

Nature Publishing Group. 2003. Coping with peer-rejection. Nature 425: 645.

Alejandro G. Farji-Brener
Lab. Ecotono
CRUB-Universdad del Comahue, Argentina
E-mail: [email protected] ; [email protected]

Editorial Polices at Ecology and Ecological Monographs

I would like to thank Dr. Farji-Brener for his letter, which addresses important issues that need discussion. In recent years, many of us have had the new and distinctly unpleasant experience of having one or another of our manuscripts rejected without review. Dr. Farji-Brener is correct in suggesting that editorial policies at ecological journals have changed; indeed, they have changed a great deal. The reason for the change is that ecology is flourishing. It has become a mainstream science, and ecological journals are magnets for some of the most important science of our time. The breadth of ecology is also increasing, and new disciplines from an ever wider sphere of the sciences appear each year. The shift of ecology from the wings to center stage of science has brought a deluge of submission to our journals, to levels that would astound an editor from even a decade ago. In 2005, 1409 manuscripts were submitted to Ecology/Ecological Monographs, which is a 14.6% increase relative to 2004. A great deal of my service as Editor-in-Chief has been dedicated to addressing these growing pains. Without countermeasures the sheer volume of submissions would consume the energy and time that we need for careful deliberation of submissions. Editing could assume the features of trying to drink from a fire hose.

The steps taken to deal with increased submissions have been expensive. Ecology and Ecological Monographs have tripled the number of editors, to about 100, in the last few years. To attract and retain the best editors, we must insure them against being overwhelmed by the job. They must have time for careful deliberation of the science in the manuscripts and the reviews. We insure this by placing limits on the number of manuscripts that editors handle. These and other changes of the last few years have paid off. Ecology and Ecological Monographs have achieved rapid publication while maintaining very high bibliometric rankings. The time, from submission to print, of articles in both journals has decreased greatly. A substantial fraction of our publications now appear within a year of submission, and the most rapid quartile appears within 9 months. A major disappointment of these necessary changes is that we have been unable to review every submission; we are a nonprofit society and simply do not have the resources to do so.

Our policy is to allow editors to decide which ms to review among the large number of submissions that we receive. Our experience is that our editors readily identify a large portion of the most exciting, interesting, cutting-edge science among the manuscripts submitted to us. At the same time, we know that we miss some good science with this policy. It is reassuring that there are many other excellent outlets for manuscripts that we choose not to review or publish.

Donald R. Strong
Ecology and Ecological Monographs

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Minutes of the ESA
Governing Board

5–6 August 2006
Memphis, Tennessee

Members present:
Nancy Grimm (President), Jerry Melillo (Past-President), Alan Covich (President-Elect), Gus Shaver (Vice President for Science), Carol Brewer (Vice President for Education and Human Resources), Richard Pouyat (VP for Public Affairs), Bill Parton (VP for Finance), David Inouye (Secretary), Shahid Naeem (Member-at-Large), Dee Boersma (Member-at-Large), Dennis Ojima (Member-at-Large), and Meg Lowman (incoming VP for Education and Human Resources). Norm Christensen (incoming President-Elect) arrived 2:30 pm.

Staff present:
Katherine McCarter (Executive Director), Cliff Duke (Director of Science), Elizabeth Biggs (Director of Finance), Fran Day (Director of Development), Jason Taylor (Director of Education), Nadine Lymn (Director of Public Affairs), Sue Silver (Editor), David Baldwin (Managing Editor), David Gooding (Associate Managing Editor)

Guests present (6 August 2006):
Steve Chaplin (Co-Chair Meetings Committee), Edward Johnson (Editor-in-Chief, ESA Bulletin), Kiyoko Miyanishi (Co-Chair, Meetings Committee), and Don Strong (Editor-in-Chief, Ecology and Ecological Monographs). Staff: Michelle Horton (Meetings Manager) and Devon Rothschild (Program Assistant)

I. ROLL CALL (9:02 am)

A) The GB unanimously adopted the proposed agenda.

B) Minutes from the May 2006 Governing Board meeting were adopted with one editorial correction.


A) Report of the President (Grimm)

President Grimm highlighted 5 areas of activity:

1) Regional activities (Regional Knowledge Partnerships), and the pilot Southeast Project (update from Alan and Jerry later on).

2) Growth in membership. On track to reach 10,000, perhaps at this meeting.

3) Education. There are many SEEDS activities for 40 student participants at this meeting. A proposal has been submitted for a SEEDS For Teachers program. Remaining funds from the Ford Foundation Mexico meeting grant are supporting five graduate students from South America to make presentations at this meeting.

4) ESA representing the ecological science community. The goal is to be a voice to funding agencies, to foster initiatives that require an integrated and coherent community effort; e.g., meetings with federal agencies (USDA’s CREES program last May). Science office has been busy (more later). A special session on Monday with representatives of USDA, NSF and NASA on a panel, and initiatives from the ecological community (LTER-based or initiated programs).

5) Publications review. A charge has been developed for Board approval at this meeting, to form an Ad-Hoc “Publications Review Committee.”

B) Report of the Executive Director and staff

1) Executive Director (McCarter)

The Annual Meeting will probably be a little smaller than expected (possibly 2800 to 3000 vs. 3200 expected), but is expected to achieve budget expectations. The new team has worked well during the build-up to the meeting. Web site redesign is proceeding, with staff reviewing and rewriting text.

2) Public Affairs (Lymn)

Annie Drinkard has been working hard to get media coverage of the meeting: reporters from the Nature office in London, Mother Jones, a local TV station, Smithsonian Magazine, etc. Board members are invited to the Rapid Response Team lunch. The King County (Washington State) executive, Ron Simms, will be speaking at the Opening Plenary. As an experiment this year, Anne Bartuska will give a talk on environmental justice at the Anointed Temple of Praise, a popular African-American church 15 miles from the Convention Center. A wetlands course on the Hill is being offered for congressional staffers for 2 days during the August recess, with a field trip on the third day to the Smithsonian’s Environmental Research Center.

3) Publications (Baldwin)

Submissions continue to increase ~10%/yr, average paper length and acceptance rates have gone down, so there has been no increase in the backlog of unpublished papers. First ever supplement for Ecology was published in conjunction with the July issue this year. A small-scale experiment with open access (one article per issue) was started with the July issue of Ecology; this is not being advertised widely yet, but is a trial to see how this works. An attempt will be made to implement “mini-subscriptions,” allowing individuals to purchase numbers of on-line articles in quantities of 1, 5, 10, 20 over a year or two (using the Geological Society’s program as a model).

4) Frontiers (Silver)

The new impact factor for the journal is 4.745, placing it second among environmental science journals and sixth among ecological journals. The September issue will be the one focused on China, and will be open access. The whole issue may be translated (abstracts already are); 300 copies will be sent to authors to distribute, and it may be made available at the May 2007 meeting in China. Other special topic issues, including one from the ESA meeting in Merida, Mexico, are coming up.

5) Administration (Biggs)

The fiscal year just ended with a positive balance. For the coming fiscal year, $50,000 less is budgeted than last year as income from the Annual Meeting will be less than Montreal, and the bump in enrollment that typically accompanies the meeting may not quite get us to 10,000 members this year.

6) Education (Taylor)

A fourth CD of Teaching Issues and Experiments in Ecologywas published this year. SEEDS (now in its 10th year) had a field trip to the Konza Prairie LTER site in June. The largest number of applications ever received came in this year for attendance at the Annual Meeting, and about 40 ESA members are serving as mentors for these students. Board members are encouraged to attend the Diversity Luncheon. The student coordinator will be leaving for a graduate program this fall and interviews are ongoing for a replacement.

7) Science (Duke)

About 350 people attended the workshop on agricultural air quality in June. The sustainability science workshop funding has been approved, for next spring in the northeast. The data registry workshop occurred last month. The NBII collaboration on a web site about pollination is proceeding. The cooperative agreement (a 3-year project) with them could lead to a variety of other collaborative efforts. In September a project will begin to produce a series of papers about ecological services provided by agricultural wetlands (NRCS collaboration); plan is to publish these as an open-access supplement to Applications. Involvement in 2007 nitrogen meeting in Brazil is proceeding (primarily publication support – maybe a special issue of Applications). Two Board members are working with Cliff on the issues of sustainability science (Shaver) and data sharing (Grimm). Another potential issue is the ecological effects of warfare (Boersma). ESA blog will be initiated with the new Web site. There was discussion about how the blog will be monitored / controlled.

8) Development (Day)

Frontiers will become a focus for funding after this Annual Meeting. SEEDS is progressing well, with an upcoming focus on ecology clubs and an international component. Collaboration with the Science office will include focusing on nitrogen and ecological effects of war. An upcoming survey will help in development of both membership and Frontiers funding. In the coming year a series of “friend-raising” events will be held around the country to try and cultivate major donors (D.C. as a prototype, possibly Boston, Atlanta, Santa Fe). Knowledge Partnerships for the Southeast are also in progress. It was noted that almost all Board members contributed to the Millennium Fund.

C) Report of the Vice President for Finance (Parton)

The unaudited year-end statement was presented. Grant revenue was more than expected, and interest and dues income were greater than anticipated. Bottom line is that we are able to add $281,000 to the unrestricted fund (reserve) category. As subscription numbers continue to erode, we may need to find a way to endow the journals to support their continued publication and to deal with publishing issues such as open access.

Investment update: the past 3 months haven’t done so well (lost ~1%), but we are still up ~7% over the past year. We have ~60% in equities (mostly in mutual funds), 40% in bonds.


A) Fiscal Year 2006–2007 ESA budget (McCarter)

1) Approval of proposed budget

The only change since the draft was presented in May is a change from 4% to 5% (2% cost of living, 3% merit) for staff salary increases. A motion is moved, seconded, and approved: The Board approves the proposed budget of $6,025,838.

2) Long-range planning grants

At-large members of the Board helped to review the proposals from Chapters and Sections. This system worked well, and will be continued with Dennis Ojima as Chair. Listing of previous awards on the Web site might be a good way to provide some guidance to future applicants.

3) Committee funds

These have typically been used for meetings of four committees each year: Science, Public Affairs, and Education, plus one other that changes from year to year. $5600 per committee is allocated. This year the Board agreed to fund a meeting of the Ad Hoc Publications Review Committee.

4) Board initiative funds

There is a list of 10 potential recipients for this $20,000 budget category (which comes from the Millennium Fund). Budgets were estimated for each of the 10 ideas, and five (totaling $20,200) were discussed as possible candidates.

The Board moved, seconded, and approved: The Board approves the budget for Board strategic funds, including:

$3000 for dissemination of Profiles of Ecologists;
$2000 for an ESA intern to write the undergraduate survey report;
$5600 for a second meeting of the Special Ad-Hoc Publications Review Committee;
$5600 for a planning meeting in Athens, Georgia for Knowledge Partnership workshops;
$5600 toward the documentation of ESA history.

President Grimm suggested that we need a standardized time of year to make and discuss these kinds of ideas, and proposed presenting ideas in November, budgeting them in May, and then approving them in August.

5) Discussion of Council presentation

VP Parton will present the budget, discuss where the income came from, major expenditures, the goal of raising $2 million (or $5 million) for a Reserve Fund.

B) Reserve funds (Parton/McCarter/Biggs)

It would be prudent to have a full year’s funding available as a reserve, particularly in view of the small size of our endowment. A fringe benefit of this would be the interest income from such a reserve once it is achieved. A motion is moved, seconded, and approved: The Board approves the goal of establishing a reserve fund of $5,000,000.

C) Conflict of interest policy (Day)

A motion is moved, seconded, and approved with one abstention: The Board approved the proposed conflict of interest policy.

D) Guidelines for evaluating corporate funders/list of corporations (Day)

A motion is moved, seconded, and approved: The Board amended the current “Corporate and Commercial Support or Donations Policy by changing point #7 to read “When soliciting funds the ESA will consider evidence of commitment to environmental sustainability, biodiversity, and development of new applications of ecological science. The Development Committee will screen prospects and periodically consult with the Governing Board on potential donors.”

E) Knowledge partnerships (Melillo/Covich/staff)

The Southeast has been chosen as a pilot region for establishing these knowledge partnerships. To develop the concept, a set of two workshops is proposed, the first a meeting to plan the program for a larger workshop. The second meeting would bring a large number of stakeholders and scientists together. The first meeting will occur this November. The proposed Center for Progressive Land Use, which may receive legacy funding with the assistance of Governor Bush, may be associated with this effort as one of a small number of core subjects for the workshops.

How will success of the program be judged? Indicators will help to attract financial support. We hope that the model from this pilot project (the Southeast) will be a model for similar efforts in other parts of the country.

F) Integrative Science for Society and the Environment (Grimm)

A draft report, “Integrative Science for Society and Environment: A Strategic Research Plan,” prepared by the Research Initiatives Subcommittee of the LTER Planning Process Conference Committee, has been unofficially submitted to NSF and is presented to the Board. This is one of two outcomes from an award made to LTER to plan for initiatives within and beyond LTER. President Grimm requests feedback from Board members about potential involvement by ESA in this effort. There is some feeling that although the general idea is good, and that this broad perspective is something that ESA should be concerned and involved with, this document may be too focused on LTER instead of the general ecological community. President Grimm will take the comments presented back to the authors. They will revise the document, and ask that ESA consider writing a letter in support of the effort; a draft letter will be presented for consideration at the November meeting.

G) Yearly public policy priorities (Pouyat/Lymn)

The list of policy priorities has been vetted by the Public Affairs Committee. The consensus is that this list and the process for keeping it current are appropriate.

H) Data sharing update (Duke/Baldwin/Grimm)

The 2004 Society Summit of 12 organizations was an effort to identify common ground in interest/need for data sharing. NSF funded three workshops, on data registries (held last month), data centers, and obstacles to data sharing. There were 16 societies and 9 other organizations represented at the July 2006 workshop. Board members were encouraged to pass on recommendations to Cliff for potential participants in the two future workshops.

I) ESA top 10 list

Is there something of this nature that ESA should do to draw attention to particular issues? Maybe published annually in Frontiers? Other suggestions include: a top 10 list of facts that ecologically literate citizen should know, a list of the top 10 papers each year about ecological science, a list of the top 10 cited ecological papers each year, top 10 ecological concepts each year. VPs are requested to take these ideas into consideration.

J) Feedback from outgoing Board members (Melillo/Brewer/Naeem/Boersma)

The outgoing members all expressed positive feelings about their service on the Board and appreciation for the quality and hard work of the staff.

K) Annual Meeting schedule

Board members reviewed their obligations during the Memphis meeting.


L) Memphis meeting (Miyanishi, guest)

The meeting is smaller, similar to what we had before Portland; meetings in the East tend to be smaller, and this is not a joint meeting. Review of proposals and abstracts went well. Only seven proposals for Organized Oral Sessions, but seven Symposium proposals ended up as OOSs too; this is the third year we have had OOSs, and Board members are encouraged to spread the word about this form of special session for future meetings. Registration might end up about 2800.

M) 2008 Meeting theme

Program Chair for the 2008 meeting, Lou Gross, proposed a theme for the meeting.Proposed theme is “Enhancing ecological thinking through research/education linkages.” There is some sentiment that this could be improved to make it more enticing. Suggestions include “Ecology for society,” and “Emerging frontiers in ecological literacy.” Lou Gross will be asked to consider these ideas and report back to the Board in November.

N) Future meetings (Chaplin, guest)

1) Annual Meeting site (2011)

Two contenders were Austin, Texas and Charlotte, North Carolina; looking for something in the Southeast, following San Jose, Milwaukee, Albuquerque, and Pittsburgh. Steve Chaplin, Katherine McCarter, Michele Horton, and Tricia Crocker visited both Austin and Charlotte, and found the former the most suitable for ESA (a casual, laid-back attitude, lots of music and restaurants within a few blocks of the meeting venue). A motion is moved, seconded, and approved: The Board approves the proposal to hold the 2011 Annual Meeting in Austin.

2) Program chair/Local host issues

We still need a Local Host Committee chair in Albuquerque, where we also met in 1997. Pittsburgh-area ecologists will meet soon to help identify a Chair for that meeting.

O) Publications charge (Grimm)

The Publications Committee was asked last year to report to the Board with questions and issues pertaining to the Society’s publications. Their ideas have been used to create a charge for a Special Ad-hoc Publications Review Committee. There is some sentiment that we should de-emphasize the issue of open electronic access, as there are not many commercial or private publications moving in this direction at present. Ideas are suggested for the categories of participants who would be useful for a Review Committee (a graduate student, a librarian, Chair of Publications Committee, a Board member, etc.). Grimm and Covich will work on appointing a committee, which will receive staff support from Liz Biggs, David Baldwin, and Sue Silver. The Publications Review Committee charge was approved with the modification that consideration of open access issues be listed as a single item rather than three items.

P) NEON update (Melillo)

NEON is evolving, and some of the evolutionary steps are substantial, such as significant structural changes (new CEO, restructured Board), and changed emphasis on experimental work. The ESA will continue to offer its assistance in communicating with the ecological community (ESA membership).

Q) Meetings with Editors-in-Chief

1) Ecology/Ecological Monographs (Strong, guest)

Major demands on his time included: looking at and assigning 1400 manuscripts to about 100 editors (some ad-hoc); recruiting and retaining editors; responding to protests by authors (about 100 last year; about 10% of decisions were reversed). The editorial process seems to be going well from his perspective and that of authors.

2) ESA Bulletin (Johnson, guest)

Johnson is still looking for ways to take advantage of the digital format (e.g., pictures, videos, models). He’d like someone who could suggest Web sites of interest to ecologists, and hopes that more use will be made of the Emerging Technologies column.

3) Frontiers in Ecology and the Environment (Silver)

There was some discussion about how we might use Frontiers to make connections to the business community, building upon its success to date with policy makers and researchers.

R) Position paper/position statement updates

1) Fire management (Ojima)

Reviews are in and back to authors. The revision must still come to the Board, then to the membership, before being issued as an official statement of the Society. There is an existing Fact Sheet on Fire. The paper in progress is essentially a review paper, and perhaps impenetrable to policy makers. We may need a category of paper in between in size (e.g., three pages), accessible to policy makers.

2) Nuclear energy (Pouyat)

Currently in revision.

3) Invasives paper (Grimm)

Barney Caton wrote a response to the paper that will be published in the ESA Bulletin, and Ed Johnson has asked whether the Board would like to respond. Grimm requested that David Lodge (author of the position paper) develop a response with the other original authors, and including Board member Dee Boersma.

S) New business

Thanks to President Grimm and the departing Board members for their service.

Meeting is adjourned at 12:30 pm on 6 August 2006.

David Inouye

11 August 2006
Memphis, Tennessee

Members present:
Alan Covich (President), Nancy Grimm (Past-President), Norm Christensen (President-Elect), Gus Shaver (Vice President for Science), Meg Lowman (Vice President for Education and Human Resources), Richard Pouyat (VP for Public Affairs), Bill Parton (VP for Finance), David Inouye (Secretary), Jayne Belnap (Member-at-Large), Dennis Ojima (Member-at-Large).

Staff present:
Katherine McCarter (Executive Director), Cliff Duke (Director of Science), Elizabeth Biggs (Director of Finance), Sue Silver (Editor), Nadine Lymn (Director of Public Affairs), Fran Day (Director of Development), David Baldwin (Managing Editor), Jason Taylor (Director of Education).

I. ROLL CALL, 8:36 am


Feedback about the meeting program and venue has been favorable. Just about 2800 people registered; he has had some feedback that people liked the smaller size. Some of the ideas brought up at the meeting have helped set the stage for the 100th anniversary. NEON appears to be back on track; we’ll see what else ESA can do to help facilitate it. One suggestion heard was to schedule more talks by icons (e.g., one each day). What could we do to make the meeting more welcoming to members who teach at small schools that don’t emphasize research? There may be a move to create an environmental justice section, and to do more at the meeting site (e.g., student volunteers working on a local project).

Organizing for the coming year: Jayne Belnap has agreed to chair the Publications Review Committee, which will also include Scott Collins and others still to be appointed. Liz Biggs, David Baldwin, and Sue Silver will serve as staff resources to the committee.

Meeting dates: 16–17 November in D.C., and the 15th for new Board members. Suggested spring meeting dates 7 and 8 May. Climate change science program office is one possibility for an agency meeting, also DOE Office of Science, Fish and Wildlife Service, USGS, TNC, etc. Next year’s Annual Meeting in San Jose, California, 4–10 August.

Anniversary preparations for the 100th year (2015). Historical Records Committee is being involved, encourages archiving of Society records. What can we and others learn from our history? The contributions of individuals coming up with important ideas from their familiarity with natural history is one important theme—how can we ensure that this will continue? And what is the future role of big science projects? Will its budgets create problems for funding smaller-scale research? Don Shure played an important role at the 75th Annual Meeting, and should be contacted regarding the 100th. A showcase exhibit on Capitol Hill and for the media would be appropriate. Last year Nadine presented a history of the Public Affairs Office; perhaps other VPs could work on similar summaries. Perhaps we could collaborate with other agencies to highlight previous efforts such as the International Geophysical Union. Maybe try for a postage stamp? Society for Environmental History might be interested in collaborating (Nancy Langston at Wisconsin might be a contact; headquarters is in Durham).

Steering committee for the Southeast Regional Knowledge Partnership will meet in early November. Members include: Michael Binford, University of Florida, Gainesville, FL; Norman Christensen, Duke University, Durham, NC; Alan P. Covich, University of Georgia, Athens, GA; Virginia Dale, Oak Ridge National Laboratory, TN; Dr. Frank S. Gilliam, Marshall University, Huntington, WV; Margaret D. Lowman, New University of Florida, Sarasota, FL; Jerry Melillo, The Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA; Robert R. Twilley, Louisiana State University, Baton Rouge, LA; Peter S. White, University of North Carolina, Raleigh, NC. An outreach effort will be an important component.

EcoSummit: This will be the third one, the first in China. The Asian Section will play an important role. Elsevier is sponsoring it, Ecological Society of China has taken a major role (e.g., in terms of financial risk). Our role is to serve on the Program Committee and help (through our membership) suggest symposia, and help organize/award student funding. Several Board members will attend the summit. In addition, Katherine McCarter and Sue Silver will exhibit at the meeting and publicize ESA’s journals. Editor of Ecological Complexity, Bai-Lian (Larry) Li, is the point person for Elsevier. They may publish a set of symposium volumes from the meeting.


This group reviews the audit, and participates in a conference call with the Auditors. Norm Christensen was appointed to the committee.

Meeting feedback

Some donors to funds other than Millennium Fund were unhappy because they weren’t invited to a reception and didn’t get DONOR tags for their name tags. Some postdocs and recently appointed faculty members asked for graded registration fees (similar to membership fees). Maybe a young-scientist network would be a good idea. The Student Section activities attracted a lot of participants and went well, although there were some complaints about the Student Mixer (expensive, not enough food). SEEDS went well; Nancy encouraged Board members to participate in the Diversity Mixer. Katherine encouraged more Board members to attend the Bagels with the Board breakfast in the future. The Public Outreach session (community presentation) on Wednesday evening featured Ann Bartuska talking at the Anointed Temple of Praise—a mixed success due to a conflict at the church (a service that evening). Press Office report: The reporter from Nature was busy sending out reports, and about 15 other media representatives were present. The Ecological Society of Mexico will have its first meeting in November; Rich Pouyat will attend, as the Society is hoping to establish a public affairs committee modeled after ours.

Meeting was adjourned at 10:34 pm on 11 August 2006.

David Inouye


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(all rights reserved, used by permission)

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Assymetric Competition in Marine Snails

Littorine snails compete indirectly by altering the traits of a shared seaweed resource. Direct grazing by smooth periwinkles (Littorina obtusata; yellow snail on left) reduced seaweed palatability for conspecifics and congeners (common periwinkles, Littorina littorea; dark snail on right). In addition, previous grazing by smooth periwinkles reduced populations of common periwinkles on outplanted seaweeds. These interactions were asymmetric, with a specialist (smooth periwinkles) competitively superior to a generalist (common periwinkles).

This photograph illustrates the article, “Asymmetric competition via induced resistance: specialist herbivores indirectly suppress generalist preference and populations” by Jeremy D. Long, Rebecca S. Hamilton, and Jocelyn L. Mitchell, tentatively scheduled to appear in Ecology 88(4), April 2007.

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Trophic Changes in a High-elevation
Riparian System

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The consequences of climate change for ecosystem structure and function remain largely unknown. A 20-year study of a high-elevation riparian system in Arizona shows substantial changes in both structure and function related to climate change. Abundances of dominant deciduous trees have declined dramatically over the 20 years, correlated with a decline in over-winter snowfall. Snowfall can affect over-winter presence of elk, whose browsing can significantly impact deciduous tree abundance. Seven species of birds were found to initiate earlier breeding associated with an increase in spring temperature across years, with minimal consequences for populations. Climate had much larger consequences for these seven bird species by affecting trophic levels below (plants) and above (predators) them. In particular, climate-related declines in deciduous vegetation led to decreased abundance of preferred bird habitat and increased nest predation rates. In addition, summer precipitation declined over time, and drier summers also were further associated with greater nest predation in all species. The net result was local extinction and severe population declines in some previously common bird species; one species increased strongly in abundance, and two species did not show clear population changes. Thus, climate can alter ecosystem structure and function through complex pathways that include direct and indirect effects on abundances and interactions of multiple trophic components.

Declining snowfall over the 20 years of study have allowed elk to remain over winter and increase browsing pressure on the habitat. Here is a typical canyon maple stand showing the browse line and lack of understory vegetation below browse height. Note the dead understory limbs from over-browsing, and the open area in the forefront. Historically, stands such as this would have a thick understory. Many maple and quaking aspen ramets exist throughout this stand, but are browsed down.


Another typical canyon maple stand showing the browse line, lack of understory vegetation, and dead limbs in the understory. Also, note again the open areas in the background, showing that conifer incursion or other plants are not crowding out the deciduous vegetation.


Maple attempts to grow in the understory each year and here is the typical response, where any new shoots are browsed down by deer and elk. The density of understory maple stems has decreased by more than an order of magnitude over the 20 years of study.
This aspen ramet is 6 years old and attempts to produce new shoots each year, but is browsed back so much that it is only 15 cm tall, whereas aspen released from browsing at this age are 5–8 m tall. The lack of aspen recruitment is leading to loss of aspen in the system; marked monitoring of 878 aspen over the past 10 years predicts it will be gone in about four more years.

The Red-faced Warbler (Cardellina rubifrons ) uses maple and fir habitats for nesting and feeding, and shows a mild decline in abundance over the 20 years of study, associated with decreased abundance of its habitat.



The Hermit Thrush (Catharus guttatus ), here feeding its normal brood size of four young, is a common species in this system. It nests and feeds in both conifers and maple. It does not have a clear population trend over the 20 years, but shows a clear decline over the last 15 years, associated with the decline in habitat and summer precipitation.

The Orange-crowned Warbler (Vermivora celata ) depends on maple for nesting and feeding. It has shown a major decline in abundance over the 20 years, associated with the decline in maple, but also associated with decreasing summer precipitation and increasing nest predation rates.

These photographs illustrate the article, “Climate correlates of 20 years of trophic changes in a high-elevation riparian system,” by Thomas E. Martin, tentatively scheduled to appear in Ecology 88(2), February 2007.

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Mosaic Patterns of Thermal Stress
in the Rocky Intertidal Zone

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Rocky intertidal mussels experience rapid fluctuations in aerial body temperature during low tide. Because of variability in tidal regimes and climatic regimes, geographic gradients in body temperature along the west coast of the United States may be much more complex than previous assumed. As a result, climate change may not lead to poleward range shifts, but instead may cause damage at a series of “hot spots.” All photos by Brian Helmuth, unless noted otherwise.

Rock intertidal zone at Santa Cruz Island, California.
Photo credit: C. Blanchette.

Rocky intertidal zone at Boiler Bay, Oregon. The intertidal zone represents the interface between the terrestrial and marine environments. Thus animals and algae in these environments (such as Boiler Bay Oregon, shown here) may be among the first ecosystems to exhibit responses to global climate change.

Strawberry Draw, Tatoosh Island, Washington State. The rocky intertidal zone has long served as an experimental laboratory for investigating the influence of climate on small-scale distribution patterns of organisms.
Mytilus. Results suggest that patterns of aerial body temperature (and in particular maximum temperature) may be complex, and thus patterns of mortality (shown here from Bodega Bay, California) may occur in unexpected locations. Photo credit: C. Harley.

Intertidal zone at Tatoosh Island, Washington.
Sensors were deployed at mid-tidal heights in mussel beds at a series of sites in order to measure geographic patterns in aerial and aquatic body temperatures (Tatoosh Island, Washington).

In their article, Helmuth et al. describe how latitudinal patterns of intertidal thermal stress may be more complex than anticipated. Using a series of biomimetic temperature loggers designed to mimic the thermal characteristics of mussels (Mytilus californianus), shown here, they measured temperature patterns at sites ranging from northern Washington to southern California.

Intertidal zone at Tatoosh Island, Washington.
These photographs illustrate the article, “Mosaic patterns of thermal stress in the rocky intertidal zone: implications for climate change,” by Brian Helmuth, Bernardo R. Broitman, Carol A. Blanchette, Sarah Gilman, Patricia Halpin, Christopher D.G. Harley, Michael J. O’Donnell, Gretchen E. Hofmann, Bruce Menge, and Denise Strickland, which appeared in Ecological Monographs 76:461–480, November 2006.

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Mangrove Forest Structure and Dynamics,
Punta Galeta, Panama

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A long-term study, lead by Dr. Wayne Sousa (University of California, Berkeley), is investigating a variety of biotic and abiotic processes that may account for the structure and dynamics of mangrove forests on the Caribbean coast of central Panama. The study forests of Punta Galeta, near the Smithsonian Tropical Research Institute’s Galeta Marine Laboratory, contain three canopy tree species, which exhibit a pattern of shoreline zonation that is typical of the region. Seaward fringing stands are monopolized by Rhizophora mangle, low intertidal stands are a nearly even mixture of R. mangle and Laguncularia racemosa, and Avicennia germinans dominates more inland stands. L. racemosa often reappears in the canopy near the upland edge, and sometimes forms small monospecific stands along the mangrove-rain forest ecotone. Note: all photos by Wayne Sousa, with exception of Image 1, which is by Anand Varma.

Fringing stand of Rhizophora mangle at seaward end of tidal gradient.

Low intertidal mixed stand of Rhizophora mangle and Laguncularia racemosa.
Upper intertidal stand of Avicennia germinans; seedlings on the forest floor are predominantly Avicennia.
Our forthcoming paper examines the roles of propagule dispersal and establishment in explaining species distributions along the tidal gradient. The propagules of mangroves are buoyant and dispersed by tidal currents and runoff following rain storms. We quantified dispersal patterns by monitoring the movements (directions and distances) of marked propagules released at different distances from the water’s edge. A separate experiment measured rates of seedling establishment at these same positions along the tidal gradient. Rabinowitz’ Tidal Sorting Hypothesis posits that zonation is the product of the (1) differential landward movement of propagules of different size by incoming tidal flow (i.e. tidal sorting) and (2) greater ability of larger propagules to establish in the deeper water characteristic of the low intertidal zone. Our observations were not consistent with this model of zonation, but revealed a different form of dispersal limitation that seems to strongly influence species’ vertical distributions and forest structure.

A small river running through a mixed stand of Rhizophora mangle and Lagunculariaracemosa. Mangrove propagules that disperse into a river channel can be transported longer distances.
Dispersing propagules of Laguncularia racemosa trapped behind a log.
These photographs illustrate the article, “Supply-side ecology in mangroves: Do propagule dispersal and seedling establishment explain forest structure?” by Wayne P. Sousa, Peter G. Kennedy, Betsy J. Mitchell, and Benjamin M. Ordóñez L., tentatively scheduled to appear in Ecological Monographs 77(1):53–76, February 2007.

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Understanding Food Chains and Food Webs, 1700–1970

Fig. 1

Some ecological ideas developed gradually and only gained coherence and details after they had become commonplace. The history of two interrelated ideas, food chains and food webs, is an example of a gradual, cumulative history. Here is a brief survey of these concepts from about 1700 to 1970 (Fig. 1).

The earliest identified food chains seem to have concerned hyper-parasitism (Egerton 2005, 2006a), which students of insects discovered in the later 1600s. But it was entrepreneurial naturalist Richard Bradley (Egerton 2006b) who generalized the concept (Bradley 1718, part 3:60–61):

…Insects which prey upon others are not without some others of lesser Rank to feed upon them likewise, and so to Infinity; for that there are Beings subsisting, which are not commonly visible may be easily demonstrated…in a Microscope.


Fig. 2

This was turned into verse by Jonathan Swift in 1733 (lines 341–344) (Fig. 2).

So, Nat’ralists observe, a Flea

Hath smaller Fleas that on him prey,

And these have smaller yet to bite ‘em,

And so proceed ad infinitum.

In Bradley’s account, food chains illustrated the balance of nature (Egerton 1973:333–335). Swift was a prominent literary figure who had a general interest in science, but his lines on fleas were meant as a swipe at lesser poets. Neither Bradley nor Swift provided an illustration, so we can help them out with this one from Alfred Elliott’s 1957 Zoology textbook (Fig. 3), though Elliott borrowed the idea for the central figures of fleas from Robert Hegner’s 1938 book, Big Fleas Have Little Fleas, or Who’s Who Among the Protozoa.

Fig. 3

Carl Linnaeus, in an ecologically important essay, “The Economy of Nature” (Linnaeus [Latin] 1749; [English] 1775:114, 1977), briefly itemized the stages of two food chains, one terrestrial, one aquatic (Egerton 2007a). There are likely other naturalists between Linnaeus and Darwin who reported on food chains, but attracted little notice. Darwin may be the first to report a food web, occasioned by the Beagle’s stop over at the rather barren island of St. Paul on 16 February 1832. He gave its location as 0° 58’ north latitude and 29° 15’ west longitude, and 540 miles from America. He found only two species of birds, the booby (a gannet) and the Noddy Tern. The latter built a simple nest with seaweed. Then follows his food web (Darwin 1839:10):

By the side of many of these nests a small flying-fish was placed; which, I suppose, had been brought by the male bird for its partner…quickly a large and active crab (Craspus), which inhabits the crevices of the rock, stole the fish from the side of the nest, as soon as we had disturbed the birds. Not a single plant, not even a lichen, grows on this island; yet it is inhabited by several insects and spiders. The following list completes, I believe, the terrestrial fauna: a species of Feronia and an acarus, which must have come here as parasites on the birds; a small brown moth, belonging to a genus that feeds on feathers; a staphylinus (Quedius) and a woodlouse from beneath the dung; and lastly, numerous spiders, which I suppose prey on these small attendants on, and scavengers of the waterfowl.

After reading this account, Rear-Admiral William Symonds told Darwin that he had seen at St. Paul crabs drag young birds from nests and eat them. Darwin added his information to this passage in the second edition (1845) of his book on the voyage of the Beagle (Edwards 1985:34).

Fig. 4

In The Origin of Species (1859:73–74), Darwin reported the most famous example of a food chain in the scientific literature (Fig. 4).It is in chapter 3 on the “Struggle for existence,” and involves humble bees (called “bumble bees” in America) pollinating red clover; though some bees were eaten by field mice, the mice, in turn, were kept in check by domestic cats. Darwin speculated that if it were not for the cats, the mice would decimate the bees, and the clover would go unpollinated, since only humble bees pollinate clover. A later, unknown commentator extended this chain further (Milne and Milne 1966:6) by suggesting that old maids commonly kept cats, that clover-fed cattle were eaten by British seamen who protected the British Empire, and that if it were not for old maids, the British Empire would fall! In other words, Darwin’s food chain became a biological version of Englishman George Herbert’s well-known admonition (1640):

For want of a nail the shoe is lost,

For want of a shoe the horse is lost,

For want of a horse the rider is lost.

In America, this admonition is attributed to Ben Franklin, who borrowed it without acknowledgement for Poor Richard’s Almanac (1757). But getting back to Darwin’s food chain, in 1947 W. L. McAtee pointed out that Darwin’s food chain dynamic lacks full validity, since we now know that honey bees also pollinate red clover and that humble bees often appropriate mouse holes, so humble bees and mice have an ambiguous relationship. In Darwin’s defense, he heavily depended on H. W. Newman’s 1850–1851 study “On the habits of the Bombinatrices” (Darwin 1975:183).

The next discussion of note for our purposes is from a remarkable German zoologist, Karl Semper. In 1877, he gave 12 lectures at the Lowell Institute in Boston, published simultaneously in English and German editions in 1881. The English title is Animal Life as Affected by the Natural Conditions of Existence. This book was the first detailed synthesis of animal ecology. In a discussion of the food of herbivores and carnivores (Semper 1881:51–52), he pointed out that when herbivores transform vegetation into flesh, there is a loss of mass due to oxidation of organic material, and that the same is true when carnivores transform the flesh of their prey into their own flesh. To illustrate this, he arbitrarily assumed a 10 to 1 ratio of food to flesh. One thousand units of plant food could only support 100 units of a herbivore, and those 100 units of herbivore could only support 10 units of a carnivore. Although his book has 106 illustrations, this generalized food chain was not illustrated. However W. E. Pequegnat’s diagram (Fig. 5) from Scientific American (1958:86) captures Semper’s concept, even to the point of using a 10 to 1 ratio. Semper wrote at a time when there was little quantified thinking in natural history. He had first trained as an engineer and then as a physiologist (Mayr 1975), and that background came to the fore in this discussion. Although his book was widely read, apparently no one carried this line of quantitative thinking any further in the 1880s or 1890s.


Fig. 5

We are used to seeing food chains or webs diagrammed. The advantages are obvious: they provide a visual panorama of detailed information. The early history of such diagrams is elusive. The bibliography on food chains and webs that Allee, Emerson, Park, Park, and Schmidt compiled (1949:514) can assist in the search. However, they did not discover the earliest ones now known, published in 1880 by Lorenzo Camerano, which are reprinted in an English translation of his article (1994:377–378). Since Camerano’s two diagrams do not resemble any known from later zoologists, it seems likely that he did not have much, if any, influence on later diagrams. Joel Cohen (1994:353–355) suggests that Camerano was influenced by diagrams for other purposes in books by Darwin and by Hermann Helmholtz, though Camerano’s diagrams do not resemble theirs. Like Semper’s, Camerano’s food webs are generalized rather than specific.

The earliest specific food web I have found (Fig. 6) is on “The boll weevil complex,” published in 1912 by Pierce, Cushman, and Hood in a USDA Bulletin. Their motive was to promote bowl weevil eradication—by encouraging its predators and parasites. Theirs may not have been the first specific diagram published, because others appeared about the same time in different biological specializations, where it is unlikely that the members of one specialization were reading the literature of other specializations.

Fig. 6

Fig. 7. Victor Shelford at age forty in 1917, associate professor at the University of Illinois. Photo courtesy of V. E. Shelford personal papers.


The following year, University of Illinois animal ecologist Victor E. Shelford (Fig. 7; photo, Croker 1991) published Animal Communities in Temperate America as Illustrated in the Chicago Region, which contained diagrams of both aquatic (Fig. 8) and land food webs (Fig. 9). There is no reason to suspect that he was influenced by the boll weevil diagram of 1912. Shelford used both of his diagrams to show how the community tends toward equilibrium, although the terrestrial community was more complex than the aquatic community, and consequently its equilibrium was more precarious. Shelford became a leading American animal ecologist (Croker 1991); his book was reprinted in 1937 and 1977.

Fig. 8

Fig. 9


The earliest known food web diagram for a marine community was drawn by Danish fishery biologist Johannes Petersen (Fig. 10) in “A preliminary result of the investigations on the valuation of the sea”(1915). He studied the Kattegat region of shallow water between eastern Denmark and Sweden (Fig. 11), an area with maximum length of 150 miles and maximum width of 90 miles. Significantly, he attempted to establish the annual productivity for this region, and his diagram indicates the thousands of tons of each group of organisms, with both a number and a proportioned rectangle (Fig. 12). In the text he stated that the eel-grass (Zostera marina) figure of 24,000,000 tons represents only the amount produced in the summer, and that the annual production is twice that. Presumably, all the other figures are annual production and not just summer production. The tons of plaice and cod are the actual commercial catch of those fish from International Fishery Statistics for 1910, and that was possibly true also for the tons of herring given, though he did not say so. The numbers given for other animals seem to be estimates. Although he indicated on his diagram that herring fed on plankton, he thought plankton was much less important than Zostera as a foundation for this food web. He concluded that the Kattegat had a “very unfavourable proportion between producers and consumers”(Petersen 1915:32). What he meant by this seems to be indicated by the following sentence in which he stated that carp ponds have “even without artificial feeding, given a yield of fish per hectare several times greater than that of the Kattegat.” Petersen reproduced the same diagram with minor alterations in his final report, “The sea bottom and its production of fish-food”(1918:23).

Fig. 10

Fig. 11

Fig. 12

His colleague, H. Blegvad, also used rectangles in his diagram of “Food of fish and principal animals in Nyborg Fjord” (1916:24) (Fig. 13)but without attempting to represent precise quantities. However, he did give quantitative data in the text of his article, which provided some sense of the quantities of organisms involved at each level.

Fig. 13

In the same year as Blegvad, the American zoologist Harold Sellers Colton published what Jonathan A. D. Fisher calls (2005:145) “possibly the first intertidal marine food web ever illustrated.” It is in Colton’s article on a carnivorous snail, Thais lapillus (now Nucella lapillus), and shows both which animals the snail eats and which animals eat the snail (Fig. 14). Colton did not indicate what inspired his diagram, but his brief bibliography does include Shelford’s book (1913). Fisher did not find references in the later relevant literature to Colton’s two articles on this snail (probably due in part to Colton’s leaving marine biology for archeology [Miller 1991]), so we do not know of any influence that his diagram exerted.

Fig. 14

Fig. 15


Charles Elton (Fig. 15) helped make such diagrams commonplace. He went on an Oxford University Arctic expedition in 1921 to Spitsbergen and took along Shelford’s book as a possible model for his own study (Elton 1966:33). However, Elton soon realized that the community he studied had a different dynamic than Shelford’s aquatic and terrestrial ones. Elton was impressed by the transfer of food from sea to land, which is reflected in his diagram (Fig. 16) published in 1923.

Fig 16

Although V. S. Summerhayes is listed as the senior author of their joint study, since he was a botanist, we can assume that Elton developed this diagram, in which plants are not emphasized. Two years later, in 1925, Elton published this much simpler Canadian food web (Fig. 17),which includes information on the lengths of animals. In 1924, English fishery biologist A. C. Hardy published a diagram (Fig. 18)on food consumed by herring at different stages of development. It bears no similarity to any diagrams previously shown, and it seems likely that he either was inspired by some unidentified example from the fisheries literature, or that he independently developed his diagram. Be that as it may, in 1927 Elton published his classic textbook, Animal Ecology, which reprinted and explained these last three diagrams by himself and Hardy. In that book Elton also introduced (1927:55) the terms “food chain” and “food cycle.” Widespread use of his book popularized the use of food web diagrams. In both Hardy’s diagram and in Elton’s for 1925, more information was conveyed than merely which animal ate which food. Hardy’s additional information was on the age of herring in relation to food, and Elton’s was on the size of the consumer in relation to food. Elton also popularized the idea of a food pyramid (1927:68–70), which concept had been implied by Semper.


Fig. 17

Fig. 18



In 1926 Germany’s leading limnologist, August Thienemann (Fig. 19) published this unique food web of lakes (Fig. 20). His 50-page article on nutrient cycles in lakes introduced into limnology the terms “producers,” “consumers,” (though Petersen 1915 [quoted above] had used both terms in marine biology) and “reducers.”

Fig. 19

Fig. 20


Thienemann’s 1926 paper and two of his other papers influenced an American postdoctoral student, Raymond Lindeman, who produced one of the most influential diagrams in the history of ecology (Fig. 21), though few if any ecologists have published similar diagrams.

Fig. 21


It appeared in his posthumous paper, “The trophic–dynamic aspect of ecology” (1942). Like Thienemann’s diagram, Lindeman’s is a generalized food web, but both men had hard specific data backing up their concepts. In that respect theirs were similar to diagrams by Shelford, Elton, and Hardy, which illustrated specific food webs, and unlike Semper’s generalized food web, which was an educated guess. In the caption to his diagram Lindeman indicated that it was similar to one he had published the previous year. A comparison of his two diagrams indicates what he learned in his year at Yale University working under Evelyn Hutchinson (Cook 1977). The 1941 diagram is identical to the 1942 diagram except it lacks the symbols for trophic levels along the side. Lindeman (1942:159) used Thienemann’s terms “producers” and “consumers,” but suggested substituting the term “decomposers” for Thienemann’s term, “reducers,” to signify that the indicated process was not just chemical, but also biological.

Fig. 22

Fig. 23


In 1943, a year after Lindeman’s 1942 diagram appeared in the journal Ecology, Harvard marine ecologist George Clarke published this conventional food web (Fig. 22), but three years later, after he had studied Lindeman’s diagram and its explanation, Clarke published his diagram (Fig. 23) in Ecological Monographs, of a marine food web that emphasizes productivity and human removal of material. It also shows Clarke’s concern for the rate of production at each trophic level.

Fig. 24

Fig. 25


The Odum brothers, Eugene and Howard Thomas, carried Lindeman’s thinking further. The Atomic Energy Committee became interested in radiation ecology (Kwa 1989:48), and Eugene Odum (Fig. 24; photo, Craige 2001) developed a program at the University of Georgia to study food chains at the Savannah River Research Facility to trace radioactive pollution (Craige 2001). By injecting plant stems with radioactive phosphorus-32, he and his colleagues traced it up the food chain to leafhoppers, beetles, and spiders (Kwa 1989:58). About 1957 the programs at Oak Ridge and Savannah River converged, with both programs using radioactive tracers to measure the flow of materials up the food chain (Kwa 1989:66). In the second edition of Eugene Odum’s famous textbook, Fundamentals of Ecology (1959:47), there is a 1949 diagram of a food chain (Fig. 25). When I saw it, I assumed that Lindeman’s influence had flowed across the Atlantic in just a few years, but when I compared it with British ecologist Erichsen Jones’ own diagram, I discovered what Odum meant when he wrote that his diagram was “redrawn” from the one by Jones: Odum added the labels to the left of the diagram as a pedagogical aid.

Fig. 26

Fig. 27

Fig. 28


Howard Thomas Odum (Fig. 26; photo from Katherine Ewel) received his graduate training under Hutchinson at Yale. (In 1954 he taught me freshman zoology at Duke.) In 1956, he produced a diagram (Fig. 27) of matter and energy flow, in steady-state flowing-water communities in Florida. At that point, the reader could still understand the diagram without special training. However, H. T. Odum continued developing his thinking along the lines of systems ecology and used symbols from electrical engineering. By 1971 he published esoteric diagrams (Fig. 28) that integrate humans into the biotic community. This was an important step towards founding several applied ecological sciences (Mitsch 1994, Hall 1995, Egerton 2007b).

Other ecologists developed food chain and food web concepts in another direction. In 1948, D. E. Howell reported finding DDT in human fat, and by 1949 biologists were reporting that fish feeding on insects killed by DDT were also being killed (Hoffmann and Surber 1949, Langford 1949).

Fig. 29

Fig. 30

Rachel Carson (Fig. 29) publicized the discovery of insecticides traveling up the food chain in ever-increasing concentrations in her best-selling book, Silent Spring (1962:110–111), as did Robert Rudd in his less-read book, Pesticides and the Living Landscape (1964). Carson did not provide diagrams, and the ones Rudd used were quite simple. Here are four (Fig. 30) of the seven diagrams in his book. DDT was the most notorious insecticide, and in 1967 George Woodwell published a diagram (Fig. 31) in Scientific American showing increased concentrations of DDT as it progressed up the food chain. By 1970, Clive Edwards constructed a much more detailed food web (Fig. 32), showing DDT pathway and concentrations from the time of spraying DDT into the air, all the way up the food chain until it became concentrated in predatory birds, mammals, and humans.

Fig. 31

Fig. 32


From simple narratives around 1700, food chain and food web concepts have been developed into progressively more sophisticated vehicles for conveying ecological ideas (Polis et al. 2004, de Ruiter et al. 2005). Lorenzo Camerano’s two 1880 diagrams of food webs had no known influence, but after the visual stimulus of diagrams became established in the early 1900s, many ecologists found creative ways to express visually their discoveries concerning food chains and webs.

Literature cited

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Bradley, R. 1718. New improvements of planting and gardening, both philosophical and practical. Edition 2, Part 3. [Edition 1, 1717–1718.] W. Mears, London, UK.

Camerano, L. 1880. Dell’ equilibrio dei viventi mercè la reciproca distruzione. Atti della Reale Accademia delle Scienze du Torino 15:393–414 + 4 plates.

Camerano, L. 1994. On the equilibrium of living beings by means of reciprocal destruction. Claudia M. Jacobi, translator. J. E. Cohen, editor. Pages 360–380 in S. A. Levin, editor. Frontiers in mathematical biology. Springer-Verlag, Berlin, Germany.

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Cohen, J. E. 1994. Lorenzo Camerano’s contribution to early food web theory. Pages 351–359 in S. A. Levin, editor. Frontiers in mathematical biology. Springer-Verlag, Berlin, Germany.

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Craige, B. J. 2001. Eugene Odum: ecosystem ecologist and environmentalist. University of Georgia Press, Athens, Georgia, USA.

Croker, R. A. 1991. Pioneer ecologist: the life and work of Victor Ernest Shelford, 1877–1968. Smithsonian Institution Press, Washington, D.C., USA.

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Darwin, C. R. 1975. Natural selection: being the second part of his big species book written from 1856 to 1858. R.C. Stauffer, editor. Cambridge University Press, Cambridge, UK.

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Edwards, A. J. 1985. Saint Paul’s Rocks: a bibliographical review of the natural history of a mid-Atlantic island. Archives of Natural History 12:31–49.

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Egerton, F. N. 2006b. A history of the ecological sciences, part 20: Richard Bradley, entrepreneurial naturalist. ESA Bulletin 87:117–127.

Egerton, F. N. 2007a. Carl Linnaeus and the economy of nature. ESA Bulletin 88, in press.

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Hoffmann, C. H., and E. W. Surber. 1949b. Effects of an aerial application of DDT on fish and fish-food organisms in two Pennsylvania watersheds. Progressive Fish-Culture 11:203–211.

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This is a revised version of a talk given at the ESA Annual Meeting in August 2006 in Memphis, Tennessee. For comments preceding the talk, I thank Robert P. McIntosh, Professor Emeritus of Biology, University of Notre Dame (now in Florida). For several references used in the revision, I thank Jonathan A. D. Fisher, Department of Biology, University of Pennsylvania, Philadelphia.

Frank N. Egerton
Department of History
University of Wisconsin-Parkside
Kenosha WI 53141
E-mail: [email protected]

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AGU Editorial: Advocacy and Integrity

The editorial below was published in the 24 October 2006 issue of EOS, the transactions of the American Geophysical Union. Although the editorial is directed at geoscientists, some of the concerns are certainly applicable to ecologists.

A few years ago Michael Mappin and I were concerned enough about the treacherous divide between advocacy, professional conduct, and education to assemble and edit a book on Environmental Education and Advocacy: Changing Perspectives of Ecology and Education, published by Cambridge University Press. Perhaps the most interesting chapter is by Joseph Des Jardins, a philosopher, in which he discusses the complicated issue of drawing both ethical and policy directives from scientific facts. I found his discussion of the “naturalistic fallacy," the assertion that scientific facts can be used as the basis of ethical statements, particularly clear. As the AGU editorial points out, as professional scientists or engineers we must remember that we are called on by society to present the best available science.

E. A. Johnson

Advocacy and Integrity

Many learned and professional societies at some time have strayed beyond the limits of what they can support from their scientific knowledge base. Unfortunately, these excursions can alter their image and hurt their credibility. However tempting it is to espouse positions that make a large fraction of members feel good, it is dangerous and in AGU’s view wrong to do so. Opposition to the Vietnam War distracted some in the 1970s. The Space Station drew opponents and proponents, in many cases on economic rather than scientific grounds. And today “climate change” is evoking extreme reactions from some industries and government entities that support the research enterprise and from individual environmental activists and skeptics across the spectrum of opinion.

AGU has not and will not articulate or support any public position on issues that extend beyond the range of available geophysical data or recognized norms of legitimate scientific debate. Positions of AGU must be based on sound science. Members are notified when the Union is preparing a position and have an opportunity to provide input. Accepted positions are publicly available. Each has a sunset limit; all must be reviewed on a regular basis. You can read the full Union policy on advocacy at ‹http://www.agu.org/sci_soc/policy/policy-advocacy.shtml

A reputation for integrity and statements based on the best available science are critical characteristics that should differentiate a scientific society from other types of organizations. Many other organizations, understandably, advocate from their heart or their pocketbook.

AGU’s goal is to encourage policy makers around the world to make wise use of the current base of scientific understanding as they develop laws, regulations, and public policy. In doing so, the Union frequently works with other scientific and professional societies. In most cases, the end product of policy makers is a political decision that can be informed by science but not directed by it.

Individual members of AGU may use the statements adopted by the Union when they communicate with local or national policy makers. It is important, however, for each of us to maintain the same level of honesty in these communications that we apply to our research. If we stray from our scientific base when speaking as scientists, we do damage to the overall integrity of science and lose the trust of policy makers and in turn of the public.

With your support, AGU can maintain its fine reputation and continue to accomplish more and more each year.

[Reprinted from EOS 87:461, 2006.]

Soroosh Sorooshian
AGU Committee on Public Affairs

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A History of the Ecological Sciences, Part 23: Linnaeus and the Economy of Nature

Carl Linnaeus (1707–1778) was a leading naturalist of the 1700s (Lindroth 1973, 1983, Morton 1981:259–276, 281–285, Goerke 1993, Broberg 2000, Spary 2002). All ecologists know he founded modern nomenclature and systematics (Larson 1971, Stafleu 1971, Mayr 1982:171–180, Eriksson 1983), but he is less well known for inventing an ecological science he called the economy of nature. He explained it in 1749, but the overly broad science of natural history, which he had pursued since childhood, was already ecological in outlook and content. In 1749 he generalized and formalized what he had been recording specifically and informally. A series of 186 essays, largely by Linnaeus, were defended by his students as dissertations for their doctoral degrees (Jackson 1913, Ramsbottom 1959:151–153, Smit 1989:118–119, Kiger et al.1999:231), and one of these was Specimen academicum de oeconomia naturae (1749), defended by Isaac J. Biberg. Linnaeus republished these dissertations in 10 volumes entitled Amoenitates Academica (Academic Pleasures, 1749–1790), though the last two volumes appeared posthumously. The Amoenitates Academica has been reprinted several times, and 19 dissertations are translated into English (Linnaeus 1775, 1781, 1977a, b). There is a helpful Index to Scientific Names of Organisms cited in Linnaean Dissertations (Kiger et al. 1999), with a guide to collected editions. Linnaeus’ earlier natural history observations are recorded in travel books and other writings.

Fig. 1. Linnaeus in his Lapland clothes (or costume). Drawn in Holland by Martin Hoffman, 1737.

All of his travel books and the dissertations are listed in B. H. Soulsby’s catalogue of Linnaeus’ works (1933:23–26, 99–151). Florence Caddy (1886–1887) provides two good maps on Linnaeus’ travels, though the caption to the one at the end of volume I is misdated 1735–1738 (read 1732–1738). Wilfrid Blunt (1971) includes maps and summaries of the trips in his biography of Linnaeus. Linnaeus’ travel books show his broad interest in plants, animals, geology (Merriam 2004), and economic uses of natural history (Linnaeus 1766, 1781:1–67, 1977, Koerner 1999, Müller-Wille 2003, Rausing 2003). David Black selected natural history extracts from Linnaeus’ books on the 1732 and 1741 trips, which he published with a map and modern illustrations by Stephen Lee (Linnaeus 1979).

Linnaeus’ first expedition was undertaken in 1732, begun on 12 May, his 25th birthday, and lasted until 10 October. He traveled north to Lapland and then west to the Norwegian coast. On the return journey he traveled in Finland down the eastern side of the Gulf of Bothinia to Åbo and then crossed to Stockholm. It was his longest journey—he estimated 633 Swedish miles or about 3800 English miles (1811, II:270, 1971)—and the subject of his longest travel book. It was also the one travel book that he illustrated. A historian of Swedish botany judged this trip “the most productive exploratory expedition ever undertaken in Sweden”(Fries 1950:18). It was sponsored by the Royal Academy of Sciences at Uppsala, which declined to publish his manuscript, and an English translation was published (1811) long before the Swedish version (1913). Linnaeus’ most recent biographer, who reads Swedish, judges some of his behavior and writings on this trip rather harshly (Koerner 1999:59–65). She says he doubled the actual distance he traveled in his report (her figure is 4500 miles) because he was to be paid per mile, that he drew a map indicating travel to places he had not visited, and that he later claimed to have stayed in Lapland much longer than he had. While I cannot check all her claims, this statement is doubly wrong: “He never passed the sixtieth degree north Latitude, which marks the Arctic Circle” (Koerner 1999:61). The Arctic Circle is actually at 66°30’, and he did cross it. He visited Jokkmokk (29 June) just north of that line, and more than half a dozen other places north of Jokkmokk (see map in Blunt 1971:41). She does not accuse him of inventing any of his natural history observations.

When he visited the cataract of the Elf-Carleby River on 13 May, he described the salmon fishery below the cataract, the foam and spray that the cataract generated, and surrounding plants. But he did not merely describe; he also pondered how species lived (Linnaeus 1811, I:13, 1971):

Oak trees grow on the summits of the surrounding rocks. At first it seems inconceivable how they should obtain nourishment; but the vapours are collected by the hills above, and trickle down in streams to their roots.

Linnaeus’s illustrations did not always represent what he discussed in most detail. For example, he made an excellent drawing of a crane fly but only recorded that he collected it at Umeå on 9 June.

Fig. 2. Crane fly (Pedicia rivosa). Linnaeus 1811, I:186, 1971.

Remarkably, he shot a hawk owl from his horse, “going on at a good rate” (Linnaeus 1811, I:204, 1971), at 12:15 am. Regrettably, it was too damaged by the shot to be stuffed, but at least he drew its picture.

Fig. 3. Hawk Owl (Surnia ulula). Linnaeus 1811, I:205, 1971.


For two insects collected at Lulea on 21 June, he provided both illustrations and discussion (Linnaeus 1811, I:233, 1971)

Fig. 4. Capricorn Beetle (Cerambyx sutor) and black fly (Culex equines). Linnaeus 1811, I: 232, 1971.

Fig. 5. Rhododendron lapponicum. Linnaeus 1811, I:301, 1971.


1. A large Capricorn Beetle, variegated with a lighter hue. (Cerambyn Sutor, the female.) The horns were longer than the body, black, consisting of ten joints, each joint ash-coloured at its base. Body black, rugged, its wing-cases besprinkled here and there with clustered dirty spots. Abdomen cylindrical, covered towards the thorax with beautiful red lice, (Acarus coleoptratorum).

2. A minute black fly, with a roundish body and white wings, (Culex equines). This infested the horses in infinite multitudes, running under the mane, and attacking them with great fierceness, being not easily driven off.

The scientific names of species he discussed or illustrated were added by the editor, James E. Smith, from Linnaeus’s Flora Lapponica (1737) and Fauna Svecica (1746). He also discussed at length a rhododendron he drew on 8 July in the Lapland Alps (Fig. 5), but only to describe it and to evaluate whether it belonged in the azalea genus (Linnaeus 1811, I:299–301, 1971).

On the Norwegian coast at Torfjorden he went fishing in a boat and caught with hook and line “plenty” of Sey-fish (Gadus virens), which he drew (Fig. 6). He found remora sticking to some of these 10-inch fish.

Fig. 6. Sey-fish (Gadus virens). Linnaeus 1811, I:342, 1971.


He also observed, collected, and drew four different kinds of medusa (jellyfish), but made no observations on their behavior or food (Linnaeus 1811, I:336–339, 1971). Two later Linnaean dissertations were on marine subjects: Noctiluca Marina (1752) on minute phosphorescent “insects,” and Natura Pelagi (1757) on fish, turtles, and cetaceans (Smit 1979:120–123).

Reindeer were important draft and milk animals to Laplanders, and Linnaeus discussed them repeatedly in Lachesis Lapponica (1811), but only illustrated bridle, harness, and antlers (Linnaeus 1811, I:103–110, 135, 1971). He illustrated reindeer themselves in the frontispiece of Flora Lapponica, 1737). Their antlers were beginning to sprout in June, initially covered by soft skin which was often bloody from mosquito bites.

Fig. 7. Frontispiece of Linnaeus, Flora Lapponica (1737), showing reindeer. The man seated in the foreground is Linnaeus. The steep mountains in the background do not represent the topography he found in Lapland.


Females have smaller antlers than males. Squirrels gnawed antlers from previous years (Linnaeus 1811, I:127–128, 1971). Linnaeus’ discussion of what reindeer eat is interesting (Linnaeus 1811, I:161–162, 1971):

The reindeer suffers great hardship in autumn, when, the snow being all melted away during summer, a sudden frost freezes the mountain Lichen (L. rangiferinus), which is his only winter food. When this fails, the animal has no other resource, for he never touches hay. His keepers fell the trees in order to supply him with the filamentous Lichens that clothe their branches; but this kind of food does not supply the place of what is natural to him. It is astonishing how he can get at his proper food through the deep snow that covers it, and by which it is protected from the severe frosts.

The reindeer feeds also on frogs, snakes, and even on the Lemming or Mountain Rat (Mus Lemmus), often pursuing the latter to so great a distance as not to find his way back again. This happened in several instances a few years ago, when these rats came down in immense numbers from the mountains.

But he also reported that they ate nothing in hot weather, when mosquitoes were very troublesome (Linnaeus 1811, I:308, 1971). Later, he commented that the Lapps were negligent not to gather Lichen rangiferinus and horsetail (Equisetum fluviatile) in summer for winter fodder (Linnaeus 1811, II:107–108, 1971). Females give birth in May and fawns grow simple antlers their first year (Linnaeus 1811, I:313, 1971). In warm weather reindeer are tormented by the bites of gadflies (Oestrus tarandi), which leave so many scars that one author mistakenly thought they were caused by smallpox. (For more details on this fly and reindeer, see Linnaeus 1739, 1746b; part of the latter is translated by Susan Novikoff in Usinger 1964:5–6.) One insect, “probably a species of Tabanus” (Linnaeus 1811, I:280–281, 1971) bores into reindeer and lays its eggs under the skin, and the young leave by the same hole. The Lapps squeeze out the larva from their pustules to lessen the reindeer’s pain. Another fly (Oestrus nasalis) lays eggs in reindeer nostrils (Linnaeus 1811, II:45, 1971). Reindeer also suffer from an epidemic disease that Laplanders called Pekke Kattiata that could be fatal (Linnaeus 1811, II:39–40, 1971). These observations were also included in a 1754 dissertation, Cervus Rheno, defended for a doctorate degree by Charles F. Hoffberg, and is translated into English (Linnaeus 1781:167–214, 1977).

On 17 July 1732, Linnaeus had a chance to see lemmings, which he described, and said they ate grass and reindeer moss. They lived mainly in the Scandinavian alps, but (Linnaeus 1811, II:19, 1971):

in some years thousands of them come down into the woodland countries, passing right over lakes, bogs, and marshes, by which great numbers perish. They are by no means timid, but look out, from their holes, at passengers, like a dog. They bring forth five or six at a birth. Their burrows are about half a quarter (of an ell?) deep.

(The parenthetic question about burrow depth was inserted by the book’s editor.) Later in the book, Linnaeus raised his estimate of their numbers from thousands to millions and admitted that “nobody knows what becomes of them” (1811, II:82–83). In a still later article (Linnaeus 1740; partly translated in Blunt 1971:60), he rejected the belief that lemmings fall from clouds.

Fig. 8. Linnaeus’ wedding portrait, 1739, by J. H. Scheffel, now at Hammarby, Linnaeus’ summer home, managed by Uppsala University. Color reproduction courtesy of Hunt Institute for Botanical Documentation.


Without publishing his travel journal, Linnaeus still publicized his achievements, and the governor of Dalecarlia province offered to fund a survey of that province. Linnaeus agreed, and seven medical students gained permission to come along at their own expense. They first traveled to Falun, the provincial capital (where Linnaeus met his future wife), and then departed on their expedition on 3 July 3 1734, taking along the governor’s two sons. Linnaeus was an organizing genius, and he delegated specialized tasks to each student: geography; climate and soils; stones, minerals and fossils; plants; animals; economics; and logistics. Every night each student added his report to whatever Linnaeus wrote. The last entries were dated 17 August, and when they returned to Falun, Linnaeus gave their Iter Dalecarlium to the governor. It was never published, but some account of the trip appeared in a Hamburg newspaper, and Linnaeus used some of their notes in later publications (Blunt 1971:76–79, Caddy 1886–1887, I:213–249).

Linnaeus did publish observations from subsequent field trips, and the book on his trip to Öland and Gotland in 1741 is also translated into English. The government (Swedish Estates of the Realm) asked him to make an economic survey, including natural history, of these islands. Accompanied by six young men, he departed from Stockholm on 15 May. It was quite cold, and Linnaeus suggested that “Spring should be measured according to climate and temperature rather than by the calendar”(Linnaeus 1973:23), and he then gave what we call phenological observations on the progress of the leaves and flowers or buds of several trees and herbs. Back in 1737 he had publicized a thermometer in the frontispiece to his Hortus Cliffortianus.

Fig. 9. Detail from frontispiece of Linnaeus, Hortus Cliffortianus (1737).


The thermometer was probably one he had obtained during three months spent in England, and he may have suggested to his friend Anders Celsius (1701–1744) that he reverse the scale he had developed, having boiling water at zero and freezing at 100 (Nordenmark 1935), because on 30 October 1758 Linnaeus wrote to a Montpellier botanist, Boissier de la Croix de Sauvages (English translation in Middleton 1966:100):

I was the first who decided to construct our thermometers in which the freezing point is 0, and the heat of boiling water 100; and this for the greenhouses of our garden.

Two subsequent dissertations were phenological: Vernatio Arborum (1753) and Calendarium Florae (1754), and are translated into English (Linnaeus 1775:133–158, 233–286, 1977).

When our explorers reached the copper smelter at Adelfors on 23 May 1741, Linnaeus noticed that the junipers looked like “trimmed cypresses”(Linnaeus 1973:34), which he attributed to smoke from the blast furnaces. Workers and residents at Adelfors complained about the air pollution. They reached Öland on 1 June, and Linnaeus made an inventory of its plants and animals. A gamekeeper told him the time of mating and the gestation periods of red and fallow deer, wild boar, and bear, which he recorded (Linnaeus 1973:48). He examined the nest of a Rook (Corvus frugilegus) containing three nestlings and numerous mites (Simulium reptans) bloated with nestlings’ blood. He counted annual rings of an oak stump and found it was 260 years old. Some rings were wider than others, which he thought was due to different severities of winters (Linnaeus 1973:58). Although modern botanists correlate annual ring width with summer moisture, this was a beginning of paleoclimatology. He knew that Francesco Redi had described 30 kinds of bird mites, which inspired Linnaeus (1973:69) to describe oystercatcher mites (Saemundssonia haematopi) and avocet mites (Vanellus vanellus). Along the seashore, he discovered that all plant species had succulent leaves, but that the majority of them growing elsewhere had ordinary dry leaves (Linnaeus 1973:72). Potentilla anserine grew on the sand and Senecio vulgaris on rotting seaweed. Cinnabar moths (Hipocrita jacobeae) were numerous on shore, and their larva ate the Senecio (Linnaeus 1973:86). He found that other plant species also had their own particular caterpillars, which he described and named, probably assisted by the entomologist Charles de Geer (Landin 1972), whom he visited at Medevi on 23 August (Linnaeus 1973:89, 199). Near the Lummelunda church he studied a marsh in which the sedge Cladium mariscus grew. This species had not previously been reported in Sweden; he emphasized the facts that cattle ate it in early spring and that it made good thatch for roofs. Since he learned that it grew in a former lake, he suggested that it be planted in Sweden’s many “sterile and useless bogs” that could not be drained (Linnaeus 1973:113). Beyond Stenkyrka, he found under stones in water a white oval leech (Hirundo [Nephelis] octoculata) that could also be found in the stomachs of small fishes, and he thought that the liver worms of sheep were probably the “spawn” of this leech, which the sheep swallowed when grazing in marshy places (Linnaeus 1973:118–119). After transcribing runic inscriptions in the Hangvar churchyard on 27 June, he commented that a white lichen (Kecabira cakcarea) grew on the limestone tombstones but not on granite ones (Linnaeus 1973: 119).

They reached Fårö Island, just north of Gotland and much smaller (see map, Linnaeus 1973:facing page 109), on 28 June. Its inhabitants hunted seals but not porpoises. They also ate eider and their eggs, but Linnaeus thought that “The time will probably come when the excellent down of these birds will save them from being shot” (Linnaeus 1973:126), but he did not explain how to collect it (possibly from their nests). He described in some detail the growth of “sandhafre” (Ammophila arenaria) on the sand dunes, and explained how it stabilized the dunes. He also found ant lions on the dunes that were “far more multicoulored than on Öland” (Linnaeus 1973:130). He referred the reader to Réaumur’s memoir on ant lions for details.

Fig. 10. Linnaeus in 1747 by Jean E. Rehn.


Five years later, from 12 June to 11 August 1746, Linnaeus traveled through West Gothland and published his findings in Wästgöte-Resa (1747). Caddy (1886–1887, II:165–206) summarized this book, turning it into a Linnaean travelogue (she followed his route). Among the translated extracts quoted by Blunt (1971:163) is this generalization:

when animals die they are converted into mould, the mould into plants. The plants are eaten by animals, thus forming the animals’ limbs, so that the earth, transmuted into seed, then enters man’s body as seed and is changed there by man’s nature into flesh, bones, nerves, etc.; and when after death the body decomposes, the natural forces decay and man again becomes that earth from which he was taken.

These thoughts were not especially original (Isaiah 40:6 “All flesh is grass.”), but they are of interest as a prelude to the 1749 dissertation on the economy of nature.

Meanwhile, in 1744, the dissertation Oratio de telluris habitabilis incremento (On the increase of the habitable earth), defended by Johann Westmann, offered a novel geological theory (Frängsmyr 1983) and explanation of how the world had become populated with species (Linnaeus 1781:71–127, 1977b): (1) God created one pair of each sexual species and one individual of each hermaphroditic species; (2) since Adam named all species, the Garden of Eden must have been a mountain island; (3) each species increased in numbers every generation; (4) as they increased, they enlarged the geographical area they inhabited; and (5) the habitable land increased as the numbers of organisms increased. To support this argument, Linnaeus had to demonstrate the potential of all species to increase their populations. He listed the numbers of seeds reported for different flowers: Helenium 3000, Zea 2000, Helianthus 4000, Papaver 3200, and Nicotina 40,320. He then calculated correctly that an annual plant that only produced two seeds per year, if preserved from animals and accidents, would have 1,048,576 descendants in 20 years. That dissertation was only one of several publications that entitle Linnaeus to be called the founder of plant geography (Hofsten 1916:243–247, Browne 1983:16–23). Du Rietz (1957a) summarized his contributions to alpine phytogeography, paludology (on which see also Du Rietz 1957b), indicator plants, plant succession, limnology, and forest geography.

A fundamental difference between Linnaeus’ conception of an ecological science and ours is that in his, biotic interrelationships were designed by God to work harmoniously and permanently and for the benefit of humanity (Hofsten 1957:90–102), whereas in ours, interrelationships evolve and can lead to extinction of species. His conception was part of a general outlook in science: for example, in astronomy, celestial bodies were unchanging in substance and orbits; and in geology, ongoing changes in the landscape were considered minor compared to the changes caused by God in the Flood of Noah. Scientists’ study of a “static” universe gradually revealed that it is not static. This even happened to Linnaeus. In his Systema Naturae (1735, 1964) he confidently claimed that all species had been created by God at the beginning and no new ones had since appeared. However, the discovery of Peloria in 1741—so similar to Linaria, yet an apparently different species—shook his belief in the constancy of species. He eventually suspected that God had created only a few species, which later hybridized to form the great variety now seen (Hagberg 1952:196–205, Hofsten 1957:65–86, Larson 1971:94–121, Bowler 1989:64–68). On 18 August 1764, he explained this idea in a letter to Johannes Burmann (in Nicolas 1963:53).

Let us suppose God made a Ranunculus [and that] this species is crossed with a Helleborus, and Aquilegia, or a Nigella in hybrid generations. Through Divine Law the descendants of these hybrids will have, as in animals, the mother’s medulla and father’s cortex. As a result, there are so many of Ranuncula with either aquilegous leaves or nigellous ones that you could not separate them into arbitrary genera…

Linnaeus’ term “oeconomia naturae” (1749) is rather similar to the contemporary term for animal physiology, “animal economy,” which involved studying how the parts contributed to the functioning of the whole. He may have implied an analogy between organs in an animal and species in a biotic community (Linnaeus 1775:39, 1977a):

By the Oeconomy of Nature we understand the all-wise disposition of the Creator in relation to natural things, by which they are fitted to produce general ends, and reciprocal uses.

Having a passion for system, Linnaeus approached the economy of nature systematically. For each of the three kingdoms—stones (and soils), plants, and animals—he discussed a cycle of propagation, preservation, and destruction.

Surveying different kinds of stones under “Propagation,” he suggested that one or more kinds had organic origins (Linnaeus 1775:51, 1977):

…testaceous bodies and petrifactions resembling plants were once real animals or vegetables; and it seems likely that shells being of a calcareous nature have changed the adjacent clay, sand, or mould into the same kind of substance. Hence we may be certain, that marble may be generated from petrifactions, and therefore it is frequently seen full of them.

Under “Preservation,” he speculated, inaccurately, on how stones are generated and augmented by water, but under “Destruction” he was more accurate in describing the actions of weather and water in the gradual erosion of rocks. He also noted that certain animals also helped erode some kinds (Linnaeus 1775:57, 1977):

[Testaceous worms]…eat away the hardest rocks. That species of shell fish called the razor shell bores thro’ stones in Italy, and hides itself within them; so that the people who eat them are obliged to break the stones, before they can come at them. The cochlea F[auna] S[vecica number] 1299. a kind of snail that lives on craggy rocks, eats, and bores through the chalky hills, as worms do through wood. This is made evident by the observations of the celebrated de Geer.

God allegedly designed living beings to both survive and regulate each other (Linnaeus 1775:40, 1977):

…all living creatures should constantly be employed in producing individuals; that all natural things should contribute and lend a helping hand to preserve every species; and lastly, that the death and destruction of one thing should always be subservient to the restitution of another.

This explicit statement was an important contribution to the balance of nature concept, though Linnaeus did not name it (Egerton 1973:335–337).

Under plant propagation, he discussed sexual reproduction, then seed dissemination (Linnaeus 1775:64–65, 1977):

Berries and other pericarps, are by nature allotted for aliment to animals, but with this condition, that while they eat the pulp they shall sow their seeds; for when they feed upon it they either disperse them at the same time, or, if they swallow them, they are returned with interest; for they always come out unhurt. It is not therefore surprising, that if a field be manured with recent mud or dung not quite rotten, various other plants, injurious to the farmer, should come up along with the grain, that is sowed.

Under “Preservation,” he claimed that God had decreed (Linnaeus 1775:67–68, 1977): “that the whole earth should be covered with plants, and that no place should be void, none barren.” He had heard of deserts but had never seen one, so he confidently asserted that they have their own unique trees and herbs (which they do, but there is still bare ground). All environments—alpine, grassland, marshes, aquatic, deserts—have characteristic species, and he discussed examples. The graesmasken moth inhibits the spread of grass, leaving room for other plants. However, plants die, and their destruction is also part of God’s plan. Black mould, which nourishes new plants, comes from dead plants, and that cycle really begins with the liverworts that grow on bare rocks; when they die, they leave mould for mosses, and as mosses die, they leave mould for herbs and shrubs. This dissertation contains one of the earliest descriptions of plant succession (which Clements, 1916:10, credited to Biberg, the defendant). Insects contribute to the death of plants by eating parts, which make them vulnerable to other hazards (Linnaeus 1775:76–80, 1977).

Under animal propagation, Linnaeus surveyed all the known reproductive habits of different species, and although he rejected spontaneous generation, he admitted that (Linnaeus 1775:89),

The laws of generation of worms are still very obscure, as we find they are sometimes produced by eggs, sometimes by offsets, just in the same manner as happens to trees.

He pointed out that smaller animals tend to produce more offspring than larger ones: mites can increase to a thousand in a few days, but elephants only produce one offspring in two years. However, some hawks are smaller than the poultry they eat, and he acknowledged that hawks layer fewer eggs, without attempting to explain why. He calculated that two pigeons breeding nine times a year could produce 14,672 young in four years, but his translator pointed out that Linnaeus had mistakenly added in the original pair to reach this figure (Linnaeus 1775:90, 1977); however, the numbers 6 and 7 were accidentally transposed in the English edition; Linnaeus’ figure should have been 14,760 (Egerton 1967:174). In Politia Naturae (Latin, 1760, cited from the English translation, 1781:162, 1977b), he added that long-lived animals propagate slowly.

Under animal preservation, he discussed which species care for their young and which do not. Among polygamous species, “males scarcely take any care of the young” (Linnaeus 1775:93, 1977a), and cuckoos lay their eggs in the nests of wagtails and hedge-sparrows. Because of the great diversity of species, God assigned each one certain places to live and certain foods to eat. Linnaeus gave a brief survey of examples from the animal kingdom, but only provided details concerning the mutualism between the bivalve, Pinna, and the crab, Pinnotheres (Linnaeus 1775:111–113, 1977a). This relationship had been reported by Aristotle (Historia Animalium 547b16–17), but had been neglected by more modern naturalists until Linnaeus’ disciple, Fredrik Hasselqvist (1722–1752), traveled to the eastern Mediterranean (where he died) and confirmed it. Since this Oeconomia naturae dissertation was published in March 1749, and Hasselqvist did not leave Stockholm until 7 August 1749 (Blunt 1971:183–185), Linnaeus obviously added these comments on Pinna and Pinnotheres before the dissertation was republished in Amoenitates Academica, volume 2 (1751), the source of Benjamin Stillingfleet’s English translation.

Linnaeus’ survey of the destruction of animals included two food chains, one terrestrial and one aquatic (Linnaeus 1775:114, 1977a):

the tree-louse lives upon plants. The fly called musca aphidivora lives upon the tree-louse. The hornet and wasp fly upon the musca aphidivora. The dragon fly upon the hornet and wasp fly. The spider on the dragon fly. The small birds on the spider. And lastly, the hawk kind on the small birds.

In like manner the monoculus delights in putrid water, the knat eats the monoculus, the frog eats the knat, the pike eats the frog, the sea calf eats the pike.

Next, he emphasized the importance of predators to prevent their prey from over-running everything, and the importance of scavengers to prevent the earth from being overwhelmed with carcasses (Linnaeus 1775:114–122, 1977a).

In 1734, while exploring Dalecarlia, Linnaeus had watched his expedition’s horses grazing certain plants and avoiding others. Both John Ray and René Réaumur had reported insects having very specific food plants (Egerton 2005:303 and 2006: ), but in the late 1740s Linnaeus and some students (eight named, plus others) ran 2314 experiments on livestock to determine their plant preferences. Their findings were reported in a dissertation entitled Pan Svecius (Latin 1749, cited from Stillingfleet translation: Linnaeus 1775:361, 1977a):

Oxen   eat
218   plants

This was one of the earliest, if not the earliest, series of experiments on an ecological question, and surely the earliest such large-scale quantitative experiments. (Stillingfleet’s translation did not include all details in the original; more is translated in Ramsbottom 1959:162–167.) The reason for so many experiments was that, unlike the insects observed by Ray and Réaumur, these mammals were not limited to eating one or two species, but nevertheless were somewhat selective. Allegedly, God’s reason to make various animal species eat different plant species was to prevent some plant species from becoming extinct due to overeating and others from becoming too abundant because they were not eaten (Linnaeus 1775:347–349, 1977a). Also, in “Oeconomy of Nature”(1775:99–100, 1977a) Linnaeus mentioned “an oeconomical experiment well known to the Dutch,” of which he perhaps learned while he was in the Netherlands in 1735,

that when eight cows have been in a pasture, and can no longer get nourishment, two horses will do very well there for some days, and when nothing is left for the horses, four sheep will live upon it.

In 1774 a dissertation comparable to Pan Svecius appeared on the subject of plants and animals eaten by chickens, ducks, and geese, Esca Avium Domesticarum (Smit 1979:122).

Linnaeus’ second most important dissertation for ecology is Politia Naturae (1760), translated into English as ”On the Police of Nature” (1781:129–166). It is on the struggle and survival of species, including humans. A pessimistic conclusion that he drew about humans seemed also to apply to some extent to plant and animal species. Unfortunately, F. J. Brand, the English translator of this dissertation, omitted it. Fortunately, Alan Blair translated it in Kurt Hagberg’s biography of Linnaeus (1952:183).

…where the population increases too much, concord and the necessities of life decrease, and envy and malignancy towards neighbours abound. Thus it is a war of all against all!

The point of the dissertation was to explain why a war of all against all (competition) did not lead to extinction. A major reason was what we call ecological diversity: each species is confined to its own “station” (habitat). Sweden had about 1300 plant species, but only about 50–100 are in any one place (Linnaeus 1781:133, 1977b). Linnaeus argued that although it is received opinion that plants were created for the use of animals, actually, animals were created to regulate plants’ abundance. As proof, he cited numerous insect species that only eat a single plant species, doves eat surplus seeds, and other birds, bats, and anteaters eat insects to prevent them from consuming all of the plants they eat, and so on.

Linnaeus had a lifelong fascination with insects. Five of the dissertations translated by Brand were on insects and their interactions with other species of plants and animals (Linnaeus 1781:309–456, 1977b). Smit (1979:125) even claimed that Linnaeus made a major contribution to entomology, as evidenced by his Fundamenta Entomologiae (1767). If so, that dissertation was important because it synthesized briefly the works of others. Linnaeus praised as “immortal” the treatise by Réaumur, whom he had met in Paris, and he once sent Réaumur the eggs of the alpicola butterfly (Papilio Apollo). Réaumur’s thank-you note is published in Linnaeus’ correspondence (Smith 1821, II:477–479). However, Linnaeus also pointed out the necessity of providing official names for the species Réaumur had studied (Linnaeus 1772:13). Despite Linnaeus’ strong interest in both insects and plants, he never fully appreciated the role insects play in pollination, except for fig trees. He first named the nectary of flowers in 1735, and he did move from a belief that bees harm flowers by collecting nectar, to a belief that they help pollinate flowers, but never realized their crucial importance for flowers that are not wind pollinated (Miall 1912:322–324, Usinger 1964:6, Lorch 1978:518, 523, Eriksson 1983:105). He did, however, appreciate the danger of accidentally introducing American insect pests when American plants were brought to Europe. In 1739 he confessed to having brought American trees from England to the Netherlands in August 1736; these harbored aphids, which multiplied in a greenhouse and then escaped into botanical gardens in Amsterdam and Leiden (1781:325, 1977b). Pehr Kalm brought pea seeds when he returned to Sweden in 1751; he discovered that they contained live Dermestes pisorum insects, which he captured. However, Linnaeus warned against the danger of artificial introductions (Linnaeus 1781:386–387, 1977).

Linnaeus, physician and sometime professor of medicine, followed Richard Bradley (Egerton 2006:124–125) in arguing that minute organisms, “even smaller than the motes dancing in a beam of light”(Linnaeus quoted in translation from Smit 1979:123) transmit contagious diseases. He developed his argument in two dissertations, Exanthemata viva (1757) and Mundus invisibilis (1767). He also discussed and described parasitic worms. His speculations (1973:118–119) about some free-living worms being a different stage of internal parasitic worms was a reasonable hypothesis, but he supported it with unverified (and incorrect) examples (Foster 1965:32, Grove 1990:4, 40, 106, 386).

Linnaeus clearly deserves a prominent place among the founders of ecology, but was this a case like Mendel’s, in which his findings were only appreciated after others had rediscovered his basic ideas? No—many of his writings and the student dissertations were reprinted and translated into other languages, and contemporary and later naturalists, including Gilbert White and Charles Darwin, read them (Stauffer 1960, Limoges 1980).

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Linnaeus, C. 1737b. Hortus cliffortianus: plantas exhibens quas in hortistam vivis quam siccis, Hartecampi in Hollandia, coluit virnobillismus & generosissimus Georgius Clifford. Amsterdam, The Netherlands.

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Stafleu, F. A. 1971. Linnaeus and the Linnaeans: the spreading of their ideas in systematic botany. Oosthoek, Utrecht, The Netherlands.

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For their assistance I thank Jean-Marc Drouin, Muséum National d’Histoire Naturelle, Paris; and Anne Marie Drouin-Hans, Université de Bourgogne.

Frank N. Egerton
University of Wisconsin-Parkside
Kenosha WI 53141
E-mail: [email protected]


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Public Affairs Perspective

Ecological Society of America
Rapid Response Team Update

Serving Society Through Ecological Expertise                                                            Autumn 2006

* At ESA's Annual Meeting in Memphis, TN...

ESA's bustling Annual Meeting Press Office

Media Activities... ESA's Public Affairs Committee and RRT
members led a well-attended and productive workshop on communicating ecology with metaphors, anecdotes, and analogies. RRT members also were interviewed by attending media about invasive species in grasslands, and the effects of changing nitrogen cycles on human health.

Congressional Staff ... ESA often hosts one or two congressional staff members at the Annual Meeting, enhancing the communication of science and establishing personal networks between scientists and policy-makers. The congressional staff member hosted in Memphis met with Agroecology RRT members as well as RRT member and Vice-President for Public Affairs Rich Pouyat.

RRT Luncheon ... ESA's second annual luncheon for RRT members highlighted the accomplishments of the RRTs over the past two years and featured a working discussion led by outgoing ESA President nancy Grimm on the future RRT structure and functions.


* Looking Ahead... ESA's Public Affairs Office is planning the next round of RRT activities, and working on enhancing and updating ESA's Expert Guide and web site. Stay tuned—we will be contacting you for your feedback and updated biographical information. Please send ideas and suggestions for future RRT opportunities and activities to Laura Lipps at [email protected].
Congressional staff members take measurements in a tidal wetland

* Providing Input to Congress...

RRT Team Member Dennis Whigham co-led a group of congressional staff on a wetlands field trip, along with colleagues and ESA members Pat Megonigal and Tom Jordan. The field study formed part of an educational course for congressional staff on the fundamentals of wetland science, sponsored by the ESA, the Society of Wetland Scientists, and the American Society for Limnology and Oceanography.

* Providing Input to the Media ...

As the 2006 hurricane season got underway, reporters called ESA's Press Office to get in touch with RRT members with expertise in hurricanes and ecosystems. If you have been in the news lately, we encourage you to submit stories to [email protected], to be posted on ESA's web site.

* Supporting Science at Federal Agencies ...

Providing Comments on Rulemaking... ESA's President, with the assistance of Aquatic Ecology RRT members, submitted comments to the Environmental Protection Agency on its proposed rule on stream mitigation. The comments are available at:

Supporting Funding for Science ... RRT members Alan Townsend joined 80 other scientists to discuss science funding with Members of Congress, as an ESA representative for the Coalition for National Science Funding, ESA's Public Affairs Office brings RRT members, students, and scientists from all career stages to D.C. for science policy training and targeted meetings with Congressional offices.

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Public Affairs Perspective

ESA Statement on Massachusetts et al. vs. U.S. Environmental Protection Agency,
“The Certainty of Global Climate Change”

In response to the U.S. Environmental Protection Agency’s (EPA) argument in the U.S. Supreme Court case, Massachusetts et al. vs. U.S. EPA et al. that it would not regulate carbon dioxide due in part to scientific uncertainty, Alan Covich, President of the Ecological Society of America, the nation’s premier scientific society of ecological scientists, expressed dismay over the agency’s disregard for the widespread scientific consensus on the facts and effects of climate change. Covich said, “ESA strongly supports the scientific arguments regarding the certainty of climate change made by Massachusetts et al. as legitimate and accurate.”

The case currently before the Supreme Court was brought by a group of states, cities, and nonprofits, which argued that, under the Clean Air Act (CAA), carbon dioxide is a pollutant and contributor to climate change, and as such, the EPA must regulate carbon dioxide emissions. EPA argues, however, that it does not have the authority to regulate carbon dioxide and even if it had, the agency would not regulate carbon dioxide due in part to scientific uncertainty. The court’s decision will determine whether the CAA mandates that EPA regulate carbon dioxide, which may be a watershed decision for U.S. action regarding climate change.

The CAA states that absolute scientific certainty is not necessary for regulation of a pollutant. Instead, the pollutant must “reasonably be anticipated to endanger public health or welfare.” [Section 202(a)(1)] In arguing that carbon dioxide cannot reasonably be anticipated to endanger public health or welfare due to scientific uncertainty, EPA chose only the points highlighting the absence of certainty from the 2001 National Research Council Report on climate change, but disregarded a general scientific consensus that climate change is indeed occurring. While certain facets of climate are less certain than others, current scientific understanding verifies and supports the following statements:

  • The physics of the greenhouse effect are well understood by scientists. Certain atmospheric gases (including carbon dioxide) absorb radiation, causing a warming effect, and that greater concentrations of these gases will lead to an increase in surface temperatures ( Energy Information Administration 2004).
  • It is certain that human activities have increased the concentrations of carbon dioxide in the atmosphere above levels in the geologically recent historical record, which has led to the recent global climate change. Carbon dioxide levels remained constant from the period 1000–1750, then increased more than 25% in the last 150 years ( International Panel on Climate Change 2001, Energy Information Administration 2004). This had caused average global surface temperatures to rise approximately 0.6ºF; global average sea level to rise 0.1–0.2 meters; and ice extent in the Arctic to decrease by about 40% ( National Assessment Synthesis Team, U.S. Global Change Research Program 2000; International Panel on Climate Change 2001).
  • It is virtually certain that greenhouse gas emissions, continued at the current rate, will cause additional warming. At the current emissions rate, carbon dioxide levels will double by 2100, leading to a likely temperature increase of 2.7º–10.4ºF ( National Assessment Synthesis Team, U.S. Global Change Research Program 2000; International Panel on Climate Change 2001).
  • Climate change will increase threats to human health, especially in developing countries and lower income populations. Further increases in surface and ocean temperatures will cause an increase in drought, flood, and hurricane intensities, disrupt agricultural production, and expand the range of disease vectors ( National Assessment Synthesis Team, U.S. Global Change Research Program 2000; International Panel on Climate Change 2001).
  • The ecological effects of climate change are already observable and have been well-documented. Ecosystem disruptions including fire, drought, invasion of species, habitat shifts and coral bleaching events result in reduced biodiversity, increased extinction risk, and loss of ecosystem services ( National Assessment Synthesis Team, U.S. Global Change Research Program 2000; International Panel on Climate Change 2001).
  • The exact timing and magnitude of climate change depend on climate inertia and future emission rates. The elevated greenhouse gases currently in the atmosphere will continue to change the climate for an unknown period, even if all emissions are capped immediately. However, capping carbon dioxide emissions will significantly increase the timing of climate stabilization ( Committee on the Science of Climate Change, National Research Council 2001; International Panel on Climate Change 2001).
  • It is likely that significant, intense, and abrupt events caused by climate change (such as local heat waves, intense droughts, or floods) will occur. The absence of a complete understanding of certain global processes may prevent predictions of events, including thermohaline circulation reduction, significant losses to Antarctic ice sheet, and altered Asian monsoons ( National Assessment Synthesis Team, U.S. Global Change Research Program 2000; International Panel on Climate Change 2001).

Although current scientific understanding, by definition, constantly changes, these statements represent the latest information and are supported by a wide group of climate scientists. Many other scientific bodies, including the American Geophysical Union (AGU), the Geological Society of America (GSA), and the joint National Academies of Science (NAS), have issued similar statements:

AGU: ‹http://www.agu.org/sci_soc/policy/positions/climate_change.shtml

GSA: ‹http://www.geosociety.org/aboutus/position10.htm

NAS: ‹http://nationalacademies.org/onpi/06072005.pdf

Said Covich, “Communication and widespread awareness of climate change information remains essential not only to conservation of natural resources, but also to the general well-being of the nation. The recognition of these facts by the Court in Massachusetts et al. vs. U.S. EPA et al. will provide significant support to the certainty of climate change that the administration cannot ignore.”

Literature cited

National Assessment Synthesis Team, U.S. Global Change Research Program. 2000. Climate change impacts on the United States. 18 October 2006. ‹http://www.usgcrp.gov/usgcrp/Library/nationalassessment/overviewfindings.htm

Committee on the Science of Climate Change, National Research Council. 2001. Climate change science: an analysis of some key questions. 18 October 2006. ‹http://newton.nap.edu/execsumm_pdf/10139.pdf

International Panel on Climate Change. 2001. Climate Change 2001: Synthesis Report. 16 October 2006. ‹http://www.grida.no/climate/ipcc_tar/vol4/english/index.htm

Energy Information Administration. 2004. Greenhouse gases, climate change, and energy. http://www.eia.doe.gov/oiaf/1605/ggccebro/chapter1.html

Contact: Nadine Lymn (202) 833-8773 x205; [email protected]

or Annie Drinkard (202) 833-8773 x211; [email protected]

Public Affairs Office
1707 H Street NW, Suite 400
Washington, DC 20006
Web: ‹www.esa.org/pao

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Note: Dr. Harold Ornes is the editor of Ecology 101. Anyone wishing to contribute articles or reviews to this section should contact him at the Office of the Dean, College of Science, Southern Utah University, 351 W. Center, Cedar City, UT 84720; (435) 586-7921; Fax: (435) 865-8550; E-mail: [email protected].

Whether you are teaching undergraduate or graduate-level field ecology, or you are a full-time researcher and in the midst of gleaning the maximum amount of information from your most recent field research sampling trip, I think you will agree that we almost never remember everything needed in the field, and likewise almost never have a perfectly smooth and uneventful field trip or season. Thanks to Carmen Wong, University of British Columbia, we can take a few moments, sit around the fire, feel her pain, and start planning for next season.

I would encourage you to share one or two of your field season horror stories with your peers through this column.

Preparing for the Field Season

I thought I was entering the field season for my Ph.D. research in good shape. I had over 5 years of experience as a consultant working on various field research projects. And, I had spent time in the field during my Master’s research ziplocking grizzly bear feces in the Yukon. So, as a seasoned field biologist embarking on my Ph.D. research, I finally had my research permit in place, lined up two great field assistants, and purchased several boxes of waterproof paper and printed colored maps of my potential study sites. But, I should have known that you can never be too prepared for fieldwork.

My first challenge was losing a field assistant on the first day. I didn’t actually lose him in the forest; rather, he took a permanent job with a consulting company. My promise of helicopter rides in the Rockies could not compare to a decent salary with health benefits. I spent the next month looking for another field assistant. It turns out the good candidates get snapped up early.

My second challenge was with my sampling design. I had wanted to use stratified random selection of stands containing whitebark pine in order to describe dynamics across large landscapes. Random selection using a vegetation inventory and GIS had worked in a pilot study the year before in another park. Using my beautiful colored maps, my remaining field assistant and I spent a week bushwhacking to mountaintops, only to find no whitebark pine where it was suppose be. It seems that I was mistaken in my assumption that I was using a vegetation inventory. An inventory did not exist and I was actually using a coarse-grain classification. Polygons indicated only a potential for containing whitebark pine … and clearly that potential was low. It turns out that having the home number of a biometrician is handy when one is revising a sampling design in the field.

My third challenge was in vexing the park wardens. We left the door of a warden cabin in the backcountry unlocked. We almost lost the privilege of using these cabins, which would have made my research impossible, but luckily the supervisory warden was understanding. It was clear, however, that a moment of carelessness can jeopardize one’s research.

So what should you walk away with after reading the above tales? Here are a few lessons I learned this summer that may help you during your next field project:

  1. Apply early for your research permits. I knew of a Masters’ student who spent the entire summer waiting for his research permit and was not able to collect any data. As an example, it takes over 6 months to go through the application process to work in the provincial parks of Alberta and British Columbia in Canada. If you want to handle wildlife or do destructive sampling, count on much longer.
  2. Make contacts with the relevant organizations . It is very likely that people are interested in your research and can contribute expertise. This summer numerous people in Parks Canada helped locate whitebark pine stands and gave ideas on how to apply my research results to management recommendations. Organizations such as protected area agencies or forestry companies are also often able to offer significant in-kind support. By contacting the right people, I was able to tag along on the survey flights of the fire crew; these flights saved us days of hiking and innumerable dollars.
  3. Understand your sampling scheme. A pilot study is critical. Pilot data can indicate the amount of variability you can expect and thus guide you on your sampling intensity. If you can’t do a pilot study, then I recommend at least trying a couple of days in the field with your supervisor. Even the best methods devised in the office will be modified in the field due to some unforeseen variable. If your supervisor can’t come out, work together through different scenarios of what could affect your sampling plan and derive alternate methods. If possible, having the home phone number of your supervisor could prove useful. Both my supervisor and the biometrician on my committee were indispensable in the first 2 weeks of my field season. Decisions about what is done in the field take only a second to make. However, the statistical consequences of that decision could be untenable down the road.
  4. Be safe . After working in the private sector and returning back to the university environment, I was appalled by how little attention most supervisors give to safety in field. There are crazy true stories of graduate students working in the backcountry with chainsaws with no means of communication and a 30-km hike to the nearest road. The most dangerous situation my field crew and I faced this summer was a bout of food poisoning. Normally, this would not have been a big deal, except we were 26 km from the nearest road and the day we hiked out was one of the hottest days of the summer. Take your responsibility as a crew leader seriously. At minimum, this means having safety protocol discussions with your field crew, regular check-ins at the end of the day, a regulation first aid kit (an Epi-pen would not be excessive), first aid training, and either a radio or a satellite phone with the relevant emergency contact numbers. If your supervisor is as smart as mine, they should provide these to you without question. Furthermore, educate yourself on the guidelines to be followed in order for compensation to be paid for injuries on the job.
  5. Treat your field assistants well. Pay your field assistants well, which means paying them at least or above the average wage for field assistants. If you cannot afford this, then try to compensate with providing room and board. Feed your field assistants well because food makes people happy. Budget at least $10/person per day and do not complain if they want to look for fresh basil in the middle of small-town Alberta. Most important, be flexible to their needs, within reason, of course. Often fieldwork requires people to work long hours and weekends on a project they probably do not love as much as you do. The least you can do is to give them flexibility in choosing their days off or ending the day early so that they can run errands, visit the bank or post office.
  6. Treat yourself well and remember your loved ones . Take time off for yourself, plan a debriefing with friends or others in your laboratory halfway during your field season, and realize that your friends and family are more important than another sample point. I lost a boyfriend this way after I spent two summers away from him.
  7. Prepare for the worst . I know this sounds pessimistic but I think it makes you prepared. Assume that everything is going to break, that it will rain … hell, that it will snow every day. That way your field season can only be better than expected.

Finally, get out there and enjoy yourself. If you aren’t enjoying what you are doing, why are you doing it?

Carmen Wong
Tree Ring Laboratory, Department of Geography
University of British Columbia
Vancouver, BC Canada
(604) 224-5292
E-mail: [email protected]

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Since the Bulletin has gone electronic, the hope is that our journal will fill the niche between the science papers of Ecology and Ecological Applications and the science/policy of Frontiers in Ecology and the Environment.

One area that has been suggested by readers is essays on papers that particularly influenced the ideas and approaches to ecology of ecologists who have contributed to that particular subdiscipline of ecology. The idea is that the essay will both identify a paper or papers that were particularly important in catalyzing future developments, at least to the writer of the essay, and will also be a personal viewpoint. The working title for this section is: “The Paper Trail.”

The first in this new series is Steward T. A. Pickett’s essay on W. S. Cooper’s classic paper on plant succession. The original paper can be viewed at: “ The fundamentals of vegetational change, ” William S. Cooper, Ecology 7(4):391–413, October 1926), Stable URL: ‹http://links.jstor.org/

The Paper Trail: W. S. Cooper’s “Fundamentals of Vegetational Change” and a Fluent Mode of Thought for Ecology

It is a bit amazing to admit that a paper that influenced me is now 80 years old! I encountered Cooper (1926) first in graduate school, but I have revisited it at intervals, and have been repeatedly stimulated and refreshed by it. In spite of the fact that I originally consulted this paper for its insights into plant succession, it is important not so much for what is says about succession, but for the analytical approach it takes. I believe the approach is one that can benefit ecologists in any specialty and at any age.

First a little background is in order. William Skinner Cooper, a professor of Botany at the University of Minnesota, was one of the most important of the first generation of American ecologists (McIntosh 1985). If, by chance, you are an American plant ecologist, you may well trace your lineage back to him (Sprugel 1980). He was the major professor of a number of active members of the second generation, who in turn trained many academic grand- and great-grandchildren. Henry J. Oosting, Murray F. Buell, and Rexford Daubenmire were especially prolific members of the second generation. This legacy is certainly a noteworthy role in the field. But when one looks at the big debate of the day—the nature of community dynamics—Cooper is conspicuous in not holding a dogmatic view about succession, or in unfailingly supporting either Clements or Gleason, whose differences have come to symbolize the debate. Instead, Cooper was guided by careful observation of communities in the field, and was remarkably free of ideology. The paper under consideration perhaps explains his open-mindedness and ability to see beyond ideology and ossified positions. Here is what “The fundamentals of vegetational change” (Cooper 1926) did for me. 1

Cooper opens with a goal of examining and constructively criticizing the basics of his discipline: “A periodic inspection of foundations is most desirable for any edifice, and particularly so when the superstructure is being continually added to, as in the development of scientific knowledge.” (Cooper 1926:391). This is challenging stuff, because at that time it had been a mere 10 years since the publication of Clements’ (1916) big book on succession, with its proliferation of terminology, and exclusion of special cases, that promoted the questionable “organismal” concept of the plant community. Of course, the assumption that communities or ecosystems are equivalent to organisms in their development and stabilization has long since been rejected (Botkin and Sobel 1975). At the time, however, it was a persuasive and widely held idea. Cooper is telling us, whatever the orthodoxy, to go back to basics, and see what should stay and what should be rejected. This is the deepest lesson of his paper, which invites each generation of ecologists to reexamine the foundations of their discipline.

What should a rigid and dogmatic classification of succession, or for that matter, any ecological phenomenon, be replaced with? Oddly enough, although the study of succession and community change was called “dynamic ecology” at the time, Cooper noted that there was a proliferation of static classification systems of succession, each type having a narrow set of specific causes associated with it. He was concerned that the rigidity of classification systems would force ecologists to put actual communities in boxes that were not appropriate, and at any rate, that the classifications did not really account for change. He said, “It is our task to give verbal expression to constantly changing phenomena in a way that will parallel their mutations as closely as humanly possible.” (Cooper 1926:396). This was a call for flexible concepts and terms that reflected the flexibility—or perhaps in today’s terms, complexity—of the world to which the concepts referred. Such flexible terms would emerge from the actual process itself, rather than emerging from an idealization of it. This meant that “We must, accordingly, rigorously exclude all stock ecological terms and phrases, and treat the phenomena in a purely descriptive manner.” (Cooper 1926:396). So clear articulation of what the core phenomenon is, separate from the assumptions about it that yield idealized patterns or processes, becomes a key intellectual activity in ecology.

The specific features that Cooper identified as fundamental in succession are less important for the story I am telling than the approach he used. However, his summary of the fundamentals proved helpful to me in sorting through the problems of succession that were becoming clear in the 1970s, when I first encountered his paper (Box 1). For example, he took as fundamental that change in vegetation is universal, and included both contemporary and paleoecological changes. Change in vegetation could be due to any cause, not just ones that were expected to make succession move “forward.” Cooper, furthermore, included causes that arose from the characteristics and interactions of plants themselves, as well as those that were external to plants. Such things as soil differences in a biome, or fire, or people were all acceptable causes to Cooper, although Clements and his adherents viewed them as exceptions to be excluded from the study of succession. He took the interaction of plants and environmental causes (including animal activities) as a system. His recognition that different causes acted on various scales seems quite prescient of contemporary interest in scale. Cooper also indicated that all causes are always acting on vegetation change, though to different degrees. It was the complex interaction of a broad array of causes that resulted in vegetation change.

A sound conception of the fundamentals of dynamic ecology must be based upon the premise of the universality of change. The field necessarily includes all vegetation of all time, and all types of vegetational change.

The vegetation of the earth is presented as a flowing braided stream. Its constituent elements branch and interweave, disappear and reappear. Vegetation as we see it today is the advancing front of this stream, in which we discern more or less definite groupings that tend to repeat themselves in space, the advancing termini of the streamlet-elements. There is a constant tendency toward merging of related elements and thus toward simplification into fewer and larger currents, which is balanced by a trend toward multiplication of the streamlet-elements through forking.

The vegetational stream is governed and directed by the interaction of factors residing in the constituent organisms and their environment. These differ greatly in the scale on which they work and in the speed of the changes they induce. Every one is forever operating upon all portions of the stream. The net change in any given portion is determined by the resultant of their activities.

Box 1. Verbatim Summary of Cooper’s Analysis of Succession Concepts. Each major section of the paper ended with a summary of the points made. This is one that laid out his views of the process of succession (Cooper 1926:398–399).

All of this was summarized in a powerful metaphor of vegetation change as a braided stream, with some channels moving swiftly and idiosyncratically, and some broad, conspicuous channels bringing smaller streams together, and moving slowly. Although the concept of climax, and of a successional “sere,” or series of repeatable stages, could be accommodated by such a metaphor, there was no compelling need for climax, or for a repeatable sequence of stages, or strict directionality of succession in such a world. Indeed, the fact of consistent instability in the vegetation of the past was a compelling reason to expect the universality of change in contemporary vegetation. One should look for changes on any time scale, and examine the complex causes that yielded a net effect of change. It was important to recognize how different causes might dominate in different situations, while seeing that the braided stream and its causes presented a “complex whole governed by groups of causes, all of which constantly influence its course.” (Cooper 1926:409).

This insight of Cooper’s concerning the broad applicability of causes, appears later in the idea that net effects in succession reflect an array of mechanisms. A hierarchical set of causes, which may act differentially at different periods of succession, is one way to apply Cooper’s insight (Pickett and Cadenasso 2005). Also important to me was the insight that “Destruction is inherent in all successions” (Cooper 1926: 404), which of course helped me recognize the broad role of natural disturbance in communities (Pickett and White 1985). This was not the only paper of Cooper’s to make that point.

Cooper’s concern with all causes and scales of vegetation dynamics was an important exemplar for me. He sought principles to explain all sorts of vegetation change, ranging from evolution on one extreme, through the grand shifts of paleoecology, to detailed contemporary community alteration. Therefore, his work suggested a broad and inclusive framework that could account for the evolutionary basis of succession (Pickett 1976), to the role of animals, herbivores, and predators (which had long been neglected by plant ecologists). The pursuit of broad, open-ended, and intentionally mutable frameworks has become one of the tasks I have set for myself, in part due to the insights offered by Cooper so long ago.

Cooper’s fluid thinking was easy for me to apply to vegetation dynamics, since that field initially stimulated his essay. However, his charge for each generation to examine the fundamentals of its concepts and frameworks echoes throughout ecology. Rigid classificatory systems of ecological phenomena, inappropriate reification of idealized concepts, and narrow conceptions of trajectories and causes, is not uncommon in ecology. Cooper’s exhortation for us all, whatever speciality we practice, to periodically examine our fundamentals, is as cogent today as when he put pen to paper over 80 years ago.

Just as the process is more than the product, so the mode of thought is more fundamental than the result thereof. Therefore, while believing thoroughly in the truth and usefulness of the ideas outlined in this paper, I am even more seriously interested in urging upon ecologists the fluent mode of thought which I have endeavored to exemplify in the formulation and presentation of these ideas. (Cooper 1926:411)

I hope that the emerging and tentative frameworks for vegetation change (Pickett and Cadenasso 2005), boundary dynamics (Cadenasso et al. 2003), landscape heterogeneity, and patch dynamics (Pickett et al. 2000), show the mark of Cooper’s influence, and at their best, rise to the level of fluid thinking for fluid phenomena. In any event, his call to reevaluate the fundamentals seems as relevant to all parts of ecology today as his original call that emerged from vegetation dynamics in 1926.

Literature cited

Botkin, D. B., and M. J. Sobel. 1975. Stability in time-varying ecosystems. American Naturalist 109:625–646.

Cadenasso M. L., S. T. A. Pickett, K. C. Weathers, and C. G. Jones. 2003. A framework for a theory of ecological boundaries. BioScience 53:750–758.

Clements, F. E. 1916. Plant succession: an analysis of the development of vegetation. Carnegie Institution of Washington, Washington, D.C., USA.

Cooper, W. S. 1926. The fundamentals of vegetational change. Ecology 7:391–413.

McIntosh, R. P. 1985. The background of ecology: concept and theory. Cambridge University Press, Cambridge, UK.

Pickett, S. T. A. 1976. Succession: an evolutionary interpretation. American Naturalist 110:107–119.

Pickett, S. T. A., and M. L. Cadenasso. 2005. Vegetation succession. Pages 172–198 in E. van der Maarel, editor. Vegetation ecology. Blackwell Publishing, Malden, Massachusetts, USA.

Pickett, S. T. A., M. L. Cadenasso, and C. G. Jones. 2000. Generation of heterogeneity by organisms: creation, maintenance, and transformation. Pages 33–52 in M. Hutchings, editor. Ecological consequences of habitat heterogeneity. Blackwell, New York, New York, USA.

Pickett, S. T. A., and P. S. White, editor. 1985. The ecology of natural disturbance and patch dynamics. Academic Press, Orlando, Florida, USA.

Sprugel, D. G. 1980. A “pedagogical genealogy” of American plant ecologists. ESA Bulletin 61:197–200.

Steward T. A. Pickett
Institute of Ecosystem Studies
Millbrook, NY 12545
E-mail: [email protected]

1 Rereading Cooper’s paper in 2006 raises two questions about style. His use of “man” rather than “people” or “humans” is something that I find jarring. While I realize that grammarians may argue that is an acceptable use, I must warn contemporary readers that they may have to read beyond that usage. Cooper’s style is otherwise just a little more formal than today’s most formal writing. It is remarkably clear writing in spite of that formality.



2006 ESA Annual Meeting

Upstart Views of Restoration Icons

A symposium organized by D. J. Larkin and J. B. Zedler (both at University of Wisconsin-Madison), and D. Falk (Laboratory of Tree-Ring Research, University of Arizona), at the 91st ESA Annual Meeting (2006) in Memphis, Tennessee, focused on “Upstart Perspectives on Restoration Icons.”

As ecological restoration efforts become better known and better studied, and as more ecologists choose to conduct research in restoration sites, the gap between applied science and theory becomes narrower. Such was the case when Margaret Palmer catalyzed a symposium for the 2002 ESA meeting in Tucson, where ecologists indicated how theory could inform restoration practice. Several speakers then expanded their remarks in a book (Falk et al. 2006) that further merged ecological theory and practice. Still, gaps will remain if the icons relied upon by restorationists are inconsistent with the latest advances in theory and understanding of how ecosystems develop. Accordingly, this symposium set out to deconstruct some icons of restoration ecology, by applying contemporary ecological theory to restoration ecology. The combined result represents a paradigm shift, away from equilibrial, predictable models and toward a world of restored ecosystems that is at least partly stochastic, time-varying, and context dependent.

One pervasive icon is a simple illustration of how ecosystem structure and function develop following some restoration effort (Fig. 1). A. D. Bradshaw (1984) depicted restoration as having a single target that is hit after substrates are modified, with ecosystem structure and function developing along the same linear pathway. His diagram continues to appear in books, papers, and even the journal Science (Dobson et al. 1997), although evidence supporting this model has never been included in those publications (Zedler and Lindig-Cisneros 2000). Our concern is not with Tony Bradshaw, who is himself an important icon of restoration ecology, for he was among the earliest ecologists to address actual restoration problems (how to vegetate mine tailings) and to urge others to recognize the value of studying restoration sites as a test of ecological understanding (e.g., Bradshaw 1987). Instead, the issue concerns those who expect restoration efforts to follow overly simple models, or who promise unrealistic pathways and outcomes.

Fig. 1. Simplification of Bradshaw’s (1984) model of restoration, showing a single target, a straight path, and a linear correlation between community structure and ecosystem function.

Because there is no complete guide to the theoretical foundations of restoration ecology, and because ecologists have much to offer in helping restoration ecology mature as a science (Zedler 2005), we convened a symposium on “Upstart Views of Restoration Icons” to highlight some of contributions of the new book (Falk et al. 2006), while asking authors and other speakers to stretch beyond previous writings to close further the gap between theory and practice. Dan Larkin solicited the speakers and organized the session; Joy Zedler moderated the symposium; and Don Falk provided the wrap-up. As a disclaimer, we did not ask speakers to critique the icon in Fig. 1; rather, we selected results from their talks that address this widely published model of how restoration works; thus, our descriptions of each presentation are not fully representative (in keeping with instructions for symposium commentaries in the ESA Bulletin).

Ten speakers (names in boldface type) provided evidence that dispels key elements of a key restoration icon (Fig. 1). Their presentations failed to support (1) a single obvious target for restoration, (2) a straight path to the target, and (3) a linear relationship between structure and function.

Upstart view No. 1. There is no single, obvious target for restoration; that is, reference conditions represent dynamic, multivariate, and nonequilibrial processes.

Bob Peet (University of North Carolina) presented a model of relationships between environmental conditions and plant communities that suggests appropriate species lists, proportions of plants/species, and sequences for introducing plants to individual restoration sites. Science-based “designer plantings” should reduce restoration efforts and costs. North Carolina’s Department of Transportation is using this approach to restore lands in anticipation of the need to mitigate impacts of future highway construction projects.

Roberto Lindig-Cisneros (University of Michoacana) added human needs to the list of constraints on the restoration target. In southern Mexico, managers agreed to allow experimental restoration of tephra (unvegetated ash that persisted 60 years following eruption of Mt. Paracutín), but only if the target could be the two native pine species that provide livelihoods (Fig. 2). In exchange, local people helped establish experimental plantings (with and without bark mulch). As expected, mulching lowered soil temperature and enhanced pine establishment, but, unexpectedly, the effect was strong only in dry years.

Fig. 2 . Forest-restoration experiment in Comunidad Indigena de Neuvo San Juan (Michoacán), where two species of native Pinus were planted with and without mulch (pine bark) in tephra (ash from the 1943–1952 eruption of Parcutín volcano) and with and without fencing to exclude herbivores. Photo by R. Lindig-Cisneros.

Denise Seliskar (University of Delaware-Lewis) demonstrated that tailoring could extend below the species level. After planting a salt marsh restoration site in Delaware (Fig. 3, photo) as an experiment to test the effects of three genotypes of Spartina alterniflora (from Maine, Delaware, and Georgia), she and others documented numerous impacts on everything from canopy height and stem density (Fig. 3, graphs), to root and rhizome distributions, edaphic chlorophyll concentration, and decomposition rates. Even the numbers of larval fish caught in pit traps differed by a factor of 2. She and Jack Gallagher then extended their research by selecting genotypes of many halophytes via tissue culture. Some genotypes are broadly tolerant of stressful field conditions, while others perform best in specific sites. “Designer genotypes” could increase survival and growth.

Fig. 3. Photo: Construction of a tidal wetland with creeks and a pool created near Lewes, Delaware, by excavating soil from abandoned agricultural upland. Three genotypes of Spartina alterniflora (from Massachusetts, Delaware, and Georgia) were planted on the marsh plain in order to study their effect on marsh function. Graphs: Among the responses were differences in canopy height and stem density of Spartina alterniflora at the site (pink bars); values at their site of origin are in yellow. See Seliskar et al. (2000).

Stuart Findlay (Institute for Ecosystem Studies, New York) gave compelling evidence that multiple-function ecosystems are unrealistic targets for wetlands. While we might aim for clean water, high productivity, high biodiversity, flood reduction, and other functions, he argued that restoration sites are context dependent and typically do not provide all the ecosystem services expected of them, and that some combinations of functions are mutually exclusive (Fig. 4). If multiple driving factors are not correlated in space or time, individual sites cannot excel in several functions simultaneously. This leads to high interannual variability in site performance and argues for a relaxation of the restoration target. Thus, plans to restore multiple functions in a watershed will require multiple restoration sites and efforts.

Fig. 4. Functions may also have complex controlling factors, as in this example, which illustrates that the capacity of a patch of submerged aquatic vegetation to ameliorate high turbidity conditions is spatially contingent. Patches within ~100 m of the 5-m depth contour have a much lower likelihood of maintaining turbidity below 40 NTU (neothelometric turbidity units). Using this water clarity criterion as a performance measure would require relaxing the criterion for sites closer to deep waters. Graph by S. Findlay.

As Don Falk concluded, the paradigm of the “single target” should shift to that of “reference dynamics” (Falk, in press), where interactions, temporal and spatial variability, and stochastic processes are emphasized. This makes defining the ecological reference more complex, difficult, and uncertain—but also more realistic.

Upstart view No. 2. Restoration does not always follow a straight path to the target; more realistically, restoration is never finished.

Katie Suding (University of California-Irvine) offered evidence that California grasslands resist restoration and follow an alternative states model (Fig. 5), with internal feedbacks that help sustain each of the dominance states, namely, the native bunch grass (which competes more strongly for N) and exotic annual grasses (which compete more strongly for light). Through innovative experimentation, she showed that soils formed under the native tended to favor the native, and that soils formed under exotics favored exotics. These and other results collectively support the alternative state model. Thus, if restorationists plant the “right” species in the “wrong” soil, the site will not necessarily favor the native vegetation.

Fig. 5. Predictions of alternative states that have direct applicability to restoration ecology: threshold patterns (at X 1 and X 2), regional coexistence (dominance of one state or dynamic regime below X 1 and one state above X 2), multiple attractors where either state could exist (solid lines between X 1 and X 2; the dotted line is a repellor), the presence of positive feedbacks (to drive the system to one or the other state), and hysteresis (recovery trajectory is different from collapse trajectory). All these predictions can be tested in a restoration context; several lines of evidence are needed. A system that demonstrates these dynamics requires a shift from single-equilibrium steady-state management to an adaptive approach that incorporates regional refugia, legacy, and priority effects, and resilience into its tool set. Graph courtesy of Suding.

Dan Larkin (University of Wisconsin) tested the importance of intertidal pools and tidal creeks to salt marsh food webs in a large experimental salt marsh (Fig. 6) and found increased feeding opportunities at high tide (more algal biomass, more invertebrates in pools) than where pools were lacking. Also, killifish fed more in areas with pools than without. Furthermore, tidal creeks enhanced use by mudsuckers. Thus, feeding opportunities were best restored where topography mimicked natural heterogeneity (creeks plus pools), but the relationships between species and microhabitat were complex.

Fig. 6. An 8-ha experimental marsh in the southwesternmost corner of California allowed comparison of areas with and without tidal creeks and with and without tidal pools. At high tide, killifishes preferentially occurred and fed in pools, where algal and invertebrate foods were most abundant. Fig. 6a shows a 1-ha “cell” with a creek branching off the tidal channel at the base of the photo; Fig. 6b shows tidal pools at the opposite side of the marsh and the dominant plant, Salicornia virginica. July 2005 photos by J. Zedler.

Holly Menninger (University of Maryland) culled the literature for examples of real evidence that substrate heterogeneity increases the diversity of taxa. In contrast to the above speakers, she found none for a dozen experiments in streams, including the work of her co-author, Margaret Palmer (Fig. 7). While the experiments being done in streams have not enhanced diversity of the taxa being explored, a focus on restoring processes in streams might help researchers figure out how to enrich diversity.

Fig. 7. Invertebrate taxa richness did not differ among heterogeneity treatments imposed by Brooks et al. (2002) in a Virginia stream. Treatments differed in the variability of streambed particle sizes in entire riffles.
Redrawn from Brooks et al. (2002).

As Don Falk summed it up, the emerging view of the postrestoration state is one of complex, and at least partly stochastic, spatially contingent systems, with nonlinear response functions to treatments, nondeterministic outcomes, and nonequilibrial properties.

Upstart view No. 3. Structure and function are not linearly related.

Shahid Naeem (Columbia University) indicated that many plant species combinations might be viable restoration targets, based on biodiversity–ecosystem function (BEF) theory, and that the relationship to species richness will differ by function. He unveiled a new 5-year experiment (Fig. 8) that is underway in Mongolia, involving >700 plots (each 6 × 6 m) that will have controlled composition and species richness for the first years, before opening plots to grazing and assessing their functional capacity (including livestock production). “Combinatorial forecasting” could lead to recommendations for specific richness levels and/or assemblages for use in restoring specific ecosystem functions.

Fig. 8. Ambitious new 5-year experiment to test Biodiversity Ecosystem Function (BEF) theory in grasslands of Inner Mongolia. Plots are 6 × 6 m; weeded by hand to control species richness. Photo by S. Naeem.

Greg Bruland (University of Hawaii) tested the ability to restore both species and functions to a former forested wetland that was farmed (and flattened in the process), and then regraded to create a large experiment in topographic heterogeneity (1.3-m vertical range). Aboveground biomass accumulated least on hummocks and most in hollows, while species richness increased from hummocks to hollows to flats (Fig. 9)—not a linear correlation between these measures of function and structure. Overall, the specialization of species to microhabitats led to high diversity at the site scale.

Fig. 9. Mean species richness (i) and aboveground biomass (ii) in three microtopographic treatments in a restored wetland. Data are means ± 1 SE; a,b,c differed using ANOVA. Redrawn from Bruland and Richardson (2005).

Joy Zedler (University of Wisconsin) filled a gap in the program with Suzanne Kercher and Andrea Herr-Turoff’s data from wet prairie mesocosms. As an invasive grass expanded, biomass increased and species richness decreased. Then, despite killing the grass with herbicide, biomass remained high where few species remained. Productivity and species richness were negatively related, contradicting the notion of a positive linear correlation; furthermore, restorability was lower where stormwater treatments (flooding and nutrients) were continued than where they ceased.

Clever theorists and talented experimentalists continue to amass impressive data sets that challenge traditional views that restoration outcomes are predictable (we achieve a specific target), defined (variation is around a predetermined mean condition), and stable (we can keep the system that way). We summarize with:

Upstart view No. 4: Restoration outcomes are unpredictable, stochastic, and nonequilibrial, and the work is never finished.

While the “Bradshaw icon” might characterize some components of some restoration projects at some times, a more dynamic paradigm for restored ecosystems would accommodate variability and even unpredictability as positive signs of healthy, functioning systems.

Literature cited

Bradshaw, A. D. 1984. Ecological principles and land reclamation practice. Landscape Planning 11:35–48.

Bradshaw, A. D. 1987. Restoration: an acid test for ecology. Pages 23–29 in W. R. Jordan III, M. E. Gilpin, and J. D. Aber. Restoration ecology: a synthetic approach to ecological research. Cambridge University Press, Cambridge, UK.

Brooks, S. S., M. A. Palmer, B. J. Cardinale, C. M. Swan, and S. Ribblett. 2002. Assessing stream ecosystem rehabilitation: limitations of community structure data. Restoration Ecology 10: 156–168.

Bruland, G. L., and C. J. Richardson. 2005. Hydrologic, edaphic, and vegetative responses to microtopographic reestablishment in a restored wetland. Restoration Ecology 13:515–523.

Dobson, A. P., A. D. Bradshaw and A. J. M. Baker. 1997. Hopes for the future: restoration ecology and conservation biology. Science 227:515–522.

Falk, D. A. In press. Process-centered restoration and reference dynamics. Journal of Nature Conservation.

Falk, D., M. Palmer, and J. B. Zedler, editors. Foundations of restoration ecology. Island Press, Washington, D.C., USA.

Seliskar, D. M, J. L. Gallagher, D. M. Burdick, and L. A. Mutz. 2002. The regulation of ecosystem functions by ecotypic variation in the dominant plant: a Spartina alterniflora salt-marsh case study. Journal of Ecology 90:1–11.

Zedler, J. B. 2005. Restoration ecology: Principles from field tests of theory. San Francisco Estuary and Watershed Science. ‹ http://repositories.cdlib.org/jmie/sfews/vol3/iss2/art4›

Zedler, J. B., and R. Lindig-Cisneros. 2001. Functional equivalency of restored and natural salt marshes. Pages 565–582 in M. Weinstein and D. Kreeger, editors. Concepts and controversies in tidal marsh ecology. Kluwer Academic, Dordrecht, The Netherlands.

Joy B. Zedler, Botany Department and Arboretum, University of Wisconsin-Madison, Madison, WI 53706, E-mail: [email protected]

Donald A. Falk, Laboratory of Tree-Ring Research, University of Arizona, Tucson, AZ 85721

Daniel J. Larkin, Botany Department, University of Wisconsin-Madison, Madison, WI 53706

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Urban Food Webs: Errata. Revised Text and Figures

Urban Food Webs: Errata. Revised Text and Figures

In the October 2006 issue of the ESA Bulletin, two errors appeared in the Symposium Review “Urban Food Webs” (pages 387–393). In the list of authors, Thomas Parker should have appeared as Tommy Parker. And in Fig. 3a (page 389), the scientific name should be spelled “Habrophlebiodes,” as shown in the corrected figure below.

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Canada Chapter Newsletter

Chapter officers
Chair: Ken Lertzman, Simon Fraser University ‹[email protected]
Vice chair: Karen Yee, University of Calgary ‹[email protected]
Secretary/Treasurer: Beatrix Beisner ‹[email protected]
Chapter Home page: ‹http://www.esa.org/canada/

Announcing the new ESA Canada Chapter web site!

Please visit our newly revamped web site which you can find at: ‹http://www.esa.org/canada/›. Here you will find items of interest to Canadian ecologists including announcements of upcoming meetings. We have also set up a blog to discuss current Canadian environmental policy issues.

We are currently working on the new look for the web site. If you are interested in making suggestions for content or design, please contact Karen Yee at: [email protected] or (403) 220-5264.

Any West Coaster interested in representing the Canada Chapter??

We are looking for someone who is willing to represent the ESA Canada Chapter at the Pacific Ecology and Evolution Conference in March 2007, which has been sponsored by our chapter. If someone is planning on attending the Conference and would like to help the chapter out, please contact Karen Yee at: [email protected] or 403-220-5264. Minimal work is required; we just want some representation at the meeting.

CSEE Meeting announcement

We are pleased to announce that the second Canadian Society of Ecology and Evolution (CSEE) Annual Meeting will be held in Toronto, 17–20 May 2007 at Victoria College, University of Toronto. There will be three plenary talks, three symposia, 132 contributed talks, two poster sessions, and rooms for evening workshops. We expect about 500 delegates. More information can be found at: ‹http://www.eeb.utoronto.ca/csee/index.html›.

In addition, CSEE is launching a contest to find a logo representing the Society, see ‹http://www.ecoevo.ca/en/findlogo.htm › . The contest is open to graduate students and postdoctoral fellows who are members of the Canadian Society for Ecology and Evolution. The First Prize is an award of $500 in support of research or travel expenses for attending the CSEE meeting.

For more information on the CSEE please see ‹http://www.ecoevo.ca/en

Respectfully submitted,
Beatrix Beisner
E-mail: ‹[email protected]

Southeastern Chapter Newsletter

Chapter officers
Chair: Frank Gilliam (2006-2008) ‹[email protected]
Vice-Chair: Neil Billington (2005-2007) ‹[email protected]
Secretary/Treasurer: Howard Neufeld (2006-2008) ‹[email protected]
Web-Master: Mark Mackenzie‹[email protected]
Chapter Home page: ‹http://www.auburn.edu/seesa/

2007 ASB Meeting

The 2007 meeting of the Association of Southeastern Biologists will be held 18–21 April 2007, at the Columbia Metropolitan Convention Center in Columbia, South Carolina, hosted by the University of South Carolina. Details about the meeting can be found at: ‹http://www.asb.appstate.edu/meeting.php› The SE Chapter of ESA will have its luncheon and meeting on Friday, 20 April, 12:30–2:00 pm. We hope to see many of you there!

Notes from the Southeastern Chapter Brown Bag Lunch, ESA Annual Meeting, 9 August 2006

Our informal meeting was held during the SE-ESA Brown Bag Lunch at the Ecological Society of America meeting, Memphis, Tennessee. Jim Luken, Chair, called the meeting to order at 12:10 p.m. and quickly passed the gavel to Frank Gilliam, 2006– 2008 Chair.

Old business

Formal fundraising for the Elsie Quarterman–Catherine Keever Award for Best Student Poster was discussed, and several options were discussed at the luncheon.

Members were encouraged to submit symposium proposals for the 2007 meetings of the Association of Southeastern Biologists and the Ecological Society of America. Proposals seeking endorsement of the Chapter should be sent to Frank Gilliam ‹[email protected]› by 13 September 2006.

Updated ESA Chapter Financial Reports were not complete at the time of the meeting, but as of June 2006, the balance of the General Fund was $6597. The fund balances of the Odum and Quarterman-Keever Awards are estimated to be $9136 and $997, respectively.

New business

Alan Covich, 2006– 2007 President of ESA, congratulated our chapter for its activity and strong involvement of both academic and agency ecologists and proposed us as a model for other regional chapters of ESA. ESA wants to establish regional chapters throughout the United States by its 100th anniversary in 2015. These chapters would form knowledge partnerships with their region and act in both transfer of knowledge and rapid response. Ideas of problems and issues in the Southeast that our chapter could address should be sent to Frank Gilliam ‹[email protected]› by 1 March 2007. The meeting was adjourned at 1:10 p.m. by Frank Gilliam, Chair.

Keeping in touch:

Check the Chapter Home page: ‹http://www.auburn.edu/seesa/› for updates and additional information. To join the Southeastern Chapter of ESA LISTSERVE, send a message to mail to:[email protected] with “subscribe scesa” in the body of the message. Please send news or announcements to ‹[email protected] › for distribution to the listserve, or to ‹[email protected]› for inclusion in the next quarterly newsletter.

Respectfully submitted

Nicole Turrill Welch, 2004–2006 Secretary / Treasurer (who provided notes from the ESA meeting in Memphis) and

Howard Neufeld, 2006–2008 Secretary / Treasurer

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Meeting Calendar

The Ecological Society of America's
92nd Annual Meeting
Joint meeting with
The Society for Ecological Restoration
August 5–10, 2007   San Jose, California

For more information visit ‹http://www.esa.org/sanjose/

Urban Wildlife Management National Conference

The conference will be held 18–20 June 2007 at the World Forestry Center, Portland, Oregon, USA. As our use of land for nonnative and nonagricultural purposes increases, traditional wildlife habitats are lost. As a result, wildlife managers, ecologists, city planners, landscape architects, and community leaders must find ways to create and maintain ecologically significant habitats in and around urban areas. This conference demonstrates management opportunities for urban habitats from small city parks in urban centers to large subdivisions with considerable open space. The roles of urban soils, vegetation, water quality, noise, open space, greenways, habitat, animal communities, and human communities will be discussed. Presenters will share proven strategies for successfully integrating increased wildlife habitat in our cities and towns.

For further information contact:

The National Arbor Day Foundation
P.O. Box 81415
Lincoln, NE 68501
(888) 448-7337
Fax: (402) 473-9556

Instructions for Contributors

DEADLINES: Contributions for publication in the Bulletin must reach the Editor’s office by the deadlines shown below to be published in a particular issue:

January (No. 1)
April (No. 2)
July (No. 3)
October (No. 4)

15 November
15 February
15 May
15 August

Please note that all material for publication in the Bulletin must be sent to the Bulletin Editor. Materials sent to any address except that of the Editor, given below, must then be forwarded to the Editor, resulting in delay in action on the manuscripts. Send all contributions, except those for Emerging Technologies, Ecology 101, Ecological education K–12, and Obituaries/Resolutions of Respect (see addresses below), to E .A. Johnson, Bulletin Editor-in-Chief, Department of Biological Sciences, University of Calgary, Calgary, Albert, T2N 1N4 Canada. Phone (403) 220-7635, Fax (403) 289-9311, E-mail: [email protected].MANUSCRIPT PREPARATION: The manuscript should be submitted as a WordPerfect or Microsoft Word (for Mac or DOS) manuscript, preferably as an e-mail message attachment to [email protected]. E-mailed photographs and diagrams must be in .tiff or .eps format. Other forms of electronic copy (text embedded in e-mail messages, diskettes sent by post) or hard copy can be submitted if absolutely necessary. If formatting could be troublesome (e.g., tables, European alphabet characters, etc.), hard copy also should be sent via fax to E. A. Johnson at (403) 289-9311, or via post. Hard-copy manuscripts should be double-spaced, with ample margins. Plain formatting must be used on hard-copy and electronic manuscripts. PLAIN FORMATTING consists of a single font of a single size, left justification throughout, line spacing the same throughout, and up to three different weights of headings. Other formats will not be accepted for publication. The author should THOROUGHLY PROOF the manuscript for accuracy, paying special attention to phone and fax numbers and web site and e-mail addresses, which are frequently incorrect.COVER PHOTOGRAPHS: The photo should illustrate ecological processes or an ecological research design. The cover of the July, 2004 issue is a good example. It helps if the colors in the photo are bright, although black and white photos are considered if they are well composed with good contrast.

If you would like to submit a digital file, submissions can be small jpegs (72 dpi) but if the image is selected for a cover the final image must be 300 dpi and at least 7 inches wide and 5 inches high. Email the file as an attachment to the Editor of the ESA Bulletin at [email protected]. Or send a single 5 x 7 or 8 x 10 photo to the Bulletin. On an accompanying photocopy, give your name, address, a photo legend up to 100 words, and, if the photo describes a paper in ESA or in another journal, the literature citation or title of the accepted manuscript. If you wish unused photos to be returned please include a self-addressed return envelope.LETTERS TO THE EDITOR AND COMMENTARIES: Please indicate if letters are intended for publication as this is not always obvious. The Bulletin publishes letters, longer commentaries, and philosophical and methodological items related to the science of Ecology. There are no page limits but authors may be asked to edit their submissions for clarity and precision. Previously published items from other sources can be republished in the Bulletin if the contributor obtains permission of the author and the copyright holder, and clearly identifies the original publication.MEETING ANNOUNCEMENTS: Submit a brief prose description of the upcoming meeting, including title, a short paragraph on objectives and content, dates, location, registration requirements, and meeting contact person’s name, street address, and phone/fax/e-mail address. Please do not submit meeting brochures in the expectation that the Editor will write the prose description; he won’t. Compare the publication deadlines above with the meeting deadlines to be sure the announcement will appear in time.

MEETING REVIEWS: The Bulletin publishes reviews of symposia and workshops at the annual ESA meeting, as well as important and appropriate meetings that are unrelated to the annual ESA meeting. The reviewer should strive for a synthetic view of the meeting or symposium outcome, i.e., how the various presentations fit or conflict with each other and with current scientific thought on the topic. Review length is open, although about four double-spaced pages should be enough to capture the essence of most meetings.

The following advisory items are provided to help focus your review.
a) Meeting title, organizer, location, sponsoring organizations?
b) What were the meeting objectives, i.e., what scientific problems was the meeting organized to solve? Who cares (i.e., what was the relevance of this scientific problem to related ones under examination)?
c) How well did the meeting meet the objectives? Were there specific papers delivered or roundtables/discussion groups that were exemplary in reaching the objectives? You may concentrate the review on only the outstanding papers to the exclusion of all others, or give a comprehensive view of all presentations/meeting activities, or examine a selection of papers that neither describes all, nor focuses on a very few.
d) What new was discussed? What previously weak hypotheses were strengthened, confirmed or supported? Were any breakthroughs, or new or innovative hypotheses presented, that forced participants to rethink current concepts?
e) Was there anything else important that the meeting accomplished that may not have been part of its explicit objectives?
f) What subjects relevant to the meeting objectives were missing or left out? Did the scientific components of the problem that were included produce a strong slant or serious void by virtue of blind spots by the organizers, failure of invitees to appear, or similar difficulties?
g) Are there plans for a proceedings issue or meeting summary document, and if so who is editing it, who is publishing it, and when is it planned to appear (i.e., where can interested folks learn more about the meeting?)

EMERGING TECHNOLOGIES: Submissions for this section should be sent to the Section Editors in charge of the section: Dr. David Inouye, Department of Biology, University of Maryland, College Park, MD 20742. E-mail: [email protected]; or Dr. Sam Scheiner, Div. of Environmental Biology, Natl. Science Foundation, 4201 Wilson Blvd., Arlington, VA 22230. E-mail: [email protected]ECOLOGY 101: Submissions should be sent to the Section Editor in charge of this section: Dr. Harold Ornes, College of Sciences, SB 310A, Southern Utah University, Cedar City, UT 84720. E-mail: [email protected]ECOLOGICAL EDUCATION K–12: Correspondence and discussions about submissions to this section should be sent to Susan Barker, Department of Secondary Education, 350 Education South,, University of Alberta, Edmonton, Alberta T6G 2G5 Canada. E-mail: [email protected]
(780) 492 5415 Fax: (780) 492 9402
Charles W. (Andy) Anderson, 319A Erickson Hall, Michigan State University, East Lansing, MI 48824 USA. E-mail: [email protected]
(517) 432-4648 Fax: (517) 432-5092FOCUS ON FIELD STATIONS: Correspondence and discussions about submissions to this section should be sent to E. A. Johnson, Bulletin Editor-in-Chief, Department of Biological Sciences, University of Calgary, Calgary, Albert, T2N 1N4 Canada. Phone (403) 220-7635, Fax (403) 289-9311, E-mail: [email protected].OBITUARIES AND RESOLUTIONS OF RESPECT: Details of ESA policy are published in the Bulletin, Volume 72(2):157–158, June 1991, and are abstracted below. The death of any deceased member will be acknowledged by the Bulletin in an Obituary upon submission of the information by a colleague to the Historical Records Committee. The Obituary should include a few sentences describing the person’s history (date and place of birth, professional address and title) and professional accomplishments. Longer Resolutions of Respect, up to three printed pages, will be solicited for all former ESA officers and winners of major awards, or for other ecologists on approval by the President. Solicited Resolutions of Respect will take precedence over unsolicited contributions, and either must be submitted to the Historical Records Committee (see ESA website) before publication in the Bulletin.


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