Instructions for Authors

Instructions for Authors
Recent updates:
 English help for authors. Contact the free service, Editorial Assistance Program, at
 Data permissions. Authors should verify permission to publish all data and be prepared to
provide written documentation for all data presented in the manuscript. Authors are
encouraged to deposit data in public repositories, when appropriate.
 Funding information required. Instructions are on the submission form in the online system
and below in the Article Formatting section.
 Ethics statement required. Instructions are on the submissoin form in the online system and
below in the Article Formatting section.
Instructions for Authors
updated May 4, 2015
The Auk: Ornithological Advances
The Condor: Ornithological Applications
The Auk: Ornithological Advances and The Condor: Ornithological Applications are published by the American Ornithologists' Union (AOU) and the Cooper Ornithological Society (COS). The journals are managed by the Central Ornithology Publication Office (COPO). Send inquires about the suitability of an article topic for The Auk or The Condor to either Editor‐in‐Chief and inquiries about other submission or publication questions to the managing editors in the COPO office ( The Journals publish original material that has not been published elsewhere. PREPARING YOUR MANUSCRIPT
1. Choose the Journal.
The Auk: Ornithological Advances is an international, peer-reviewed journal that publishes
original research and scholarship advancing the fundamental scientific knowledge of bird species
and of broad biological concepts (e.g., ecology, evolution, behavior, physiology, genetics) through
studies of bird species. Articles often introduce or employ innovative empirical and theoretical
approaches and analyses.
The Condor: Ornithological Applications publishes original research and scholarship in peerreviewed articles that address ornithological applications in two ways: the application of
scientific theory and methods to the conservation, management, and ecology of birds; and the
application of ornithological knowledge to conservation and management policy and other issues
of importance to society. The journal aims to reach both research ornithologists and practitioners.
The Auk: Ornithological Advances
 Evolutionary history and paleontology
 Systematics and nomenclature
 Behavioral ecology
 Foraging strategies and tactics
 Population biology and ecology
 Molecular ecology
 Community and landscape ecology
Migration and orientation
Spacing patterns and habitat use
Genetics and genomics
Epigenetics, and evolutionary development
Physiology and biochemistry
Morphology and anatomy
Integrative and cross-disciplinary studies
Theoretical and methodological advances
Thematic reviews and opinion pieces
The Condor: Ornithological Applications
 Population biology, including threats to bird populations
 Conservation genetics
 Community and landscape ecology
 Ecosystem-level influences of birds
 Effects of habitat alteration and fragmentation
 Avian responses to climate change
 Anthropogenic effects on genetics, behavior, or physiological processes
 Biology of avian diseases and disease transmission by birds
 Birds in urban or agricultural settings
 Sociological and economic studies related to birds or the discipline of ornithology
 Integrative and cross-disciplinary studies
 Theoretical and methodological advances in practice
 Evaluations of science relevant to issues in conservation and management
 Thematic reviews and opinion pieces
2. Choose the Manuscript Type.
Research Article. All manuscripts that are submitted for peer review should be submitted as
Research Articles in PeerTrack. For manuscripts that might appear in special sections, such as
Perspectives, Commentaries, and Reviews, please contact the Editor-in-Chief first, then upload
your submission as a Research Article, with the section type written on the title page. Abstracts
are required for all Perspectives, Commentaries, and Reviews, as well as for Research Articles.
Review papers summarize research and are a synthesis of existing data, with the promise of
having a broad influence on the ornithological community. Please discuss with the Editor-inChief before submission.
Commentaries are brief papers that comment on articles previously published or opinion pieces
on some aspect of ornithology (especially the process or application or ornithology), or a
reconsideration of a topic in ornithology without extensive review.
Perspectives are papers that accompany an article (of any type) published at the same time. They
are invited by the Editor-in-Chief.
Editorials are written by Editors.
Book reviews, In Memoriam essays, and News and Notes go to the Book Review Editor, the In
Memoriam Editor, or to, respectively. Specify which journal you are
submitting a non-research article submission to: The Auk: Ornithological Advances or The
Condor: Ornithological Applications.
3. Optional Cover Letter. In general, cover letters are not necessary. The following
information is not needed in a cover letter: The submission system has a specific field to enter it:
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* conflict of interest statement
* results/data/figures that have been published elsewhere and the explanation
* non-plagiarism statement
* reviewers you would like to suggest
* reviewers you would prefer not to have review your manuscript
* resubmission information. In the cover letter, explain how the current submission
differs from the previously submitted version and why it should be considered again for this
journal. Please provide the previous manuscript number if available, and address the concerns
one by one from the previous decision letter and reviews.
* ownership of data. Authors should verify that they have permission to publish all data
included in the manuscript and should be prepared to provide written documentation if requested.
Authors are encouraged to review all contractual agreements involved in the study when
determining ownership of the data. Authors are encouraged to deposit their data in public
repositories, when appropriate.
* other comments
Other information that is not needed in a cover letter, because you can include it in the
manuscript itself:
* name the code of ethics followed during research in an Ethics Statement inserted in the
Acknowledgments section of your manuscript.
* note computer code and statistical procedures in the manuscript.
4. Supplemental Data and Materials. Upload supplemental files at the time of article
submission online. Supplemental material may include original and derived datasets, source code
for simulation models, multimedia files (such as sound files, videos) and details about and
software for unusual statistical analyses. Supplemental materials should not include details about
methods, results, or additional essential figures; any such content should be incorporated into the
manuscript as text, tables, or figures, or even an Appendix. Please name and cite all
supplemenary files this way: Supplemental Material Appendix A or Supplemental Material
Table S1 or Supplemental Material Figure S1. Combine supplemental material into one file when
For small pieces of supplemental material (whether essential or not to the manuscript), you
do not need to upload them as supplemental files. Instead, include short tables and figures that will
each fit on one page in an Appendix within the manuscript. Create an Appendix heading at the
end of the manuscript and insert the text and any figure captions and tables. Figures and Tables
within Appendices (not in Supplemental Material files) can be numbered following the regular
figures and tables, so for instance the first figure in Appendix A could titled “Appendix
A Figure 5”, even though it may be the only figure in Appendix A.
5. English. Authors whose native language is not English are encouraged to enlist the aid of a
native English speaker to review the manuscript for clarity and correct usage. Manuscripts that
do not meet linguistic standards may be returned without review. There is a free service available
from the Association for Field Ornithologists, called the Editorial Assistance Program: Contact: You are welcome to use this service no matter what journal you are
submitting to.
6. Format your Manuscript.
* Page limit. 25,000 words (about 70 pages when double-spaced with 12 pt. Times New
Roman type).
* Double-space all text, including figure captions and literature cited, using 12-pt. Times
New Roman or similar typeface. Margins should be 1 inch all around, on pages of 8.5 by 11 inch
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size. Do not justify the right margin (choose left-justified, not full-justified). Do not include line
numbers, headers, or footers. Insert page numbering for all pages.
* Order the sections of your manuscript in this way: Title page, Abstract with Keywords,
Introduction, Methods, Results, Discussion, Acknowledgments, Literature Cited, Figure Captions
(use capital letters for figure parts: A, B, etc. on the figure itself and (A), (B), etc. in the figure
caption), Appendix (short such as one page), and Tables go at the very end of your Word file. If
you have a Conclusion section, list it under a subheading of Conclusion within the Discussion
section. Tables and Figures may be in separate files. Supplemental data must be in
one or more separate files. Avoid long appendices within the manuscript itself; put such material
in a Supplemental Material file. (Small amounts of supplemental material can be included in the
manuscript if each item will fit on one printed 8.5 x 11 inch page.)
* Figure captions should accompany all figures. If you include figures in your Word file,
be sure to put the caption under each figure. If you submit separate figure files, please put the
caption under each figure as well and within the manuscript file, in a group at the end of the
Word file (as mentioned in the previous paragraph on ordering the article’s sections).
* File formats. For the manuscript: Word (.doc or .docx), WordPerfect (.wpd), Rich-text
format (.rft), or LaTeX (.tex). For tables: Word (.doc or .docx), Excel (.xls or .cvs), or Turbo
(.tds). For figures and images: .doc, .jpeg, .tif, .gif, .eps, or .ppt (or figures may be included in the
manuscript). Final figures will need to be in .tif, .eps, or .pdf format. For supplemental data: .doc,
.html, .mpeg, .xls, source code, .zip. Video files may be submitted in the following formats:
Quicktime, MPEG, and AVI. Audio files may be submitted in the following formats: MP3,
AAC, and WMA. Resolution should be 600 dpi for illustrations and 1200 dpi for line art. Put
figure numbers on each figure file (they will be cropped out). For a one-column figure the width
is 3.5" and for a two-column figure the width is 7".
* See the Ornithology Style Sheet and the formatted article below for additional style
items including author byline and footnotes, table format, reference format, etc., and for
scientific terminology. For items not on the style sheet below, see Scientific Style and Format:
The CBE Manual for Authors, Editors, and Publishers, Fifth, Sixth, or Seventh Editions
(Council of Biology Editors, Cambridge University Press). Also see the Merriam-Webster
Collegiate Dictionary.
1. Go to or and
read about each journal on the home page. Then choose the appropriate journal and you will be
taken to the login page. Internet Explorer 11 is incompatible with the submissions platform, so we
suggest using Firefox or Chrome.
If you received an email with your username and password, please use that to log in. Otherwise,
search for your name on the login page, and if you do not find it, then register as a new author.
* Corresponding author. The corresponding author is responsible for the submission of the
manuscript and all correspondence with staff and editors, from submission through publication.
An acknowledgment letter will be sent to the corresponding author once the staff has ensured that
the submission adheres to the requirements and is ready to be sent to the Editor-in- Chief. A
decision letter will ultimately be sent to the corresponding author and all coauthors; and if the
manuscript is accepted there will be further correspondence during the publication process.
* Updating author profiles. Authors are responsible for modifying their profile to keep
the editors and staff informed of changes in their contact information. Before submitting a
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manuscript, please be sure your profile information is current. (After logging in, choose "Update
My Information.") Please notify your coauthors to update their profiles as well prior to
2. Complete the following steps where information is gathered and where you upload and
approve your files. You can save and exit at any time in the process and come back later to
where you left off by logging in again as an author and choosing the Incomplete Submission link.
Basic information: Enter the information for article type (Research Article), title (limit
of 25 words), short title (running head limit of 8 words), coauthors' information, abstract (limited
to 300 words), keywords (up to 8 keywords, separated by commas), and topic classifications
(select or search for from 1 to 8 words/phrases).
Blinded submissions: You may submit your paper as blinded or unblinded. You will
have this choice at the last step of the submission process at the Attach Files stage, where you
will be asked to choose item type: Unblinded Submission or Blinded Submission, before you
upload your manuscript file. If you choose a blinded submission, please make sure to remove
identifying information from all sections of your manuscript file, such as in headers and footers
and in the Acknowledgments. The title page of this file should only have the article title. You
will also need to upload a separate and complete Title page file (choose “Title Page for Blinded
Review” file type), which includes the title, author names and author affiliations, and
corresponding author email. This separate, complete title page will not be shown to reviewers.
Additional information:
Required information. On the additional information page, answer several questions
requiring answers about conflict of interest, any part of the submission that is previously
published material, whether all coauthors agree to the submission, a non-plagiarism statement
(submissions will be checked with CrossCheck and iThenticate), funding information, and ethics
protocols used.
Optional information. There are boxes for optional information such as comments to the
editor, suggested reviewers, and suggested non-reviewers. There is also a choice of languages for
your Abstract to be translated into: Please choose Spanish, Portuguese, French, or Other. If you
choose Other, please include the foreign language abstract yourself in your manuscript. If you
upload your own foreign-language Abstract, please use only the scientific names for birds, not
common names. For example, the abstract text “Blue Jays (Cyanocitta cristata) build nests in
trees…’ would be translated into Spanish as “Cyanocitta cristata hacen sus nidos en árboles…”
Uploading files: Upload each file, re-order them if necessary, wait for the system to build
a merged PDF file (of all the files except Supplemental data), follow the prompt to Submissions
Waiting for Approval, view the merged PDF, and then approve it. This approves your merged
PDF and finalizes your submission.
After you approve your manuscript submission, you are finished with the submission process and
no longer have access to modify files or the information about your manuscript. The manuscript
will enter the submission queue, and you and your coauthors will receive a confirmation email
with the assigned manuscript number. The publication office will contact you if there are any
issues with your files. If not, the Editor-in-Chief will receive your submission for consideration.
You can access the status of your manuscript at any time by logging in and selecting Submissions
Being Processed in the New Submissions box. Under current status, you can see the stage of
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your manuscript: Incomplete; With the Editor; Under Review; Revise; Completed, Accept; or
Completed, Reject. You can use Send Email if you need to correspond with the publication
For papers that had a previous decision of major revisions or minor revisions, there will be a onemonth deadline to submit a revision. You may request an extension from the Editorial Office at We realize that some major revisions may take a full two months,
while some minor revisions can sometimes take only one week. Authors will be contacted after
two months if we have not heard from you about your revision.
Decision letter and copyedited manuscript. After your manuscript is accepted for publication,
carefully review the information in the decision letter. Shortly thereafter, you will receive a
copyedited manuscript and perhaps a request for higher-resolution figures. Your accepted
manuscript will have been copyedited to conform to scientific, technical, stylistic, and
grammatical standards. Please review the changes and answer the queries and any request for
new figures or other files. Then return the manuscript promptly to
Proofs. Next you will receive a PDF proof, copyright forms, reprint forms, and an invoice for
page charges. Please return your proofs, copyright forms, and a statement that all authors agree
with the final content and format of the article as soon as possible. Any delay in returning the
proofs will delay the publication date of your paper. Normally, papers will be published online
about two weeks after you return your article proof.
Page charges. Because the journals are published by nonprofit ornithological societies, we
request your support of the journal through page charges of $100 per published page, which will
be billed after your article publishes online. Discounts and waivers are available for authors
unable to pay page charges, especially for non-U.S. authors, according to international protocols.
Apply at
Copyright form. Each author will receive a copyright form to sign. There will be a place on the
form to indicate that the author is a federal employee.
Reprints. Authors will receive reprint information when the quarterly print issue is published.
Permission to use previously published material. If your article contains material (e.g., a table,
graph, diagram, illustration, photo, or section of text) that was previously published by someone
else (or published by you in a publication that does not give authors the right to republish
materials from their own articles without obtaining permission), you must obtain written
permission from the copyright-holder to republish that material. This restriction applies not only to
material that you intend to reproduce in its original form, but also, for example, to modified
artwork or graphs. The written (unrestricted) permission must be forwarded to the publication
office at The source should be listed in your paper in the Literature Cited
section. In each figure, illustration, table, or block of text that uses previously copyrighted
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material, a citation in one of the following forms should appear: "From Jones (1979)" or
"Modified from Jones (1979)" or "Redrawn from Jones (1979)."
Embargo. Authors are free to post their articles on their own website or their institution’s
website and to promote their work once they receive the final published PDF article. Let us know
when your work is cited in the media by emailing
Open Access. The Journals’ open access policy includes the following:
 Authors can distribute their own article as soon as it is published online, using the final
PDF that the publication office sends them.
 Journal articles are open access 24 months after publication of the quarterly printed issue.
 Authors can arrange for immediate open access of their article by paying a fee of $2,000,
or $1,500 for members of the American Ornithologists’ Union or the Cooper
Ornithological Society. Contact for this option.
-------------------------------------------------------------------------------------------------------------------ORNITHOLOGY STYLE SHEET
a priori and a posteriori. Not italicized because they are in the English dictionary.
Abbreviations. Minimize the use of nonstandard abbreviations or acronyms that readers must
memorize to follow your paper. Spell out any abbreviations at first usage with the abbreviation in
About. Use ~before numbers instead of about: ~90%, not about 90%.
Acknowledgments. List your funding sources here. If authors want to mention themselves,
intials are sufficient: K.W.H. would like to thank…. . Editor names and reviewer names are not
appropriate in this section: Editors and reviewers are doing their job and do not need to be
thanked individually.
Abstract. Maximum word count is 300. Avoid long lists of common methods or discursive
explanations of what you set out to accomplish. Abstracts should provide a brief summary of the
research, including the purpose, methods, results, and major conclusions. Do not include
citations in the Abstract. Authors are encouraged to submit a technically competent foreign
language abstract, or else the Journal will provide one in Spanish, Portuguese, or French. When
you submit your paper, you are asked which of the three languages you would like your Abstract
translated into.
Affiliation. See Author names.
And/or. May be used where appropriate.
Antarctic. Capitalized.
APPENDIX section. If more than one appendix, label APPENDIX A, APPENDIX B. Only
include short appendices in the paper itself. Upload long appendices as Supplemental Appendix
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files. Tables within appendices that are in the main text should follow the numbering of other
tables in the paper. So an appendix table citation in the text might be: “Table 5 in the Appendix”.
Approximately. Use ~before numbers instead of approximately: ~90%, not approximately
Arctic. Capitalized.
Author names. List authors with superscripted numbers to indicate affiliations at the time the
research was conducted. List institutional affiliations under the authors’ names. Include the email
address of the corresponding author with an asterisk before it, and put an asterisk after the
author’s name in the byline after the last affiliation superscripted number. Do not superscript the
asterisk (an asterisk is already a superscript). Above the corresponding author email address,
include any other footnotes such as
These authors contributed equally to the paper.
This author is deceased.
The corresponding author information is placed last.
Individuals listed as authors should have played a significant role in designing or carrying out the
research, writing the manuscript, or providing extensive guidance on the execution of the project.
Those individuals whose role was limited to providing materials, financial support, or review
should be recognized in the Acknowledgments section.
Biogeographical realms. These regions are capitalized: Neotropic and Neotropical, Antarctic,
Arctic, Holarctic, Palearctic, and Nearctic.
Bonferroni correction. (not Bonferroni Correction)
Boolean. (not boolean)
Capitalization. Proper names:
Hairy, Downy, and Ivory-billed woodpeckers (when multiple groups are named together). Hairy
Woodpecker. Down Woodpecker. Ivory-billed Woodpecker. (when named alone) Mississippi
and Missouri rivers. (when named together) Mississippi River. (when named alone). Lakes Erie
and Superior. Etc.
chi-square. (not Chi-square)
Citation order. Lists of Literature Cited citations within the text of the manuscript should be in
chronological order.
Companies and commercial product names. Use this style for products, companies, and
company location: Predation MP3 Game Caller (Western Rivers, Lexington, Tennessee, USA).
No trademark or registered trademark symbols. No Inc. or Co. on the company name.
Cover art. Photos may be submitted for cover art. They need not be figures from a submitted
article. Make sure they are high resolution for the large format of the journal covers. Send to
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Data. This is a plural noun, carrying a plural verb: Data were too few to assess significance.
Dataset. Dataset is one word.
Dates. Use American dating (September 29, 1992).
Day of the year. Refer to the ordinal date (number) 1 to 366. [This is not the Julian date.]
Deceased authors. See Author names.
Decimals. No naked decimals except with caliber: .410 shotgun. Otherwise, 0.17. Probability
rounded to two decimal places unless P < 0.01, in which case round to three decimal places; use
P < 0.001 as the smallest P-value.
DISCUSSION section. It is useful to start the Discussion with a statement that summarizes the
main results. The Discussion should develop the significance and importance of the Results and
set them into a framework of previous research. The Discussion should follow logically from the
Results. Additional statistical tests and results are usually inappropriate here and should be
presented in the Results section, except in unusual cases.
Document format. Page size of 8.5 x 11 inch format, double-spaced throughout, one inch
margins, left-justified.
doi. doi numbers will be provided by the publisher, doi: 10.1650/cond.2013.xxxxx or doi:
e.g., (for example) takes a comma and is roman.
Email. One word, email.
Ethics statements and guidelines. In the Acknowledgments section, you may state any Ethics
guidelines that you followed.
Equations. Center long equations on the page. Indicate where long equations should have a line
break. Use MathType to create equations (it is an add-on program to Word). Put spaces around
operators such as = , + , etc. Use bold and italics where appropriate for symbols (see Symbols).
Figures. Cite each figure in the text in numerical order (except for Appendix figures, see below).
Spell out the word Figure in citations and figure captions (Figure 1, Figures 2 and 3, Figure 1A,
1B). Figure citations from another work should use the word “figure” with lowercase “f” such as
(figure 2 in Smith 1980). Figures should be simple and easily comprehended without reference to
the manuscript text. Once accepted, a paper’s figures must be submitted as high- resolution
figures of 600 dpi in .tif, .eps, or .pdf formats (as reproduction of PowerPoint or Word figures is
not reliable). Figure captions should not repeat information already presented in text or tables.
Use capital letters for figure parts in the figure caption: (A), (B), etc. Bold the letters but not the
parentheses. For sound spectrograms (sonograms), use the actual tracing if it is sharp, clear, and
relatively short. If intensity differences are not important, then submit a high-contrast
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digital image that meets the above specifications. Label all axes, use sentence case labels (only
the first word is capitalized unless it is a proper noun). You can group related illustrations as
panels into a single figure file (Figure 1 would include 1A, 1B, 1C, 1D) so that they can be
placed together on the same page/screen. Mark each section of the figure A, B, C. If necessary,
you may submit each part of a figure as a separate file as long as it is clear how to combine the
parts into one figure for publication. When mixing figure citations in the text of your manuscript
with reference citations, use a semicolon: . . . text text text (Figure 1, Figure 2A and 2B; Jones
and Johnson 1978).
Appendix figure numbering should follow from the figure numbers in the rest of the
manuscript. So the first figure in Appendix A may be Figure 5. It should be cited as “Figure 5 in
Appendix A”, or “Appendix A Figure 5”.
Footnotes. No footnotes in the text. Put footnote-type information in parentheses in the text.
Footnotes may be used in tables; include them after the table itself.
Gene or amino acid sequences. Must be deposited in GenBank or an equivalent repository and
the accession numbers reported in Methods.
Holarctic. Capitalized.
flush left and bold. Second-level headings should be flush left and bold in title case (each word
capitalized), third-level headings are bold in sentence case (only the first word is capitalized) with
a period at the end, run in to the paragraph indented, and fourth-level headings are the same as
third-level headings except they are italic instead of bold. Text immediately following an H1
heading or a H2 heading should not be indented.
Hyphens. Do not use one hyphen to imply the rest of a word unless you use the second hyphen
as well. For instance, do not use inter- and intrasexual, as they are not parallel. Correct usage
would be “inter- and intra-sexual”. To avoid the problem, use intersexual and intrasexual, for
instance. Use hyphens only when necessary for meaning.
i.e. (that is) takes no comma after it and is not italicized.
Internet. Internet is capitalized.
INTRODUCTION section. It should provide the aims and significance of the research and
place it within the framework of existing work. Limit the use of citations; in general there a few
points that cannot be supported by three or fewer citations. Long lists of citations are seldom
required and detract from the readability of the manuscript.
Italics. Limit the extent to which italics are used for emphasis. Foreign words are italicized if
they do not appear in the American English dictionary (Merriam-Webster, Merriam-Webster
Collegiate, or Webster’s Third New International Dictionary Unabridged).
Keywords. One to 8 keywords. List after the Abstract. Put the word “Keywords” in italics.
Keywords need not be in alphabetical order. Follow the author’s order (which may be in order of
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Latin terms. Leave roman if they are in the American English dictionary (Merriam-Webster,
Merriam-Webster Collegiate, or Webster’s Third New International Unabridged). Latin terms
and other non-English words that do not appear in the American English dictionary are to be
Latitude and longitude. N 139°, W 64.15°, or 139°N, 64.15°W. Be consistent.
Literature Cited. Only cite references in the text that are listed in the Literature Cited section,
and vice versa. Lists of citations within the text of the manuscript should be in chronological
order. Do not alphabetize or rearrange the list other than chronologically. Cite 2014 articles from
The Auk and The Condor this way: The Auk: Ornithological Advances, and The Condor:
Ornithological Applications, as these are the new names of the Journals. For articles published in
2013 and earlier, cite as The Auk, and The Condor.
Within the text, cite references this way: Darwin and Huxley (1993), or (Darwin and
Huxley 1993), (Zar 1973, Giles 1994a, 1994b). For citations of three or more authors: (Ricklefs
et al. 1999). In the Literature Cited section, list references alphabetically and in the following
Ankney, C. D., and R. T. Alisauskas (1991). The use of nutrients by breeding waterfowl.
Proceedings of the International Ornithological Congress 20:2170–2176.
Avery, M. L. (1995). Rusty Blackbird (Euphagus carolinus). In The Birds of North America
200, (F. B. Gill and A. Poole, Editors). Academy of Natural Sciences, Philadelphia, PA,
USA, and American Ornithologists’ Union, Washington DC, USA. doi:10.2173/bna.599
Darley, J. A. (1968). The social organization of breeding Brown-headed Cowbirds. Ph.D.
dissertation, University of Western Ontario, London, ON, Canada.
Greenberg, R., C. Elphick, J. Nordby, C. Gjerdrum, H. Spautz, W. G. Shriver, B. Schmeling, B.
Olsen, P. Marra, N. Nur, and M. Winter (2006). Flooding and predation: Trade-offs in the
nesting ecology of tidal-marsh sparrows. In Terrestrial Vertebrates of Tidal Marshes:
Evolution, Ecology, and Conservation (R. Greenberg, J. E. Maldonado, S. Droege, and
M. V. MacDonald, Editors). Studies in Avian Biology 32:96–109.
Greenberg, R., J. E. Maldonado, S. Droege, and M. V. McDonald (Editors) (2006). Terrestrial
Vertebrates of Tidal Marshes: Evolution, Ecology, and Conservation. Studies in Avian
Biology 32.
Lafferty, K. D., A. P. Dobson, and A. M. Kuris (2006). Parasites dominate food web links.
Proceedings of the National Academy of Sciences USA 103:11211–11216.
National Audubon Society 2010. The Christmas Bird Count historical results. [Note: last date accessed is not necessary, as URLs
are checked at copyedit before publishing.] [Reprinted style…. .]
Peterson, J. M. C. (1988). Rusty Blackbird, Euphagus carolinus. In The Atlas of Breeding Birds
in New York State (R. F. Andrle and J. R. Carroll, Editors). Cornell University Press,
Polačiková, L., F. Takasu, B. G. Stokke, A. Moksnes, E. Røskaft, P. Cassey, M. E. Hauber, and
T. Grim (2013). Egg arrangement in avian clutches covaries with the rejection of foreign
eggs. Animal Cognition Online First. doi:10.1007/s1007101306151
Powell, G. V. N. (1985). Sociobiology and adaptive significance of interspecific foraging flocks
in the Neotropics. In Neotropical Ornithology (P. A. Buckley, M. S. Foster, E. S. Morton,
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R. S. Ridgely, and F. G. Buckley, Editors). Ornithological Monographs 36.
Ralph, C. J., G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt (Editors) (1995). Ecology and
conservation of the Marbled Murrelet. USDA Forest Service General Technical Report
Ringelman, K. M., and M. J. Stupaczuk (2013). Dabbling ducks increase nest defense after
partial clutch loss. The Condor 115:290-297.
SAS Institute. 1990. SAS-STAT user’s guide. Version 6, 4th edition. SAS Institute, Cary, NC,
Spector, D. A. (1992). Wood-warbler song systems: A review of paruline singing behaviors. In
Current Ornithology 9 (D. M. Power, Editor). Plenum Press, New York, NY, USA. pp.
Svensson-Coelho, M., J. G. Blake, B. A. Loiselle, A. S. Penrose, P. G. Parker, and R. E. Ricklefs
(2013). Diversity, prevalence, and host specificity of avian Plasmodium and
Haemoproteus in a Western Amazon assemblage. Ornithological Monographs 76:1–47.
Wilson, S., E. M. Anderson, A. S. G. Wilson, D. F. Bertram, and P. Arcese (2013). Citizen
science reveals an extensive shift in the winter distribution of migratory Western Grebes.
PLoS ONE 8:e65408.
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Table 1. Wintering locations in South America of Red-eyed Vireos (n = 10) migrating from
northwestern Pennsylvania. Values are means (with SD in parentheses), and n is the number of
days used to estimate location. Letters correspond to maps in Figure 2.
Bird Latitude
N 1.39° (2.90)
W 64.15° (0.98)
N 0.56° (2.05)
W 64.15° (0.98)
S 3.54° 92.99)
W 69.00° (1.11)
S 3.80° (2.56)
W 65.2° (0.70)
N 1.52° (2.61)
W 59.15° (0.66)
N 1.08° (2.30)
W 62.42° (0.63)
S 0.55° (3.13)
W 69.93° (0.94)
N 3.27° (2.12)
W 62.87° (0.91)
N 7.24° (2.24)
W 64.38° (0.71)
S 0.64° (2.45)
W 60.62° (0.83)
S 3.01° (1.80)
W 63.33° (0.73
N 1.81° (1.73)
W 63.70° (0.52)
Individual changed locations during seasons; listed in chronological order.
Not depicted in Figure 2.
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names of birds, and their order of presentation in the manuscript, including figures and tables,
Page 16
should follow:
North America and Middle America: the 7th edition of the American Ornithologists’ Union
Checklist of North American Birds and its supplements (
South America: AOU South American Classification Committee Checklist for South American
Birds (
Outside the Americas: Avibase Clements Checklist (
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Page 17
also see below).
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----------------------------------------------Red-eyed Vireo migration
Prolonged spring migration in the Red-eyed Vireo (Vireo olivaceus)
Paul A. Callo,1* Eugene S. Morton,2,3# and Bridget J. M. Stutchbury3a#
Department of Biology, Mary Baldwin College, Staunton, Virginia, USA
Hemlock Hill Field Station, Cambridge Springs, Pennsylvania, USA
Department of Biology, York University, Toronto, Canada
Current address: Yale University, New Haven, Connecticut, USA
These authors contributed equally to the paper.
This author is deceased.
* Corresponding author:
[if 2 corresponding authors, list name, email address; name, email address:
* Corresponding authors: Paul Callo,; Eugene Morton,]
Submitted November 12, 2012; Accepted February 15, 2013; Published April 28, 2013 [these
dates will be supplied by the journal publisher]
Page 18
We used archival geolocators to track the migration of Red-eyed Vireos (Vireo
olivaceus), abundant forest songbirds with significantly increasing breeding-population
trends, to identify important stopover and wintering regions. All individuals from a
single breeding site (n = 10) wintered in northwestern South America, an extensively
forested region, and in spring used a consistent route, crossing the Gulf of Mexico from
the Yucatan to Louisiana. . . .
Keywords: frugivory, geolocators, geologgers, migration, Red-eyed Vireo, stopovers,
Vireo olivaceus
Migración Prolongada de Primavera en Vireo olivaceus
Usamos geolocalizadores para rastrear la migración de Vireo olivaceu, un ave canora de bosque
abundante con tendencias a incrementar su población reproductiva, para identificar regioanes
importantes de parada e invernada. Todos los individuos de un único sitio de reproducción (n =
10) pasaron el invierno en el noroccidente de Sur América, una région con bosques extensos. En
la primavera, las aves usaron una ruta común, cruzando el golfo de México desde Yucatán hasta
Luisiana. . . .
Palabras clave: …….
Widespread and long-term effects on populations of songbirds that migrate to the tropics for the
northern winter are driven by both breeding-ground productivity and mortality during migration
and the nonbreeding season (Terbrough 1980, Sherry and Holmes 1995, Faaborg et al. 2010).
Data on the timing of migration, routes taken, stopover locations and durations, and overwintering
locations are needed to permit an informed assessment of conservation needs and for projecting
future population trends. For most Western Hemisphere songbirds, banding recovery records that
link breeding and tropical wintering sites are too infrequent to answer these and other questions.
However, tracking of small birds for a full year is now possible using light- level geolocators
(Stutchbury et al. 2009), which make it feasible to map migration routes and destinations of
breeding populations.
We used data from light-level geolocators (Mk20S, 0.6 g; British Antarctic Survey [BAS])
deployed on male Red-eyed Vireos (n = 26) between June 3 and June 17, 2011, and retrieved
between 26 May and 9 June 2012 (n = 10) at the 150-ha Hemlock Hill Field Station in
northwestern Pennsylvania (41.8°N, 79.9°W). The site is covered by mature mixed-deciduous
forest with scattered Eastern Hemlocks (Tsuga canadensis). Individuals were captured by use of
a targeted playback of Red-eyed Vireo song and a 6-m mist net. A taxidermic mount of a male
Red-eyed vireo was used in most instances. Geolocators were attached to birds using a leg-loop
harness made of a 2.5-mm Teflon ribbon (Stutchbury et al. 2011).
Page 19
Wintering Locations and Migration Routes [second level heading]
All Red-eyed Vireos from the Hemlock Hill breeding population wintered in a similar region in
northwestern South America that represented an area of ~15% of the total winter range (Table 1
and Figure 1). Average distance between individuals (all pairwise comparisons, n = 45) was 712
± 300 km (mean ± SD), and average nearest-neighbor distance was 286 ± 142 km (n = 10). Most
individuals (8 of 10) occupied a single wintering region, but two individuals (Figure 2E, 2I) first
occupied a winter site from late October to the beginning of December before moving ~40 km
westward to their final wintering region, where they stayed for 4 months.
The spring migration route was similar among all 10 individuals as birds migrated through
Central America to the Yucatan Peninsula (Figure 2) .
Stopovers and rate of migration [third level heading]. Spring migration, from start to
finish, averaged 46 days (range 39–52 days), and with stopovers, migration rate averaged 146
km day−1 (Table 2). However, most of the spring migration consisted of stopover days, and
individuals covered the journey of ~6,600 km in only 13 days of flight. Migration rate and
stopover duration varied greatly among different stages of the journey (Table 2 and Figure 2).
Red-eyed vireos had prolonged stopovers in Colombia (18.6 ± 4.9 days [all durations reported as
means ± SD]; range: 12–27 days) immediately after beginning spring migration. Spring
migration rate through South America was slow, averaging 72 km−1 day, and increased
significantly as birds traveled through Central America (mean = 178 km day−1) and completed
their journey across the Gulf of Mexico and through the United States to the breeding site (mean
= 310 km−1 day; one way ANOVA, F = 33.5, df = 2 and 27, P < 0.0001; Table 2). Most birds
also had a shorter stopver (6.3 ± 3.3 days) in Central Nicaragua.
Fourth-level heading. All birds remained at the breeding site throughout August, but the
onset of fall migration in September was unknown because birds could have moved south with
no change in longitude compared with the breeding site. Average arrival date at the wintering
site was October 22 (range: October 14 to November 4).
Red-eyed Vireos from this population all overwintered in northwestern South America (Figure 1)
in either the Amazon or Orinoco River basins. These river basins are perhaps the most pristine
region in South America, with >90% forest cover (Fraser et al. 2012). Two of the 10 Red-eyed
Vireos (Figure 2E, 2I) changed locations during the winter season, both to the southwest of their
initial site, but over relatively short distances (400 km). Intratropical migration has also been
documented using geologgers for Veeries (Catharus fuscescens; Heckscher et al. 2011; 5 of 5)
and Purple Martins (Progne subis; Fraser et al. 2012; 63 of 95), but both of these species move
over long distances (average movement >500 km) from site to site within South America. Little
is known about Red-eyed Vireos’ behavior on their wintering grounds (Cimprich et al. 2000), but
they appear to have high social tolerance, typical of highly frugivorous species while not breeding.
Vireos often occur in groups of conspecifics as well as mixed-species flocks in the tropical forest
canopy and edge, and are largely silent (Ridgely and Tudor 1989, Ridgely and Greenfield 2001).
Spring migration featured a prolonged stopover (18.6 ± 4.9 days) in Colombia soon after
departure from winter sites (Figure 2 and Table 2). Very long spring stops do not occur in Purple
Martins or Wood Thrushes (Fraser et al. 2012, Stanley et al. 2012) but have been documented
with geolocators in Swainson’s Thrush (Catharus ustulatus; Delmore et al. 2012). Swainson’s
Thrushes breeding in inland British Columbia, and wintering in South America, had long spring
Page 20
We thank L. Welch and J. Silverton for assistance with field work and E. Jones for statistical
assistance. We also thank O. Love. [Copyeditors will remove any editors or reviewers who are
thanked here, except “anonymous reviewers”.]
Funding statement: This research was funded by the Natural Sciences and Engineering Research
Council of Canada and by grants from Mary Baldwin College [include grant numbers]. None of
the funders had any input into the content of the manuscript. None of the funders required their
approval of the manuscript before submission or publication.
[Required: A list of funders, a statement about whether any funders had input into the content of
the manuscript, and a statement about whether any funders required their approval of the
manuscript before submission or publication.]
Ethics statement: [Required: List of protocols used, license numbers, etc.] [For example:
Ethics statement: This research was conducted in compliance with the Guidelines to the Use of Wild Birds in Research. LITERATURE CITED
Bayly, N. J., C. Gómez, K. A. Hobson, A. M. González, and K. V. Rosenberg (2012). Fall
migration of the the Veery (Catharus fuscescens) in northern Colombia: Determining the
energetic importance of a stopover site. The Auk 129:449–453.
Cimprich, D. A., F. R. Moore, and M. P. Guilfoyle (2000). Red-eyed Vireo (Vireo olivaceus). In
The Birds of North America, no. 527 (A. Poole and F. Gill, Editors), Birds of North
America, Philadelphia, PA, USA. doi:10.2173/bna.599
Cooper, N. W., M. T. Murphy, L. J. Redmond, and A. C. Dolan (2011). Reproductive correlates
of spring arrival date in the Eastern Kingbird Tyrannus tyrannus. Journal of Ornithology
Delmore, K. E., J. W. Fox, and D. E. Irwin (2012). Dramatic intraspecific differences in
migratory routes, stopover sites and wintering areas, revealed using light-level
geolocators. Proceedings of the Royal Society of London, Series B 279:4582–4589.
Faaborg, J., R. T. Holmes, A. D. Anders, K. L. Bildstein, K. M. Dugger, S. A. Gauthreaux, Jr., P.
Heglund, K. A. Hobson, A. E. Jahn, D. H. Johnson, et al. (2010). Conserving migratory
land birds in the New World: Do we know enough? Ecological Applications 20:398–418.
The Appendix may contain text and/or tables. Avoid long appendices, or upload as supplemental
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Supplemental Tables and Figures should be named Supplemental Material Table S1 and
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Appendix in the manuscript Word file.
Page 21
Figure 1. Wintering locations in South America of Red-eyed Vireos (n = 10) tracked with
geolocators from one breeding population in northwestern Pennsylvania (inset). Typical standard
deviation in latitude and longitude for mean location is shown with lines for one bird (also see
Table 1).
Figure 2. Estimated migration routes, timing, and destination for individual male Red-eyed Vireos
(n = 9) (A–I) tracked with geolocators from the Hemlock Hill, Pennsylvania, breeding population,
2011 to 2012. Dashed lines indicate periods where locations are uncertain because of equinox
periods or low-confidence sunrise-sunset transitions. The individual maps are arranged according
to time of departure from South America from earliest (A) to latest (I). One bird was omitted
because of space constraints (departed March 31, arrived May 8).
[Figures may be embedded in the manuscript or uploaded as separate files. Figure labels, axis
labels, and captions should be consistent. Make sure your figure files have part labels on them
(A, B, C, etc.) if there is more than one part. Put all figure parts into one figure file.]
Table 1. Wintering locations in South America of Red-eyed Vireos (n = 10) migrating from
northwestern Pennsylvania. Values are means (with SD in parentheses), and n is the number of
days used to estimate location. Letters correspond to maps in Figure 2.
N 1.39° (2.90)
W 64.15° (0.98)
N 0.56° (2.05)
W 64.15° (0.98)
S 3.54° 92.99)
W 69.00° (1.11)
S 3.80° (2.56)
W 65.2° (0.70)
N 1.52° (2.61)
W 59.15° (0.66)
N 1.08° (2.30)
W 62.42° (0.63)
S 0.55° (3.13)
W 69.93° (0.94)
N 3.27° (2.12)
W 62.87° (0.91)
N 7.24° (2.24)
W 64.38° (0.71)
S 0.64° (2.45)
W 60.62° (0.83)
S 3.01° (1.80)
W 63.33° (0.73
N 1.81° (1.73)
W 63.70° (0.52)
Individual changed locations during seasons; listed in chronological order.
Not depicted in Figure 2.
Page 22
Table 2. Spring migration distance, duration, and rate, and cumulative duration of stopovers in
South America, Central America, and the United States (including the Gulf of Mexico crossing
for Red-eyed Vireos (n = 10) migrating from northern South America to northwestern
Pennsylvania. Values are means (± SD).
[Any Supplemental Material should be cited in the manuscript, but not included in the
manuscript file. Upload as separate file(s) as Supplemental Material file type. Small amounts of
supplemental material should be incorporated into the paper itself, such as small figures and
tables that will altogether take up fewer than 2-3 pages, and where each item does not take up
more than 1 page of 8.5 x 11 inches.
Citations for Supplemental Material should look like this:
(see Supplemental Material); (see Supplemental Material Table S1 and Figure S1)
South America
Central America
Gulf crossing and Start-to-finish
United States
Distance (km)
1,636 ± 252
2,150 ± 234
2,848 ± 195
6,631 ± 397
Duration (days)
23.3 ± 4.7
13.1 ± 3.3
9.7 ± 2.2
45.9 ±4.6
Rate (km day−1)
72.4 ± 17.5
178.4 ± 66.9
310.0 ± 89.3
145.9 ±18.4
Stopovers (days)
18.6 ± 4.9
9.2 ± 3.3
3.5 ± 2.0
33.4 ± 4.8
Citations for Appendices within the manuscript (not a Supplemental file) should look like this:
(see Appendix Figure 5); (Appendix Table 7) ]
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