Bulletin of the Peabody Museum of Natural History

The Bulletin of the Peabody Museum of Natural History, Yale University, in publication since 1925, publishes original research in the disciplines represented by the collections of the Yale Peabody Museum’s curatorial divisions.

The Bulletin of the Peabody Museum of Natural History (ISSN 0079-032X; eISSN 2162-4135) is published twice a year, in April and October. Abstracts and full text of the electronic version of articles, beginning with Volume 47 (2006), are available online worldwide to institutional subscribers of BioOne Complete, the full-text database of more than 200 titles from over 150 society and independent publishers in the biological, ecological, and environmental sciences.

BioOne Complete

Through BioOne Complete the Bulletin of the Peabody Museum of Natural History is both discoverable in and interlinked with leading academic resources. Bulletin is archived through BioOne’s participation in Portico and LOCKSS. In addition, by inclusion in BioOne Complete, Bulletin is available at no cost to more than 2,500 institutions in the developing world through Research4Life. Publication is supported by the Theodore and Ruth Wilmanns Lidz Endowment Fund for Excellence in Scholarly Publications, dedicated to the dissemination of scholarly research and study of the world and its cultures.

Faculty, postdocs, affiliates, and students working with curatorial divisions are encouraged to submit their research for publication in the Bulletin of the Peabody Museum of Natural History. Manuscripts must be original and not previously published or simultaneously submitted elsewhere. Publication is in English only. Interested authors should contact the executive editor before submitting a manuscript.

Peer Review

All manuscripts are sent out for external peer review by one or more referees, as well as being reviewed by the executive editor in consultation with the curatorial editor-in-chief and one or more members of the appropriate Yale Peabody Museum curatorial division. Authors are encouraged to recommend and provide email addresses for up to 3 suitable reviewers.

Submissions are returned for revision with referee comments and recommendations. Revised manuscript require new and complete electronic files (including revised figures, if any). Manuscripts may go through a single or several revision cycles. Failure to make required changes can result in rejection. Unrequested changes without written justification may be treated as a new submission if substantial, or may result in rejection. Delays may result if materials are not in proper form. Keep copies of all materials submitted.

Manuscripts are copyedited for Bulletin style and format once peer review is complete and revisions incorporated. Submissions are published approximately in the order in which they are accepted for publication. A digital object identifier (DOI) is assigned with online publication.

Supplementary Files

Bulletin accepts supplementary files meant to accompany published articles. Files must be declared on submission of manuscripts and made available for peer review. Supplementary material may include, but is not limited to, multimedia such as video or audio clips, datasets, tables, and additional figures. Specialist filetypes can often be accommodated, but keep in mind that only users with the relevant software will be able to download such files. Total file size is typically limited to 50MB. A DOI will be registered for supplementary material when posted, associating it with the source article. There is no cost for supplementary material hosting.

Page Proofs

Copyedited and formatted PDF page proofs are provided for online review to the corresponding author through a vendor online proofing system. The corresponding author is responsible for the timely return of these proofs, to include comments from all co-authors. These proofs are for checking printer’s errors only and for resolving queries or missing information. Only trivial revisions can be made at this stage. Extensive revision, including changes in page composition, will not be accepted unless the author is willing to cover the costs of any such revisions.

Manuscript Preparation

Attention to Bulletin editorial policies and manuscript and figure preparation recommendations will substantially expedite production of your submission. Carefully review the Bulletin manuscript preparation requirements given here. Delays may result if materials are not in proper form. Include our author information form with full contact information, email addresses, and ORCiD identifiers for all contributors with your submission. Designate a corresponding author and indicate who should receive editorial correspondence and page proofs if it is not the lead author.

Manuscripts are copyedited for Bulletin style and format after all reviewer revisions have been incorporated and a final revised and complete manuscript has been received. Keep copies of all submitted files. The Bulletin is not responsible for lost or corrupted files.


To enhance online accessibility, the Bulletin recommends an ORCiD identifier (Open Researcher and Contributor ID) be provided for all listed authors, along with funding source information, including award numbers. Provide this information on our author information form with submission of your manuscript.

Page Charges

There are currently no page charges. Authors are encouraged to support the Bulletin with a voluntary donation toward the cost of publication.

Type Material & Vouchers

Whenever possible, all primary name-bearing types (such as holotypes, neotypes, and lectotypes) based on Yale Peabody Museum specimens designated in the Bulletin should be deposited in the appropriate collection at the Yale Peabody Museum. Secondary types (such as paratypes) should be deposited in major institutional collections. Voucher specimens from ecological or related work should be deposited in whole or major part in the appropriate collection at the Yale Peabody Museum.


All published articles are copyrighted in the name of the Peabody Museum of Natural History on behalf of Yale University. To protect its publications and facilitate wide distribution and archiving, it is the policy of the Yale Peabody Museum to require that contributors assign their copyright interests in their work, including electronic rights, and rights in all renewals and extensions of copyright. The Yale Peabody Museum in turn grants contributors the right to re-use their work provided that all such use is for the personal noncommercial benefit of the author(s), with attribution to Bulletin. Completed and signed agreements, provided by the lead author on behalf of all co-authors, are due with submission of revised accepted manuscripts.


Authors are responsible for obtaining and submitting written documentation of rights granted from the copyright holders, at the authors’ expense, for any previously published or copyrighted materials used in their articles not owned by the authors. Full credit and attribution for such materials must be included in captions or the acknowledgments. It is the responsibility of the authors to make copyright holders aware that Bulletin articles are made available online after publication. Such permission statements should grant both print and electronic worldwide rights in perpetuity. Download our sample permission request letter here.

For guidance on rights usage and permissions see the Chicago Manual of Style, 17th ed., Chapter 4.

All Google Maps and Google Earth imagery must be at high resolution and have attribution. Follow Google’s requirements. Rights clearance documentation is not required, but the original credit and copyright information shown on the images must be retained (at a size legible in the final version) or included in a caption.

Complimentary Copies

Each author receives 1 printed copy of the Bulletin of the Peabody Museum of Natural History issue containing his or her contribution(s), on publication. Contributors also receive a free PDF file of their published paper for the personal, noncommercial use of the author(s).

Submit manuscripts to the executive editor only as digital files (including figures and supplementary files, if any) by electronic mail or through an online cloud service for files larger than 20MB. Contact the publications manager for assistance.

Include all illustrative materials (charts, drawings, and photographs) as separate image files with your initial submission, so that resolution and compatibility can be assessed. Follow Bulletin requirements for figures or contact the publications manager for guidance. Bulletin reserves the right to reject manuscripts that do not conform to these requirements.


Article titles should be short (no more than 15 words), precise, and straightforwardly descriptive, succinctly conveying the manuscript’s content. Names of new taxa should not be used in the title. Avoid abbreviations, acronyms, and hyphenated terms and excessive punctuation.

Author Affiliations

On the first page of the manuscript file provide the full name and institutional affiliation (with city, state or province, and country, but not street address) for each contributor. Designate a corresponding author with an email address.


Provide an informative abstract of up to 300 words that conforms to the requirements of BioAbstracts. Do not use abbreviations or acronyms in the abstract unless widely understood (e.g., DNA, pH). Do not include citations. Authors have the option of also including foreign language abstracts as appropriate.


Provide a list of up to 10 keywords or terms, not used in the title, following the abstract that give the central topics of the article.


The first mention of an animal or plant in the text should include the full scientific name and other pertinent attribution. Bulletin recommends including a reference for these in the Literature Cited. Italicize all generic and specific names. Nomenclature should conform to the International Code of Zoological Nomenclature for zoological studies and the International Code of Botanical Nomenclature for botanical research.


Write with precision, clarity, and economy for an audience that could include students, early career professionals, and interested nonspecialists. Write in the active voice and first person. Using the passive voice sparingly and avoid jargon. Main headings and subheadings should be explicit, descriptive, and short. Hierarchical levels should be parallel and consistent. Avoid more than 4 outline levels. Use at least 2 headings in each level. Do not use all capital letters for titles, headings, or subheadings. Manuscripts should contain these main sections:

Materials and Methods
Discussion or Conclusion
Appendices (if present)
Literature Cited

Style & Format

Bulletin uses The Chicago Manual of Style, 17th edition, as its primary style manual. For specific formatting not discussed here, consult the latest edition of Scientific Style and Format, for specific examples in the sciences, and the Society for American Archaeology Style Guide for archaeology and anthropology conventions. We strongly recommend examining a recent Bulletin issue for formatting guidance. Manuscripts will be copyedited to Bulletin style and for clarity and readability. Bulletin reserves the right to adjust style and format to meet the specifications of Yale Peabody Museum publications.

Italicize only species names and non-English terms; do not underline, bold, or use other text formatting. Flag special characters, accents, and symbols, particularly in graphic files, that require a special font with a note both in the manuscript and in your submission letter.

Avoid abbreviations and acronyms unless well established and used widely in the scientific literature (for example, DNA, pH). Define parenthetically less known acronyms and abbreviations at first use, but if only used a few times spell these terms out instead. Avoid the excessive use of Latin terms and abbreviations.

Institutional abbreviations and acronyms, such as catalog number prefixes, should be listed and defined in the “Materials and Methods” section. Alternatively, reference the Standard Symbolic Codes for Institutional Resource Collections in Herpetology and Ichthyology, as appropriate.

Follow the International System of Units (SI) for measurements. Provide metric equivalents when non-SI units are necessary. Use the 24-hour clock (0930, 2315), give temperature in degrees Celsius (32 °C) and format dates as 16 June 2021.

Spell out numbers from one to nine (except in a series with other numbers over 10); use numerals for 10 and above and very large numbers (250 million, 2.2 billion, except one million). Use numerals for zero and one with units of measure, but spell out if used alone in text. Use math symbols only in equations, not in prose (by not x; is not =; more than 40 objects not >40 objects or 40+ objects).

Complex mathematical formulas should be created using the equation editor in Microsoft Word, with MathType, or provided as separate vector or raster image files (see Bulletin requirements given here).

For statistical notation, use an italicized, uppercase N for the number of subjects or participants in a total sample (N = 328); use an italicized, lowercase n in reference to a portion of the sample (n = 42).

Designate Yale Peabody Museum specimen and object numbers with the Museum’s 6-digit format and divisional prefix, as follows: YPM PREFIX 000000. Use full numbers in ranges separated by an en dash. For example, YPM ANT 017898; YPM VPPU 016235, 016245; YPM HERR 003873–003881.

Do not use footnotes in text or figures, but incorporate these as parentheticals into the text or in captions. Use lowercase letter footnotes in tables, especially to designate source materials.


Tabular material should be typed separately as separate electronic tab-delimited word processing files and not placed into the main text; spreadsheet files are also acceptable. Avoid nested tables. Number tables consecutively using Arabic numerals, separate from figures. All tables must be cited in the main text. Do not use embedded graphics, vertical lines, vertical writing, or all capital letters. Do not submit tables as image files.

For guidance on designing data in standard table formats for see The Chicago Manual of Style, 17th edition, Chapter 3.


Captions are to provide all explanatory text for figures and tables. Submit captions double-spaced in a clearly labeled separate electronic file (not attached to illustrations or tables), with a separate paragraph devoted to each figure or table number. Provide a short introductory phrase or a formal title for each figure or table; do not place titles, legends, and footnotes in the image itself.

All figures and tables must be cited in the main text and numbered consecutively using Arabic numerals according to first mention. Number tables separately from figures. Do not split tables and figures into parts.

Attribution and credit for illustrations or photographs that are not your own must be provided in the caption, or listed in the acknowledgments. Acknowledge sources for figures based on other work with “From Smith 2001” for work substantially reprinted as first published, or with “Modified from Smith 2001” for work adapted from previously published material with changes.


Include information on granting agencies, and on individuals and institutions that have lent or provided access to materials, specimens, financial support, field assistance or editorial review, among others, in a separate Acknowledgments section at the end of the main text. Provide grant numbers when available. Spell out names of individuals, institutions, and agencies.

In-text Citations & Literature Cited

The Bulletin follows the name-year system recommended in the Chicago Manual of Style, 17th edition; see specific examples listed there. We recommend consulting a recent issue for guidance.

List citations in the text chronologically from earliest to latest; citations in the same year are listed alphabetically. For more than 3 authors, use “et al.” For articles with different multiple authors in the same year, list as many authors as necessary to distinguish the citations (Gupta, Collinson et al. 2006; Gupta, Michels et al. 2006). Specific pages and figures should use the format: Mayr (1963:236, fig. 4); Mayr (1963, tbl. 4).

Every citation in the text must be referenced in the reference list, including internet sources, repositories, software, and databases. Provide DOIs (digital object identifiers with the format https://doi.org/…) or persistent URLs for online references; include date of access and availability information for hard-to-find sources.

List references in the Literature Cited alphabetically by author surname, then chronologically from earliest to latest. References with identical lists of authors in the same year are designated as 2010a, 2010b, etc. Provide the names of all authors, editors, and translators as appropriate (use “et al.” for names beyond the 10th). Do no capitalize or use small capitals for author names.

Spell out journal titles in full. Use lowercase sentence format for journal article titles, upper and lowercase title case for book titles; do not italicize.

Unpublished reports, manuscripts in preparation, or other sources that are not accessible to researchers are cited directly in the text and not in the Literature Cited. Cite personal knowledge as: A.B. Smith, pers. obs.; A.B. Smith, pers. comm., 2 January 2020; or A.B. Smith, unpubl. data; A.B. Smith, in prep. Include forthcoming articles in the Literature Cited as “in press” only if under contract to be published.

Provide all illustrations, charts, and photographs at the time of submission as digital files scanned or prepared to professional standards. For best practices on designing figures see The Chicago Manual of Style, 17th edition, Chapter 3.

Authors are responsible for providing high-resolution electronic image files of acceptable quality. Prepare figures at just above 1:1 reproduction size to fit the width of the publication page (6 inches; 15 centimeters). For technical guidance see the recommendations at KnowledgeWorks Global Ltd. Digital Art Support.

Acceptable raster file formats are TIF, JPG, and PSD (Adobe Photoshop). Minimum raster requirements at 1:1 reproduction size:

Monochrome (black-and-white only) images such as line graphs and drawings: 1200 ppi

Color or grayscale photographs (RGB, CMYK, and grayscale halftones: 300 ppi

Combination halftones images with both photographs and screens, text, lines, or illustrations: 600 ppi (800 ppi preferred)

Acceptable vector file formats are EPS and AI (Adobe Illustrator).

Recommended for charts, text added to photos, and illustrations.

Vector files that include photographs must have all images embedded (not linked).

Embedded images must follow the resolution requirements for raster files.

Use a commonly available standard sans serif font such as Myriad Pro, Helvetica, or Arial. Use only bold uppercase letters for labels. Be consistent in the style and placement of labels throughout all figures; do not use special graphic effects such as shadows or outlining. If special fonts (e.g., for symbols on maps) are used, convert all fonts to outlines and save as EPS.

Other formats (PNG, GIF, SVG) and charts and illustrations created in Microsoft Excel and other Office programs are not accepted. Do not submit final images as a figures placed into a Microsoft document.

Authors may choose to submit color figures as RGB (best for online viewing) or CYMK (best for print) color mode. Note that RGB-to-CMYK and CYMK-to-RGB conversion by the vendor may shift some colors in the final reproduction.

In preparing color figures, especially charts and maps, we recommend optimizing accessibility for users with color vision impairment, including people with color blindness, through Photoshop soft-proofing and Color Universal Design.

Figures created from specialized software should be exported for print publication to meet these requirements as recommended by the software vendor. Original camera images should be taken at the highest resolution setting and size available, preferably as TIF files. Scanned materials should meet the raster image requirements above, and use a descreening filter if scanned from a published print source.

Guidance on creating maps to professional standards is available at ESRI’s ArcGIS Blog and the Axis Maps Cartography Guide.

Bulletin of the Peabody Museum of Natural History is available online worldwide to institutional subscribers of the full-text database BioOne Complete and is provided as an exchange journal to libraries and institutions worldwide through the Yale University Library.

The Bulletin is also available as a separate print subscription. Subscriptions are for the calendar year and include shipping.

Annual Subscription Rates

$145.00 (US addresses)
$175.00 (non-US addresses)

All prices are in US dollars. All orders must be prepaid. Payment is due by January 31 of the subscription year; there are no refunds for mid-year cancellations.

Checks must be in US dollars drawn on a US bank and payable to Yale University.

Institutional purchase orders are accepted.

We accept VISA and MasterCard. Please fax or call with your order and do not send payment information by email.

Contact us for wire transfer instructions. There is a US$35.00 fee for wire transfers in addition to the subscription fee.

Individual print copies of past issues can be purchased, while supplies last, with our order form. Please allow 2 to 4 weeks for the processing of your order.

Single Issue Price

$48.50 (plus shipping)

Individual articles are available for purchase through BioOne Complete.

Returns and claims for missing issues purchased directly from Yale Peabody Museum Publications are accepted within 6 months.

Returns and claims for defective or damaged copies must be made on receipt. Contact us in advance for authorization and instructions.

The Bulletin of the Peabody Museum of Natural History was published as a monograph series from 1926 through 1999 (Volumes 1–46). Bulletin became a journal in 2006, beginning with Volume 47.

Archived monographs, along with the Museum’s former Postilla series of short papers, are available as free downloadable files on EliScholar, a digital platform for scholarly publishing provided by Yale University Library.

Editorial Board

Jacques A. Gauthier
Curatorial Editor-in-Chief
Professor of Earth & Planetary Sciences
Curator of Vertebrate Paleontology & Vertebrate Zoology, Reptiles

Derek E.G. Briggs
G. Evelyn Hutchinson Professor of Earth & Planetary Sciences
Curator-in-Charge, Invertebrate Paleontology

Eric J. Sargis
Professor of Anthropology, Ecology & Evolutionary Biology
Curator of Vertebrate Zoology, Mammals

David K. Skelly
Frank R. Oastler Professor of Ecology
Director of the Peabody Museum of Natural History
Curator of Vertebrate Zoology, Amphibians

Tim White
Director of Collections & Research


Patrick Sweeney Executive Editor: Peabody Bulletin,
Rosemary Volpe Publications Manager,
Shannon Giacobbe Editorial Assistant,
+1 203 432 3154 shannon.giacobbe@yale.edu

Mailing address:

Publications Office
Yale Peabody Museum of Natural History

P.O. Box 208118
New Haven CT 06520-8118 USA