Trophic energy flows increase in more diverse communities of coastal food webs: Testing the vertical diversity hypothesis

Noboru Okuda^{1, 5}, Zin’ichi Karube², Yoichiro Sakai³, Tomohiro Takeyama⁴, Ichiro Tayasu⁵

Chikage Yoshimizu⁵, Toshi Nagata⁶

¹Research Center for Inland Seas, Kobe University, e-mail:nokuda@people.kobe-u.ac.jp;²Department of Biotechnology and Chemistry, Kindai University, e-mail:karube@hiro.kindai.ac.jp;³Lake Biwa Environmental Research Institute, e-mail:sakai-y@lberi.jp;⁴Department of Biosphere-Geosphere Science, Okayama University of Science, e-mail:takeyama@big.ous.ac.jp;⁵Research Institute for Humanity & Nature, National Institutes for the Humanities, e-mail:ichirotayasu@chikyu.ac.jpto IT, chikage@chikyu.ac.jpto CY; ⁶Atmosphere and Ocean Research Institute, The University of Tokyo, e-mail:nagata@aori.u-tokyo.ac.jp

Statement of authorship: NO took a leading role in designing the study, data analysis and writing the manuscript; ZK, SY, and TT collected the field data; IT, CY, and TN conducted the stable isotope analysis.

Data accessibility statement: Should this manuscript be accepted, unpublished data supporting the results will be archived in Dryad and the data are available fromhttps://doi.org/10.5061/dryad.rn8pk0pbk.

A short running title: Testing vertical diversity hypothesis

Keywords: Benthic macroinvertebrate, Biodiversity–Ecosystem functioning, Food chain length, Horizontal diversity, Integrated trophic position, Multitrophic system, Size-based food web, Stable isotope analysis, Trophic transfer efficiency, Vertical diversity

The type of article: Letter

The number of words in the abstract: 149 words

The number of words in the main text: 4999 words

The number of words in each text box: 386 words for supplementary boxes

The number of references: 84

The number of figures: 4

The number of tables: 0 (6 supplementary tables)

The number of text boxes: 0 (1 supplementary boxes)

Correspondence author: Noboru Okuda

Research Center for Inland Seas, Kobe University, Rokkodai 1-1, Kobe, P.O. BOX 657-8501, JAPAN

Tel +81 78 803 5733; Fax +81 78 803 5733; e-mail:nokuda@people.kobe-u.ac.jp