Diel vertical migration promotes zooplankton horizontal patchiness

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ORIGINAL ARTICLE

Diel vertical migration promotes zooplankton horizontal patchiness Bingzhang Chen1,2,3   · Eiji Masunaga4 · Sherwood Lan Smith5 · Hidekatsu Yamazaki1,6 Received: 18 September 2019 / Revised: 29 July 2020 / Accepted: 30 August 2020 © The Author(s) 2020

Abstract Spatial patchiness of plankton enhances fishery production and carbon export in the ocean. While diel vertical migration (DVM) has been identified as an important factor contributing to vertical patchiness, its effect on horizontal patchiness has never been investigated. We use a simple individual-based zooplankton model to examine the effect of DVM on the horizontal patchiness of four zooplankton groups with differing DVM patterns in a two-dimensional ocean circulation model. We find that zooplankton horizontal patchiness can be induced by two mechanisms: (1) in stratified waters, DVM can synchronize zooplankton vertical positions with the horizontal current velocities that drive them, resulting in horizontal patchiness; and (2) migrating zooplankton tend to aggregate in deep waters when they encounter sea bottom. Due to these mechanisms, zooplankton horizontal patchiness may be ubiquitous in the ocean, enhancing secondary production and fisheries. Keywords  Internal wave · Individual based model · Patchiness · Stratification · Seamount · Dispersion

1 Introduction Spatial patchiness of plankton is a key factor enhancing fishery production and carbon export in the ocean (Okubo and Levin 2001; Franks 2005; Benoit-Bird and McManus 2012; Woodson and Litvin 2015). The formation of plankton

Electronic supplementary material  The online version of this article (https​://doi.org/10.1007/s1087​2-020-00564​-4) contains supplementary material, which is available to authorized users. * Bingzhang Chen [email protected] 1



Department of Ocean Sciences, Tokyo University of Marine Science and Technology, Tokyo, Japan

2



Department of Mathematics and Statistics, University of Strathclyde, Glasgow, UK

3

Hong Kong Branch of Southern Marine Science and Engineering Guangdong Laboratory (Guangzhou), Hong Kong University of Science & Technology, Hong Kong, China

4

Global and Local Environment Co‑Creation Institute, Ibaraki University, Mito, Japan

5

Research Institute for Global Change, Earth Surface System Research Center, Japan Agency for Marine-Earth Science and Technology, Yokosuka, Japan

6

Alpha Hydraulic Engineering Consultants Co. Ltd., Chuoh‑ku, Tokyo 104‑0045, Japan







patchiness requires some mechanisms for plankton to overcome turbulent diffusion (Abraham 1998). These mechanisms can be broadly classified into two categories: physical and biological. For example, Woodson and Litvin (2015) have shown that convergent currents at ocean fronts create hotspots of plankton, which in turn attract active, larger animals such as fish, sea birds and marine mammals. However, Folt and Burns (1999) argued that physical mechanisms alone cannot fully explain patterns of plankton patchiness. They proposed four biological mechanisms tha