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Highly Diverse Mesophotic Reef Fish Communities in Raja Ampat, West Papua 3 4 5 Dominic A bioRxiv preprint doi: https://doi.org/10.1101/640490; this version posted January 15, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 1 Title: 2 Highly diverse mesophotic reef fish communities in Raja Ampat, West Papua 3 4 5 Dominic A. Andradi-Brown1,2*, Angela J. E. Beer3,4, Luigi Colin1, Hastuti3,5, Catherine E. I. 6 Head1,6, Nur Ismu Hidayat3, Steven J Lindfield7, Catherine R. Mitchell1,2, Defy N. Pada3, 7 Nikola M. Piesinger1, Purwanto8, Gabby N. Ahmadia2 8 9 1Department of Zoology, University of Oxford, Oxford, OX1 3PS, UK 10 2Ocean Conservation, World Wildlife Fund–US, 1250 24th St. NW, Washington, D.C., 20037, 11 USA 12 3Conservation International Indonesia, Sorong, Papua Barat, 98414, Indonesia 13 4School of Environment and Sustainability, Royal Roads University, Victoria, British 14 Columbia, V9B 5Y2, Canada 15 5 Marine Science, Hasanuddin University, Makassar, Sulawesi Selatan, 90245, Indonesia 16 6Institute of Zoology, Zoological Society of London, Regent’s Park, London, NW1 4RY, UK 17 7Coral Reef Research Foundation, PO Box 1765 Koror, PW, 96940, Palau 18 8Universitas Papua, Manokwari, Papua Barat, 98314, Indonesia 19 20 *Corresponding author: [email protected] bioRxiv preprint doi: https://doi.org/10.1101/640490; this version posted January 15, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 21 Abstract 22 Mesophotic coral ecosystems (MCEs; reefs 30-150 m depth) are poorly studied, with 23 existing research heavily geographically biased away from the most species-rich reef 24 regions. Yet, MCEs are of high interest because of their unique species and potential to act 25 as refuges from the impacts of fishing. Using baited remote underwater video systems, we 26 surveyed reef fish communities from depths of 2 to 85 m throughout the Raja Ampat 27 archipelago in West Papua, Indonesia – an area considered the heart of the Coral Triangle 28 where coral reef biodiversity is greatest. Here we show—similar to shallow reefs—Raja 29 Ampat MCEs are exceptionally diverse, with 152 fish species recorded at depths greater 30 than 40 m. We found that fish communities were highly depth driven, with declines in fish 31 abundance at increased depth. In contrast to previous studies elsewhere in the world, we 32 found that the proportion of planktivores declined across the shallow reef to MCE depth 33 gradient. While greater human population density correlated with lower reef fish biomass, 34 we did not observe differential impacts based on depth, and so found no evidence that 35 MCEs provide a depth refuge from fishing. Our results expand and contrast some previously 36 established MCE-depth patterns in fish communities, highlighting the need for future MCE 37 studies within the Coral Triangle region. bioRxiv preprint doi: https://doi.org/10.1101/640490; this version posted January 15, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 38 Introduction 39 Mesophotic coral ecosystems (MCEs)—reefs from approximately 30-150 m depth—are 40 of increased conservation and research interest because of their unique species 41 assemblages (Rocha et al. 2018; Pinheiro et al. 2019) and potential to act as refuges from 42 shallow reef impacts (Hinderstein et al. 2010; Loya et al. 2016). MCEs are found in tropical 43 and sub-tropical waters globally, and contain a rich array of biodiversity including corals, 44 fishes, sponges, and algae (Kahng et al. 2010; Baker et al. 2016). MCEs have been poorly 45 documented compared to shallower coral reefs because of sampling limitations at greater 46 depths. With advances in survey technology there has been a greater understanding of 47 MCEs, especially in regions such as the Caribbean, Red Sea, Australia, and Hawaii (Turner et 48 al. 2017; Laverick et al. 2018). The Coral Triangle (comprising: Indonesia, Malaysia, Papua 49 New Guinea, the Philippines, Timor Leste, and the Solomon Islands) hosts the greatest 50 shallow reef biodiversity globally (Veron et al. 2009). Yet—despite this diversity—there is 51 comparatively little research effort in the Coral Triangle (Fisher et al. 2011) especially for 52 MCEs (Turner et al. 2017; Laverick et al. 2018). Coral Triangle MCE surveys have been mostly 53 limited to several sites in the Philippines, where highly diverse MCEs have been identified 54 (Cabaitan et al. 2019; Pinheiro et al. 2019). 55 Based on ecological community composition, MCEs can be divided into upper-MCEs 56 (approximately 30-60 m depth) and lower-MCEs (approximately 60-150 m depth) (Loya et 57 al. 2016). Coral reef fish species richness, abundance, and biomass has broadly been 58 reported to decline across the depth gradient from shallow reefs to MCEs (Kahng et al. 59 2010; 2014). While reef fish communities are variable between geographical locations, 60 some patterns have begun to emerge. For example, habitat complexity is an important 61 predictor of fish abundance and biomass on MCEs (Boland and Parrish 2005; Brokovich et al. 62 2008; Bryan et al. 2013). Herbivorous fish—though present in upper-MCEs—often decline in 63 abundance with increased depth and can be near-absent from lower-MCEs (Thresher and 64 Colin 1986; Brokovich et al. 2008; Rocha et al. 2018). While planktivores, in contrast, often 65 increase as a proportion of the fish community on deeper reefs (Thresher and Colin 1986; 66 Brokovich et al. 2008; Rocha et al. 2018). 67 MCEs may act as fish refuges because shallow reefs maybe more exposed to habitat 68 damage and overfishing, or MCEs may be located at greater distances from shore and so be bioRxiv preprint doi: https://doi.org/10.1101/640490; this version posted January 15, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 69 subject to fewer human impacts (Bridge et al. 2013). Greater biomass of fisheries target 70 species has been observed on MCEs compared to adjacent shallow reefs in many locations 71 (Bejarano et al. 2014; Lindfield et al. 2016; Andradi-Brown et al. 2017). This supports the 72 idea that some MCEs may be exposed to lower fisheries pressures than adjacent shallow 73 reefs. Many small-scale fisheries gears are highly depth specific, with differential impacts 74 across small depth gradients (Ashworth and Ormond 2005; Tyler et al. 2009; Goetze et al. 75 2011). Small-scale fisheries gears that reach MCEs require additional cost and effort 76 compared to those used in the shallows (Wood et al. 2006; Lindfield et al. 2014). Yet, in 77 some regions MCEs may support important fisheries – for example, in Papua New Guinea 78 mesophotic species comprise large a proposition of some subsistence fish catches 79 (Longenecker et al. 2017; 2019). Testing potential MCE refuge effects is challenging, 80 however, as fish abundance, biomass, and community composition transitions with depth 81 (Thresher and Colin 1986; Brokovich et al. 2008; Rocha et al. 2018; Pinheiro et al. 2019) and 82 with benthic composition and structural complexity (Boland and Parrish 2005; Gratwicke 83 and Speight 2005; Bridge et al. 2012). Despite some potential protection by virtue of their 84 depth, MCEs are still vulnerable to many human impacts including some fisheries, 85 sedimentation, and global stressors such as coral bleaching and increased tropical storm 86 frequency and intensity (Andradi-Brown et al. 2016a; Rocha et al. 2018). With few—though 87 increasing—MCE studies in the Coral Triangle, human impacts on Coral Triangle MCEs 88 remain poorly understood. 89 Raja Ampat, located in West Papua, Indonesia, and at the center of the Coral Triangle 90 contains extensive reefs, as well as mangrove forests and seagrass beds (Mangubhai et al. 91 2012). Raja Ampat reefs contain the greatest richness of scleractinian corals (Veron et al. 92 2009) and fishes (Allen and Erdmann 2009) recorded in a single region – with over 550 93 scleractinian coral species and over 1,400 fish species (Mangubhai et al. 2012). Given Raja 94 Ampat’s unique diversity, there have been several focused efforts to build regional species 95 lists. For example, Allen and Erdmann (2009) combined detailed field surveys with other 96 records to produce a fish species list for the region. Scleractinian corals have been recorded 97 to 160 m depth within Raja Ampat, though reefs beyond the depths of conventional scuba 98 diving remain poorly characterized (Mangubhai et al. 2012). Unlike many reefs globally, the 99 shallow reefs of Raja Ampat have not faced widespread mortality from bleaching events and bioRxiv preprint doi: https://doi.org/10.1101/640490; this version posted January 15, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. 100 shallow reef scleractinian coral cover remained stable between 2010-2016 at approximately 101 30% (Ahmadia et al. 2017). 102 Local people living within Raja Ampat have high marine resource dependence, with 103 approximately two-thirds reliant on marine fish for over half their dietary protein needs, 104 and approximately a quarter reliant on marine capture fisheries as their primary occupation 105 (Ahmadia et al.
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