{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,2]],"date-time":"2026-04-02T11:15:53Z","timestamp":1775128553339,"version":"3.50.1"},"reference-count":141,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2023,12,8]],"date-time":"2023-12-08T00:00:00Z","timestamp":1701993600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Mar. Sci."],"abstract":"<jats:sec><jats:title>Introduction<\/jats:title><jats:p>The production of resting cysts is a key dispersal and survival strategy of many dinoflagellate species. However, little is known about the role of suspended cysts in the benthic nepheloid layer (BNL) in the initiation and decline of planktonic populations.<\/jats:p><\/jats:sec><jats:sec><jats:title>Methods<\/jats:title><jats:p>In September 2019, sampling of the dinoflagellate cyst community at different water depths in the water column and in the bottom sediments, and studies of spatio-temporal changes in physical properties (temperature, salinity, density and suspended sediment concentration), were carried out along a land-sea transect off Figueira da Foz (NW Portugal) to investigate the dinoflagellate cyst distribution and the factors (physical and biological) affecting it. A clustering analysis was used to compare the BNL and sediment cyst records with the cyst rain recorded by a sediment trap at a fixed station. Furthermore, Lagrangian particle experiments enabled simulating cyst trajectories in the BNL 5 and 10 days before sampling and assessing cross-shore, vertical and alongshore transport within the studied region.<\/jats:p><\/jats:sec><jats:sec><jats:title>Results<\/jats:title><jats:p>A well-developed BNL was present during the survey, which covered a change from active (14<jats:sup>th<\/jats:sup> of September) to relaxed (19<jats:sup>th<\/jats:sup> of September) upwelling conditions. Organic-walled dinoflagellate cysts were dominant in all samples, although calcareous dinoflagellate cysts consistently occurred (at low abundances). High proportions of full cysts were observed in the BNL, of which a significant portion was viable as shown by excystment experiments. Moreover, BNL cyst records collected on the 19<jats:sup>th<\/jats:sup> of September along the land-sea transect were similar to the sediment trap cyst record but greatly differed from sediment cyst records. The heterotrophic small spiny brown cysts (SBC) and cysts of the autotrophic yessotoxin-producer <jats:italic>Protoceratium reticulatum<\/jats:italic> notably increased during the survey, in the BNL and in the water column above.<\/jats:p><\/jats:sec><jats:sec><jats:title>Discussion<\/jats:title><jats:p>The comparison of the BNL, surface sediment and sediment trap cyst records supported that the main origin of cysts in the BNL was the recent production in the water column. The spatial coincidences in the distribution of cysts and vegetative cells of <jats:italic>Protoceratium reticulatum<\/jats:italic> also supported that full cysts in the water column were being produced in surface waters. New data evidenced the presence of a significant reservoir of viable cysts in the BNL that have the potential to seed new planktonic blooms. Furthermore, back-track particle modelling evidenced that alongshore advection was the main physical mechanism controlling cyst dynamics in the BNL during most part of the survey period, being particularly intense in coastal stations (&amp;lt;100 m depth). Consequently, the sediment cyst signal is a mixture of locally and regionally produced cysts. We provide multi-disciplinary data evidencing that cysts recently formed in the photic zone can be laterally advected within the studied region through the BNL, contributing to a better understanding of the role of the BNL in cyst dynamics and tracing the seed sources of the new blooms.<\/jats:p><\/jats:sec>","DOI":"10.3389\/fmars.2023.1270343","type":"journal-article","created":{"date-parts":[[2023,12,8]],"date-time":"2023-12-08T12:30:36Z","timestamp":1702038636000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":6,"title":["New physical and biological evidence of lateral transport affecting dinoflagellate cyst distribution in the benthic nepheloid layer along a land-sea transect off Figueira da Foz (Atlantic Iberian margin)"],"prefix":"10.3389","volume":"10","author":[{"given":"Iria","family":"Garc\u00eda-Moreiras","sequence":"first","affiliation":[]},{"given":"Melissa","family":"Hatherly","sequence":"additional","affiliation":[]},{"given":"Karin","family":"Zonneveld","sequence":"additional","affiliation":[]},{"given":"Jesus","family":"Dubert","sequence":"additional","affiliation":[]},{"given":"Rita","family":"Nolasco","sequence":"additional","affiliation":[]},{"given":"Ana Isabel","family":"Santos","sequence":"additional","affiliation":[]},{"given":"Anabela","family":"Oliveira","sequence":"additional","affiliation":[]},{"given":"Teresa","family":"Moita","sequence":"additional","affiliation":[]},{"given":"Paulo B.","family":"Oliveira","sequence":"additional","affiliation":[]},{"given":"Jorge M.","family":"Magalh\u00e3es","sequence":"additional","affiliation":[]},{"given":"Ana","family":"Amorim","sequence":"additional","affiliation":[]}],"member":"1965","published-online":{"date-parts":[[2023,12,8]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1016\/S0399-1784(99)80055-3","article-title":"Water column and recent sediment data on diatoms and coccolithophorids, off Portugal, confirm sediment record of upwelling events","volume":"22","author":"Abrantes","year":"1999","journal-title":"Oceanol. Acta"},{"key":"B2","doi-asserted-by":"publisher","first-page":"849","DOI":"10.1016\/j.hal.2008.04.007","article-title":"Renewal time and the impact of harmful algal blooms on the extensive mussel raft culture of the Iberian coastal upwelling system (SW Europe)","volume":"7","author":"\u00c1lvarez-Salgado","year":"2008","journal-title":"Harmful Algae"},{"key":"B3","unstructured":"AmorimA.\n          LisbonUniversity of Lisbon, PortugalDinoflagellate cyst distribution along the coast of Portugal. PhD thesis2001"},{"key":"B4","doi-asserted-by":"publisher","first-page":"572","DOI":"10.2216\/i0031-8884-40-6-572.1","article-title":"Gymnodinium catenatum-like cysts (Dinophyceae) in recent sediments from the coast of Portugal","volume":"40","author":"Amorim","year":"2002","journal-title":"Phycologia"},{"key":"B5","first-page":"89","article-title":"Dinoflagellate blooms related to coastal upwelling plumes off Portugal","volume-title":"Harmful Algae 2002","author":"Amorim","year":"2004"},{"key":"B6","first-page":"133","article-title":"On a Lingulodinium polyedrum bloom in Set\u00fabal bay, Portugal","author":"Amorim","year":"2001"},{"key":"B7","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1111\/j.1529-8817.1980.tb03013.x","article-title":"Effects of temperature conditioning on development and germination of Gonyaulax tamarensis (Dinophyceae) hypnozygotes","volume":"16","author":"Anderson","year":"1980","journal-title":"J. Phycol."},{"key":"B8","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1021\/bk-1984-0262.ch011","article-title":"Shellfish toxicity and dormant cysts in toxic dinoflagellate blooms","volume-title":"Seafood Toxins","author":"Anderson","year":"1984"},{"key":"B9","doi-asserted-by":"crossref","first-page":"1000","DOI":"10.4319\/lo.1985.30.5.1000","article-title":"Sinking characteristics of dinoflagellate cysts 1","volume":"30","author":"Anderson","year":"1985","journal-title":"Limnol. Oceanogr."},{"key":"B10","first-page":"3","article-title":"HABs in a changing world: a perspective on harmful algal blooms, their impacts, and research and management in a dynamic era of climatic and environmental change","author":"Anderson","year":"2014"},{"key":"B11","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1111\/j.1529-8817.1988.tb00085.x","article-title":"The unique micro reticulate cyst of the naked dinoflagellate Gymnodinium catenatum","volume":"24","author":"Anderson","year":"1988","journal-title":"J. Phycol."},{"key":"B12","doi-asserted-by":"publisher","first-page":"6","DOI":"10.1016\/j.dsr2.2013.10.002","article-title":"Alexandrium fundyense cysts in the Gulf of Maine: long-term time series of abundance and distribution, and linkages to past and future blooms","volume":"103","author":"Anderson","year":"2014","journal-title":"Deep Sea Res. Part II: Topical Stud. Oceanogr."},{"key":"B13","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1016\/0302-3524(79)90098-7","article-title":"The seeding of two red tide blooms by the germination of benthic Gonyaulax tamarensis hypnozygotes","volume":"8","author":"Anderson","year":"1979","journal-title":"Estuar. Coast. Mar. Sci."},{"key":"B14","doi-asserted-by":"publisher","first-page":"2522","DOI":"10.1016\/j.dsr2.2005.06.014","article-title":"Alexandrium fundyense cyst dynamics in the Gulf of Maine","volume":"52","author":"Anderson","year":"2005","journal-title":"Deep Sea Res. Part II: Topical Stud. Oceanogr."},{"key":"B15","doi-asserted-by":"publisher","first-page":"340","DOI":"10.4319\/lo.1987.32.2.0340","article-title":"The effects of darkness and anaerobiosis on dinoflagellate cyst germination","volume":"32","author":"Anderson","year":"1987","journal-title":"Limnol. Oceanogr."},{"key":"B16","doi-asserted-by":"publisher","first-page":"46","DOI":"10.5670\/oceanog.2017.109","article-title":"Harmful algal blooms in fjords, coastal embayments, and stratified systems: Recent progress and future research","volume":"30","author":"Berdalet","year":"2017","journal-title":"Oceanography"},{"key":"B17","first-page":"287","article-title":"Cyst germination of two dinoflagellate species from Galicia (NW Spain)","volume":"54","author":"Blanco","year":"1990","journal-title":"Sci. Mar."},{"key":"B18","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1093\/plankt\/17.1.165","article-title":"Cyst production in four species of neritic dinoflagellates","volume":"17","author":"Blanco","year":"1995","journal-title":"J. Plankton Res."},{"key":"B19","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1093\/plankt\/17.2.283","article-title":"The distribution of dinoflagellate cysts along the Galician (NW Spain) coast","volume":"17","author":"Blanco","year":"1995","journal-title":"J. Plankton Res."},{"key":"B20","doi-asserted-by":"publisher","first-page":"472","DOI":"10.2216\/i0031-8884-36-6-472.1","article-title":"The use of sodium polytungstate for the separation and concentration of living dinoflagellate cysts from marine sediments","volume":"36","author":"Bolch","year":"1997","journal-title":"Phycologia"},{"key":"B21","doi-asserted-by":"publisher","first-page":"215","DOI":"10.2216\/i0031-8884-30-2-215.1","article-title":"The resting cyst of the red-tide dinoflagellate Alexandrium minutum (Dinophyceae)","volume":"30","author":"Bolch","year":"1991","journal-title":"Phycologia"},{"key":"B22","doi-asserted-by":"publisher","first-page":"1095","DOI":"10.1016\/j.scitotenv.2019.01.314","article-title":"Seasonal and multi-annual trends of bivalve toxicity by PSTs in Portuguese marine waters","volume":"664","author":"Botelho","year":"2019","journal-title":"Sci. Total Environ."},{"key":"B23","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1093\/plankt\/16.5.513","article-title":"The effects of temperature, growth medium and darkness on excystment and growth of the toxic dinoflagellate Gymnodinium catenatum from northwest Spain","volume":"16","author":"Bravo","year":"1994","journal-title":"J. Plankton Res."},{"key":"B24","doi-asserted-by":"publisher","first-page":"11","DOI":"10.3390\/microorganisms2010011","article-title":"Towards an ecological understanding of dinoflagellate cyst functions","volume":"2","author":"Bravo","year":"2014","journal-title":"Microorganisms"},{"key":"B25","doi-asserted-by":"publisher","first-page":"222","DOI":"10.1016\/j.dsr2.2009.09.004","article-title":"Bloom dynamics and life cycle strategies of two toxic dinoflagellates in a coastal upwelling system (NW Iberian Peninsula)","volume":"57","author":"Bravo","year":"2010","journal-title":"Deep Sea Res. Part II: Topical Stud. Oceanogr."},{"key":"B26","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1080\/09670260600810360","article-title":"Resting cysts of the toxigenic dinoflagellate genus Alexandrium in recent sediments from the Western Mediterranean coast, including the first description of cysts of A. kutnerae and A. Peruvianum","volume":"41","author":"Bravo","year":"2006","journal-title":"Eur. J. Phycol."},{"key":"B27","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/j.pocean.2018.12.007","article-title":"Dinoflagellate cyst production in the Cariaco Basin: a 12.5 year-long sediment trap study","volume":"171","author":"Bringu\u00e9","year":"2019","journal-title":"Prog. Oceanogr."},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1016\/j.hal.2019.101728","article-title":"Cyst-forming dinoflagellates in a warming climate","volume":"91","author":"Brosnahan","year":"2020","journal-title":"Harmful Algae"},{"key":"B29","doi-asserted-by":"publisher","first-page":"2829","DOI":"10.1002\/lno.10664","article-title":"Bloom termination of the toxic dinoflagellate Alexandrium catenella: vertical migration behaviour, sediment infiltration, and benthic cyst yield","volume":"62","author":"Brosnahan","year":"2017","journal-title":"Limnol. Oceanogr."},{"key":"B30","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/j.1365-2419.2007.00447.x","article-title":"The influence of diel vertical migration on zooplankton transport and recruitment in an upwelling region: estimates from a coupled behavioral-physical model","volume":"17","author":"Carr","year":"2008","journal-title":"Fish. Oceanogr."},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1016\/j.pocean.2020.102480","article-title":"Fixed-point time series, repeat survey and high-resolution modelling reveal event scale responses of the Northwestern Iberian upwelling","volume":"190","author":"Cordeiro","year":"2021","journal-title":"Prog. Oceanogr."},{"key":"B32","doi-asserted-by":"publisher","first-page":"770","DOI":"10.1016\/j.hal.2006.03.006","article-title":"Downwelling and dominance of autochthonous dinoflagellates in the NW Iberian margin: the example of the R\u00eda de Vigo","volume":"5","author":"Crespo","year":"2006","journal-title":"Harmful Algae"},{"key":"B33","first-page":"43","article-title":"Sedimentary dynamics of the Mondego estuary","volume-title":"Aquatic Ecology of the Mondego River Basin. Global Importance of Local Experience","author":"Cunha","year":"2002"},{"key":"B34","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1016\/0034-6667(76)90010-5","article-title":"Cyst formation, sedimentation, and preservation: factors affecting dinoflagellate assemblages in recent sediments from Trondheimsfjord, Norway","volume":"22","author":"Dale","year":"1976","journal-title":"Rev. Palaeobot. Palynol."},{"key":"B35","first-page":"69","article-title":"Dinoflagellate resting cysts:\u201d benthic plankton\u201d","volume-title":"Survival strategies of the algae","author":"Dale","year":"1983"},{"key":"B36","first-page":"1249","article-title":"Dinoflagellate cyst ecology: modelling and geological applications","volume-title":"Palynology: principles and applications","author":"Dale","year":"1996"},{"key":"B37","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.hal.2014.10.001","article-title":"Coupling planktonic and benthic shifts during a bloom of Alexandrium catenella in southern Chile: Implications for bloom dynamics and recurrence","volume":"40","author":"Diaz","year":"2014","journal-title":"Harmful algae"},{"key":"B38","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1016\/j.hal.2009.12.002","article-title":"Bloom dynamics of Dinophysis acuta in an upwelling system: in situ growth versus transport","volume":"9","author":"Escalera","year":"2010","journal-title":"Harmful Algae"},{"key":"B39","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1357\/002224008787536790","article-title":"Cross-shelf structure of coastal upwelling: A two-dimensional extension of Ekman\u2019s theory and a mechanism for inner shelf upwelling shut down","volume":"66","author":"Estrade","year":"2008","journal-title":"J. Mar. Res."},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.3390\/rs13040675","article-title":"Assessing phytoplankton bloom phenology in upwelling-influenced regions using ocean color remote sensing","volume":"13","author":"Ferreira","year":"2021","journal-title":"Remote Sens."},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0197627","article-title":"Transient response of the Northwestern Iberian upwelling regime","volume":"13","author":"Ferreira Cordeiro","year":"2018","journal-title":"PloS One"},{"key":"B42","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1023\/A:1021309222459","article-title":"Coastal upwelling, primary production and mussel growth in the R\u00edas Baixas of Galicia","volume":"484","author":"Figueiras","year":"2002","journal-title":"Hydrobiologia"},{"key":"B43","doi-asserted-by":"publisher","first-page":"219","DOI":"10.3354\/meps076219","article-title":"Microplankton assemblages in three Rias Baixas (Vigo, Arosa and Muros, Spain) with a subsurface chlorophyll maximum: their relationships to hydrography","volume":"76","author":"Figueiras","year":"1991","journal-title":"Marine. Ecol. Prog. Ser."},{"key":"B44","doi-asserted-by":"publisher","first-page":"370","DOI":"10.1111\/j.1529-8817.2005.04150.x","article-title":"Sexual reproduction and two different encystment strategies of Lingulodinium polyedrum (dinophyceae) in culture","volume":"41","author":"Figueroa","year":"2005","journal-title":"J. Phycol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"13","DOI":"10.3354\/ame01206","article-title":"New life-cycle stages of Gymnodinium catenatum (Dinophyceae): laboratory and field observations","volume":"52","author":"Figueroa","year":"2008","journal-title":"Aquat. Microbial. Ecol."},{"key":"B46","doi-asserted-by":"publisher","first-page":"645","DOI":"10.1016\/j.protis.2018.06.001","article-title":"Quantitative response of Alexandrium catenella cyst dormancy to cold exposure","volume":"169","author":"Fischer","year":"2018","journal-title":"Protist"},{"key":"B47","first-page":"31","article-title":"Climatological space and time variation of the Portuguese coastal upwelling","volume":"5","author":"Fi\u00faza","year":"1982","journal-title":"Oceanol. Acta"},{"key":"B48","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1093\/plankt\/fbz026","article-title":"Mixotrophic protists and a new paradigm for marine ecology: where does plankton research go now","volume":"41","author":"Flynn","year":"2019","journal-title":"J. Plankton Res."},{"key":"B49","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1016\/0272-7714(88)90093-5","article-title":"Influence of upwelling relaxation on dinoflagellates and shellfish toxicity in Ria de Vigo, Spain","volume":"27","author":"Fraga","year":"1988","journal-title":"Estuar. Coast. Shelf Sci."},{"key":"B50","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/S0025-3227(01)00126-8","article-title":"Influence of lateral particle advection and organic matter degradation on sediment accumulation and stable nitrogen isotope ratios along a productivity gradient in the Canary Islands region","volume":"177","author":"Freudenthal","year":"2001","journal-title":"Mar. Geol."},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.3389\/fmars.2021.699483","article-title":"Environmental factors affecting spatial dinoflagellate cyst distribution in surface sediments off aveiro-figueira da foz (atlantic iberian margin)","volume":"8","author":"Garc\u00eda-Moreiras","year":"2021","journal-title":"Front. Mar. Sci."},{"key":"B52","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1016\/j.epsl.2017.11.008","article-title":"Global assessment of benthic nepheloid layers and linkage with upper ocean dynamics","volume":"482","author":"Gardner","year":"2018","journal-title":"Earth Planet. Sci. Lett."},{"key":"B53","doi-asserted-by":"publisher","first-page":"2585","DOI":"10.1016\/j.csr.2005.08.003","article-title":"Dinoflagellate cysts in recent marine sediments from Thermaikos Gulf, Greece: effects of resuspension events on vertical cyst distribution","volume":"25","author":"Giannakourou","year":"2005","journal-title":"Continent. Shelf Res."},{"key":"B54","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1080\/00318884.2021.1930796","article-title":"Cyst-theca relationships of Spiniferites bentorii, S. hyperacanthus, S. ramosus, S. scabratus and molecular phylogenetics of Spiniferites and Tectatodinium (Gonyaulacales, Dinophyceae)","volume":"60","author":"Gu","year":"2021","journal-title":"Phycologia"},{"key":"B55","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1080\/01916122.2019.1580627","article-title":"Taxonomic revision, phylogeny, and cyst wall composition of the dinoflagellate cyst genus Votadinium Reid (Dinophyceae, Peridiniales, Protoperidiniaceae)","volume":"44","author":"Gurdebeke","year":"2019","journal-title":"Palynology"},{"key":"B56","first-page":"191","article-title":"Phytoplankton Manual. VI","volume-title":"Monographs on Oceanic Methodology","author":"Hasle","year":"1978"},{"key":"B57","first-page":"1197","article-title":"Modern dinoflagellate cysts and their biological affinities","volume-title":"Palynology: Principles and Applications","author":"Head","year":"1996"},{"key":"B58","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1007\/BF00350743","article-title":"Mass encystment and sinking of dinoflagellates during a spring bloom","volume":"116","author":"Heiskanen","year":"1993","journal-title":"Mar. Biol."},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1029\/2007JC004220","article-title":"Internal wave detection using the moderate resolution imaging spectroradiometer (MODIS)","volume":"112","author":"Jackson","year":"2007","journal-title":"J. Geophys. Res.: Oceans"},{"key":"B60","doi-asserted-by":"publisher","first-page":"108","DOI":"10.5670\/oceanog.2012.46","article-title":"The generation of nonlinear internal waves","volume":"25","author":"Jackson","year":"2012","journal-title":"Oceanography"},{"key":"B61","doi-asserted-by":"publisher","first-page":"133","DOI":"10.3354\/ame040133","article-title":"Feeding by phototrophic red-tide dinoflagellates: five species newly revealed and six species previously known to be mixotrophic","volume":"40","author":"Jeong","year":"2005","journal-title":"Aquat. Microbial. Ecol."},{"key":"B62","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1016\/S0079-6611(02)00010-1","article-title":"Recent sedimentation and sedimentary budgets on the western Iberian shelf","volume":"52","author":"Jouanneau","year":"2002","journal-title":"Prog. Oceanogr."},{"key":"B63","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.ecss.2003.07.001","article-title":"Encystment of Zygabikodinium lenticulatum (Dinophyceae) during a summer bloom of dinoflagellates in the southern Benguela upwelling system","volume":"59","author":"Joyce","year":"2004","journal-title":"Estuarine Coast. Shelf Sci."},{"key":"B64","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1016\/j.pocean.2009.07.047","article-title":"Impact of particle aggregation on vertical fluxes of organic matter","volume":"83","author":"Karaka\u015f","year":"2009","journal-title":"Prog. Oceanogr."},{"key":"B65","doi-asserted-by":"publisher","first-page":"2543","DOI":"10.1016\/j.dsr2.2005.06.009","article-title":"Suspended Alexandrium spp. hypnozygote cysts in the Gulf of Maine","volume":"52","author":"Kirn","year":"2005","journal-title":"Deep Sea Res. Part II: Topical Stud. Oceanogr."},{"key":"B66","first-page":"57","article-title":"Effects of cyst resuspension on germination and seeding of two bloom-forming dinoflagellates in the Baltic Sea","volume-title":"Marine Ecology Progress Series","author":"Kremp","year":"2001"},{"key":"B67","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1146\/annurev-fluid-011212-140701","article-title":"Internal wave breaking and dissipation mechanisms on the continental slope\/shelf","volume":"46","author":"Lamb","year":"2014","journal-title":"Annu. Rev. Fluid Mech."},{"key":"B68","doi-asserted-by":"publisher","first-page":"629","DOI":"10.2216\/11-16.1","article-title":"Buried alive\u2013germination of up to a century-old marine protist resting stages","volume":"50","author":"Lundholm","year":"2011","journal-title":"Phycologia"},{"key":"B69","first-page":"101","article-title":"On the variability of internal waves on the western Portuguese shelf","author":"Magalhaes","year":"2022"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1016\/j.csr.2021.104396","article-title":"Using SAR imagery to survey internal solitary wave interactions: A case study off the Western Iberian shelf","volume":"220","author":"Magalhaes","year":"2021","journal-title":"Continent. Shelf Res."},{"key":"B71","volume-title":"Technical guide for modern dinoflagellate cyst study","author":"Matsuoka","year":"2000"},{"key":"B72","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1016\/j.pocean.2015.11.006","article-title":"New insights into the Western Iberian Buoyant Plume: Interaction between the Douro and Minho River plumes under winter conditions","volume":"141","author":"Mendes","year":"2016","journal-title":"Prog. Oceanogr."},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1016\/j.marmicro.2019.101773","article-title":"A review of rare, poorly known, and morphologically problematic extant marine organic-walled dinoflagellate cyst taxa of the orders Gymnodiniales and Peridiniales from the Northern Hemisphere","volume":"159","author":"Mertens","year":"2020","journal-title":"Mar. Micropaleontol."},{"key":"B74","doi-asserted-by":"publisher","first-page":"102426","DOI":"10.1016\/j.hal.2023.102426","article-title":"An unprecedented bloom of Lingulodinium polyedra on the French Atlantic coast during summer 2021","volume":"125","author":"Mertens","year":"2023","journal-title":"Harmful Algae"},{"key":"B75","first-page":"299","article-title":"Development of toxic dinoflagellates in relation to upwelling patterns off Portugal","volume-title":"Toxic phytoplankton blooms in the sea","author":"Moita","year":"1993"},{"key":"B76","article-title":"Dynamics of Dinophysis acuta, D. acuminata, D. tripos and Gymnodinium catenatum during an upwelling event off the northwest coast of Portugal","author":"Moita","year":"2001"},{"key":"B77","doi-asserted-by":"publisher","first-page":"S125","DOI":"10.1016\/S1146-609X(03)00011-0","article-title":"Distribution of chlorophyll a and Gymnodinium catenatum associated with coastal upwelling plumes off central Portugal","volume":"24","author":"Moita","year":"2003","journal-title":"Acta Oecol."},{"key":"B78","first-page":"76","article-title":"Protoceratium reticulatum bloom in NW Iberia mid-shelf waters","author":"Moita","year":"2022"},{"key":"B79","first-page":"5","article-title":"Checklist of phytoplankton species off Portugal: 70 years, (1929-1998) of studies","volume":"18","author":"Moita","year":"1999","journal-title":"Portugaliae Acta Biol. S\u00e9r. B Sist"},{"key":"B80","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1007\/BF00993643","article-title":"Modulating effect of cold-dark storage on excystment in Alexandrium pseudogonyaulax (Dinophyceae)","volume":"127","author":"Montresor","year":"1996","journal-title":"Mar. Biol."},{"key":"B81","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-03641-0_2","article-title":"The Climate of Portugal","volume-title":"Landscapes and Landforms of Portugal. World Geomorphological Landscapes","author":"Mora","year":"2020"},{"key":"B82","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.marmicro.2017.05.004","article-title":"Atlas of modern dinoflagellate cyst distributions in the Black Sea Corridor: from Aegean to Aral Seas, including Marmara, Black, Azov and Caspian Seas","volume":"134","author":"Mudie","year":"2017","journal-title":"Mar. Micropaleontol."},{"key":"B83","doi-asserted-by":"publisher","first-page":"513","DOI":"10.1002\/iroh.19960810404","article-title":"Recruitment of planktonic dinoflagellates: importance of benthic resting stages and resuspension events","volume":"81","author":"Nehring","year":"1996","journal-title":"Internation. Rev. der gesamten Hydrobiol. und Hydrogr."},{"key":"B84","doi-asserted-by":"publisher","first-page":"2641","DOI":"10.1038\/s41598-018-19833-w","article-title":"Independent estimates of marine population connectivity are more concordant when accounting for uncertainties in larval origins","volume":"8","author":"Nolasco","year":"2018","journal-title":"Sci. Rep."},{"key":"B85","doi-asserted-by":"publisher","first-page":"1178","DOI":"10.1029\/2019PA003606","article-title":"Transport bias by ocean currents in sedimentary microplankton assemblages: Implications for paleoceanographic reconstructions","volume":"34","author":"Nooteboom","year":"2019","journal-title":"Paleoceanogr. Paleoclimatol."},{"key":"B86","unstructured":"NunesP.\n          LisbonUniversity of LisbonSpatial-Temporal distribution of phytoplankton biomass during a coastal upwelling episode obtained from remote sensing and in 2021"},{"key":"B87","first-page":"105","article-title":"Evidence of upwelling separation from the coast off NW Portugal","author":"Nunes","year":"2022"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1016\/j.csr.2019.103987","article-title":"Phytoplankton distribution and physical processes off NW Iberia during two consecutive upwelling seasons","volume":"190","author":"Oliveira","year":"2019","journal-title":"Continent. Shelf Res."},{"key":"B89","first-page":"139","article-title":"Mid-shelf internal wave activity off Figueira da Foz in September 2019","author":"Oliveira","year":"2020"},{"key":"B90","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1016\/j.pocean.2009.07.030","article-title":"Summer diatom and dinoflagellate blooms in Lisbon Bay from 2002 to 2005: Pre-conditions inferred from wind and satellite data","volume":"83","author":"Oliveira","year":"2009","journal-title":"Prog. Oceanogr."},{"key":"B91","doi-asserted-by":"publisher","first-page":"759","DOI":"10.1016\/j.csr.2008.08.004","article-title":"Surface temperature, chlorophyll and advection patterns during a summer upwelling event off central Portugal","volume":"29","author":"Oliveira","year":"2009","journal-title":"Continent. Shelf Res."},{"key":"B92","first-page":"273","article-title":"Dynamics of Nepheloid layers associated with internal wave activity off Figueira da Foz","author":"Oliveira","year":"2020"},{"key":"B93","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/j.margeo.2007.04.017","article-title":"Sedimentary particle distribution and dynamics on the Nazar\u00e9 canyon system and adjacent shelf (Portugal)","volume":"246","author":"Oliveira","year":"2007","journal-title":"Mar. Geol."},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1016\/j.csr.2023.105027","article-title":"Nepheloid layer dynamics overview of the Portuguese continental shelf","volume":"261","author":"Oliveira","year":"2023","journal-title":"Continent. Shelf Res."},{"key":"B95","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/S0079-6611(02)00006-X","article-title":"Nepheloid layer dynamics in the northern Portuguese shelf","volume":"52","author":"Oliveira","year":"2002","journal-title":"Prog. Oceanogr."},{"key":"B96","doi-asserted-by":"publisher","first-page":"238","DOI":"10.1016\/j.jmarsys.2007.05.016","article-title":"Variability of river plumes off Northwest Iberia in response to wind events","volume":"72","author":"Otero","year":"2008","journal-title":"J. Mar. Syst."},{"key":"B97","doi-asserted-by":"publisher","first-page":"621","DOI":"10.1016\/j.dsr.2004.11.005","article-title":"Winter upper ocean circulation in the Western Iberian Basin\u2014Fronts, Eddies and Poleward Flows: an overview","volume":"52","author":"Peliz","year":"2005","journal-title":"Deep sea Res. Part I: Oceanogr. Res. papers"},{"key":"B98","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/s0924-7963(02)00076-3","article-title":"Fronts, jets, and counterflows in the Western Iberian upwelling system","volume":"35","author":"Peliz","year":"2002","journal-title":"J. Mar. Syst."},{"key":"B99","doi-asserted-by":"publisher","first-page":"1","DOI":"10.7773\/cm.v35i1.1406","article-title":"Environmental factors influencing the variability of Lingulodinium polyedrum and Scrippsiella trochoidea (Dinophyceae) cyst production","volume":"35","author":"Pe\u00f1a-Manjarrez","year":"2009","journal-title":"Cienc. Marinas"},{"key":"B100","first-page":"611","article-title":"Dinoflagellate reproduction","volume-title":"The Biology of Dinoflagellates","author":"Pfiester","year":"1987"},{"key":"B101","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/j.dsr2.2012.11.001","article-title":"Spatial and temporal variability of Alexandrium cyst fluxes in the Gulf of Maine: relationship to seasonal particle export and resuspension","volume":"103","author":"Pilskaln","year":"2014","journal-title":"Deep Sea Res. Part II: Topical Stud. Oceanogr."},{"key":"B102","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1016\/j.dsr2.2013.05.021","article-title":"Benthic nepheloid layers in the Gulf of Maine and Alexandrium cyst inventories","volume":"103","author":"Pilskaln","year":"2014","journal-title":"Deep Sea Res. Part II Topical Stud. Oceanogr."},{"key":"B103","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1016\/j.pocean.2010.02.002","article-title":"The physical oceanography of upwelling systems and the development of harmful algal blooms","volume":"85","author":"Pitcher","year":"2010","journal-title":"Prog. Oceanogr."},{"key":"B104","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1046\/j.1529-8817.2002.01206.x","article-title":"Islandinium brevispinosum sp. nov. (Dinoflagellata), a new organic-walled dinoflagellate cyst from modern estuarine sediments of New England (USA)","volume":"38","author":"Pospelova","year":"2002","journal-title":"J. Phycol."},{"key":"B105","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1016\/j.marmicro.2011.03.003","article-title":"High-resolution sediment trap study of organic-walled dinoflagellate cyst production and biogenic silica flux in Saanich Inlet (BC, Canada)","volume":"80","author":"Price","year":"2011","journal-title":"Mar. Micropaleontol."},{"key":"B106","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1016\/j.margeo.2007.04.019","article-title":"Evidence of sediment resuspension by nonlinear internal waves on the western Portuguese mid-shelf","volume":"246","author":"Quaresma","year":"2007","journal-title":"Mar. geol."},{"key":"B107","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.marmicro.2012.11.001","article-title":"Operational taxonomy and (paleo-)autecology of round, brown, spiny dinoflagellate cysts from the Quaternary of high northern latitudes","volume":"98","author":"Radi","year":"2013","journal-title":"Mar. Micropaleontol."},{"key":"B108","volume-title":"R: A Language and Environment for Statistical Computing","year":"2013"},{"key":"B109","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1016\/j.pocean.2007.04.021","article-title":"Physical oceanography of the western Iberia ecosystem: latest views and challenges","volume":"74","author":"Relvas","year":"2007","journal-title":"Prog. Oceanogr."},{"key":"B110","doi-asserted-by":"publisher","first-page":"213","DOI":"10.3354\/ame036213","article-title":"Factors regulating the recruitment of cyanobacterial and eukaryotic phytoplankton from littoral and profundal sediments","volume":"36","author":"Rengefors","year":"2004","journal-title":"Aquat. Microbial. Ecol."},{"key":"B111","doi-asserted-by":"publisher","first-page":"969","DOI":"10.1007\/s10530-011-0132-6","article-title":"Reconstructing the history of an invasion: the toxic phytoplankton species Gymnodinium catenatum in the Northeast Atlantic","volume":"14","author":"Ribeiro","year":"2012","journal-title":"Biol. Invasions"},{"key":"B112","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.csr.2013.04.008","article-title":"Cross-shelf variability in the Iberian Peninsula Upwelling System: Impact of a mesoscale filament","volume":"59","author":"Rossi","year":"2013","journal-title":"Continent. Shelf Res."},{"key":"B113","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.hal.2017.07.008","article-title":"The life history of the toxic marine dinoflagellate Protoceratium reticulatum (Gonyaulacales) in culture","volume":"68","author":"Salgado","year":"2017","journal-title":"Harmful Algae"},{"key":"B114","first-page":"141","article-title":"ADCP backscattering patterns in a mid-shelf water column (Figueira da Foz \u2013 W Portugal): Interaction between physical\/biological forcing and particle dynamics","author":"Santos","year":"2022"},{"key":"B115","doi-asserted-by":"publisher","first-page":"101982","DOI":"10.1016\/j.seares.2020.101982","article-title":"Phytoplankton communities in two wide-open bays in the Iberian upwelling system","volume":"167","author":"Santos","year":"2021","journal-title":"J. Sea Res."},{"key":"B116","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/0025-3227(71)90070-3","article-title":"Normal and accelerated sinking of phytoplankton in the sea","volume":"11","author":"Smayda","year":"1971","journal-title":"Mar. Geol."},{"key":"B117","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1093\/plankt\/23.5.447","article-title":"Community assembly in marine phytoplankton: application of recent models to harmful dinoflagellate blooms","volume":"23","author":"Smayda","year":"2001","journal-title":"J. Plankton Res."},{"key":"B118","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/S1568-9883(02)00010-0","article-title":"Turbulence, watermass stratification and harmful algal blooms: an alternative view and frontal zones as \u201cpelagic seed banks\u201d","volume":"1","author":"Smayda","year":"2002","journal-title":"Harmful Algae"},{"key":"B119","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/S1385-1101(02)00219-8","article-title":"Strategies of marine dinoflagellate survival and some rules of assembly","volume":"49","author":"Smayda","year":"2003","journal-title":"J. Sea Res."},{"key":"B120","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/j.pocean.2010.02.006","article-title":"Dinoflagellate blooms in upwelling systems: seeding, variability, and contrasts with diatom bloom behaviour","volume":"85","author":"Smayda","year":"2010","journal-title":"Prog. Oceanogr."},{"key":"B121","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1111\/j.1440-1835.2005.tb00362.x","article-title":"Organic-walled dinoflagellate cyst production in relation to upwelling intensity and lithogenic influx in the Cape Blanc region (off north-west Africa)","volume":"53","author":"Susek","year":"2005","journal-title":"Phycological Res."},{"key":"B122","unstructured":"SuzukiR.\n            TeradaY.\n            ShimodairaH.\n          Hierarchical Clustering with P-Values via Multiscale Bootstrap Resampling, V. 2.2-0. Cran repository2022"},{"key":"B123","doi-asserted-by":"publisher","first-page":"4980","DOI":"10.1002\/2015JC010758","article-title":"On the year-to-year changes of the Iberian Poleward Current","volume":"120","author":"Teles-MaChado","year":"2015","journal-title":"J. Geophys. Res.: Oceans"},{"key":"B124","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1016\/S0079-6611(02)00013-7","article-title":"Processes in the benthic boundary layer at the Iberian continental margin and their implication for carbon mineralization","volume":"52","author":"Thomsen","year":"2002","journal-title":"Prog. Oceanogr."},{"key":"B125","doi-asserted-by":"publisher","first-page":"678","DOI":"10.3390\/w14050678","article-title":"Formation mechanisms and characteristics of the marine nepheloid layer: A review","volume":"14","author":"Tian","year":"2022","journal-title":"Water"},{"key":"B126","doi-asserted-by":"publisher","first-page":"241","DOI":"10.3354\/meps112241","article-title":"Upwelling-downwelling sequences in the generation of red tides in a coastal upwelling system","volume":"112","author":"Tilstone","year":"1994","journal-title":"Mar. Ecol. Prog. Series. Oldendorf"},{"key":"B127","doi-asserted-by":"publisher","first-page":"23","DOI":"10.3354\/meps205023","article-title":"Diatom dynamics in a coastal ecosystem affected by upwelling: coupling between species succession, circulation and biogeochemical processes","volume":"205","author":"Tilstone","year":"2000","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"B128","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/j.pocean.2010.02.003","article-title":"The distribution and impacts of harmful algal bloom species in eastern boundary upwelling systems","volume":"85","author":"Trainer","year":"2010","journal-title":"Prog. Oceanogr."},{"key":"B129","doi-asserted-by":"publisher","first-page":"57","DOI":"10.3354\/ame027057","article-title":"Zooplankton fecal pellets, marine snow and sinking phytoplankton blooms","volume":"27","author":"Turner","year":"2002","journal-title":"Aquat. Microbiol. Ecol."},{"key":"B130","doi-asserted-by":"publisher","DOI":"10.1016\/j.marmicro.2019.101814","article-title":"An overview and a brief description of common marine organic-walled dinoflagellate cyst taxa occurring in surface sediments of the Northern Hemisphere","volume":"159","author":"van Nieuwenhove","year":"2020","journal-title":"Mar. Micropaleontol."},{"key":"B131","first-page":"7","article-title":"Sampling strategies","volume-title":"Phytoplankton Manual. Monographs on Oceanic Methodology","author":"Venrick","year":"1978"},{"key":"B132","doi-asserted-by":"publisher","first-page":"9531","DOI":"10.1029\/2019JC015619","article-title":"Bottom boundary layer and particle dynamics in an upwelling affected continental margin (NW Iberia)","volume":"124","author":"Villacieros-Robineau","year":"2019","journal-title":"J. Geophys. Res.: Oceans"},{"key":"B133","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1016\/S0079-6611(02)00004-6","article-title":"Winter dynamics on the northern Portuguese shelf: Part 2: bottom boundary layers and sediment","volume":"52","author":"Vitorino","year":"2002","journal-title":"Prog. Oceanogr."},{"key":"B134","doi-asserted-by":"publisher","first-page":"87","DOI":"10.4319\/lo.2000.45.1.0087","article-title":"Mass sedimentation of picoplankton embedded in organic aggregates","volume":"45","author":"Waite","year":"2000","journal-title":"Limnol. Oceanogr."},{"key":"B135","doi-asserted-by":"publisher","first-page":"2229","DOI":"10.1016\/S0967-0645(00)00023-0","article-title":"(Palaeo-)ecological significance, transport and preservation of organic-walled dinoflagellate cysts in the Somali Basin, NW Arabian Sea","volume":"47","author":"Zonneveld","year":"2000","journal-title":"Deep-Sea Res. II"},{"key":"B136","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1016\/j.dsr.2018.06.003","article-title":"Transport of organic-walled dinoflagellate cysts in nepheloid layers off Cape Blanc (N-W Africa)","volume":"139","author":"Zonneveld","year":"2018","journal-title":"Deep-Sea Res. Part I"},{"key":"B137","doi-asserted-by":"publisher","DOI":"10.3389\/fmars.2022.915755","article-title":"Dinoflagellate cysts production, excystment and transport in the upwelling off Cape Blanc (NW Africa)","volume":"1317","author":"Zonneveld","year":"2022","journal-title":"Front. Mar. Sci."},{"key":"B138","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1080\/09670262.2021.1885066","article-title":"Export flux succession of dinoflagellate cysts and planktonic foraminifera in an active upwelling cell off Cape Blanc (NW Africa)","volume":"57","author":"Zonneveld","year":"2021","journal-title":"Eur. J. Phycol."},{"key":"B139","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1080\/01916122.2014.990115","article-title":"A determination key for modern dinoflagellate cysts","volume":"39","author":"Zonneveld","year":"2015","journal-title":"Palynology"},{"key":"B140","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/j.marmicro.2008.01.015","article-title":"Preservation and organic chemistry of late Cenozoic organic-walled dinoflagellate cysts: a review","volume":"68","author":"Zonneveld","year":"2008","journal-title":"Mar. Micropaleontol."},{"key":"B141","doi-asserted-by":"crossref","first-page":"202","DOI":"10.1111\/j.1529-8817.2009.00799.x","article-title":"Seasonal variability of the organic-walled dinoflagellate cyst production in the coastal upwelling region off Cape Blanc (Mauritania): a five-year survey","volume":"46","author":"Zonneveld","year":"2010","journal-title":"J. Phycology"}],"container-title":["Frontiers in Marine Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmars.2023.1270343\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,12,12]],"date-time":"2023-12-12T16:36:04Z","timestamp":1702398964000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmars.2023.1270343\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,12,8]]},"references-count":141,"alternative-id":["10.3389\/fmars.2023.1270343"],"URL":"https:\/\/doi.org\/10.3389\/fmars.2023.1270343","relation":{},"ISSN":["2296-7745"],"issn-type":[{"value":"2296-7745","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,12,8]]},"article-number":"1270343"}}