{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,11,3]],"date-time":"2025-11-03T19:28:42Z","timestamp":1762198122203,"version":"build-2065373602"},"reference-count":37,"publisher":"MDPI AG","issue":"4","license":[{"start":{"date-parts":[[2025,11,1]],"date-time":"2025-11-01T00:00:00Z","timestamp":1761955200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000780","name":"European Union\u2019s Horizon 2020 Research and Innovation Programme","doi-asserted-by":"publisher","award":["818173"],"award-info":[{"award-number":["818173"]}],"id":[{"id":"10.13039\/501100000780","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Phycology"],"abstract":"<jats:p>Codium tomentosum is a native green seaweed of high ecological and commercial interest along the Atlantic coast of Portugal. Although recently introduced into aquaculture via vegetative propagation, its cultivation remains limited. Production still relies primarily on wild-harvested biomass, including the supply of starting material for aquaculture. Understanding the species\u2019 reproductive biology, particularly the timing of sexual reproduction, is important to support sustainable biomass production and reduce pressure on wild populations. In this context, the present study aimed to assess the seasonal reproductive pattern of a natural C. tomentosum population from the Agu\u00e7adoura shore (Northern Portugal). The reproductive status of C. tomentosum was assessed monthly between September 2020 and August 2022. Gametogenesis was initiated in summer, with gametangia persisting through autumn and winter, followed by a marked reduction or complete absence in spring, coinciding with thallus regression observed in the field. Male and female gametangia differed significantly in length (247.34 \u00b1 36.22 \u00b5m and 288.70 \u00b1 28.39 \u00b5m, respectively). Fertile individuals, yielding viable gametes, were identified between August and March across the years. Male gametes measured 4.32 \u00b1 0.46 \u00b5m in length, and female gametes were significantly larger (19.42 \u00b1 0.40 \u00b5m). This study confirms that C. tomentosum exhibits a well-defined reproductive period on the Northern Portuguese coast, extending from late summer to early spring. These findings are expected to contribute to the responsible management of natural populations and guide strategies for the collection of starting material, ultimately ensuring the long-term sustainable production of this species.<\/jats:p>","DOI":"10.3390\/phycology5040066","type":"journal-article","created":{"date-parts":[[2025,11,3]],"date-time":"2025-11-03T18:21:46Z","timestamp":1762194106000},"page":"66","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Reproduction Pattern of a Codium tomentosum Population from the Northern Portuguese Coast"],"prefix":"10.3390","volume":"5","author":[{"ORCID":"https:\/\/orcid.org\/0009-0007-7673-005X","authenticated-orcid":false,"given":"Teresa Cunha","family":"Pacheco","sequence":"first","affiliation":[{"name":"Department of Biology, Faculty of Sciences, University of Porto, Rua do Campo Alegre s\/n, 4169-007 Porto, Portugal"},{"name":"CIIMAR\/CIMAR-LA, Interdisciplinary Centre of Marine and Environmental Research, University of Porto, Terminal de Cruzeiros do Porto de Leix\u00f5es, 4450-208 Matosinhos, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8846-3314","authenticated-orcid":false,"given":"Maria Francisca","family":"S\u00e1","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Sciences, University of Porto, Rua do Campo Alegre s\/n, 4169-007 Porto, Portugal"},{"name":"CIIMAR\/CIMAR-LA, Interdisciplinary Centre of Marine and Environmental Research, University of Porto, Terminal de Cruzeiros do Porto de Leix\u00f5es, 4450-208 Matosinhos, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9231-0553","authenticated-orcid":false,"given":"Isabel","family":"Sousa-Pinto","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Sciences, University of Porto, Rua do Campo Alegre s\/n, 4169-007 Porto, Portugal"},{"name":"CIIMAR\/CIMAR-LA, Interdisciplinary Centre of Marine and Environmental Research, University of Porto, Terminal de Cruzeiros do Porto de Leix\u00f5es, 4450-208 Matosinhos, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1038-9502","authenticated-orcid":false,"given":"Gon\u00e7alo Silva","family":"Marinho","sequence":"additional","affiliation":[{"name":"Department of Biology, Faculty of Sciences, University of Porto, Rua do Campo Alegre s\/n, 4169-007 Porto, Portugal"},{"name":"CIIMAR\/CIMAR-LA, Interdisciplinary Centre of Marine and Environmental Research, University of Porto, Terminal de Cruzeiros do Porto de Leix\u00f5es, 4450-208 Matosinhos, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2025,11,1]]},"reference":[{"key":"ref_1","first-page":"275","article-title":"Influence of different habitats on the chemical constituents of Codium tomentosum","volume":"58","author":"Mansour","year":"2018","journal-title":"Egypt. J. Bot."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1016\/j.marenvres.2018.07.008","article-title":"Spatio-temporal dynamics of Codium populations along the rocky shores of N and NW Spain","volume":"140","author":"Olabarria","year":"2018","journal-title":"Mar. Environ. Res."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"102079","DOI":"10.1016\/j.seares.2021.102079","article-title":"Temporal and spatial variation of seaweed biomass and assemblages in Northwest Portugal","volume":"174","author":"Melo","year":"2021","journal-title":"J. Sea Res."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1016\/j.algal.2015.09.020","article-title":"Decoding bioactive polar lipid profile of the macroalgae Codium tomentosum from a sustainable IMTA system using a lipidomic approach","volume":"12","author":"Melo","year":"2015","journal-title":"Algal Res."},{"key":"ref_5","first-page":"293","article-title":"A dichotomous species of Codium (Bryopsidales, Chlorophyta) is colonizing Northern Chile","volume":"77","author":"Santelices","year":"2004","journal-title":"Rev. Chil. Hist. Nat."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1007\/s00227-018-3329-8","article-title":"The ability to incorporate functional plastids by the sea slug Elysia viridis is governed by its food source","volume":"165","author":"Rauch","year":"2018","journal-title":"Mar. Biol."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Rey, F., Cartaxana, P., Melo, T., Calado, R., Pereira, R., Abreu, H., Domingues, P., Cruz, S., and Ros\u00e1rio Domingues, M. (2020). Domesticated populations of Codium tomentosum display lipid extracts with lower seasonal shifts than conspecifics from the wild-relevance for biotechnological applications of this green seaweed. Mar. Drugs, 18.","DOI":"10.3390\/md18040188"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1285","DOI":"10.1080\/10807039.2012.708253","article-title":"Bioaccumulation of Key Metals and Other Contaminants by Seaweeds from the Egyptian Mediterranean Sea Coast in Relation to Human Health Risk","volume":"19","year":"2013","journal-title":"Hum. Ecol. Risk Assess."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1007\/s10811-020-02289-9","article-title":"Effects of photoperiod and light spectra on growth and pigment composition of the green macroalga Codium tomentosum","volume":"33","author":"Marques","year":"2021","journal-title":"J. Appl. Phycol."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Marques, R., Moreira, A., Cruz, S., Calado, R., and Cartaxana, P. (2022). Controlling Light to Optimize Growth and Added Value of the Green Macroalga Codium tomentosum. Front. Mar. Sci., 9.","DOI":"10.3389\/fmars.2022.906332"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1111\/raq.12580","article-title":"The underexplored potential of green macroalgae in aquaculture","volume":"14","author":"Moreira","year":"2022","journal-title":"Rev. Aquac."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Giossi, C.E., Cruz, S., Rey, F., Marques, R., Melo, T., do Ros\u00e1rio Domingues, M., and Cartaxana, P. (2021). Light Induced Changes in Pigment and Lipid Profiles of Bryopsidales Algae. Front. Mar. Sci., 8.","DOI":"10.3389\/fmars.2021.745083"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"102492","DOI":"10.1016\/j.bcab.2022.102492","article-title":"Phytochemical and anti-inflammatory properties of green macroalga Codium tomentosum","volume":"45","author":"Rabecca","year":"2022","journal-title":"Biocatal. Agric. Biotechnol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/s11274-019-2740-8","article-title":"The microbiome of Codium tomentosum: Original state and in the presence of copper","volume":"35","author":"Gall","year":"2019","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_15","unstructured":"Per\u00e9z Llor\u00e9ns, J.L. (2018). Those Curious and Delicious Seaweeds: A Fascinating Voyage from Biology to Gastronomy, Informa UK Limited."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/S0304-3770(02)00036-0","article-title":"Formation and growth of filamentous thalli from isolated utricles with medullary filaments of Codium fragile spongy thalli","volume":"73","author":"Nanba","year":"2002","journal-title":"Aquat. Bot."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1007\/s10811-007-9265-5","article-title":"Cultivation of the green alga, Codium fragile (Suringar) Hariot, by artificial seed production in Korea","volume":"20","author":"Hwang","year":"2008","journal-title":"J. Appl. Phycol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"91","DOI":"10.2216\/i0031-8884-8-2-91.1","article-title":"Reproduction and early development in Codium fragile (Suringar) Hariot: Chlorophyceae","volume":"8","author":"Borden","year":"1969","journal-title":"Phycologia"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1515\/BOT.2003.043","article-title":"Gametangium-like Structures as Propagation Buds in Codium edule Silva (Bryopsidales, Chlorophyta)","volume":"46","author":"Chang","year":"2003","journal-title":"Bot. Mar."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1111\/j.1440-1835.2012.00640.x","article-title":"Female gametogenesis and female gamete germination in the anisogamous green alga Codium fragile subsp. novae-zelandiae (Bryopsidophyceae, Chlorophyta)","volume":"60","author":"Miravalles","year":"2012","journal-title":"Phycol. Res."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"8567","DOI":"10.1002\/ece3.5379","article-title":"Contrasting seasonal and spatial distribution of native and invasive Codium seaweed revealed by targeting species-specific eDNA","volume":"9","author":"Muha","year":"2019","journal-title":"Ecol. Evol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"533","DOI":"10.3390\/phycology4040029","article-title":"Sexual Propagation in the Green Seaweed Codium tomentosum\u2014An Emerging Species for Aquaculture","volume":"4","author":"Pacheco","year":"2024","journal-title":"Phycology"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.marenvres.2014.09.006","article-title":"Coexistence of congeneric native and invasive species: The case of the green algae Codium spp. in Northwestern Spain","volume":"101","author":"Rojo","year":"2014","journal-title":"Mar. Environ. Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1111\/j.0022-3646.1972.00147.x","article-title":"Seasonal patterns of reproduction in New York. Populations of Codium fragile (sur.) Hariot subsp. tomentosoides (Van Goor) Silva","volume":"8","author":"Churchill","year":"1972","journal-title":"J. Phycol."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Ara\u00fajo, R., V\u00e1zquez Calder\u00f3n, F., S\u00e1nchez L\u00f3pez, J., Azevedo, I.C., Bruhn, A., Fluch, S., Garcia Tasende, M., Ghaderiardakani, F., Ilmj\u00e4rv, T., and Laurans, M. (2021). Current Status of the Algae Production Industry in Europe: An Emerging Sector of the Blue Bioeconomy. Front. Mar. Sci., 7.","DOI":"10.3389\/fmars.2020.626389"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1557","DOI":"10.1007\/s10811-025-03489-x","article-title":"Seasonal variation in the composition and antioxidant potential of Codium tomentosum and Ulva lacinulata produced in a land-based integrated multi-trophic aquaculture system","volume":"37","author":"Sousa","year":"2025","journal-title":"J. Appl. Phycol."},{"key":"ref_27","first-page":"1","article-title":"Characterization and multivariate analysis of Patella intermedia, Patella ulyssiponensis and Patella vulgata from Povoa de Varzim (Northwest Portugal)","volume":"21","author":"Cabral","year":"2003","journal-title":"Iberus"},{"key":"ref_28","first-page":"1","article-title":"Effects of light intensity and artificial aeration on growth and photosynthetic physiology of marine invasive green alga Codium fragile from Bohai Sea, China","volume":"rs.3","author":"Ding","year":"2022","journal-title":"Res. Sq."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1007\/BF00387009","article-title":"Growth Patterns of Codium fragile ssp. tomentosoides in Response to Temperature, Irradiance, Salinity, and Nitrogen Source","volume":"50","author":"Hanisak","year":"1979","journal-title":"Mar. Biol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.aquabot.2018.09.002","article-title":"Life history traits of Laminaria ochroleuca in Portugal: The range-center of its geographical distribution","volume":"152","author":"Pereira","year":"2019","journal-title":"Aquat. Bot."},{"key":"ref_31","unstructured":"L\u00fcning, K. (1991). Seaweeds: Their Environment, Biogeography, and Ecophysiology, John Wiley & Sons."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"3318","DOI":"10.1016\/j.apenergy.2010.11.027","article-title":"Macroalgae: Raw material for biodiesel production","volume":"88","author":"Maceiras","year":"2011","journal-title":"Appl. Energy"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"37","DOI":"10.24310\/abm.v39i1.2584","article-title":"Codium tomentosum var. mucronatum et son epiphyte Aglaothamnion pseudobyssoides, deux nouvelles esp\u00e8ces d\u2019algues benthiques pour la phycoflore du Maroc. Codium tomentosum var. mucronatum and Aglaothamnion pseudobyssoides, two new records for Morocco","volume":"39","author":"Hassoun","year":"2014","journal-title":"Acta Bot. Malacit."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1515\/BOT.2004.062","article-title":"Reproduction in the green macroalga Codium (Chlorophyta): Characterization of gametes","volume":"47","author":"Prince","year":"2004","journal-title":"Bot. Mar."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"370","DOI":"10.2216\/10-25.1","article-title":"Male gametogenesis ultrastructure of Codium fragile subsp. novae-zelandiae (Bryopsidophyceae, Chlorophyta)","volume":"50","author":"Miravalles","year":"2011","journal-title":"Phycologia"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"201611","DOI":"10.1098\/rsos.201611","article-title":"A comparative test of the gamete dynamics theory for the evolution of anisogamy in Bryopsidales green algae","volume":"8","author":"Togashi","year":"2021","journal-title":"R Soc. Open Sci."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Gallardo, T. (2015). Marine Algae: General Aspects (Biology, Systematics, Field and Laboratory Techniques). Marine Algae: Biodiversity, Taxonomy, Environmental Assessment, and Biotechnology, CRC Press.","DOI":"10.1201\/b17540-2"}],"container-title":["Phycology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2673-9410\/5\/4\/66\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,11,3]],"date-time":"2025-11-03T19:02:46Z","timestamp":1762196566000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2673-9410\/5\/4\/66"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,1]]},"references-count":37,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["phycology5040066"],"URL":"https:\/\/doi.org\/10.3390\/phycology5040066","relation":{},"ISSN":["2673-9410"],"issn-type":[{"value":"2673-9410","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,1]]}}}