{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T19:50:55Z","timestamp":1787687455452,"version":"build-2784847793"},"reference-count":74,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2025,1,9]],"date-time":"2025-01-09T00:00:00Z","timestamp":1736380800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"French Agency for AlUla Development (AFALULA)"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Intensive surveys recently conducted in AlUla County (northwest Saudi Arabia) have made it possible to compile an exhaustive list of the forty-eight bryophyte species (six liverworts and forty-two mosses) known in this region and compare it with an updated checklist of bryophytes from Saudi Arabia, which now counts 31 liverworts and 135 mosses. The ecology, taxonomy, and distribution of each taxon are provided and discussed in a systematic catalog. Although particularly arid and lacking permanent watercourses, AlUla County has proven to be surprisingly rich in bryophytes. Some remarkable species are even confined here, such as Riella affinis, Acaulon triquetrum, Microbryum rectum, and Syntrichia rigescens. Two endemic species were also observed. An updated checklist of Arabian bryophyte endemics is provided. Specific recommendations regarding the conservation of bryophyte heritage are outlined.<\/jats:p>","DOI":"10.3390\/plants14020170","type":"journal-article","created":{"date-parts":[[2025,1,9]],"date-time":"2025-01-09T07:00:47Z","timestamp":1736406047000},"page":"170","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Bryophyte Flora of AlUla County (Saudi Arabia)\u2014Distribution, Ecology, and Conservation"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-2689-523X","authenticated-orcid":false,"given":"Vincent","family":"Hugonnot","sequence":"first","affiliation":[{"name":"SARL P\u00e9pin Hugonnot, 43380 Blassac, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Florine","family":"P\u00e9pin","sequence":"additional","affiliation":[{"name":"SARL P\u00e9pin Hugonnot, 43380 Blassac, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Freedman","sequence":"additional","affiliation":[{"name":"Royal Commission for AlUla, AlUla 43544, Saudi Arabia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,1,9]]},"reference":[{"key":"ref_1","unstructured":"Chaudhary, S.A. (2001). Flora of the Kingdom of Saudi Arabia, Ministry of Agriculture and Water."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Ghazanfar, S.A., and Fisher, M. (1998). Bryophytes and Lichens. Vegetation of the Arabian Peninsula, Springer-Science+Business Media.","DOI":"10.1007\/978-94-017-3637-4"},{"key":"ref_3","first-page":"1","article-title":"Bryophyte Flora of the Arabian Peninsula and Socotra","volume":"55","year":"2000","journal-title":"Bryophyt. Bibl."},{"key":"ref_4","first-page":"157","article-title":"The First Records of Bryophytes from Saudi Arabia: Studies in Arabian Bryophytes","volume":"8","author":"Frey","year":"1982","journal-title":"Lindbergia"},{"key":"ref_5","first-page":"1","article-title":"Liverworts, Mosses and Hornworts of Southwest Asia: Marchantiophyta, Bryophyta, Anthocerotophyta, 2nd enlarged and revised revision","volume":"149","author":"Frey","year":"2020","journal-title":"Nova Hedwig. Beih."},{"key":"ref_6","first-page":"17","article-title":"An Annotated Checklist of Saudi Arabian Mosses","volume":"10","author":"Taha","year":"2019","journal-title":"Egypt. Acad. J. Biolog. Sci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1127\/nova_hedwigia\/2020\/0574","article-title":"Bryophyte Novelties from the Arabian Peninsula and Socotra Island","volume":"110","author":"Frey","year":"2020","journal-title":"Nova Hedwig."},{"key":"ref_8","first-page":"91","article-title":"Bryum radiculosum Brid. New to Arabian Peninsula with New Bryaceae Additions to South Hejaz Region in Saudi Arabia","volume":"11","author":"Taha","year":"2020","journal-title":"Egypt. Acad. J. Biolog. Sci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1127\/nova_hedwigia\/2022\/0671","article-title":"Reappraisal of Barbula trifaria var. desertorum (J. Froehl.) S. Agnew (Pottiaceae, Bryophyta), Based on Morphological and Molecular Evidence","volume":"114","author":"Cano","year":"2022","journal-title":"Nova Hedwig."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"24899","DOI":"10.11609\/jott.8609.16.3.24899-24909","article-title":"New Plant Records for the Flora of Saudi Arabia","volume":"16","author":"Alzahrani","year":"2024","journal-title":"J. Threat. Taxa"},{"key":"ref_11","unstructured":"Hugonnot, V., P\u00e9pin, F., and Freedman, J. (Nova Hedwig., 2025). New Bryophyte Species for Saudi Arabia and the Arabian Peninsula from AlUla County, Nova Hedwig., submitted."},{"key":"ref_12","unstructured":"Holyoak, D.T. (2021). European Bryaceae: A Guide to the Species of the Moss Family Bryaceae in Western and Central Europe and Macaronesia, Pisces Publications."},{"key":"ref_13","first-page":"37","article-title":"Bryophytes of the Arabian Peninsula and Socotra: Floristics, Phytogeography and Definition of the Xerothermic Pangaean Element","volume":"46","author":"Frey","year":"1988","journal-title":"Nova Hedwig."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1179\/jbr.1984.13.1.25","article-title":"Studies in Arabian Bryophytes 3: Crossidium asirense (Pottiaceae), a New Species from Asir Mountains (Saudi Arabia)","volume":"13","author":"Frey","year":"1984","journal-title":"J. Bryol."},{"key":"ref_15","first-page":"1","article-title":"Conspectus Bryophytorum Orientalum et Arabicorum: An Annotated Catalogue of the Bryophytes in Southwest Asia","volume":"39","author":"Frey","year":"1991","journal-title":"Bryophyt. Bibl."},{"key":"ref_16","first-page":"113","article-title":"An Annotated List of Fissidens Species from the Yemen Arab Republic and Sultanate of Oman, with F. laxetexturatus nov. spec. Studies in Arabian Bryophytes 7","volume":"45","year":"1987","journal-title":"Nova Hedwig."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1111\/jse.12735","article-title":"A Multilocus Phylogeny of the Moss Genus Didymodon and Allied Genera (Pottiaceae): Generic Delimitations and Their Implications for Systematics","volume":"60","author":"Cano","year":"2022","journal-title":"J. Syst. Evol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"241","DOI":"10.7872\/cryb\/v39.iss2.2018.241","article-title":"The Didymodon tophaceus Complex (Pottiaceae, Bryophyta) Revisited: New Data Support the Subspecific Rank of Currently Recognized Species","volume":"39","author":"Blockeel","year":"2018","journal-title":"Cryptogam. Bryol."},{"key":"ref_19","first-page":"69","article-title":"Gymnostomiella (Pottiaceae, Bryophyta) Revisited: New Insights Revealed by a Morphometric Analysis","volume":"138","author":"Werner","year":"2010","journal-title":"Nova Hedwig. Beih."},{"key":"ref_20","first-page":"1","article-title":"Identity of the Endemic African Species Gymnostomiella monodii P. de la Varde (Pottiaceae, Musci) and New Records for Macaronesia (Cape Verde)","volume":"39","author":"Stow","year":"2017","journal-title":"J. Bryol."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Ghazanfar, S.A., and Fisher, M. (1998). Biogeography and Introduction to Vegetation. Vegetation of the Arabian Peninsula, Springer.","DOI":"10.1007\/978-94-017-3637-4"},{"key":"ref_22","first-page":"119","article-title":"A Desert Bryophyte Synusia from the Jabal Tuwayq Mountain Systems (Central Saudi Arabia) with the Description of Two New Crossidium Species (Pottiaceae)","volume":"45","author":"Frey","year":"1987","journal-title":"Nova Hedwig."},{"key":"ref_23","first-page":"269","article-title":"Two Remarkable Hygrophytic Mosses New to the Bryophyte Flora of Yemen (Studies in Arabian Bryophytes 25)","volume":"44","author":"Ochyra","year":"1999","journal-title":"Fragm. Flor. Geobot."},{"key":"ref_24","first-page":"27","article-title":"Tortula rigescens Broth. et Geh. (Pottiaceae): A Remarkable Species New to the Moss Flora of Jordan","volume":"14","author":"Frey","year":"1988","journal-title":"Lindbergia"},{"key":"ref_25","first-page":"643","article-title":"New Taxonomic Data on a Circum-Tethyan Group of Syntrichia (Pottiaceae, Bryophyta): The S. caninervis Complex","volume":"27","author":"Gallego","year":"2002","journal-title":"Syst. Bot."},{"key":"ref_26","first-page":"47","article-title":"A Taxonomic Study of the Genus Syntrichia Brid. (Pottiaceae, Musci) in the Mediterranean Region and Macaronesia","volume":"98","author":"Gallego","year":"2005","journal-title":"J. Hattori Bot. Lab."},{"key":"ref_27","first-page":"127","article-title":"Targionia hypophylla L. ssp. linealis (Marchantiidae, Hepaticae), a New Subspecies from Saudi Arabia, and Further Discoveries. Studies in Arabian Bryophytes 17","volume":"57","author":"Frey","year":"1993","journal-title":"Nova Hedwig."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1002\/biuz.960280410","article-title":"W\u00fcstenmoose: Anpassungen und \u00dcberlebensstrategien im T\u00e4glichen Kampf mit der Sonne","volume":"28","author":"Frey","year":"1998","journal-title":"BIUZ"},{"key":"ref_29","first-page":"359","article-title":"Bryum nanoapiculatum (Bryaceae, Musci): A Species New to the Moss Flora of the Arabian Peninsula","volume":"47","author":"Ochi","year":"1988","journal-title":"Nova Hedwig."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"401","DOI":"10.2307\/3242906","article-title":"Pseudaloina (Pottiaceae, Musci): A New Genus from Yemen","volume":"85","year":"1982","journal-title":"Bryologist"},{"key":"ref_31","first-page":"101","article-title":"Fissidens arabicus (Fissidentaceae): A New Species from the Asir Mountains (Saudi Arabia). Studies in Arabian Bryophytes 6","volume":"44","author":"Pursell","year":"1987","journal-title":"Nova Hedwig."},{"key":"ref_32","first-page":"27","article-title":"Riccia crenatodentata (Marchantiales) sp. nov. aus Arabien. Studien \u00fcber Bryophyten aus Arabien II","volume":"46","author":"Volk","year":"1988","journal-title":"Nova Hedwig."},{"key":"ref_33","first-page":"330","article-title":"Muscineae. In I.B. Balfour, Botany of Socotra","volume":"31","author":"Mitten","year":"1888","journal-title":"Trans. R. Soc. Edinb."},{"key":"ref_34","first-page":"467","article-title":"New Records, Additions and a New Species, Sematophyllum socotrense Buck (Sematophyllaceae, Musci) to the Bryophyte Flora of Socotra (Yemen). Studies in Arabian Bryophytes 19","volume":"61","author":"Mies","year":"1995","journal-title":"Nova Hedwig."},{"key":"ref_35","first-page":"157","article-title":"Bryophyta Nova (18 & 19)","volume":"7","author":"Dixon","year":"1934","journal-title":"Ann. Bryol."},{"key":"ref_36","unstructured":"M\u00fcller, C. (1901). Genera Muscorum Frondosorum, Eduard Kummer."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1179\/006813869804146664","article-title":"Bryophytes from Socotra","volume":"5","author":"Townsend","year":"1969","journal-title":"Trans. Br. Bryol. Soc."},{"key":"ref_38","first-page":"345","article-title":"Tortula mucronifera (Pottiaceae, Musci), a New Xerophytic Species of the Arabian Peninsula and Jordan","volume":"59","author":"Frey","year":"1994","journal-title":"Nova Hedwig."},{"key":"ref_39","first-page":"378p","article-title":"Genera of the Pottiaceae: Mosses of Harsh Environments","volume":"32","author":"Zander","year":"1993","journal-title":"Bull. Buffalo Soc. Nat. Sci."},{"key":"ref_40","first-page":"4306","article-title":"Bryophyte Development and Simple Structural Features","volume":"73","author":"Hofmeister","year":"2021","journal-title":"J. Exp. Bot."},{"key":"ref_41","first-page":"228","article-title":"The Biology of Bryophytes: Evolution, Simplicity, and Adaptation","volume":"34","author":"Rensing","year":"2022","journal-title":"Plant Cell"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"205","DOI":"10.3372\/wi.33.33122","article-title":"Erpodium glaziovii (Erpodiaceae, Bryopsida) and Further Novelties from the Arabian Peninsula. Additions to the Bryophyte Flora of the Arabian Peninsula and Socotra 4","volume":"33","author":"Ochyra","year":"2003","journal-title":"Willdenowia"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1127\/nova.hedwigia\/73\/2001\/253","article-title":"Two Tropical Species New to the Bryophyte Flora of the Arabian Peninsula. Additions to the Bryophyte Flora of the Arabian Peninsula and Socotra 2","volume":"73","author":"Buck","year":"2001","journal-title":"Nova Hedwig."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Shaw, A.J., and Goffinet, B. (2000). Physiological Ecology of Bryophytes. Bryophyte Biology, Cambridge University Press.","DOI":"10.1017\/CBO9781139171304"},{"key":"ref_45","unstructured":"Goffinet, B., and Shaw, A.J. (2005). Mosses in Harsh Environments. Bryophyte Biology, Cambridge University Press. [2nd ed.]."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1890\/1540-9295(2003)001[0181:TWAYFD]2.0.CO;2","article-title":"The World at Your Feet: Desert Biological Soil Crusts","volume":"1","author":"Belnap","year":"2003","journal-title":"Front. Ecol. Environ."},{"key":"ref_47","first-page":"1","article-title":"Biological Soil Crusts: Underappreciated Contributors to Dryland Ecosystem Resilience","volume":"113","author":"Zundel","year":"2013","journal-title":"Biol. Environ. Proc. R. Ir. Acad."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1639\/0007-2745(1997)100[13:PARBOS]2.0.CO;2","article-title":"Phenology and Reproductive Biology of Syntrichia caninervis (Pottiaceae) in the Mojave Desert","volume":"100","author":"Stark","year":"1997","journal-title":"Bryologist"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1575","DOI":"10.1242\/jeb.02179","article-title":"Constraints of Tolerance: Why Are Desiccation-Tolerant Organisms So Small or Rare?","volume":"209","author":"Alpert","year":"2006","journal-title":"J. Exp. Biol."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Vanderpoorten, A., and Goffinet, B. (2009). Introduction to Bryophytes, Cambridge University Press.","DOI":"10.1017\/CBO9780511626838"},{"key":"ref_51","first-page":"275","article-title":"Chemical Evolution of a Travertine-Depositing River System: The Rapolano Terme Travertine System, Central Italy","volume":"9","author":"Drysdale","year":"2003","journal-title":"Aquat. Geochem."},{"key":"ref_52","first-page":"119","article-title":"Tufa Deposits from China","volume":"86","author":"Pentecost","year":"2001","journal-title":"Quat. Int."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/S0012-8252(96)00030-X","article-title":"A Review of Tufa and Travertine Deposits of the World","volume":"41","author":"Ford","year":"1996","journal-title":"Earth-Sci. Rev."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Goudie, A.S. (2013). Arid and Semi-Arid Geomorphology, Cambridge University Press.","DOI":"10.1017\/CBO9780511794261"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1071\/SR9940389","article-title":"Microbiotic Soil Crusts: A Review of Their Roles in Soil and Ecological Processes in the Rangelands of Australia","volume":"32","author":"Eldridge","year":"1994","journal-title":"Aust. J. Soil Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"390","DOI":"10.2307\/3899593","article-title":"Effects of Grazing on the Vegetation of the Blackbrush Association","volume":"40","author":"Jeffries","year":"1987","journal-title":"J. Range Manag."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1639\/0007-2745(2007)110[595:DIBAR]2.0.CO;2","article-title":"Desiccation-Tolerance in Bryophytes: A Review","volume":"110","author":"Proctor","year":"2007","journal-title":"Bryologist"},{"key":"ref_58","unstructured":"Zohary, M. (1973). Geobotanical Foundations of the Middle East, Fischer Verlag."},{"key":"ref_59","unstructured":"Hadley, D.G. (1987). Explanatory Notes to the Geological Map of the Sahl Al Matran Quadrangle, Saudi Arabian Deputy Ministry for Mineral Resources. Sheet 26C."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Rasul, N., and Stewart, I. (2015). Introduction to the Red Sea: Its Origin, Structure, and Environment. The Red Sea, Springer. Springer Earth System Sciences.","DOI":"10.1007\/978-3-662-45201-1"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1016\/j.lithos.2019.02.007","article-title":"Geodynamic Setting of Upper Miocene to Quaternary Alkaline Basalts from Harrat al \u2018Uwayrid (NW Saudi Arabia): Constraints from K-Ar Dating, Chemical and Sr-Nd-Pb Isotope Compositions, and Petrological Modeling","volume":"330\u2013331","author":"Altherr","year":"2019","journal-title":"Lithos"},{"key":"ref_62","unstructured":"Al-Bassam, A.M., Kahlil, A.R., and Kassem, O.M. (2014). Using Updated DurovPwin Program for Hydro-Chemical Data Processing: Case Study of Al-Ula Area, Saudi Arabia. Recent Adv. Environ. Sci. Geosci., 75\u201381."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"692","DOI":"10.1111\/cobi.12508","article-title":"Incorporating Geodiversity into Conservation Decisions","volume":"29","author":"Cromer","year":"2015","journal-title":"Conserv. Biol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"618","DOI":"10.1111\/cobi.12505","article-title":"The Theory Behind the Challenges of Conserving Nature\u2019s Stage in a Time of Rapid Change","volume":"29","author":"Lawler","year":"2025","journal-title":"Conserv. Biol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.ecolind.2018.03.068","article-title":"The Role of Geodiversity in Providing Ecosystem Services at Broad Scales","volume":"91","author":"Alahuhta","year":"2018","journal-title":"Ecol. Indic."},{"key":"ref_66","unstructured":"CMEP (2023). Inventory and Vegetation of AlUla Flora Project, Centre for Middle Eastern Plants, Part of the Royal Botanic Garden Edinburgh, Royal Commission for AlUla."},{"key":"ref_67","unstructured":"Hausleiter, A., Eichmann, R., and Al-Najem, M. (2018). Tayma Oasis (Saudi Arabia) and Its Surroundings\u2014A First Synthesis of the Flora, Vegetation, Natural Resources, and Floral History. Tayma I. Archaeological Exploration, Palaeoenvironment, Cultural Contact, Archaeopress."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Braun-Blanquet, J. (1964). Pflanzensoziologie, Grundz\u00fcge der Vegetationskunde, Springer. [3rd ed.].","DOI":"10.1007\/978-3-7091-8110-2"},{"key":"ref_69","first-page":"211","article-title":"Soziologie und Lebensstrategien Epiphytischer Bryophyten in Israel und Jordanien","volume":"61","author":"Frey","year":"1995","journal-title":"Nova Hedwig."},{"key":"ref_70","unstructured":"Hugonnot, V., P\u00e9pin, F., and Freedman, J. (Nova Hedwig., 2025). Bryophyte Communities from AlUla County (Saudi Arabia), Nova Hedwig., submitted."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"300","DOI":"10.11646\/phytotaxa.454.4.8","article-title":"A New Combination in the Genus Molendoa (Pottiaceae, Bryophyta)","volume":"454","author":"Brinda","year":"2020","journal-title":"Phytotaxa"},{"key":"ref_72","unstructured":"Rubasinghe, S.C.K. (2011). Phylogeny and Taxonomy of the Complex Thalloid Liverwort Family Cleveaceae Cavers. [Ph.D. Thesis, University of Edinburgh]."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"297","DOI":"10.5252\/cryptogamie-bryologie2019v40a23","article-title":"A Complete Description and Conservation Assessment of Riella affinis Howe & Underwood (Riellaceae, Sphaerocarpales) New to Continental Europe","volume":"40","author":"Puche","year":"2019","journal-title":"Cryptogam. Bryol."},{"key":"ref_74","first-page":"271","article-title":"Entosthodon commutatus Durieu et Mont. (Funariaceae, Bryopsida), New to Europe and Morocco","volume":"31","author":"Ros","year":"2010","journal-title":"Cryptogam. Bryol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/2\/170\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T10:25:46Z","timestamp":1759919146000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/2\/170"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,1,9]]},"references-count":74,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2025,1]]}},"alternative-id":["plants14020170"],"URL":"https:\/\/doi.org\/10.3390\/plants14020170","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,9]]}}}