Maryam Kokabi and Morteza Yousefzadi* Checklist of the marine macroalgae of Iran

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1 Botanica Marina 2015; 58(4): Maryam Kokabi and Morteza Yousefzadi* Checklist of the marine macroalgae of Iran DOI /bot Received 1 January, 2015; accepted 16 June, 2015; online first 10 July, 2015 Abstract: In this study, a revised checklist of macroalgae of Iran with an updated nomenclature and taxonomy has been compiled based on published records. Using currently accepted names, 309 species and infraspecific taxa of macroalgae have been identified to date, including 78 Chlorophyta (within 15 families), 70 Ochrophyta (Phaeophyceae; within 7 families) and 161 Rhodophyta (within 30 families). The brown alga Sargassum with 25 taxa was the most diverse genus, and the Rhodomelaceae (Rhodophyta) with 36 taxa was the most species-rich family. The Cheney ratio of 3.4 and the species composition of brown seaweeds suggest that the Iranian marine algal flora is warm-temperate. Sørensen similarity indices were used to compare the marine algal flora of Iran with that of Saudi Arabia within the Persian Gulf and the Sultanate of Oman within the Gulf of Oman. The algal diversity of the Iranian coast within the Gulf of Oman is less than that within the Persian Gulf, and this difference is attributed to undercollecting. Given that this is the first inclusive checklist of macroalgae of Iran, covering the coast lying within the Persian Gulf and the Gulf of Oman, it could serve as a foundation for future phycological and biogeographical studies of the taxa in the country and the region. Keywords: Gulf of Oman; Iran; macroalgae; Persian Gulf; Sørensen similarity index. Introduction Iran is a semi-arid country in southwestern Asia, extending from latitude 25 to 40 North and longitude 44 to 63 East, with a large variation in climate from north to south (Abbaspour et al. 2009). To the north, Iran borders the largest closed body of water in the world, the Caspian Sea, with a shoreline approximately 800 km long. In the south, *Corresponding author: Morteza Yousefzadi, Faculty of Marine Science and Technology, Department of Marine Biology, Hormozgan University, Bandar Abbas, Iran, Morteza110110@gmail.com Maryam Kokabi: Young Researchers and Elite Club, Mashhad Branch, Islamic Azad University, Mashhad, Iran Iran has a coastline of approximately 5000 km lying within the Persian Gulf and the Gulf of Oman (Dibajnia et al. 2012), to which it is connected through the narrow Strait of Hormuz. The Persian Gulf is a semi-enclosed and evaporative marginal sea with a mean depth of about 35 m and dotted by many small islands (Hassanzadeh et al. 2011, Kourosh Niya et al. 2013). In comparison, the Gulf of Oman is connected to the Indian Ocean and provides direct access to open seas (Pak and Farajzadeh 2007). Compared with those in the north, the water boundaries in the south of Iran lie within the sub-tropical high-pressure zone, and thus are characterized by low precipitation and high aridity. Along this coastal area a variety of ecosystems, such as mangrove forests, rocky shores and coral reefs (Pak and Farajzadeh 2007, Abuzinada et al. 2008), provide suitable habitats for seaweed growth (Figure 1). Seaweeds or marine benthic macroalgae form a rather heterogeneous group of primary producers, including three major divisions based on the nature of their pigments and other fundamental characteristics: Chlorophyta (green algae), Ochrophyta (Phaeophyceae; brown algae) and Rhodophyta (red algae). Seaweeds are recognized to be resources of high ecological and economic value because they provide a wide assortment of bioactive compounds that could be added to food, medicine, and to biofertilizers (Hong et al. 2007, Dhargalkar and Verlecar 2009, Koch et al. 2013). However, there is no comprehensive listing of the seaweed species that have been recorded in Iran. Furthermore, knowledge of the marine seaweed diversity of Iran is largely inaccessible to researchers because most data are scattered in regional papers or reports (often in Persian), and the recent checklist of the Persian Gulf by John and Al-Thani (2014) covers only the Persian Gulf coast of Iran. Therefore, the objective of this study was to collect and update all the information on macroalgal diversity in Iran in a comprehensive checklist accessible to the international community. History of seaweed study in Iran The history of seaweed study in Iran dates back to 1845, when Endlicher and Diesing reported the names of six species of brown and red algae from Khark (also Kharg) and Kish Islands in the Persian Gulf. In 1939, Børgesen reported 77 species of marine algae from Kish Island and

2 308 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran Figure 1: Coastline of Iran and neighbouring countries, showing the islands and areas mentioned in the text. Bushehr province. Several papers were published in the 20th century on species of marine algae in the Persian Gulf, but most of them covered the Arabian side of the Gulf (e.g. Nizamuddin and Gessner 1970, Basson 1978, De Clerck and Coppejans 1996). Consequently, there was a wide gap in records of Iranian marine macroalgae between 1939 and 1996, when Sohrabipour and Rabiei (1996, 1999) published the very first checklist on the marine algae of the Hormozgan province of Iran. They listed 144 species of seaweeds, including 36 Chlorophyta, 33 Ochrophyta (Phaeophyceae) and 75 Rhodophyta in the Persian Gulf (Sohrabipour and Rabiei 1999). In 2004, Sohrabipour et al. reported 119 species of seaweeds, including 29 Chlorophyta, 31 Phaeophyceae and 59 Rhodophyta in the Persian Gulf, Lengeh area of Iran. Sohrabipour and Rabiei (2007, 2008b) also published a checklist of 62 Chlorophyta and 74 Rhodophyta from the Iranian coast of the Persian Gulf and the Gulf of Oman. In 2011, Gharanjik and Rohani-Ghadikolaei reported 150 species of Iranian seaweeds, including 39 Chlorophyta, 40 Phaeophyceae and 71 Rhodophyta. Recent taxonomic studies on the seaweeds of Iran have been conducted by Sohrabipour and Rabiei, on Gracilaria (Rabiei et al. 2008) and Gracilariopsis (Bellorin et al. 2008, Sohrabipour and Rabiei 2008a). In addition, Shams et al. (2013) made the first reports of five species of Sargassum from Iran. The ecological features of Iranian seaweeds have been studied to a small extent. In 2012, Fatemi et al. studied the seaweed biomass from intertidal rocky shores of Qeshm Island. The authors collected and identified a total of 73 species of marine algae from the southern and the northern parts of the island. Dadolahi-Sohrab et al. (2012) noted seasonal changes in seaweed diversity on the northern coast of the Persian Gulf (Bushehr Province). Seasonal variation in macroalgal assemblages was also observed along the eastern coasts of Qeshm Island (Persian Gulf; authors unpublished data). Interest in studies of the potential economic importance of seaweeds has significantly increased in recent years. The agarophyte taxa, especially the Iranian endemic species Gracilariopsis persica, has received more attention than other local seaweeds (Salehi et al. 2011, Saeidniaa et al. 2012, Matinfar et al. 2013, Yousefi et al. 2013, Heydari et al. 2014). Aside from studies focusing on the commercial potential of Iranian seaweeds, some researchers have investigated the pharmaceutical features of Iranian seaweeds to find novel medicines (Zandi et al. 2007, Derakhshesh et al. 2011, Ghannadi et al. 2013, Jassbi et al. 2013, Kokabi et al. 2013, Yousefzadi et al. 2014). Methods of checklist production This checklist of the seaweed species of Iran is based on an exhaustive bibliographical search in which both local reports and international publications were consulted for species records. All taxon names and their classification were verified by reference to AlgaeBase

3 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran 309 Table 1: Species names and authorities of Chlorophyta of Iran. Boodleaceae 1 Cladophoropsis fasciculata (Kjellman) Wille = Cladophoropsis sundanensis Reinbold [b, c, d, g] 2 Cladophoropsis membranacea (C. Agardh) Børgesen [a, b, c, d, e, g] Bryopsidaceae 3 Bryopsis corymbosa J. Agardh = Bryopsis implexa De Notaris [a, d, g] 4 Bryopsis hypnoides J.V. Lamouroux [a, d, g] 5 Bryopsis pennata J.V. Lamouroux [c, d, e, g] 6 Bryopsis pennata var. minor J. Agardh = Bryopsis pennatula J. Agardh [b, d, e, g] 7 Bryopsis pennata var. secunda (Harvey) Collins et Hervey [b, c, d, g] 8 Bryopsis plumosa (Hudson) C. Agardh [b, c, d, e, f, g] Caulerpaceae 9 Caulerpa brachypus Harvey [d] 10 Caulerpa chemnitzia (Esper) J.V. Lamououx = Caulerpa peltata J. V. Lamouroux [b, c, d, e, g] 11 Caulerpa cupressoides (Vahl) C. Agardh [e] 12 Caulerpa faridii Nizamuddin [e] 13 Caulerpa fastigiata Montagne [c, d, g] 14 Caulerpa manorensis Nizamuddin [a, b, d, g] 15 Caulerpa mexicana Sonder ex Kützing = Caulerpa crassifolia (C. Agardh) J. Agardh [a, d, g] 16 Caulerpa racemosa var. macrophysa (Sonder ex Kützing) W.R. Taylor [b, c, d, g] 17 Caulerpa scalpelliformis (Turner) C. Agardh [d, e] 18 Caulerpa sertularioides (S.G. Gmelin) M. Howe [a, b, c, d, e, f, g] 19 Caulerpa sertularioides (S.G. Gmelin) M. Howe forma farlowii (Weber-Van Bosse) Børgesen [b, c, d, f, g] 20 Caulerpa taxifolia (Vahl) C. Agardh [b, c, d, e, f, g] Cladophoraceae 21 Chaetomorpha aerea (Dillwyn) Kützing [a, b, c, d, g] 22 Chaetomorpha antennina (Bory) Kützing [d, e, g] 23 Chaetomorpha brachygona Harvey [b, d, g] 24 Chaetomorpha californica F.S. Collins [b, c, d, g] 25 Chaetomorpha vieillardii (Kützing) M.J. Wynne = Chaetomorpha crassa sensu auctorum (C. Agardh) Kützing [d] 26 Chaetomorpha gracilis (Kützing) Kützing [b, e, g] 27 Chaetomorpha linum (O.F. Muller) Kützing [a, c, d, g] 28 Chaetomorpha spiralis Okamura [e] 29 Cladophora albida (Nees) Kützing = Cladophora magdalenae Harvey [b, d, g] 30 Cladophora aokii Yamada [e] 31 Cladophora oligoclada Harvey [e] 32 Cladophora coelothrix Kützing [a, d, g] 33 Cladophora echinus (Biasoletto) Kützing [a, c, d, g] 34 Cladophora flexuosa (O.F. Müller) Kützing = Cladophora gracilis (Griffiths) Kützing [c, d, g] 35 Cladophora herpestica (Montagne) Kützing = Cladophoropsis javanica (Kützing) P.C. Silva = Cladophoropsis zollingerii (Kützing) Reinbold [a, d, g] 36 Cladophora koeiei Børgesen [b, c, d, g] 37 Cladophora nitellopsis Børgesen [a, b, c, d, g] 38 Cladophora radiosa (Suhr) Kützing [a, b, d, g] 39 Cladophora rugulosa G. Martens [e] 40 Cladophora sericioides Børgesen [a, b, d, g] 41 Cladophora vagabunda (Linnaeus) van den Hoek = Cladophora fascicularis (Mertens ex C. Agardh) Kützing [b, d, e, g] 42 Rhizoclonium riparium (Roth) Harvey = Rhizoclonium implexum (Dillwyn) Kützing [b, d, e, g] 43 Rhizoclonium tortuosum (Dillwyn) Kützing = Chaetomorpha capillaris (Kützing) Børgesen [a, c, d, g] Codiaceae 44 Codium bartlettii C.K. Tseng et W.J. Gilbert [e] 45 Codium cylindricum Holmes [e] 46 Codium flabellatum P.C. Silva ex Nizamuddin [e] 47 Codium fragile (Suringar) Hariot [d, e, g] 48 Codium geppiorum O.C. Schmidt [e] 49 Codium indicum S.C. Dixit = Codium iyengarii Børgesen [e] 50 Codium papillatum C.K.Tseng et W.J. Gilbert [a, b, c, d, e, g] 51 Codium repens P. Crouan et H. Crouan [e] 52 Codium subtubulosum Okamura [e]

4 310 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran Table 1 (continued) Derbesiaceae 53 Derbesia marina (Lyngbye) Kjellman [d, g, h] Dichotomosiphonaceae 54 Avrainvillea amadelpha (Montagne) A. Gepp et E.S. Gepp [d] 55 Avrainvillea calathina Kraft et Olsen-Stojkovich [a, b, c, e, g] 56 Avrainvillea erecta (Berkeley) A. Gepp et E.S. Gepp [e] Halimedaceae 57 Halimeda discoidea Decaisne [d] 58 Halimeda tuna (Ellis and Solander) Lamouroux [e] Phaeophilaceae 59 Phaeophila dendroides (P.l. Crouan et H.M. Crouan) Batters [a, d, g] Polyphysaceae 60 Acetabularia calyculus J.V. Lamouroux [b, c, d, g] 61 Parvocaulis clavatus (Yamada) S. Berger, U. Fettweiss, S. Gleissberg, L.B. Liddle, U. Richter, H. Sawitzky et G.C. Zuccarello = Acetabularia clavata Yamada [a, b, d, g] 62 Parvocaulis parvulus (Solms-laubach) S. Berger, U. Fettweiss, S. Gleissberg, L.B. Liddle, U. Richter, H. Sawitzky et G.C. Zuccarello = Polyphysa parvula (Solms-laubach) Schnetter et Bule Meyer = Acetabularia parvula Solms-Laubach = Acetabularia moebiii Solms-laubach [a, b, g] Siphonocladaceae 63 Dictyosphaeria cavernosa (Forskål) Børgesen [a, b, c, d, f, g] 64 Siphonocladus feldmannii Børgesen [d, g] Ulotrichaceae 65 Spongomorpha arcta (Dillwyn) Kützing [e] Ulvaceae 66 Ulva californica Wille [a, b, d, g] 67 Ulva clathrata (Roth) C. Agardh = Enteromorpha clathrata (Roth) Greville J. Agardh [a, b, c, d, e, g] 68 Ulva compressa Linnaeus = Enteromorpha compressa (Linnaeus) Nees [a, b, c, d, g] 69 Ulva flexuosa Wulfen = Enteromorpha flexuosa (Wulfen) J. Agardh [a, c, d, e, f, g] 70 Ulva intestinalis Linnaeus = Enteromorpha intestinalis (Linnaeus) Link [b, d, f, g] 71 Ulva lactuca Linnaeus = Ulva fasciata Delile [a, b, c, d, e, f, g] 72 Ulva linza Linnaeus = Enteromorpha linza (Linnaeus) J. Agardh [b, d, g] 73 Ulva prolifera O.F. Muller = Enteromorpha prolifera (O.F. Muller) J. Agardh [a, c, d, g] 74 Ulva reticulata Forsskål [d, g] 75 Ulva rigida C. Agardh [d, e] Ulvellaceae 76 Ulvella viridis (Reinke) R. Nielsen, C.J.O. Kelly et B. Wysor = Entocladia viridis Reinke = Endoderma viride (Reinke) Lagerh = Acrochaete viridis (Reinke) Nielsen [a, d, g] Valoniaceae 77 Valonia utricularis (Roth) C. Agardh [d] 78 Valoniopsis pachynema (G. Martens) Børgesen [e] References: a: Silva et al. (1996); b: Sohrabipour and Rabiei (1999); c: Sohrabipour et al. (2004); d: Sohrabipour and Rabiei (2007); e: Gharanjik and Rohani-Ghadikolaei (2011); f: Kokabi et al. (2014); g: John and Al-Thani (2014); h: Sohrabipour and Rabiei (2005). ( and updated when necessary (Guiry and Guiry 2014). This website contains the latest taxonomic and floristic publications on algae and the taxonomy and nomenclature of the entries are updated regularly. The checklist is arranged in alphabetical order of families under three different algal groups, the Chlorophyta, Ochrophyta (Phaeophyceae) and Rhodophyta. The species are also listed alphabetically under each family. For some taxa, the taxonomic synonyms are presented after an equal sign and references to the original publications are indicated in brackets as follows: Current name and Author = Synonym and Author [Reference]. To determine the relative tropical or temperate affinities of the algal flora, Cheney s floristic ratio was applied. This ratio is (R+C)/P, where R is the number of taxa of Rhodophyta, C the number of taxa of Chlorophyta, and P the number of species belonging to the Ochrophyta, Class Phaeophyceae. The ratio characterizes the geographic nature of an algal flora (Kaldy et al. 1995, Augytė 2011, Augytė and Shaughnessy 2014). A ratio of < 3.0 indicates a cold-water flora, whereas ratios of > 6.0 indicate a tropical flora; intermediate values represent a mixed floristic affinity (e.g. warm-temperate; Kaldy et al. 1995, Witman and Roy 2009).

5 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran 311 Table 2: Species names and authorities of Ochrophyta (Phaeophyceae) of Iran. Acinetosporaceae 1 Feldmannia indica (Sonder) Womersley et A. Bailey = Hincksia indica (Sonder) J. Tanaka [e] 2 Feldmannia irregularis (Kützing) Hamel [a, b, c, g] 3 Feldmannia mitchelliae (Harvey) H.S. Kim = Hincksia mitchelliae (Harvey) P.C. Silve [a, b, c, g] Bachelotiaceae 4 Bachelotia antillarum (Grunow) Gerloff [a, b, c, g] Chordariaceae 5 Stilophora iranica Børgesen [a, g] 6 Tinocladia crassa (Suringar) Kylin [e] Dictyotaceae 7 Canistrocarpus cervicornis (Kützing) De Paula et De Clerck = Dictyota cervicornis Kuetzing = Dictyota indica Sonder et Kützing = Dictyota divaricata (J. Agardh) J. Agardh [a, b, c, e, g] 8 Dictyopteris australis (Sonder) Askenasy [a, e, g] 9 Dictyota ciliolata Sonder ex Kützing [a, b] 10 Dictyota dichotoma (Hudson) J.V. Lamouroux = Dictyota volubilis Kützing [a, b, c, e, f, g] 11 Dictyota friabilis Setchell [e, g] 12 Dictyota implexa (Desfontaines) J.V. Lamouroux [a, b, c, g] 13 Lobophora variegata (J.V. Lamourax) Womersley ex E.C. Oliveira [a, b, c, e, g] 14 Padina australis Hauck [a, b, c, e, f, g] 15 Padina boergesenii Allenderet Kraft [a, b, c, e, g] 16 Padina boryana Thivy = Padina tenuis Bory [a, b, e, g] 17 Padina distromatica Hauck [e] 18 Padina dubia Hauck [e] 19 Padina glabra Gaillard [e] 20 Padina gymnospora (Kützing) Sonder = Padina crassa Yamada [a, b, e, g] 21 Padina minor Yamada [c, g] 22 Padina pavonica (Linnaeus) Thivy [b, c, e, g] 23 Padina tetrastromatica Hauck [a, b, c, e, f, g] 24 Spatoglossum asperum J. Agardh [e] 25 Spatoglossum dichotomum C.K. Tseng et B. Lu [e] 26 Spatoglossum variabile Figari et De Notaris [a, b, c, e, g] 27 Stoechospermum polypodioides (J.V. Lamouroux) J. Agardh = Stoechospermum marginatum (C. Agardh) Kützing [a, b, c, e, g] Sargassaceae 28 Hormophysa cuneiformis (J.F. Gmelin) P.C Silva [a, b, c, g] 29 Nizamuddinia zanardinii (Schiffner) P.C. Silva = Sargassum yemenense forma monstrosum Zanardini [e] 30 Polycladia indica (Thivy et Doshi) Draisma, Ballesteros, F. Rousseau et T. Thibaut = Cystoseira indica (Thivy et Doshi) Mairh [e] 31 Polycladia myrica (S.G. Gmelin) Draima, Ballesteros, F. Rousseau et T. Thibaut = Cystoseira myrica (S.G. Gmelin) C. Agardh [a, b, c, e, g] 32 Sargassum acinaciforme Montagne [a, g] 33 Sargassum angustifolium C. Agardh [a, b, c, e, g] 34 Sargassum aquifolium (Turner) C. Agardh = Sargassum crassifolium J. Agardh = Sargassum binderi Sonder ex J. Agardh [a, c, g, i] 35 Sargassum assimile Harvey [e] 36 Sargassum baccularia (Mertens) C. Agardh [i] 37 Sargassum boveanum J. Agardh [a, b, c, g] 38 Sargassum boveanum J. Agardh var. aterrimum Grunow [a, g, i] 39 Sargassum fluitans (Børgesen) Børgesen [a, b, c, g] 40 Sargassum gemmiphorum C.K. Tseng et B. Lu [i] 41 Sargassum glaucescens J. Agardh [e] 42 Sargassum henslowianum C. Agardh [i] 43 Sargassum herbaceum Kützing [a, g] 44 Sargassum ilicifolium (Turner) C. Agardh = Sargassum cristaefolium C. Agardh [e, g] 45 Sargassum latifolium (Turner) C. Agardh [a, b, c, g] 46 Sargassum longifructum C.K. Tseng et B. Lu [i] 47 Sargassum oligocystum Montagne [c, g] 48 Sargassum persicum Kützing [a, g] 49 Sargassum spinuligerum Sonder [i] 50 Sargassum swartzii J. Agardh = Sargassum acutifolium Greville [a, e, g] 51 Sargassum tenerrimum J. Agardh [e] 52 Sargassum tenuissimum (Endlicher et Diesing) Grunow = Sargassum vulgare C. Agardh var. tenuissimum Endlicher et Diesing [a, g]

6 312 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran Table 2 (continued) 53 Sargassum virgatum C. Agardh [a, g] 54 Sargassum vulgare C. Agardh [a, b, c, g] 55 Sargassum vulgare var. angustifolium (Turner) C. Agardh [a, g] 56 Sargassum vulgare var. latifolium Endlicher et Diesing [a, g] 57 Sirophysalis trinodis (Forsskal) Kützing = Cystoseira trinodis (Forskål) C. Agardh [a, b, c, e, g] 58 Stephanocystis neglecta (Setchell et N.L. Gardner) Draisma, Ballesteros, F. Rousseau et T. Thibaut = Cystoseira neglecta Setchell et Gardner [a, b, c, g] 59 Turbinaria conoides (J. Agardh) Kützing [b, e, g] 60 Turbinaria ornata (Turner) J. Agardh [e] Scytosiphonaceae 61 Colpomenia sinuosa (Mertens ex Roth) Derbes et Solier [a, b, c, e, f, g] 62 Iyengaria stellata (Børgesen) Børgesen [a, b, c, e, f, g] 63 Jolyna laminarioides S.M. Guimarães [e] 64 Rosenvingea floridana (W.R. Taylor) W.R. Taylor [a, b, g] 65 Rosenvingea intricata (J. Agardh) Børgesen [c, e, g] 66 Rosenvingea orientalis (J. Agardh) Børgesen [e] 67 Scytosiphon dotyi M.J. Wynne = Scytosiphon lomentaria f. cylindricus nanus Tokida [c, g] Sphacelariaceae 68 Sphacelaria novae-hollandiae Sonder [a, b, g] 69 Sphacelaria rigidula Kützing [a, b, c, g] 70 Sphacelaria tribuloides Meneghini [a, b, g] References: a: Silva et al. (1996); b: Sohrabipour and Rabiei (1999); c: Sohrabipour et al. (2004), e: Gharanjik and Rohani-Ghadikolaei (2011); f: Kokabi et al. (2013), 2014); g: John and Al-Thani (2014); i: Shams et al. (2013). Using Sørensen similarity indices (Magurran 1988, Nguyen et al. 2013), the Iranian seaweed flora was compared with those of the Persian Gulf coast of Saudi Arabia and also the Sultanate of Oman (including the Gulf of Oman and Arabian Sea coastlines of this country). These two countries have longer coastlines along the Persian Gulf and the Gulf of Oman than the other neighbouring countries (Figure 1), and they are comparable with the Iranian coastlines from the diversity standpoint. The floristic data for all neighbouring countries were obtained from AlgaeBase and John and Al-Thani (2014). A Sørensen index of 1.0 indicates complete similarity (when the two sets of species are identical) and 0 if the sites have no species in common (Magurran 1988). Ochrophyta (Class Phaeophyceae) To date, a total of 70 taxa of Ochrophyta (Phaeophyceae), from 7 families, have been reported from the Iranian coastline of the Persian Gulf and the Gulf of Oman (Table 2). The most diverse family among brown algae is the Sargassaceae with 33 taxa, followed by the Dictyotaceae with 21 species. Furthermore, the genera Sargassum and Padina have the most taxa (25 and 10 species, respectively) among the brown algae of Iran (Table 2). The species Ectocarpus cryptophilus Børgesen is rejected here because it is recorded only once from Iran in 1970 (John and Al-Thani 2014). Results Chlorophyta A total of 78 taxa belonging to the Chlorophyta, within 15 families, are presented in this checklist (Table 1). The family Cladophoraceae contained the most species (23), followed by the Caulerpaceae with 12 species. Moreover, Cladophora and Caulerpa were the most diverse genera among the Chlorophyta with 13 and 12 taxa, respectively (Table 1). Rhodophyta A total of 161 taxa belonging to the Rhodophyta, within 30 families, have been reported from the Iranian coastlines bordering the Persian Gulf and the Gulf of Oman (Table 3). The most species-rich family among the red algae was the Rhodomelaceae with 36 taxa, followed by the Gracilariaceae with 16 species. Furthermore, Gracilaria with 14 and Hypnea with 10 were the most species-rich genera in the Rhodophyta of Iran (Table 3).

7 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran 313 Table 3: Species names and authorities of Rhodophyta of Iran. Acrochaetiaceae 1 Acrochaetium robustum Børgesen [c, g, m] 2 Acrochaetium savianum (Meneghini) Nägeli = Audouinella saviana (Meneghini) Woelkerling [m] Ahnfeltiaceae 3 Ahnfeltia plicata (Hudson) E.M. Fries [a, g] Bonnemaisoniaceae 4 Asparagopsis taxiformis (Delile) Trevisan [a, g] Callithamniaceae 5 Aglaothamnion cordatum (Børgesen) Feldman-Mazoyer = Callithamnion cordatum Børgesen [a, b, c, g, n] 6 Aglaothamnion hookeri (Dillwyn) Maggs et Hommersand [e] 7 Crouania attenuata (C. Agardh) J. Agardh [a, b, g] Ceramiaceae 8 Antithamnion cruciatum (C. Agardh) Nägeli = Antithamnion cruciatum var. radicans (J. Agardh) F.S. Collins [a, g] 9 Centroceras clavulatum (C. Agardh) Montagne [a, b, e, f, g, m, n] 10 Ceramium cimbricum H.E. Petersen = Ceramium fastigiatum (Wulfen ex Roth) Harvey [a, b, c, g, n] 11 Ceramium cimbricum f. flaccidum (H.E. Petersen) G. Funari et Serio = Ceramium fastigiatum f. flaccidum H.E. Petersen [c, g, m] 12 Ceramium cruciatum F.S. Collins et Hervey [a, g] 13 Ceramium diaphanum (Lightfoot) Roth = Ceramium tenuissimum (Roth) Areschoug [e] 14 Ceramium luetzelburgii O.C. Schmidt [a, g] 15 Ceramium manorense P. Anand [b, g, m] 16 Ceramium maryae Weber-van Bosse [a, g] 17 Ceramium tenerrimum (Martens) Okamura [e] 18 Ceramium truncatum H.E. Petersen [e] 19 Corallophila bella (Setchell et Gardner) R.E. Norris = Centroceras bellum Setchel et Gardner [a, b, c, g, m] 20 Corallophila kleiwegii Weber-van Bosse = Corallophila apiculata (Yamada) R. Norris = Centroceras apiculatum Yamada [a, b, c, g, i] 21 Gayliella flaccida (Harvey ex Kützing) T.O. Cho et L.J. McIvor = Ceramium flaccidum (Harvey ex Kützing) Ardissone [a, b, c, g] Champiaceae 22 Champia compressa Harvey [c, e, g, m] 23 Champia compressa var. scindica Harvey [a, b, e, g] 24 Champia globulifera Børgesen [a, b, c, g, m] 25 Champia indica Børgesen [a, b, g, m] 26 Champia kotschyana (C. Agardh) Harvey [a, e, g] 27 Champia parvula (C. Agardh) Harvey [a, b, c, n, g, m] 28 Champia zonata (J. Agardh) J. Agardh [e] Corallinaceae 29 Amphiroa fragilissima (Linnaeus) J.V. Lamouroux [b, g, m] 30 Hydrolithon boreale (Foslie) Chamberlain [a, g] 31 Hydrolithon farinosum (J.V. Lamouroux) D. Penrose et Y.M. Chamberlain = Fosliella farinosa (J.V. Lamouroux) M. Howe [a, b, c, g, m, n] 32 Jania adhaerens J.V. Lamouroux [a, b, c, e, g, m] 33 Jania pumila J.V. Lamouroux [a, g] 34 Jania rubens (Linnaeus) J.V. Lamouroux [a, b, c, g, m] 35 Jania tenella (Kützing) Grunow [c] 36 Pneophyllum fragile Kützing = Melobesia lejolisii Rosanoff [a, g] Cystocloniaceae 37 Hypnea aspera Kützing = Hypnea boergesenii T. Tanaka [e, m] 38 Hypnea charoides J.V. Lamouroux [e, m] 39 Hypnea cornuta (Kützing) J. Agardh [a, b, c, e, g, m, n] 40 Hypnea ecklonii Suhr [e] 41 Hypnea flagelliformis Greville ex J. Agardh [a, b, g] 42 Hypnea hamulosa (Esper) Lamouroux [e] 43 Hypnea musciformis (Wulfen) J.V. Lamouroux [a, b, e, g, m] 44 Hypnea pannosa J. Agardh [a, b, c, e, g, m] 45 Hypnea spinella (C. Agardh) Kützing = Hypnea cervicornis J. Agardh [a, b, c, e, f, g, m, n] 46 Hypnea valentiae (Turner) Montagne [a, b, e, g, m] Dasyaceae 47 Dasya anastomosans (Weber-van Bosse) M.J. Wynne = Dasya pilosa (Weber-van Bosse) A. Millar [e] 48 Dasya baillouviana (S.G. Gmelin) Montagne = Dasya pedicellata (C. Agardh) C. Agardh [a, b, c, g] 49 Dasya ocellata (Grateloup) Harvey [a, g] 50 Heterosiphonia crispella (C. Agardh) M.J. Wynne = Heterosiphonia crispella var. laxa (Børgesen) M.J. Wynne [a, g, m]

8 314 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran Table 3 (continued) Delesseriaceae 51 Apoglossum spathulatum (Sonder) Womersley et Shepley = Hypoglossum spathulatum (Sonder) Kützing [a, g] 52 Myriogramme okhaensis Børgesen [a, g] 53 Taenioma nanum (Kützing) Papenfuss [a, b, g] Erythrotrichiaceae 54 Erythrotrichia carnea (Dillwyn) J. Agardh [a, b, c, g, m, n] 55 Erythrocladia irregularis Rosenvinge [a, b, c, m] 56 Sahlingia subintegra (Rosenvinge) Kornmann = Erythrocladia subintegra Rosenvinge [a, g] Furcellariaceae 57 Furcellaria lumbricalis (Hudson) J.V. Lamouroux = Furcelaria fastigiata (Turner) J.V. Lamouroux [b, g] Galaxauraceae 58 Actinotrichia fragilis (Forsskål) Børgesen [e] 59 Dichotomaria obtusata (J. Ellis et Solander) Lamarck [m] 60 Galaxaura rugosa (Ellis et Solander) J.V. Lamouroux = Galaxaura lapidescens (Ellis et Solander) J.V. Lamouroux [a, b, c, g] Gelidiaceae 61 Gelidium chilense (Montagne) Santelices et Montalva [a, b, e, g, m] 62 Gelidium crinale (Hare ex Turner) Gaillon [a, c, g, m] 63 Gelidium micropterum Kützing [e] 64 Gelidium pusillum (Stackhouse) Le Jolis [a, b, c, e, g, m, n] 65 Gelidium pusillum var. pulvinatum (C. Agardh) Feldmann [a, g] Gelidiellaceae 66 Gelidiella acerosa (Forsskål) Feldmann et G. Hamel [a, b, c, e, g, m, n] 67 Gelidiella myrioclada (Børgesen) Feldmann et G. Hamel [c, e, g] 68 Gelidiella ramellosa (Kützing) Feldmann et G.Hamel [e] Gigartinaceae 69 Chondracanthus acicularis (Roth) Fredericq = Gigartina acicularis (Roth) J.V. Lamouroux [b, e, g] 70 Chondrus ocellatus Holmes [e] Gracilariaceae 71 Gracilaria arcuata Zanardini [e, m] 72 Gracilaria armata (C. Agardh) Greville [m] 73 Gracilaria canaliculata Sonder = Gracilaria crassa Harvey ex J. Agardh [a, b, c, n, g, m] 74 Gracilaria corticata (J. Agardh) J. Agardh [a, b, c, e, n, g, m] 75 Gracilaria foliifera (Forskål) Børgesen [a, b, c, e, n, g, m] 76 Gracilaria gracilis (Stackhouse) Steentoft, L.M. Irvine et W.F Farnham [e] 77 Gracilaria mammillaris (Montagne) Howe = Gracilaria veleroae Dawson [m] 78 Gracilaria millardetii (Montagne) J. Agardh [e] 79 Gracilaria pulvinata Skottsberg = Gracilaria pygmaea V.J. Chapman [e, m] 80 Gracilaria robusta Setchell [m] 81 Gracilaria salicornia (C. Agardh) Dawson = Corallopsis cacalia J. Agardh [a, b, c, n, g, m] 82 Gracilaria spinulosa (Okamura) C.F. Chang et B.M. Xia [e] 83 Gracilaria textorii (Suringar) De Toni [e, m] 84 Gracilaria vieillardii P.C. Silva [m] 85 Gracilariopsis longissima (S.G. Gmelin) M. Steentoft, L.M. Irvine et W.F. Farnham [a, b, c, n, g, l] 86 Gracilariopsis persica A.M. Bellorin, J. Sohrabipour et E.C. Oliveira [f, g, j] Halymeniaceae 87 Corynomorpha prismatica (J. Agardh) J. Agardh [g, o] 88 Grateloupia comorinii Børgesen [a, g] 89 Grateloupia filicina (J.V. Lamouroux) C. Agardh [m] 90 Grateloupia somalensis Hauck [e, m] 91 Halymenia dilatata Zanardini [e] 92 Halymenia porphyraeformis Parkinson [e, m] Liagoraceae 93 Dermonema pulvinatum (Grunow ex Holmes) K.C. Fan [e] 94 Dermonema virens (J. Agardh) Pedroche et Avila Ortiz [e] 95 Helminthocladia australis Harvey [e, m] 96 Liagora distenta (Mertens ex Roth) J.V. Lamouroux [c, g] 97 Liagora filiformis K.C. Fan et W.H. Li [e]

9 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran 315 Table 3 (continued) Lomentariaceae 98 Ceratodictyon variabile (J. Agardh) R.E. Norris = Gelidiopsis variabilis (Greville ex J. Agardh) F. Schmitz [m] 99 Lomentaria corallicola Børgesen [a, g] 100 Lomentaria divaricata (Durant) M.J. Wynne = Lomentaria baileyana (Harvey) Farlow [a, b, g] Phyllophoraceae 101 Ahnfeltiopsis pygmaea (J. Agardh) P.C. Silva et DeCew [e] Rhodomelaceae 102 Acanthophora muscoides (Linnaeus) Bory [a, b, e, g, m] 103 Acanthophora nayadiformis (Delile) Papenfuss [a, e, g] 104 Acanthophora spicifera (M. Vahl) Børgesen [a, b, c, e, g, m, n] 105 Chondria arcuata Hollenberg [c, g] 106 Chondria bernardii P. Dangeard [a, b, c, g, n] 107 Chondria cornuta Børgesen [a, b, e, g, m] 108 Chondria dasyphylla (Woodward) C. Agardh [a, b, c, g, m] 109 Chondria nidifica Harvey [a, b, g] 110 Chondria oppositiclada E.Y. Dawson [a, b, g] 111 Chondria seticulosa (Forsskål) C. Agardh = Chondria hypnoides Børgesen [a, g] 112 Chondrophycus glandulifer (Kützing) Lipkin et P.C. Silva = Laurencia glandulifera (Kützing) Kützing [a, g] 113 Chondrophycus undulatus (Yamada) Garbary et Harper = Laurencia undulata Yamada [e] 114 Digenea simplex (Wulfen) C. Agardh [a, b, c, g, m] 115 Herposiphonia secunda (C.Agardh) Ambronn [c, g, m] 116 Herposiphonia secunda f. tenella (C. Agardh) M.J. Wynne [a, b, c, g] 117 Laurencia dendroidea J. Agardh = Laurencia majuscula (Harvey) A.H.S. Lucas [a, c, g, m] 118 Laurencia filiformis (C. Agardh) Montagne [e, m] 119 Laurencia intricata J.V. Lamouroux [a, b, c, g, m] 120 Laurencia obtusa (Hudson) Lamouroux [a, c, e, g, n] 121 Laurencia platyclada Børgesen [e] 122 Laurencia snyderae E.Y. Dawson [b, c, e, g, m, n] 123 Leveillea jungermannioides (Hering et Martens) Harvey [a, b, c, e, f, g, m] 124 Lophocladia lallemandii (Montagne) F. Schmitz [a, g] 125 Lophosiphonia obscura (C. Agardh) Falkenberg = Lophosiphonia subadunca (Kützing) Falkenberg [a, g] 126 Melanothamnus somalensis Bornet et Falkenberg [e] 127 Neosiphonia ferulacea (Suhr ex J. Agardh) S.M. Guimaräes et M.T. Fujii = Polysiphonia ferulacea Suhr ex J. Agardh [a, g] 128 Osmundea pedicularioides (Børgesen) G. Funari, Serio et Cormaci = Laurencia pedicularioides Børgesen [a, b, g] Palisada perforata (Bory) K.W. Nam = Chondrophycus papillosus 129 (C. Agardh) Garbary et Harper = Laurencia papillosa (C. Agardh) Greville [a, b, c, e, f, g, m] 130 Palisada thuyoides (Kützing) Cassano, Sentíes, Gil-Rodríguez et M.T. Fujii = Laurencia paniculata (C. Agardh) J. Agardh [e] 131 Polysiphonia brodiei (Dillwyn) Sprengel [a, g] 132 Polysiphonia codicola Zanardini ex Frauenfeld [c, g, n] 133 Polysiphonia crassicollis Børgesen [a, c, g] Polysiphonia denudata (Dillwyn) Greville ex Harvey = Polysiphonia variegata 134 (C. Agardh) Zanardini [a, b, g, m] 135 Polysiphonia kampsaxii Børgesen [a, g] 136 Polysiphonia scopulorum var. villum (J. Agardh) Hollenberg = Lophosiphonia villum (J. Agardh) Setchell et Gardner [a, b, c, g, m] 137 Tolypiocladia glomerulata (C. Agardh) F. Schmitz = Roschera glomerulata (C. Agardh) Weber-van Bosse [a, b, g, m] Rhodymeniaceae 138 Botryocladia leptopoda (J. Agardh) Kylin [b, e, g, m] 139 Rhodymenia sp. [b, c, g] Sarcomeniaceae 140 Cottoniella filamentosa (M.A. Howe) Børgesen [e] 141 Platysiphonia delicata (Clemente) Cremades = Platysiphonia miniata (C. Agardh) Børgesen [b, g, m] Scinaiaceae 142 Scinaia carnosa (Kützing) J. Agardh [e] 143 Scinaia fascicularis (Børgesen) Huisman [e] 144 Scinaia furcellata (Turner) J. Agardh [e] 145 Scinaia hatei Børgesen [m] 146 Scinaia moniliformis J. Agardh [e] 147 Scinaia tsinglanensis C.K. Tseng [b, c, g, m]

10 316 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran Table 3 (continued) Sebdeniaceae 148 Sebdenia flabellata (J. Agardh) P.G. Parkinson [e, m] Solieriaceae 149 Sarconema filiforme (Sonder) Kylin [a, b, c, n, e, f, g, m] 150 Solieria anastomosa P.W. Gabrielson et Kraft [b, c, g] 151 Solieria dura (Zanardini) F. Schmitz [e] 152 Solieria filiformis (Kützing) Gabrielson [b, c, n, g, m] 153 Solieria robusta (Greville) Kylin [a, b, c, n, e, g, m] 154 Solieria tenuis J. Zhang et E. Xia [e] 155 Wurdemannia miniata (Sprengel) Feldmann et G. Hamel [c, g, m] Spyridiaceae 156 Spyridia filamentosa (Wulfen) Harvey [a, b, c, g, m] Stylonemataceae 157 Chroodactylon ornatum (C. Agardh) Basson = Asterocytis ornata (C. Agardh) Hamel [a, b, c, n, g] 158 Stylonema alsidii (Zanardini) K. Drew [a, b, c, g] Wrangeliaceae 159 Anotrichium tenue (C. Agardh) Nägeli = Griffithsia tenuis C. Agardh [a, b, g, m] 160 Griffithsia globulifera Harvey ex Kützing [a, g] 161 Gymnophycus sp. [g] References: a: Silva et al. (1996); b: Sohrabipour and Rabiei (1999); c: Sohrabipour et al. (2004); e: Gharanjik and Rohani-Ghadikolaei (2011); f: Kokabi et al. (2014); g: John and Al-Thani (2014); j: Bellorin et al. (2008); k: Rabiei et al. (2008); l: Sohrabipour and Rabiei (2008a); m: Sohrabipour and Rabiei (2008b); n: Rabiei et al. (2005); o: Sohrabipour and Rabiei (2006). Diversity of marine macroalgae in Iran compared to the neighbouring countries Iran, with a total of 309 taxa, appears to be the most diverse among the countries surrounding the Persian Gulf and the Gulf of Oman (Figure 2). The Sørensen similarity index was applied to examine the similarity of seaweeds along the Iranian coasts of the Persian Gulf and the Gulf of Oman (Table 4). Furthermore, the diversity of the Iranian seaweeds was compared with those of the Persian Gulf coast of Saudi Arabia and also the Sultanate of Oman (including the Gulf of Oman and Arabian Sea coastlines of this country; Table 5). The indices were lower than 0.5 (Cs < 0.5) for all of these comparisons, but comparing the Persian Gulf coast of Iran with Saudi Arabia resulted in a higher coefficient (Cs = 0.57; Table 6). Table 4: Comparison of the species composition of the seaweed flora of Iran on the Persian Gulf coasts with that on the Gulf of Oman coasts. Numbers of species Rhodophyta Ochrophyta Chlorophyta Iranian coastlines Number of taxa Common Sørensen similarity index Persian Gulf Gulf of Oman 192 Table 5: Comparison of species composition of the seaweed flora of Iran with those of two neighbouring countries. Iran Persian Gulf (Iran) Gulf of Oman (Iran) Saudi Arabia Sultanate of Oman Region Bahrain Kuwait Qatar UAE Neighbouring countries N a = 309 N b N a+b Cs Saudi Arabia Sultanate of Oman Figure 2: Numbers of species of green, brown and red seaweeds in Iran, on the Persian Gulf and the Gulf of Oman coasts of Iran, and in neighbouring countries. N a, number of species in Iran; N b, number of species in each neighbouring country; N a + b, number of species in common with Iran; Cs, Sørensen similarity index = (2 N a + b )/(N a +N b ).

11 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran 317 Table 6: Comparison of species composition of the seaweed flora of the Persian Gulf coasts of Iran with the seaweed flora of Saudi Arabia, and that of the Gulf of Oman coast of Iran with the seaweed flora of the Sultanate of Oman. Neighbouring countries N a N b N a+b Cs Saudi Arabia Sultanate of Oman N a, number of species on the corresponding coast of Iran; N b, number of species in each neighbouring country; N a + b, number of species in common with Iran; Cs, Sørensen similarity index = (2 N a + b )/(N a +N b ). Discussion In a changing world in which anthropogenic impacts are dramatically altering the biota of the Earth, the task of discovering and documenting biodiversity has been given an increased sense of urgency (De Clerck et al. 2013). This paper is the most comprehensive checklist of marine macroalgae for the biogeographic region that includes the Persian Gulf and the Gulf of Oman coastlines of Iran. A total of 309 taxa (78 Chlorophyta, 70 Phaeophyceae and 161 Rhodophyta) are listed in this paper, compiled from both regional and international publications on the macroalgae from the coastal waters of Iran. The results indicate that Sargassum (Ochrophyta, Phaeophyceae) with 25 taxa was the most diverse genus, and that the Rhodomelaceae (Rhodophyta) with 36 taxa was the most species-rich family among the macroalgae of Iran. Globally, the relative abundance of species of red, brown and green algae varies depending on latitude linked to sea temperature. The species richness of brown seaweeds increases toward higher latitudes and colder regions, whereas green and red seaweeds increase from the polar regions to the tropics (in the northern hemisphere; Lee 2008, Witman and Roy 2009). Based on this information, the ratio of (R+C)/P and other similar ratios (e.g. C/P) (Witman and Roy 2009) change with latitude. In this study, the Cheney s ratio of 3.4 for the Iranian seaweed flora indicates a mixed algal flora with strong cold-water affinity, but classifying the seaweed flora of Iran as cold-temperate might be misleading. It is clear that the reduction of species diversity of brown algae in warm waters does not apply to all groups. The Dictyotales (e.g. the genus Padina) (Witman and Roy 2009, Ni-Ni-Win et al. 2011, Silberfeld et al. 2013) and some genera belonging to the Fucales, such as Sargassum, are more speciesrich in the tropics, whereas the Laminariales, many other members of the Fucales, and other brown algal groups are generally more diverse in temperate or cold waters (Witman and Roy 2009). Given this information, it is important to consider the composition of brown seaweeds when determining the climatic affinities of an algal flora based on Cheney s ratio. Given that 50% of the brown algae reported from Iran belong to the genera Sargassum and Padina (Table 2), we suggest that the seaweed flora of this coastal region should be regarded as warm-temperate. Because Iran has a longer coastline along the Persian Gulf and the Gulf of Oman (5000 km) than its neighbouring countries surrounding these two water bodies, it was not surprising that the number of taxa reported for the entire northern coastlines of the Persian Gulf and the Gulf of Oman would be greater than that for the neighbouring countries (Figure 2). However, the total number of seaweeds reported from the Sultanate of Oman (223 taxa; AlgaeBase) is greater than that of the northern coastline of the Gulf of Oman (192 taxa). The Sørensen similarity index between these two floras is also low (Cs < 0.5; Table 6). These results indicate that the species composition of the two areas is different, and this may be because the Sultanate of Oman also has a shoreline on the northern Arabian Sea. If it was possible to compare the seaweed flora of only the Gulf of Oman coastline of the Sultanate of Oman, the results might be different. Although we cannot ignore the fact that the shoreline of the Sultanate of Oman on the northern Arabian Sea probably accounts for the higher seaweed diversity in this country than in Iran (Wynne 2000), we are of the opinion that the lower number of seaweed species on the Iranian coastline of the Gulf of Oman and the low similarity to neighbouring countries (Cs = 0.38) are probably the result of the relatively low number of taxonomic studies, especially along the northern coast of the Gulf of Oman. The lower number of species along this coast than along the Persian Gulf coast of Iran (215 taxa) supports this idea. The Iranian coastline of the Persian Gulf is bordered by three provinces (Khuzestan, Bushehr and Hormozgan from west to east) with many academic and research centres studying marine algae and other marine organisms. In comparison, the Iranian coastline of the Gulf of Oman is bordered by only one province (Sistan and Baluchestan) with few research centres. After comparing the number of publications on Iranian seaweeds along these two coasts, it becomes evident that the marine algal flora of the Persian Gulf coast of Iran is more intensively studied than the coast of the Gulf of Oman. Considering the less extreme oceanological conditions along the Gulf of Oman coast of Iran and less urbanization (in contrast to the Persian Gulf), it is very probable that many more species are waiting to be discovered in this area.

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14 320 M. Kokabi and M. Yousefzadi: Marine macroalgae of Iran Morteza Yousefzadi Faculty of Marine Science and Technology, Department of Marine Biology, Hormozgan University, Bandar Abbas, Iran, Morteza Yousefzadi is an Associate Professor of Plant Biology at Hormozgan University, Bandar Abbas, Iran. His recent research has concentrated on aquaculture and identification of seaweeds to provide raw material for applications in nanoscience, discovery of anticancer and antifouling properties, and bioremediation.