Seaweeds of the Greek Coasts: Rhodophyta Excluding Ceramiales

ISSN: 0001-5113 ACTA ADRIAT., ORIGINAL SCIENTIFIC PAPER AADRAY 60(1): 3 - 24, 2019

Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales

Konstantinos TSIAMIS1,2* and Panayotis PANAYOTIDIS1

1 Hellenic Centre for Marine Research (HCMR), Institute of Oceanography, Anavyssos 19013, Attica, Greece

2 European Commission, Joint Research Centre (JRC), Ispra, Italy

*Corresponding author: [email protected]

An updated checklist of the red seaweeds (Rhodophyta), excluding the order Ceramiales, of the Greek coasts is provided, based on literature records, critically reviewed from present-day taxonomic and nomenclatural aspects. The total number of genera, species and infraspecific taxa currently accepted is 78, 138 and 4 respectively. The occurrence of each taxon in the North Aegean, South Aegean and Ionian Seas is given. Knowledge gaps are pointed, with 57 taxa pending confirmation of their presence marked. Moreover, 13 excludenda and 12 inquirenda taxa are briefly discussed. In overall, a solid updated baseline of the Rhodophyta (excluding Ceramiales) taxa occurence in Greece is provided, critical for future tailor-targeting seaweed studies.

Key words: Aegean Sea, red algae, Rhodophyta, checklist, Ionian Sea

INTRODUCTION & GEISSLER (1974). Later on, ATHANASIADIS (1987) compiled a critically reviewed catalogue Phycological studies on marine macroalgae of marine seaweeds, but concerning only the have been carried out along the Greek coasts Aegean Sea. In addition, the annotated check- since the early 19th century (SIBTHORP, 1813; lists of the Mediterranean seaweed flora by GREVILLE, 1826; BORY, 1832), resulting in several RIBERA et al. (1992), GALLARDO et al. (1993) and new species and genera and in numerous records GÓMEZ GARRETA et al. (2001) included seaweeds of common Mediterranean taxa. However, the occurring in Greece. major part of these studies has not been updated Aiming to update the knowledge of the in a modern context, while the provided data in Greek marine seaweed flora, the present work the classical publications are usually too limited is focusing exclusively in the red seaweeds to allow unequivocal identification, frequently (Rhodophyta) excluding the order Ceramiales resulting in taxonomic confusion (TSIAMIS & Oltmanns, aiming to deliver a solid updated PANAYOTIDIS, 2016a). and annotated baseline of the Greek taxa, criti- A first attempt to produce a checklist of the cal for future tailor-targeting studies. Our work Greek marine seaweeds was made by DIANNE- constitutes the final part in a series that previ- LIDIS (1950), followed by the work of GERLOFF ously treated the Phaeophyceae, the Ulvophy- 4 ACTA ADRIATICA, 60(1): 3 - 24, 2019 ceae and the Ceramiales (Rhodophyta) of the Taxa are listed alphabetically. For each taxon Greek coasts (TSIAMIS et al., 2013a, 2014; TSIAMIS applied synonyms to previous records from & PANAYOTIDIS, 2016a). Greece are provided. The distribution of each accepted taxon is given for the three biogeo- MATERIAL AND METHODS graphic regions: North Aegean, South Aegean and Ionian Sea (Fig. 1). Due to space limitation, From the early 19th century until 2017, 13 only one reference is given for each region, giv- PhD theses and about 150 research papers ing priority to publications that include descrip- have been published on seaweeds from Greece. tions and/or illustration(s). Additional references Master and Bachelor Degree theses as well as are available by the authors on request. conference contributions have not been taken into account for this study. Records of rhodo- RESULTS phytes in all other publications have been critically reviewed from present-day taxonomic and The present checklist recognizes at least nomenclatural aspects, following the on-line 142 Rhodophyta species and infraspecific taxa data provided by SILVA (2018) and GUIRY & GUIRY (excluding Ceramiales), within 78 genera, to (2018), unless otherwise specified in the “Notes”. occur in Greece (Table 1). In addition, there are The checklist has been compiled following 57 taxa pending confirmation of their presence the scheme in TSIAMIS & PANAYOTIDIS (2016a): on the Greek coasts (Table 2). 1. accepted taxa: there is at least one documented record from Greece, in the form of Taxa Excludenda description and/or illustration(s), 2. pending confirmation of their presence: Bangia atropurpurea (Mertens in Roth) undocumented records or records with no C. Agardh sufficient documentation to permit accurate Based on MULLER et al. (2003) molecular identification under modern standards, work, Bangia atropurpurea is apparently a 3. excludenda: misidentifications, freshwater species, distinct from B. fuscopurpu- 4. inquirenda: taxa not typified, with uncertain rea found in the marine environment. Therefore, taxonomic application. the several Greek records (e.g. ANAGNOSTIDIS, 1968; NIKOLAIDIS & HARITONIDIS, 1990; MALEA & HARITONIDIS, 2001), which all lack sufficient documentation, should be excluded from the Greek marine flora, possibly referring to B. fuscopurpurea.

Bangia thaerasiae Bory Originally described by BORY (1832: 334) based on specimens collected from Santorini Island (South Aegean Sea), but was later proven to belong to a diatom by DE TONI (1897: 12).

Chondrus crispus Stackhouse = Chondrus norvegicus (Gunnerus) Lam- ouroux = Gymnogongrus norvegicus (Gunnerus) J. Fig. 1. Accepted Rhodophyta algal taxa (excluding Cera- Agardh miales) within each biogeographic region of the Greek This North Atlantic alga does not occur in the seas Mediterranean Sea and the Greek records from Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 5 the Aegean Sea (RAULIN, 1869; TSEKOS et al., we prefer to cite the several Greek records 1982) and Ionian Sea (TSEKOS & HARITONIDIS, (e.g. GIACCONE, 1968a; DIAPOULIS et al., 1986; 1977), which all lack documentation, should be NIKOLAIDIS & HARITONIDIS, 1990), which all lack considered as misidentifications, possibly refer- documentation, within the list of taxa exclu- ring to a Gymnogongrus species. denda for the Greek flora.

Colaconema caespitosum (J. Agardh) Liagora ceranoides Lamouroux Jackelman, Stegenga & Bolton 0This species has been reported from the 0= Rhodothamniella codii (Crouan frat.) J. Peloponnese by BORY (1832: 329) but actually Feldmann referring to L. viscida (see also ATHANASIADIS, The single undocumented record by DIAPOUA - 2016: 137), and from Rhodes Island (South AegeB - LIS et al. (1986, as Rhodothamniella codii) from an Sea) by REINBOLD (1898) but correspoding the Saronikos Gulf (South Aegean Sea) should to a misidentification of Ganonema farinosum be a misidentification, possibly referring to (HAMEL, 1931: 310). The record by DIANNELIDIS Colaconema codicola (Børgesen) Jackelman, et al. (1977) from Rhodes Island, which lacks Stegenga & Bolton (see also BIDOUX & MAGNE, documentation, should be also treated as a misi- 1989; VERLAQUE et al., 2015: 139). dentification since this species does not seem to occur in the Eastern Mediterranean Sea. Dumontia contorta (S.G. Gmelin) Ruprecht = Dumontia incrassata (O.F. Müller) Lam- Mastocarpus stellatus (Stackhouse in ouroux Withering) Guiry The single undocumented record by TSEKOS 0= Gigartina stellata (Stackhouse in Wither- & HARITONIDIS (1977) from the Ionian Sea should ing) Batters be a misidentification, since this circumpolar The single record by GERLOFF & GEISSLER species does not occur in the Mediterranean Sea. (1974) from the Ionian Sea is without justifica- tion and should be a misidentification, since this Gigartina pistillata (S.G. Gmelin) Stackhouse species does not occur in the Eastern Mediter- The Greek records from the North Aegean ranean Sea. Sea (HARITONIDIS & TSEKOS, 1975) and Ionian Sea (HARITONIDIS & TSEKOS, 1976; TSEKOS et Nemalion multifidum (Lyngbye) Chauvin al., 1982; HARITONIDIS et al., 1986), which all lack 0This species does not seem to occur in descriptions or illustrations, should be excluded the Mediterranean Sea and the Greek records from the Greek flora as misidentifications, since from the Ionian Sea (HARITONIDIS & TSEKOS, this species does not seem to occur in the East- 1976; TSEKOS & HARITONIDIS, 1977; TSEKOS et al., ern Mediterranean Sea. 1982), which all lack descriptions or illustrations, should be excluded from the Greek flora, possi- Iridaea cordata (Turner) Bory bly referring to Nemalion lubricum (see also LE This South America species is not known to GALL & SAUNDERS, 2010). occur in the Mediterranean Sea and the undocumented Greek record by TSEKOS et al. (1982) from Phyllophora crispa (Hudson) Dixon the North Aegean Sea should be considered as a 0= Phyllophora epiphylla (O.F. Müller) Bat- misidentification. ters = Phyllophora rubens auctorum Kallymenia reniformis (Turner) J. Agardh Based on ATHANASIADIS (2016: 524-528) this = Fucus reniformis Turner species does not occur in the Mediterranean Since most of the Mediterranean records of Sea and the numerous records reported from the this NE Atlantic species should refer to Mer- basin should refer to Phyllophora spiralis (= P. edithia microphylla (ATHANASIADIS, 2016: 650), nervosa). Thus, the several Greek records (e.g. 6 ACTA ADRIATICA, 60(1): 3 - 24, 2019

POLITIS, 1928; KOUSSOURIS, 1976; DIAPOULIS & Erythrotrichia simplex Dangeard nom. inval. HARITONIDIS, 1984), which all lack documenta- Entity of uncertain taxonomic status since tion, should be excluded, probably referring to no type has been designated. There are sev- P. spiralis. eral records of this species from Greece (e.g. DIAPOULIS & HARITONIDIS, 1987a; CHRYSSOVER- Phyllophora pseudoceranoides (S.G. Gmelin) GIS, 1995) but in all cases lacking documentation. Newroth & Taylor ex Dixon & Irvine = Phyllophora membranifolia (Goodenough Goniolithon subtenellum Foslie & Woodward) Endlicher nom. illeg. = Lithothamnion subtenellum (Foslie) Lem- There are no documented records of this oine North Atlantic species from the Mediterranean Based on ATHANASIADIS (2016: 289), the taxo- Sea, and the Greek records from the Aege- nomic position of this species is uncertain, since an Sea (CANDARGY, 1899; ANAGNOSTIDIS, 1968; it has been associated with both Lithophyl- NIKOLAIDIS & HARITONIDIS, 1990) probably cor- lum orbiculatum (Foslie) Foslie and Hydroli- respond to misidentifications. thon samoënse (Foslie) Keats & Chamberlain. In Greece, Goniolithon subtenellum has been Taxa Inquirenda reported by GIACCONE (1968a) from the Aegean Amphiroa articulata (Bory) Athanasiadis Sea but without documentation. The species is = Nullipora articulata Bory accepted by CORMACI et al. (2017) as Lithophyl- Erected by BORY (1832: 206) based on plants lum orbiculatum (Foslie) Foslie. collected from Sapience islet (Peloponnese). ATHANASIADIS (2016: 292) placed the species Grateloupia neglecta (Bory) Kützing under the genus Amphiroa based on Bory’s pro- = Gelidium neglectum Bory tologue and plates. However, in the absence of Originally described from the Peloponesse the type specimen and a detailed diagnosis of it by BORY (1832: 324), it has been associated we agree with CORMACI et al. (2017) to place the with Grateloupia filicina (DE TONI, 1905). Since species within the the taxa inquirenda. Bory’s original material has never been re- examined its taxonomic status remains unclari- Bangia sericea Bory fied (see also GARGIULO et al., 2013: 6). This taxon was erected by BORY (1832: 334) based on plants collected from the South Pelo- Halymenia rhodymenioides Ercegović ponnese coasts. Based on him and following A rare and deep-water species (below 60 studies (PREDA, 1908-09) the species is close m depth) described by ERCEGOVIĆ (1949) from to Conferva atropurpurea (= Bangia atropurpu- Jabuka (Adriatic Sea), but with obscure taxo- rea), although this taxon is considered nowdays nomic status. It was later reported by SCH- to be a freshwater species (see taxa excludenda). NETTER & SCHNETTER (1981) from Kephallonia We prefer to cite Bangia sericea within the list Island (Ionian Sea), but lacking documentation. of taxa inquirenda until the type specimen is re- investigated. Lithophyllum fasciculatum auctorum = ? Lithothamnion fasciculatum sensu Are- Chylocladia pygmaea (Funk) Kylin schoug The specific status of this taxon remains The binomial Lithophyllum fasciculatum unclear and requires taxonomic re-investigation (Lamarck) Foslie is based on Millepora fascicula- (ALONGI et al., 2008: 149). In Greece, it was report- ta Lamarck, which according to BASSO et al. (2004) ed by CATRA & GIARDINA (2009) from Karpathos its type collection is heterogenous, containing Island, but without documentation. two disticnt plants: one belonging to an unde- scribed species of Clathromorphum, and the other belonging to a another Lithothamnion spe- Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 7 cies. The sporadic Greek records from the North uncertain, since the type has not been desig- Aegean (DIANNELIDIS, 1950) and South Aegean nated (BRODIE et al., 2007). The latter authors Sea (POLITIS, 1928) are without documentation state that it could correspond to Porphyra olivii and should be considered as douptful (see also (=Pyropia koreana). Based on ATHANASIADIS ATHANASIADIS, 1987: 40). (2016) it could represent a synonym of Po. leucostica (= Py. leucosticta). In Greece, the Lithothamnion hauckii Rothpletz species has been reported by KATSIKOPOULOS = Lithothamnion mamillosum Hauck nom. (1939) and HARITONIDIS & TSEKOS (1974), but illeg. without documentation. = Neogoniolithon mamillosum (Hauck) Setchell & Mason DISCUSSION Species of uncertain taxonomic status (WOE- LKERLING et al., 1998), but frequently reported The first checklist of Greek seaweeds (DIAN- from the Greek coasts (e.g. DIANNELIDIS et al., NELIDIS, 1950) included 44 currently accepted 1977; SCHNETTER & SCHNETTER, 1981; TSEKOS species and infraspecific taxa of Rhodophyta et al., 1982); however, none of the Greek records excluding Ceramiales. Later on, GERLOFF & contain any description or illustrations. GEISSLER (1974) listed 81, while ATHANASIADIS (1987) reported 122 taxa for the Aegean Sea only. Peyssonnelia codana (Rosenvinge) Levring Based on our study, the total number has pres- sensu Verlaque ently increased to 142 confirmed taxa, following the several new studies during the recent years Although the type material of Peyssonne- (e.g. TSIAMIS et al., 2010; TSIAMIS, 2012; CATRA & lia codana belongs to P. dubyi, Mediterranean ALONGI, 2013; TSIAMIS & PANAYOTIDIS, 2016b). plants differ from the latter (VERLAQUE, 1978). Combined with the annotated list of the Greek The single Greek record from Sithonia (North Ceramiales taxa (TSIAMIS & PANAYOTIDIS, Aegean Sea) by ATHANASIADIS (1987: 50-51, 2016a), the total number of accepted Rhodophyta as Peyssonnelia sp.) is in good agreement taxa along the Greek coasts is 262. with Verlaque’s concept, showing spermatangial The distribution of these 142 non-Cer- structures distinct from those of P. dubyi (ATHA- amiales along the Greek coasts has as follows: NASIADIS, 2016: 675). 120 taxa have been found in the N. Aegean, 120 taxa in the S. Aegean and 111 taxa in the Ionian Pneophyllum zonale (Crouan frat.) Sea (Fig. 1). Chamberlain The Greek marine flora seems to host = Melobesia zonalis (Crouan frat.) Foslie far less non-Ceramiales Rhodophyta taxa com- Previously considered as a synonym of Pne- paring with the neighboring Italian coasts, ophyllum coronatum, but the type specimen of where 290 taxa have been recorded (FURNARI P. zonale’s basionym (Hapalidium coronatum et al., 2010). This difference definitely reflects Crouan frat.) does not match with P. coronatum. the limited studies that have been hitherto con- Hence, the species should be included within the ducted in Greece, and generally in the eastern taxa inquirenda (see also CORMACI et al., 2017). Mediterranean Sea, with several coastal regions In Greece, P. zonale has been reported from and islands still remaining poorly surveyed, par- the Aegean Sea by ATHANASIADIS (1987, the ticularly in the sublittoral and circalittoral zones. description possibly matches with P. coronatum) All taxa reported in the current study and GIACCONE (1968a, as Melobesia zonalis, have been critically reviewed from present-day without documentation). taxonomic and nomenclatural aspects, provid- ing also all synonyms ever cited in the literature Porphyra atropurpurea (Olivi) De Toni of Greek seaweeds, minimizing thus the vast The taxonomic position of this species is taxonomic confusion of the Greek records. Still, 8 ACTA ADRIATICA, 60(1): 3 - 24, 2019 too many taxa (57) are pending confirmation in We conclude that there are still major gaps Greece. This is due to the lack of documenta- in the documentation of the marine flora of the tion, since most records have been given in form Aegean and Ionian Seas, and surveys in the of species names without deposition of material unexplored areas and particularly in deeper in public herbaria. Several other taxa are treated habitats will further increase the number of taxa. as excludenda or inquirenda (13 and 12 taxa Our study can offer a solid updated baseline respectively), which underlies the taxonomic of the current knowledge of Rhodophyta taxa difficulties, particularly concerning the identifi- (excluding Ceramiales) in Greece, critical for cation of members of the genera Acrochaetium, future tailor-targeting seaweed studies. Gracilaria, Porphyra, and the non-articulated calcified taxa of the order Corallinales (e.g. Fos- ACKNOWLEDGEMENTS liella, Goniolithon, Lithophyllum, Lithotham- nion, Pneophyllum, Titanoderma). We are grateful to the reviewers for their comments.

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Received: 22 June 2018 Accepted: 15 May 2019 Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 13

Table 1. Accepted Rhodophyta algal taxa (excluding Ceramiales) in the North and South Aegean, and the Ionian Seas. For each taxon a basic reference is provided together with applied synonyms in previous Greek records

Taxa North Aegean South Aegean Ionian Sea DIAPOULIS & DIAPOULIS & Acrochaetium crassipes (Børgesen) Børgesen ATHANASIADIS, 1987 HARITONIDIS, HARITONIDIS, = Audouinella crassipes (Børgesen) Garbary 1987a 1987b Acrochaetium humile (Rosenvinge) Børgesen ATHANASIADIS, 1987 = Audouinella humilis (Rosenvinge) Garbary Acrochaetium mediterraneum (Levring) Athanasiadis TSIRIKA & DIAPOULIS & = Audouinella mediterranea (Levring) Ballesteros nom. LAZARIDOU, 1994 HARITONIDIS, HARITONIDIS, 1984 inval. 2005 Acrochaetium microscopicum (Nägeli ex Kützing) Nägeli TSIRIKA & CHRYSSOVERGIS & = Audouinella microscopica (Nägeli ex Kützing) LAZARIDOU, 1994 HARITONIDIS, PANAYOTIDIS, 1995 Woelkerling 2005 Acrochaetium secundatum (Lyngbye) Nägeli TSIRIKA & = Audouinella secundata (Lyngbye) Dixon DIANNELIDIS, 1953 LAZARIDOU, 1994 HARITONIDIS, = Acrochaetium virgatulum (Harvey) Batters 2005 = Audouinella virgatula (Harvey) Dixon Acrodiscus vidovichii (Meneghini) Zanardini SCHNETTER & = Chondrus vidovichii Meneghini ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 = Cryptonemia vidovichii (Meneghini) Zanardini Acrosymphyton purpuriferum (J. Agardh) Sjöstedt DIANNELIDIS et SCHNETTER & = Dudresnaya purpurifera J. Agardh ATHANASIADIS, 1987 al., 1977 SCHNETTER, 1981 = Helminthiopsis purpurifera (J. Agardh) Papenfuss KOUSSOURIS et Amphiroa beauvoisii Lamouroux BITIS, 1988 al., 1973 Amphiroa cryptarthrodia Zanardini SCHNETTER & LAZARIDOU, 1994 = Amphiroa verruculosa Kützing nom. illeg. (1) SCHNETTER, 1981 CATRA & Amphiroa fragilissima (Linnaeus) Lamouroux GIARDINA, 2009 Amphiroa rigida Lamouroux ATHANASIADIS, 1987 LAZARIDOU, 1994 BITIS, 1988 SARTONI & DE Amphiroa rubra (Philippi) Woelkerling ATHANASIADIS, 1987 BIASI, 1999 PANAYOTIDIS & Asparagopsis armata Harvey CHRYSSOVERGIS, CATRA & CHRYSSOVERGIS, = Falkenbergia rufolanosa (Harvey) Schmitz 1995 ALONGI, 2012 1998 Asparagopsis taxiformis (Delile) Trevisan TSIAMIS, 2012 TSIAMIS, 2012 TSIAMIS, 2012 ATHANASIADIS, DIANNELIDIS et SCHNETTER & Bangia fuscopurpurea (Dillwyn) Lyngbye 1987(2) al., 1977 SCHNETTER, 1981

Beckerella dentata (Kützing) Athanasiadis = Beckerella mediterranea H. Huvé GIACCONE, 1968b HUVE, 1962 = Ptilophora mediterranea (H. Huvé) Norris = Phyllophora aegei Giaccone nom. inval.

Bonnemaisonia asparagoides (Woodward) C. Agardh DIAPOULIS et al., ATHANASIADIS, 1987 = Hymenoclonium serpens (Crouan frat.) Batters 1986 Bonnemaisonia clavata Hamel ATHANASIADIS, 1987 Botryocladia botryoides (Wulfen) J. Feldmann = Botryocladia uvaria (C. Agardh) Kylin nom. illeg. SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Chrysymenia uvaria (C. Agardh) J. Agardh nom. illeg. SCHNETTER, 1981 = Gastroclonium uvarium (C. Agardh) Kützing nom. illeg. CATRA & TSIAMIS & Botryocladia madagascariensis G. Feldmann GIARDINA, 2009 VERLAQUE, 2011 14 ACTA ADRIATICA, 60(1): 3 - 24, 2019

DIAPOULIS & Calosiphonia vermicularis (J. Agardh) Schmitz ATHANASIADIS, 1987 HARITONIDIS, 1987a SCHNETTER & Caulacanthus ustulatus (Mertens ex Turner) Kützing ATHANASIADIS, 1987 SCHNETTER, 1981 Champia parvula (C. Agardh) Harvey ATHANASIADIS, 1987 LAZARIDOU, 1994 BITIS, 1988 Chondracanthus acicularis (Roth) Fredericq DIAPOULIS & DIANNELIDIS et = Gigartina acicularis (Roth) Lamouroux BITIS, 1988 HARITONIDIS, 1984 al., 1977 = Sphaerococcus acicularis (Roth) C. Agardh Chondracanthus teedei (Mertens in Roth) Kützing = Chondroclonium teedei (Mertens in Roth) Kützing DIANNELIDIS et ANAGNOSTIDIS, 1968 BITIS, 1988 = Gigartina teedei (Mertens in Roth) Lamouroux al., 1977 = Sphaerococcus teedei (Mertens in Roth) C. Agardh TSIRIKA & Choreonema thuretii (Bornet) Schmitz ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, 2005 Chroodactylon ornatum (C. Agardh) Basson = Asterocytis ornata (C. Agardh) Hamel SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Asterocytis ramosa (Thwaites in Harvey) Gobi ex SCHNETTER, 1981 Schmitz SCHNETTER & Chrysymenia ventricosa (Lamouroux) J. Agardh ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 Chylocladia verticillata (Lightfoot) Bliding = Chylocladia squarrosa (Kützing) Thuret = Chylocladia kaliformis (Withering) Greville in W.J. CATRA & Hooker ATHANASIADIS, 1987 BITIS, 1988 GIARDINA, 2009 = Gastroclonium kaliforme (Withering) Ardissone = Lomentaria kaliformis var. squarrosa (Kützing) J. Agardh Colaconema daviesii (Dillwyn) Stegenga DIAPOULIS & = Acrochaetium daviesii (Dillwyn) Nägeli ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, = Audouinella daviesii (Dillwyn) Woelkerling 1987b CATRA & SCHNETTER & Contarinia peyssonneliiformis Zanardini ATHANASIADIS, 1987 GIARDINA, 2009 SCHNETTER, 1981 Corallina caespitosa Walker, Brodie & Irvine WALKER et al., 2009 Corallina elongata Ellis & Solander (3) ATHANASIADIS, ATHANASIADIS, 1987 BITIS, 1988 = Corallina mediterranea Areschoug 1987 Corallina officinalis Linnaeus DIANNELIDIS, 1953 LAZARIDOU, 1994 BITIS, 1988 Cryptonemia lomation (Bertoloni) J. Agardh SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Sphaerococcus lactuca C. Agardh nom. illeg. SCHNETTER, 1981 Dudresnaya verticillata (Velley in Withering) Le Jolis TSIAMIS et al., SCHNETTER & ATHANASIADIS, 1987 = Dudresnaya coccinea (C. Agardh) Crouan frat. 2013b SCHNETTER, 1981 Erythrotrichia carnea (Dillwyn) J. Agardh ATHANASIADIS, 1987 LAZARIDOU, 1994 BITIS, 1988 = Erythrotrichia ceramicola (Lyngbye) Areschoug Ganonema farinosum (Lamouroux) Fan & Wang TSIAMIS, 2012 TSIAMIS, 2012 TSIAMIS, 2012 = Cladostephus dubium Bory TSIAMIS et al., Gastroclonium clavatum (Roth) Ardissone ATHANASIADIS, 1987 BITIS, 1988 2013b CATRA & SCHNETTER & Gelidiella lubrica (Kützing) J. Feldmann & Hamel ATHANASIADIS, 1987 GIARDINA, 2009 SCHNETTER, 1981 DIAPOULIS & Gelidium crinale (Turner) Gaillon ANAGNOSTIDIS, 1968 HARITONIDIS, BITIS, 1988 = Acrocarpus crinalis (Turner) Kützing 1987a Gelidium pectinatum Montagne (4) SCHNETTER & TSIAMIS, 2012 LAZARIDOU, 1994 = Gelidium bipectinatum Furnari nom. illeg. SCHNETTER, 1981 Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 15

TSIAMIS & Gelidium pulchellum (Turner) Kützing PANAYOTIDIS, 2016b Gelidium pusillum (Stackhouse) Le Jolis ATHANASIADIS, 1987 LAZARIDOU, 1994 BITIS, 1988 = Gelidium intricatum Lamouroux Gelidium spinosum (S.G. Gmelin) Silva SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Gelidium latifolium (Greville) Bornet SCHNETTER, 1981 TSEKOS & Gracilaria bursa-pastoris (S.G. Gmelin) Silva DIANNELIDIS et DIANNELIDIS, 1953 HARITONIDIS, = Gracilaria compressa (C. Agardh) Greville al., 1977 1977 Gracilaria corallicola Zanardini ATHANASIADIS, 1987 GIACCONE, 1968a Gracilaria dura (C. Agardh) J. Agardh DIANNELIDIS et DIANNELIDIS, 1953 GRUNOW, 1861 = Sphaerococcus durus C. Agardh al., 1977 SCHNETTER & Grateloupia filicina (Lamouroux) C. Agardh ATHANASIADIS, 1987 SCHNETTER, 1981 SCHNETTER & Gymnogongrus griffithsiae (Turner) Martius ATHANASIADIS, 1987 COPPEJANS, 1974 SCHNETTER, 1981 Halarachnion ligulatum (Woodward) Kützing ATHANASIADIS, 1987 GIACCONE, 1968a TSIAMIS, 2012 Halymenia elongata C. Agardh = Halymenia trigona auctorum ATHANASIADIS, 1987 GIACCONE, 1968b = ? Halymenia cavernicola Giaccone nom. inval. Halymenia floresii (Clemente) C. Agardh SCHNETTER & = Halymenia floresii var./f. macroptera Kützing ATHANASIADIS, 1987 GIACCONE, 1968a SCHNETTER, 1981 = Halymenia floresii var./f. tripinnata Kützing Halymenia latifolia Crouan frat. ex Kützing ATHANASIADIS, 1987 Hildenbrandia rubra (Sommerfelt) Meneghini SCHNETTER & ATHANASIADIS, 1987 = Hildenbrandia prototypus Nardo SCHNETTER, 1981 Hydrolithon boreale (Foslie) Chamberlain = Melobesia farinosa f. callithamnioides Foslie = Hydrolithon farinosum f. callithamnioides (Foslie) Serio TSEKOS & DIANNELIDIS et = Melobesia solmsiana Falkenberg DIANNELIDIS, 1953 HARITONIDIS, al., 1977 = Fosliella farinosa var. solmsiana (Falkenberg) Taylor 1977 = Melobesia farinosa var. solmsiana (Falkenberg) Lemoine = Melobesia callithamnioides sensu Falkenberg Hydrolithon farinosum (Lamouroux) Penrose & Chamberlain var. farinosum (5) ATHANASIADIS, 1987 LAZARIDOU, 1994 BITIS, 1988 = Melobesia farinosa Lamouroux = Fosliella farinosa (Lamouroux) Howe TSIAMIS & Hypnea anastomosans Papenfuss, Lipkin & Silva VERLAQUE, 2011 Hypnea cornuta (Kützing) J. Agardh TSIAMIS, 2012 ATHANASIADIS, SCHNETTER & Hypnea musciformis (Wulfen) Lamouroux ATHANASIADIS, 1987 1987 SCHNETTER, 1981 Hypnea spinella (C. Agardh) Kützing TSIAMIS, 2012 TSIAMIS, 2012 = ? Hypnea cervicornis auctorum TSIAMIS & Hypnea valentiae (Turner) Montagne VERLAQUE, 2011 Irvinea boergesenii (J. Feldmann) Wilkes, McIvor & TSIRIKA & Guiry ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, = Botryocladia boergesenii J. Feldmann 2005 CHRYSSOVERGIS, NIZAMUDDIN & Jania adhaerens Lamouroux BITIS, 1988 1995 LEHNBERG, 1970 16 ACTA ADRIATICA, 60(1): 3 - 24, 2019

DIAPOULIS & SCHNETTER & Jania longifurca Zanardini ex Zanardini DIANNELIDIS, 1953 HARITONIDIS, SCHNETTER, 1981 1987a Jania rubens (Linnaeus) Lamouroux var. rubens ATHANASIADIS, 1987 LAZARIDOU, 1994 BITIS, 1988 = Corallina rubens Linnaeus Jania rubens var. corniculata (Linnaeus) Yendo(6) SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Jania corniculata (Linnaeus) Lamouroux SCHNETTER, 1981 Jania virgata (Zanardini) Montagne = Corallina virgata Zanardini = Haliptilon virgatum (Zanardini) Garbary & Johansen SCHNETTER & = Corallina virgata var. attenuata (Kützing) Schiffner & ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 Schussnig = Corallina virgata var. penicillata Schiffner = Corallina granifera Ellis & Solander Liagora viscida (Forsskål) C. Agardh NIZAMUDDIN & SCHNETTER & ATHANASIADIS, 1987 = Liagora cladoniaeformis Bory LEHNBERG, 1970 SCHNETTER, 1981 Lithophyllum cystoseirae (Hauck) Heydrich = Melobesia cystoseirae Hauck = Titanoderma cystoseirae (Hauck) Woelkerling, TSIRIKA & Chamberlain & Silva (7) ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, = Dermatolithon cystoseirae (Hauck) Huvé 2005 = Dermatolithon papillosum var. cystoseirae (Hauck) Lemoine TSIRIKA & Lithophyllum dentatum (Kützing) Foslie ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, 2005 DIAPOULIS & Lithophyllum incrustans Philippi LAZARIDOU, 1994 BITIS, 1988 HARITONIDIS, 1984 Lithophyllum papillosum (Zanardini ex Hauck) Foslie SCHNETTER & = Dermatolithon papillosum (Zanardini ex Hauck) Foslie ATHANASIADIS, 1987 SCHNETTER, 1981 = Goniolithon papillosum (Zanardini ex Hauck) Foslie Lithophyllum pustulatum (Lamouroux) Foslie = Melobesia pustulata Lamouroux = Dermatolithon pustulatum (Lamouroux) Foslie = Dermatolithon hapalidioides (Crouan frat.) Foslie CATRA & SCHNETTER & ATHANASIADIS, 1987 = Titanoderma pustulatum (Lamouroux) Nägeli GIARDINA, 2009 SCHNETTER, 1981 = Titanoderma hapalidioides (Crouan frat.) Price, John & Lawson = Titanoderma verrucatum (Lamouroux) Chamberlain Lithophyllum racemus (Lamarck) Foslie BALBINI & BRESSAN, = Lithothamnion crassum Philippi LEMOINE, 1915 GIACCONE, 1968c 1997 = Lithophyllum crassum (Philippi) Heydrich nom. illeg. Lithophyllum stictaeforme (Areschoug) Hauck = Lithophyllum expansum f. stictaeformis (Areschoug) TSIRIKA & Foslie ATHANASIADIS, GIACCONE, 1968a HARITONIDIS, = Lithophyllum frondosum (Dufour) Furnari, Cormaci & 1987(8) 2005 Alongi = Pseudolithophyllum expansum f. stictaeformis Philippi Lithophyllum trochanter (Bory) Huvé ex Woelkerling (9) = Nullipora trochanter Bory NIKOLAIDIS & ATHANASIADIS, BORY, 1832 = Titanoderma trochanter (Bory) Benhissoune, HARITONIDIS, 1990 2016 Boudouresque, Perret-Boudouresque & Verlaque Lithothamnion corallioides (Crouan frat.) Crouan frat. = Lithothamnion solutum (Foslie) Foslie BALBINI & BRESSAN, SCHNETTER & LEMOINE, 1915 = Lithophyllum solutum (Foslie) Lemoine 1997 SCHNETTER, 1981 = Nullipora informis (Lamarck) Schweigger (10) Lithothamnion minervae Basso = Lithothamnion fruticulosum auctorum LEMOINE, 1915 GIACCONE, 1968c = ? Lithothamnion fruticulosum f. crassiusculum Foslie Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 17

TSIRIKA & CATRA & Lomentaria chylocladiella Funk ATHANASIADIS, 1987 HARITONIDIS, GIARDINA, 2009 2005 DIAPOULIS & GERLOFF & Lomentaria clavellosa (Lightfoot ex Turner) Gaillon ATHANASIADIS, 1987 HARITONIDIS, GEISSLER, 1974 1987b Lomentaria compressa (Kützing) Kylin GIACCONE, 1968a BITIS, 1988 Lomentaria ercegovićii Verlaque, Boudouresque, Meinesz, Giraud & Marcot-Coqueugniot LAZARIDOU, 1994 TSIAMIS, 2012 = Lomentaria tenera Ercegović nom. illeg. Melobesia membranacea (Esper) Lamouroux DIAPOULIS & = Epilithon membranaceum (Esper) Heydrich ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, = Lithothamnion membranaceum (Esper) Foslie 1987b = Hapalidium roseolum Kützing Meredithia microphylla (J. Agardh) J. Agardh SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Kallymenia microphylla J. Agardh SCHNETTER, 1981 Mesophyllum expansum (Philippi) Cabioch & Mendoza = Lithophyllum expansum Philippi ATHANASIADIS & = Crodelia expansa (Philippi) Kylin LAZARIDOU, 1994 HUVE, 1962 NETO, 2010 = Lithothamnion expansum (Philippi) Foslie = Pseudolithophyllum expansum (Philippi) Lemoine Mesophyllum macedonis Athanasiadis ATHANASIADIS, 1999 Mesophyllum philippii (Foslie) Adey SCHNETTER & LEMOINE, 1915 = Lithothamnion philippii Foslie SCHNETTER, 1981 Metapeyssonnelia feldmannii Boudouresque, Coppejans CATRA & & Marcot ALONGI, 2013 Nemalion lubricum Duby TSIAMIS et al., = Nemalion elminthoides auctorum (11) ATHANASIADIS, 1987 BITIS, 1988 2013b = Alcyonidium nemalion auctorum KOUSSOURIS et SCHNETTER & Nemastoma dichotomum J. Agardh var. dichotomum ATHANASIADIS, 1987 al., 1973 SCHNETTER, 1981 Nemastoma dichotomum var. biasolettianum (Kützing) RODRIGUEZ- TSIAMIS, 2012 Rodríguez-Prieto, Verlaque & Vergés PRIETO et al., 2004 Neogoniolithon brassica-florida (Harvey) Setchell & DIAPOULIS & Mason SCHNETTER & ATHANASIADIS, 1987 HARITONIDIS, = Neogoniolithon notarisii (Dufour) Hamel & Lemoine SCHNETTER, 1981 1987a = Spongites notarisii (Dufour) Athanasiadis Neurocaulon foliosum (Meneghini) Zanardini ex Kützing TSIRIKA & = Cryptonemia forbesii Harvey in W.J. Hooker LAZARIDOU, 1994 HARITONIDIS, = Constantinea reniformis sensu Postels & Ruprecht 2005 = Neurocaulon reniforme auctorum TSIRIKA & Parviphycus antipae (Celan) Santelices ATHANASIADIS, 1985 LAZARIDOU, 1994 HARITONIDIS, = Gelidiella antipae Celan 2005 Peyssonnelia armorica (Crouan frat.) Weber van Bosse CATRA & ATHANASIADIS, 1985 = Cruoriella armorica Crouan frat. GIARDINA, 2009 TSIRIKA & Peyssonnelia bornetii Boudouresque & Denizot ATHANASIADIS, 1985 LAZARIDOU, 1994 HARITONIDIS, 2005 TSIAMIS et al., Peyssonnelia coriacea J. Feldmann 2010a Peyssonnelia crispata Boudouresque & Denizot ATHANASIADIS, 1985 LAZARIDOU, 1994 CATRA & Peyssonnelia dubyi Crouan frat. ATHANASIADIS, 1985 GIARDINA, 2009 18 ACTA ADRIATICA, 60(1): 3 - 24, 2019

DIAPOULIS & CATRA & Peyssonnelia harveyana J. Agardh ATHANASIADIS, 1987 HARITONIDIS, GIARDINA, 2009 1987b Peyssonnelia heteromorpha (Zanardini) Athanasiadis SCHNETTER & = Peyssonnelia polymorpha (Zanardini) Schmitz in ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 Falkenberg Peyssonnelia immersa Maggs & Irvine ATHANASIADIS, 1987 Peyssonnelia orientalis (Weber-van Bosse) Cormaci & LAZARIDOU, 1994 Furnari SCHNETTER & Peyssonnelia rosa-marina Boudouresque & Denizot ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 Peyssonnelia rubra (Greville) J. Agardh SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Zonaria rubra Greville SCHNETTER, 1981 Peyssonnelia squamaria (S.G. Gmelin) Decaisne ex J. Agardh SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Padina squamaria (S.G. Gmelin) Lamouroux ex Bory SCHNETTER, 1981 = Padina squamaria var. sanguinea Bory nom. illeg. TSIAMIS et al., Peyssonnelia stoechas Boudouresque & Denizot TSIAMIS, 2012 2010 Phyllophora spiralis (Ginanni) Athanasiadis = Phyllophora nervosa (Lamouroux) Greville(12) SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Sphaerococcus nervosus (Lamouroux) C. Agardh SCHNETTER, 1981 = Phyllophora crispa auctorum Phymatolithon calcareum (Pallas) Adey & McKibbin ex Woelkerling & Irvine BALBINI & BRESSAN, SCHNETTER & = Lithothamnion calcareum (Pallas) Areschoug LEMOINE, 1915 1997 SCHNETTER, 1981 = Nullipora calcarea (Pallas) Lamarck = Phymatolithon polymorphum (Linnaeus) Foslie DIAPOULIS & Phymatolithon lenormandii (Areschoug) Adey HARITONIDIS, 1978 LEMOINE, 1915 HARITONIDIS, = Lithothamnion lenormandii (Areschoug) Foslie 1987b SCHNETTER & Platoma cyclocolpum (Montagne) Schmitz ATHANASIADIS, 1987 COPPEJANS, 1974 SCHNETTER, 1981 Plocamium cartilagineum (Linnaeus) Dixon (13) DIAPOULIS & = Plocamium coccineum var. uncinatum (C. Agardh) J. NIKOLAIDIS & SCHNETTER & HARITONIDIS, Agardh HARITONIDIS, 1990 SCHNETTER, 1981 1987a = Fucus coccineus Hudson Pneophyllum confervicola (Kützing) Chamberlain = Hapalidium phyllactidium Kützing nom. nov. illeg. CATRA & = Fosliella minutula (Foslie) Ganesan comb. inval. ATHANASIADIS, 1987 GRUNOW, 1861 GIARDINA, 2009 = Pneophyllum confervicola f. minutulum (Foslie) Chamberlain Pneophyllum coronatum (Rosanoff) Penrose ATHANASIADIS, 1987 = Pneophyllum rosanoffii Chamberlain Pneophyllum fragile Kützing = Melobesia lejolisii Rosanoff CHRYSSOVERGIS, SCHNETTER & LAZARIDOU, 1994 = Fosliella lejolisii (Rosanoff) Howe 1995 SCHNETTER, 1981 = Pneophyllum lejolisii (Rosanoff) Chamberlain Polystrata compacta (Foslie) Denizot ex Athanasiadis ATHANASIADIS, 1987 Porphyrostromium ciliare (Carmichael in WJ Hooker) TSIAMIS et al., Wynne 2010 DIAPOULIS & Predaea ollivieri J. Feldmann ATHANASIADIS, 1987 HARITONIDIS, TSIAMIS, 2012 1987a Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 19

Pterocladiella capillacea (S.G. Gmelin) Santelices & Hommersand = Gelidium capillaceum (S.G. Gmelin) Meneghini ATHANASIADIS, DIANNELIDIS, 1953 BITIS, 1988 = Gelidium corneum var. pinnatum (Hudson) Greville 1987 = Pterocladia capillacea (S.G. Gmelin) Bornet = Pterocladia pinnata (Hudson) Papenfuss Pterocladiella melanoidea (Schousboe ex Bornet) DIAPOULIS & Santelices & Hommersand ATHANASIADIS, 1987 LAZARIDOU, 1994 HARITONIDIS, = Gelidium melanoideum Schousboe ex Bornet 1987b = Pterocladia melanoidea (Schousboe ex Bornet) Dawson Pyropia elongata (Kylin) Neefus & Brodie BRODIE et al., 2007 = Porphyra rosengurttii Coll & Cox Pyropia koreana (Hwang & Lee) Hwang, Choi, Oh & Lee BRODIE et al., 2007 = Porphyra olivii Orfanidis, Neefus & Bray Rhizophyllis squamariae (Meneghini) Kützing(14) CATRA & = Contarinia squamariae (Meneghini) Denizot comb. ATHANASIADIS, 1987 GIARDINA, 2009 inval. Rhodochaete pulchella Thuret ex Bornet ATHANASAIDIS, 1987 = Rhodochaete parvula Thuret ex Bornet Rhodophyllis divaricata (Stackhouse) Papenfuss SCHNETTER & = Rhodophyllis appendiculata J. Agardh ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 = Rhodophyllis bifida (De Candolle) Kützing Rhodymenia ardissonei (Kuntze) J. Feldmann SCHNETTER & DIANNELIDIS, 1953 LAZARIDOU, 1994 = Rhodymenia corallicola sensu Ardissone SCHNETTER, 1981 DIAPOULIS & Sahlingia subintegra (Rosenvinge) Kornmann CHRYSSOVERGIS, LAZARIDOU, 1994 HARITONIDIS, = Erythrocladia subintegra Rosenvinge 1995 1987b

Sarconema scinaioides Børgesen TSIAMIS, 2012

Schmitzia neapolitana (Berthold) Silva ATHANASIADIS, 1987 TSIAMIS, 2012

Schottera nicaeensis (Lamouroux ex Duby) Guiry & TSEKOS & Hollenberg ATHANASIADIS, 1987 GIACCONE, 1968a HARITONIDIS, = Petroglossum nicaeense (Lamouroux ex Duby) Schotter 1977 TSIAMIS et al., Sebdenia dichotoma Berthold 2013c

Sebdenia rodrigueziana (J. Feldmann) Codomier ex ATHANASIADIS, 1987 Athanasiadis Sphaerococcus coronopifolius Stackhouse SCHNETTER & = Rhynchococcus coronopifolius (Stackhouse) Kützing ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 = Haematocelis fissurata Crouan frat. Spongites fruticulosus Kützing = Spongites ramulosus (Philippi) Kützing = Lithothamnion fruticulosum (Kützing) Foslie RÖSLER et al., TSEKOS et al., = Lithothamnion fruticulosum f. clavulatum Foslie ATHANASIADIS, 1987 2016 1982 = Lithothamnion fruticulosum f. kuetzingii Foslie = Lithothamnion fruticulosum f. ramulosum (Philippi) Foslie Stylonema alsidii (Zanardini) Drew = Goniotrichum alsidii (Zanardini) Howe SCHNETTER & ATHANASIADIS, 1987 LAZARIDOU, 1994 = Goniotrichum elegans (Chauvin) Zanardini SCHNETTER, 1981 = Goniotrichum elegans var. alsidii (Zanardini) Zanardini DIAPOULIS & Stylonema cornu-cervi Reinsch CHRYSSOVERGIS, LAZARIDOU, 1994 HARITONIDIS, = Goniotrichum cornu-cervi (Reinsch) Hauck 1995 1987b 20 ACTA ADRIATICA, 60(1): 3 - 24, 2019

Tenarea tortuosa (Esper) Lemoine ATHANASIADIS, HARITONIDIS & = Lithophyllum tortuosum (Esper) Foslie TSIAMIS, 2012 1995 TSEKOS, 1976 = Tenarea undulosa Bory nom. illeg. SCHNETTER & Thuretella schousboei (Thuret) Schmitz ATHANASIADIS, 1987 SCHNETTER, 1981 Tricleocarpa fragilis (Linnaeus) Huisman & Townsend = Galaxaura oblongata (Ellis & Solander) Lamouroux SCHNETTER & = Tricleocarpa oblongata (Ellis & Solander) Huisman & ATHANASIADIS, 1987 LAZARIDOU, 1994 SCHNETTER, 1981 Borowitzka = Galaxaura adriatica Zanardini Verlaquea lacerata (J. Feldmann) Le Gall & Vergés ATHANASIADIS, 1987 = Kallymenia lacerata J. Feldmann

SCHNETTER & Wurdemannia miniata (Sprengel) J. Feldmann & Hamel TSIAMIS, 2012 LAZARIDOU, 1994 SCHNETTER, 1981

1. Synonymy based on ROSAS-ALQUICIRA et al. (2010). 2. Reported by ATHANASIADIS (1987) as Bangia atropurpurea. 3. Synonym of Ellisolandia elongata (Ellis & Solander) Hind & Saunders based on GUIRY & GUIRY (2018). However, we follow ATHANASIADIS (2016), who quenstions the raise of Ellisolandia since no morphological or anatomical differences were indicated between Ellisolandia and Corallina, but their separation was based on the differentiation of several gene sequences. 4. Gelidium pectinatum Montagne contained in its protologue the citation of Fucus serra Gmelin and thus was considered as illegitimate. Excluding F. serra, a new combination was proposed, as G. bipectinatum (see FURNARI et al., 1999). However, based on ATHANADIADIS (2016: 405), F. serra corresponds to a crinoid animal and hense the name G. pectinatum Montagne should be re-instated, while G. bipectinatum should be considered as a superfluous substitute. 5. The generic attribution of Hydrolithon farinosum is uncertain in the absence of molecular data (RÖSLER et al., 2016). To be mentioned that ATHANASIADIS (2016: 272) treated the genus Hydrolithon as distinct from Fosliella. Here, we tentatively follow CORMACI et al. (2017) in considering Fosliella and Hydrolithon as congeneric. 6. Based on ROSAS-ALQUICIRA et al. (2011) the taxonomic status of this species is uncertain since the type specimen has not been studied under modern standards. Treated as taxonomicaly valid by CORMACI et al. (2017). 7. The dinstinction between the genera Lithophyllum and Titanoderma is controversial. Here we follow CORMACI et al. (2017) in considering them congeneric, with Lithophyllum having nomenclatural priority. ATHANASIADIS (2016) treats them as distinct. 8. Misidentified as Lithophyllum expansum (ATHANASIADIS, 2016: 325). 9. We follow CORMACI et al. (2017) giving nomenclature priority to Lithophyllum trochanter instead of Titanoderma trochanter. We also agree with them that records (including the ones from Greece) of Lithophyllum byssoides (Lamarck) Foslie [=Lithothamnion byssoides (Lamarck) Philippi, =Goniolithon byssoides (Lamarck) Foslie] which predate WOELKERLING (1998) paper should be referred to as Lithophyllum trochanter. 10. Placed in synonym with Lithothamnion corallioides by ATHANASIADIS (2016: 215). 11. Based on LE GALL & SAUNDERS (2010) Mediterranean records of Nemalion elminthoides (Velley) Batters correspond to N. lubricum, while the former is found in the Atlantic Ocean. 12. We follow ATHANASIADIS (2016: 536-537) in considering Phyllophora nervosa as a synonym of P. spriralis. 13. Based on the molecular work of SAUNDERS & LEHMKUHL (2005), world-wide reports of this NE Atlantic species possibly represent other species of the genus. To note that Greek plants are much smaller in size than the NE Atlantic plants (see also ATHANASIADIS, 1987: 56). 14. We follow ATHANASIADIS (2016: 685) in giving priority to Rhizophyllis squamariae instead of Contarinia squamariae based on the species zonate tetrasporangia. Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 21

Table 2. Rhodophyta algal taxa (excluding Ceramiales) pending confirmation in Greece, based on undocumented records. For each taxon a basic reference is provided together with applied synonyms in previous Greek records

Taxa Greece DIAPOULIS & HARITONIDIS, Acrochaetium molinieri Coppejans & Boudouresque 1987a Botryocladia chiajeana (Meneghini) Kylin CATRA & GIARDINA, 2009 Botryocladia microphysa (Hauck) Kylin TSIRIKA & HARITONIDIS, 2005 Catenella caespitosa (Withering) Irvine TSEKOS et al., 1982 = Catenella opuntia (Goodenough & Woodward) Greville Chondrymenia lobata (Meneghini) Zanardini PETKOFF, 1943 Chylocladia pelagosae Ercegović CATRA & GIARDINA, 2009 Colaconema leptonema (Rosenvinge) Alongi, Cormaci & G.Furnari CHRYSSOVERGIS, 1995 = Audouinella leptonema (Rosenvinge) Garbary Colaconema savianum (Meneghini) Nielsen = Acrochaetium savianum (Meneghini) Nägeli CHRYSSOVERGIS, 1995 = Audouinella saviana (Meneghini) Woelkerling = Callithamnion pygmaeum Kützing Cruoria cruoriiformis (Crouan frat.) Denizot CATRA & GIARDINA, 2009 = Cruoria purpurea Crouan frat. nom. illeg. Cryptonemia tunaeformis (Bertolini) Zanardini HUVE, 1962 Erythrocladia irregularis Rosenvinge (1) TSIRIKA & HARITONIDIS, 2005 SCHNETTER & SCHNETTER, Erythrotrichia bertholdii Batters 1981 Erythrotrichia investiens (Zanardini) Bornet SCHNETTER & SCHNETTER, = Bangia investiens Zanardini 1981

Feldmannophycus rayssiae (J. Feldmann & G. Feldmann) Augier & Boudouresque LAZARIDOU, 1994 (2)

Furcellaria lumbricalis (Hudson) Lamouroux HARITONIDIS & TSEKOS, 1976 = Furcellaria fastigiata (Turner) Lamouroux Gelidium corneum (Hudson) Lamouroux (3) HARITONIDIS & TSEKOS, 1976 = Sphaerococcus corneus (Hudson) Stackhouse Gelidium minusculum (Weber-van Bosse) R.E. Norris CATRA & GIARDINA, 2009 = Gelidium pusillum var. minusculum Weber-van Bosse SCHNETTER & SCHNETTER, Gelidium spathulatum (Kützing) Bornet 1981 Gelidium spinosum var. hystrix (J. Agardh) Furnari SCHNETTER & SCHNETTER, = Gelidium latifolium var. hystrix (J. Agardh) Hauck 1981 Gloiocladia repens (C. Agardh) Sánchez & Rodríguez-Prieto HUVE, 1962 = Fauchea repens (C. Agardh) Montagne & Bory Gracilaria armata (C. Agardh) Greville ORFANIDIS et al., 2001 = Sphaerococcus armatus C. Agardh Gracilaria gracilis (Stackhouse) Steentoft, Irvine & Farnham (4) CHRISTIA et al., 2011 Gracilaria longa Gargiulo, De Masi & Tripodi MALEA & HARITONIDIS, 2001 Gracilariopsis longissima (S.G. Gmelin) Steentoft, Irvine & Farnham (4) = Gigartina confervoides (Wiggers) Lamouroux = Gracilaria confervoides (Wiggers) Greville CHRISTIA et al., 2011 = Gracilaria verrucosa (Hudson) Papenfuss = Sphaerococcus confervoides (Linnaeus) C. Agardh nom. illeg. Gymnogongrus crenulatus (Turner) J. Agardh BORY, 1832 = Chondrus norvegicus sensu Lamouroux SCHNETTER & SCHNETTER, Helminthora divaricata (C. Agardh) J. Agardh 1981 22 ACTA ADRIATICA, 60(1): 3 - 24, 2019

Huismaniella ramellosa (Kützing) G. Boo & S. Boo ORFANIDIS et al., 2001 = Gelidiella ramellosa (Kützing) J. Feldmann & Hamel Hydrolithon cruciatum (Bressan) Chamberlain TSIRIKA & HARITONIDIS, 2005 Hydrolithon farinosum var. chalicodictyum (Taylor) Serio CATRA & GIARDINA, 2009 Hypnea furnariana Cormaci, Alongi & Dinaro CATRA & GIARDINA, 2009 Jania squamata (Linnaeus) Kim, Guiry & Choi = Corallina squamata Linnaeus DIANNELIDIS et al., 1977 = Haliptilon squamatum (Linnaeus) Johansen, Irvine & Webster Liagora distenta (Mertens ex Roth) Lamouroux CATRA & GIARDINA, 2009 Lithophyllum byssoides (Lamarck) Foslie = Lithophyllum lichenoides Philippi GIACCONE 1968a = Lithophyllum tortuosum f. cristatum (Meneghini) Foslie Lithophyllum corallinae (Crouan frat.) Heydrich DIANNELIDIS, 1953 = Melobesia corallinae Crouan frat. Lithothamnion crispatum Hauck DIANNELIDIS et al., 1977 = Lithophyllum crispatum (Hauck) Hauck SCHNETTER & SCHNETTER, Lithothamnion sonderi Hauck 1981 DIAPOULIS & HARITONIDIS, Lomentaria articulata (Hudson) Lyngbye var. articulata 1987a Lomentaria articulata var. linearis Zanardini CATRA & GIARDINA, 2009 = Lomentaria linearis (Zanardini) Zanardini ex Kützing Lomentaria clavaeformis Ercegović TSIRIKA & HARITONIDIS, 2005 Lomentaria uncinata Meneghini in Zanardini PETKOFF, 1943 = Chylocladia uncinata (Meneghini in Zanardini) Kützing Mesophyllum lichenoides (Ellis) Lemoine CHRYSSOVERGIS, 1995 = Lithothamnion lichenoides (Ellis) Foslie Millerella pannosa (J. Feldmann) Boo & Gall DIAPOULIS & HARITONIDIS, = Gelidiella pannosa (J. Feldmann) J. Feldmann & Hamel 1987a = Gelidiella tenuissima J. Feldmann & Hamel nom. illeg. SCHNETTER & SCHNETTER Naccaria wiggii (Turner) J. Agardh 1981 SCHNETTER & SCHNETTER, Peyssonnelia inamoena Pilger 1981 Phyllophora herediae (Clemente) J. Agardh BORY, 1832 = Sphaerococcus herediae (Clemente) C. Agardh Phyllophora sicula (Kützing) Guiry & Irvine SCHNETTER & SCHNETTER, = Phyllophora palmettoides J. Agardh 1981 Porphyra dioica Brodie & Irvine(5) HARITONIDIS, 1978 = Porphyra laciniata auctorum Porphyra linearis Greville TSIAMIS et al., 2013b Porphyra purpurea (Roth) C. Agardh (5) DIANNELIDIS et al., 1977 Porphyra umbilicalis Kützing(5) = Ulva umbilicalis Linnaeus DIANNELIDIS et al., 1977 = Porphyra umbilicalis (Linnaeus) J. Agardh nom. illeg. Pyropia leucosticta (Thuret) Neefus & Brodie (5) DIANNELIDIS et al., 1977 = Porphyra leucosticta Thuret Rhodochorton purpureum (Lightfoot) Rosenvinge DIAPOULIS & HARITONIDIS, = Rhodochorton rothii auctorum 1987b Rhodymenia delicatula Dangeard CATRA & GIARDINA, 2009 Rhodymenia ligulata Zanardini CATRA & GIARDINA, 2009 Rhodymenia pseudopalmata (Lamouroux) Silva GIACCONE, 1968a = Rhodymenia palmetta (Stackhouse) Greville Tsiamis and Panayotidis.: Seaweeds of the Greek coasts: Rhodophyta excluding Ceramiales... 23

Schizymenia dubyi (Chauvin ex Duby) J. Agardh TSIRIKA & HARITONIDIS, 2005 = Schizymenia minor (J. Agardh) J. Agardh NIKOLAIDIS & HARITONIDIS, Scinaia furcellata (Turner) J. Agardh 1990

1. The description and illustration by LAZARIDOU (1994: 44) should refer to Sahlingia subintegra. 2. LAZARIDOU (1994: 72) description cannot allow safe identification due to the lack of reproductive structures. 3. The occurrence of this species in the Mediterranean Sea is douptul, while there is a high chance of confusion with the common Mediterranean species Gelidium spinosum (FURNARI et al., 1999: 35). 4. Greek records might refer to Gracilaria gracilis and/or Gracilariopsis longissima, pending investigation of new specimens under modern taxonomic concept. 5. The occurrence of this species in the Mediterranean requires confirmation (BRODIE et al., 2007; CORMACI et al., 2017).

24 ACTA ADRIATICA, 60(1): 3 - 24, 2019

Morske alge grčke obale: Rhodophyta bez Ceramiales

Konstantinos TSIAMIS1,2* and Panayotis PANAYOTIDIS1

*Kontakt e-pošta: [email protected]

SAŽETAK

U radu autori prikazuju ažurirani popis crvenih morskih algi grčkih obala (Rhodophyta), isklju- čujući red Ceramiales, koji se temelji na literaturnim zapisima, te je kritički pregledan s današnjih taksonomskih i nomenklaturnih aspekata. Ukupan broj rodova, vrsta i infraspecifičnih taksona koji su trenutno prihvaćeni je 78, 138 i 4, respektivno. Prikazana je pojava svake svojte u sjevernom i južnom Egejskom moru i u Jonskom moru. Uoče- ne su praznine u znanju, tj. s 57 taksona koji čekaju potvrdu njihove prisutnosti. Štoviše, ukratko se raspravlja o 13 isključenih i 12 ispitivanih svojti. U cjelini, osigurana je čvrsta ažurirana osnovica pojavljivanja Rhodophyta (isključujući Ceramiales) u Grčkoj, što je presudno za buduće studije usmjerene na ciljane vrste.

Ključne riječi: Egejsko more, crvene alge, Rhodophyta, popis vrsta, Jonsko more