Mediterranean Marine Science

Vol. 11, 2010

Updated records and range expansion of alien marine macrophytes in Greece (2009)

TSIAMIS K. Hellenic Centre for Marine Research, Institute of Oceanography, P.O. Box 712, P.C. 19013, Anavyssos, Attiki MONTESANTO B. Department of Biology, Section of Ecology & Systematics, University of Athens, 15784 Panepistimiopolis, Athens PANAYOTIDIS P. Hellenic Centre for Marine Research, Institute of Oceanography, P.O. Box 712, P.C. 19013, Anavyssos, Attiki KATSAROS C. Athens University, Faculty of Biology, Department of Botany, Panepistimiopolis 15784, Athens VERLAQUE M. UMR 6540, DIMAR, COM, CNRS, Université de la Méditerranée F13288 Marseille https://doi.org/10.12681/mms.91

Copyright © 2010

To cite this article:

TSIAMIS, K., MONTESANTO, B., PANAYOTIDIS, P., KATSAROS, C., & VERLAQUE, M. (2010). Updated records and range expansion of alien marine macrophytes in Greece (2009). Mediterranean Marine Science, 11(1), 61-80. doi:https://doi.org/10.12681/mms.91

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Mediterranean Marine Science Research Article Indexed in WoS (Web of Science, ISI Thomson) The journal is available on line at http://www.medit-mar-sc.net

Updated records and range expansion of alien marine macrophytes in Greece (2009)

K. TSIAMIS1,2, B. MONTESANTO1, P. PANAYOTIDIS2, C. KATSAROS3 and M. VERLAQUE4

1 Athens University, Faculty of Biology, Department of Ecology & , Panepistimiopolis 15784, Athens, Hellas 2 Hellenic Centre for Marine Research, Institute of Oceanography, P.O.Box 712, Anavissos, 19013, Hellas 3 Athens University, Faculty of Biology, Department of Botany, Panepistimiopolis 15784, Athens, Hellas 4 UMR 6540, CNRS, Université de la Méditerranée, Parc scientifique et Technologique de Luminy, F13288 Marseille cedex 9, France

Corresponding author: [email protected]

Received: 2 September 2009; Accepted: 21 January 2010; Published on line: 26 February 2010

Abstract

In the present study the list of alien marine macrophytes already recorded on Greek coasts has been revised in the light of recent studies and new observations. In comparison to 2008, the total number consists of 32 taxa, and the classification as established, casual and debatable species has been modified, with a total of 14, 5 and 13 species respectively. An interesting increase in established species from 9 taxa in 2008 to 14 taxa in 2009 is noted. With 23 taxa listed, Rhodobionta is the best represented group, followed by Chlorobionta (4 taxa) and Chromobionta (4 taxa), while seagrasses (Streptobionta) are represented by only one species. Several new records, one new entry and two putative additions are considered here, while two other taxa previously assumed introduced are excluded from the list of aliens.

Keywords: Alien; Invasive species; Marine macrophytes; Greece; Mediterranean.

Introduction functions, and even result in negative so- cioeconomic effects in coastal areas The introduction of alien marine species (BOUDOURESQUE & VERLAQUE, is considered to be one of the greatest threats 2005; WALLENTINUS & NYBERG, 2007). to the marine environment at the dawn of Alien species have puzzled the scientif- the 21st century (SCHMITZ & SIMBER- ic community over recent decades, since LOFF, 1997; CARLTON, 2000). Alien there has been an uncontested increase in species, especially those showing invasive introductions worldwide, especially in behavior, can induce biodiversity losses, al- (CARLTON, 2002; STREFTARIS et al., terations to an ecosystems’ structure and 2005; GALIL, 2008, 2009). This increase is

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | attributed mainly to intensive human activ- are many gaps in our knowledge of alien ma- ities in the marine environment (shipping, rine macrophytes in Greece; this is due to the aquaculture, fisheries, tourism, etc.) since immensely long coastline and lack of research the 1960’s (BOUDOURESQUE & programs focusing on the subject. VERLAQUE, 2005). Therefore, today there The aim of the present study is to con- are several scientific programs focusing on tribute to the knowledge of alien marine macro- alien species, such as CIESM: atlas of exot- phytes in the Greek seas, since Greece rep- ic species, www.ciesm.org/atlas, and DAISIE: resents an important geographical link be- Delivering Alien Invasive Species Invento- tween the Levantine Sea, the Black Sea, the ries for Europe, http://www.europe-aliens.org/. Adriatic Sea and the Western Mediterranean Among alien species, alien marine macro- basin. To the last published list of alien ma- phytes represent one of the most important rine macrophytes from the Greek coasts groups (SCHAFFELKE et al., 2006), espe- (TSIAMIS et al., 2008), we add new records, cially in Europe and Australia (WALLE- pointing out revised species and debatable NTINUS, 2002; HEWITT et al., 2007). In cases, as well as giving detailed maps of their the , more than 110 taxa current range expansion in the Greek seas. of alien marine macrophytes have been hith- erto recorded, most of which originate from Methods the Indo-Pacific Ocean, and are introduced into the Mediterranean Sea through aqua- The list reported in the review paper by culture, shipping, and via the Suez Canal TSIAMIS et al. (2008, Table 1) was used as (BOUDOURESQUE & VERLAQUE, primary basis for the present work. Ac- 2002; CORMACI et al., 2004; HEWITT et cording to that list, 31 alien macrophytic al., 2007; MINEUR et al., 2007; VERLAQUE species had been recorded on Greek coasts et al., 2007, in press; ZENETOS et al., 2008). until 2007, of which 9 were classified as ‘es- Several alien marine macrophytes, such as tablished’, 8 as ‘casuals’ and 14 as ‘debat- Caulerpa taxifolia (M. Vahl) C. Agardh able’ cases. Since then, field samplings fo- (THIBAUT & MEINESZ, 2004) and Cauler- cusing on alien marine macrophytes have pa racemosa var. cylindracea (Sonder) Ver- been carried out along various Greek coasts, laque, Huisman et Boudouresque (KLEIN in both the Ionian and the Aegean Seas, & VERLAQUE, 2008), exhibit invasive be- aiming at an updated expansion range of havior and have a severe impact not only on alien macrophytes. Consequently, new the native algal and/or macrophytic com- entries and additional records of alien taxa munities but also on human activities have been included in this study, giving an (SCHAFFELKE & HEWITT, 2007; updated list with records up to August 2009. WILLIAMS & SMITH, 2007). Location, date, and collection data for the In Greece, there has been a sharp in- new records are given for each established crease in alien marine macrophytes over the alien species. In addition, the list has been last four decades (TSIAMIS et al., 2008), as critically reviewed and several records have well as the augmentation of other alien ma- been revised. For example, some previously rine organisms, such as fish and benthic in- assumed debatable taxa have now been vertebrates (ZENETOS et al., 2005; CORSINI- moved to the established category, while FOKA & ECONOMIDIS, 2007). However, some other records are now considered as there is still information missing and there excluded from the list of aliens. Moreover,

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Table 1 Alien marine macrophytes in Greece classified as established (E) and casual (C). Taxon: CHL= Chlorobionta, CHR= Chromobionta, RHO= Rhodobionta, STR= Streptobionta. Taxa in Bold correspond to new entries for the list and to changes in the status since TSIAMIS et al. (2008, Table 1). D=debatable.

Group Species Status 2008 2009 RHO Antithamnionella elegans (Berthold) J.H. Price et D.M. John DE RHO Asparagopsis armata Harvey a EE RHO Asparagopsis taxiformis (Delile) Trevisan de Saint-Léon DE RHO Bonnemaisonia hamifera Hariot b CC CHL Caulerpa racemosa var. cylindracea (Sonder) Verlaque, E E Huisman et Boudouresque CHL Caulerpa racemosa var. lamourouxii f. requienii (Montagne) CE Weber van Bosse RHO Ceramium strobiliforme G.W. Lawson et D.M. John C C RHO Chondria curvilineata F.S. Collins & Hervey c DC CHL Codium fragile (Suringar) Hariot subsp. fragile d EE CHR peregrina Sauvageau E E STR Halophila stipulacea (Forsska l) Ascherson E E RHO Hypnea cornuta (Kützing) J. Agardh CE RHO Hypnea spinella (C. Agardh) Kützing E E RHO Laurencia caduciramulosa Masuda et Kawaguchi DC RHO Lophocladia lallemandii (Montagne) F. Schmitz E E RHO Neosiphonia harveyi (J. Bailey) M.-S. Kim, H.-G. Choi, - C Guiry & G.W. Saunders RHO Sarconema scinaioides B rgesen CE CHR Stypopodium schimperi (Buchinger ex Kützing) E E Verlaque et Boudouresque RHO Womersleyella setacea (Hollenberg) R.E. Norris E E a Most records are based only on a ‘Falkenbergia’ life-history phase arbitrarily attributed to "F. rufolanosa", the sporophyte of A. armata. b Only the tetrasporophyte phase was found (SKOUFAS & TSIRIKA, 2003). c Misidentified as C. collinsiana M.A. Howe (ATHANASIADIS, 1987). Although ATHANASIADIS’s material was not re-examined, his description agrees with C. curvilineata and not with C. collinsiana. d Synonym of Codium fragile subsp. tomentosoides (van Goor) P.C. Silva. alien species considered previously not in- (TSIAMIS et al., 2008), alien macrophytes troduced as well as unpublished records by have been grouped into four broad cate- regional experts have been also included in gories according to their establishment suc- this study, thus giving a more precise view cess, namely ‘established’ (permanent, self- of alien marine macrophytes in Greece. maintaining populations), ‘casual’ (record- Following the same classification ed only once), ‘debatable’ (insufficient

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | information, highly probable records re- category differs with 14 established (E), 5 quiring confirmation, and cryptic introduc- casual (C) and 13 debatable (D) species tions) and ‘excluded’ (native species, misiden- (Tables 1 & 2). With 23 taxa listed, tifications). Updated range expansion maps Rhodobionta is the best represented group, are given for the established and casual alien followed by Chlorobionta (4 taxa) and Chro- taxa, taking into account both our own and mobionta (4 taxa), while seagrasses (Strep- already published records for each taxon tobionta) are represented by only one species. (only some references of interest are cited One new entry (C) and two putative addi- due to space limitations). On the other hand, tions (D) are considered here, while three debatable and excluded species are briefly other taxa previously assumed casuals discussed, with, however, special attention (TSIAMIS et al., 2008) are now considered paid to new entries and revised cases. as debatable. Moreover, there are also two taxa previously assumed introduced and Results are now excluded from the list of aliens. Distribution records and range expansion By August 2009, the total number of are given for each established (Fig. 1) alien marine macrophytes recorded on and casual (Fig. 2) alien marine macrophyte Greek coasts (32 taxa) had slightly increased in Greece. Location, date, and collecting when compared to TSIAMIS et al. (2008) data for the new records are given for each (31 taxa); moreover, the proportion of each established alien species (Table 3).

Table 2 Alien marine macrophytes in Greece classified as debatable. Taxon: CHL= Chlorobionta, CHR= Chromobionta, RHO= Rhodobionta. Taxa in Bold correspond to new entries for the list. TAXON SPECIES COMMENT RHO Acanthophora nayadiformis (Delile) Papenfuss native (or) cryptic introduction RHO Anotrichium okamurae Baldock ? native (or) cryptic introduction RHO Antithamnionella spirographidis (Schiffner) needs confirmation E.M. Wollaston RHO Ceramium bisporum D.L. Ballantine needs confirmation RHO Chondria coerulescens (J. Agardh) Falkenberg native (or) cryptic introduction RHO Chondria pygmaea Garbary & Vandermeulen highly probable presence, needs observation RHO Ganonema farinosum (J.V. Lamouroux) native (or) cryptic introduction K.C. Fan et Yung C. Wang RHO Hypnea valentiae (Turner) Montagne a needs confirmation CHR Padina boryana Thivy b needs confirmation RHO Polysiphonia atlantica Kapraun et J.N. Norris high risk of confusion with Mediterranean species RHO Polysiphonia fucoides (Hudson) Greville native (or) cryptic introduction CHR Pylaiella littoralis (Linnaeus) Kjellman c needs confirmation CHL Ulva fasciata Delile native (or) cryptic introduction a from Casual to Debatable compared to TSIAMIS et al., 2008 b from Casual to Debatable compared to TSIAMIS et al., 2008 c from Casual to Debatable compared to TSIAMIS et al., 2008

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Antithamnionella elegans Asparagopsis armata

Asparagopsis taxiformis Caulerpa racemosa var. cylindracea

Caulerpa racemosa var. lamourouxii f. requienii Codium fragile subspp. fragile Fig. 1: (continued)

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Colpomenia peregrina Halophila stipulacea

Hypnea cornuta Hypnea spinella

Lophocladia lallemandii Sarconema scinaioides Fig. 1: (continued)

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Stypopodium schimperi Womersleyella setacea

Fig. 1 (continued): Distribution records and range expansion of established alien marine macrophytes in Greece until 2009. Both old (published) and new records are given.

Fig. 2: Casual alien marine macroalgae in Greece until 2009.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Table 3 Data for the new records of established alien marine macrophytes in Greece up to August 2009.

Taxon Location Depth Substrate Date Collector Antithamnionella Neoi Epivates, Thermaikos Gulf 1m epiphyte July 2007 K. Tsiamis elegans Parga port 1m epiphyte October 2008 K. Tsiamis Selonda, Epidauros Gulf 1m epiphyte January 2009 K. Tsiamis *Antikyra, Korinthiakos Gulf 1m epiphyte October 2006 K. Tsiamis *Vourvourou, Sithonia 2m epiphyte August 2007 K. Tsiamis *Keri, Zakinthos Isl. 1m epiphyte September 2008 K. Tsiamis *Nies, Pagasitikos Gulf 2m epiphyte September 2008 K. Tsiamis Parga port 1m epiphyte / rocky October 2008 K. Tsiamis *Sxoinias, Attiki 1m epiphyte November 2008 K. Tsiamis Asparagopsis Selonda, Epidauros Gulf 1-3m epiphyte / rocky January 2009 K. Tsiamis taxiformis *Vourlias, Argolikos Gulf 1m epiphyte January 2009 K. Tsiamis * = only the Nauplio port, Argolikos Gulf 1m rocky February 2009 K. Tsiamis tetrasporophytic *Skala Kydonion, Lesvos Isl. 1m epiphyte March 2009 K. Tsiamis phace found Koroni port 1m rocky April 2009 G. Bardamaskos Pilos Port 1m rocky April 2009 G. Bardamaskos Aliveri, Evoikos Gulf 1-2m epiphyte / rocky June 2009 K. Tsiamis Porto Coufo, Sithonia 1-3m rocky June 2009 K. Tsiamis Gerolimenas, Messiniakos Gulf 1-2m rocky June 2009 K. Tsiamis Gyaros Isl. 1-15m rocky July 2009 M. Salomidi Near Pyrgi, Chios Isl. 2m rocky August 2009 S. Katsanevakis Kimolos Isl., port 1m rocky July 2006 M. Salomidi Karpathos Isl., port 1m rocky August 2006 A. Zenetos Agia Paraskevi, Skiathos Isl. 1m rocky August 2006 K. Tsiamis Korthi, Andros Isl. 1m epiphyte June 2008 K. Tsiamis Moudros bay, Limnos Isl. 1m rocky August 2008 S. Orfanidis Poros, Leukada Isl. 1m rocky August 2008 K. Tsiamis Nies, Pagasitikos Gulf 2m sandy / rocky September 2008 K. Tsiamis Parga port 1m rocky October 2008 K. Tsiamis Caulerpa Sxoinias, Attiki 1-15m rocky November 2008 K. Tsiamis racemosa var. Skala Kydonion, Lesvos Isl. 1-3m rocky March 2009 K. Tsiamis cylindracea Aliveri, Evoikos Gulf 1m rocky June 2009 K. Tsiamis Tristinika, Sithonia 1m rocky June 2009 K. Tsiamis Gyaros Isl. 1-15m rocky July 2009 M. Salomidi Kaldera, Santorini Isl. 1-3m Rocky / August 2009 K. Tsiamis epiphyte Rinia, Dilos Isl., 1-15m rocky August 2009 S. Katsanevakis Tragonisi, Mikonos Isl. 1-15m rocky August 2009 S. Katsanevakis Aspro & Stroggili, Syros Isl. 1-15m rocky August 2009 S. Katsanevakis Tinos Isl., south coasts 1-15m rocky August 2009 S. Katsanevakis (continued)

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Table 3 (continued)

Taxon Location Depth Substrate Date Collector Caulerpa Sapienza Isl., Messiniakos Gulf 50m rocky April 2009 K. Tsiamis racemosa var. Ladiko, Rhodes Isl. 2-3m rocky July 2009 K. Tsiamis lamourouxii Codium fragile Glyfada, Saronikos Gulf 0,5m rocky July 2009 K. Tsiamis subsp. fragile Apollakia, Rhodes Isl. 1m rocky June 1994 P. Panayotidis Colpomenia Andros Isl., port 1m rocky June 2008 K. Tsiamis peregrina Aliveri, Evoikos Gulf 1m epiphyte June 2009 K. Tsiamis Vouliagmeni, Saronikos Gulf 0,2m rocky August 2009 K. Tsiamis Halophila Aliveri, Evoikos Gulf 3m sandy June 2009 K. Tsiamis stipulacea Elinta, Chios Isl. 5m sandy August 2009 S. Katsanevakis Hypnea cornuta Agios Kosmas, Saronikos Gulf 0,2m epiphyte June 2007 K. Tsiamis Agios Kosmas, Saronikos Gulf 0,2m sandy August 2009 K. Tsiamis Hypnea spinella Agios Kosmas, Saronikos Gulf 0,5m epiphyte June 2006 K. Tsiamis Selonda, Epidauros Gulf 0,5m epiphyte January 2009 K. Tsiamis Tsigrado, Milos Isl. 1m rocky July 2007 K. Tsiamis Lophocladia Koukouras, Messiniakos Gulf 1m rocky July 2008 K. Tsiamis lallemandii Naxos Isl., port 1m rocky October 2008 V. Roussis Istron, Crete Isl. 1m rocky October 2008 V. Roussis Red beach, Santorini 1m rocky August 2009 K. Tsiamis Sarconema Agios Kosmas, Saronikos Gulf 0,2m rocky-sandy September 2008 K. Tsiamis scinaioides Korthi, Andros Isl. 2m rocky June 2008 K. Tsiamis Istron, Crete Isl. 1m rocky September 2008 V. Roussis Aigina Isl., Saronikos Gulf 1-26m rocky September 2008 M. Salomidi Vourlias, Argolikos Gulf 1m rocky January 2009 K. Tsiamis Stypopodium Glaronisi, Gyaros Isl. 1-15m rocky July 2009 M. Salomidi schimperi Legraina, Attiki 2-5m rocky July 2009 K. Tsiamis Thirasia, Santorini Isl. 0,5m rocky August 2009 K. Tsiamis Chios Isl., west coasts 0-10m rocky August 2009 S. Katsanevakis Rinia Isl, Cyclades 0-15m rocky August 2009 S. Katsanevakis Tragonisi, Mikonos Isl. 0-15m rocky August 2009 S. Katsanevakis Aspro & Stroggili, Syros Isl. 0-15m rocky August 2009 S. Katsanevakis Womersleyella Rhodes Port, Rhodes Isl. 1m epiphyte January 2006 K. Tsiamis setacea

New entries and putative additions Anotrichium furcellatum J. Agardh, a species described from Amalfi in Naples (AGARDH, Anotrichium okamurae Baldock ? 1842, as Griffithsia furcellata), seemed to be In 1939, the Mediterranean range of expanding so rapidly that FELDMANN-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | MAZOYER & MESLIN (1939) queried Neosiphonia harveyi (J. Bailey) M.-S. Kim, whether the species now present was real- H.-G. Choi, Guiry & G.W. Saunders ly the one described by J. Agardh. The sce- This species was first described as Polysi- nario of expansion of this species in the phonia harveyi from the NW Atlantic (USA, Mediterranean Sea resembles that of As- Connecticut, BAILEY, 1848). It is present- paragopsis taxiformis and Caulerpa race- ly considered to be widespread in the world, mosa. In the present case two taxa have and is regarded as introduced in the Atlantic probably been confused under the name Ocean and the Mediterranean Sea (MAGGS of A. furcellatum: the native taxon described & STEGENGA, 1999; CHOI et al., 2001; from Naples and a species (A. okamurae) VERLAQUE et al., in press). Based on an originally described from Japan, probably analysis of rbcL sequences, MCIVOR et al. introduced from the Pacific at an unknown (2001) concluded that Japan is its native re- time and exhibiting an invasive behaviour gion. In the Mediterranean Sea, Neosipho- (A. okamurae originally described from Ja- nia harveyi is more common in the western pan; OKAMURA 1934, as Monospora tenuis) basin than in the eastern basin. In Greece, (VERLAQUE et al., in press). In Greece, the species has been reported only once where A. furcellatum was frequently record- from the Gulf of Thessaloniki (C. MAGGS, ed in the Ionian and Aegean Seas, three pers. com. in VERLAQUE et al., in press) hypotheses have to be considered: either (Table 1, casual category). we have the native taxon, or the introduced species, or both taxa. Pending new infor- Revised species, now considered as debatable mation, the Greek records of A. furcellatum are provisionally attributed to A. okamurae There are three species, originally con- ? as a debatable case (Table 2). sidered as casuals (TSIAMIS et al., 2008), which have been revised and now consid- Chondria pygmaea Garbary & Vandermeulen ered as debatable: (not yet observed but presence highly prob- able) Hypnea valentiae (Turner) Montagne GARBARY & VANDERMEULEN Hypnea valentiae has been originally (1990) first described Chondria pygmaea described from the Red Sea (TURNER, from Eilat in the Red Sea as an epiphyte on 1809, as Fucus valentiae). Its introduction the seagrass Halophila stipulacea (Forsska l) into the Mediterranean Sea was unequiv- Ascherson. In the Mediterranean Sea, C. ocally identified for the first time in 1996 pygmaea was found among the epiflora of in France (Thau Lagoon), and succes- the introduced H. stipulacea each time it was sively in Italy (Venice and Mar Piccolo di studied (CORMACI et al., 1992; ALONGI Taranto) (VERLAQUE et al., in press). In et al., 1993; KASHTA & PIZZUTO, 1995; Greece, H. valentiae was first recorded in RINDI et al., 1999). Considering the wide 1894 from Rhodes Island (REINBOLD, distribution of the introduced seagrass in 1898), but it turned out to be a misidenti- Greece (Fig. 1), the occurrence of C. pyg- fication of H. cornuta (Kützing) J. Agardh. maea in this region is highly probable. Pend- More specifically, REINBOLD (1898) wrote: ing confirmation by observation, we provi- ‘Die exemplare zeigten die für diese art sionally range C. pygmaea in the debatable characteristischen spinulae stellulaeformes.’, category (Table 2). that is: ‘The specimens bear the charac-

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | teristic stellate spines of the species’. Such Island in the N. Aegean Sea (CANDARGY, stellate spines, which never occur in H. 1899) is probably based on a misidentifica- valentiae, are typical of H. cornuta. How- tion. We agree with their remark. ever, there are several other records of H. valentiae from Greece (BOGDANOS Neosiphonia sphaerocarpa (B rgesen) M.S. & DIAPOULIS, 1984; DIAPOULIS & Kim et I.K. Lee HARITONIDIS, 1987), but in the absence Originally described from St. Thomas of detailed specimen descriptions and the (Virgin Islands) in the Caribbean Sea complex taxonomy of the genus we cannot (B RGESEN, 1918, as Polysiphonia sphae- consider these data as sufficient to estab- rocarpa) and presently distributed world- lish the presence of H. valentiae in Greece. wide in warm seas, Neosiphonia sphaerocarpa Therefore, pending new observations, we was first recorded in the Mediterranean Sea consider H. valentiae as debatable in Greece. in 1970 from Tunisia (BEN ALAYA, 1970, misspelled P. sphaerocaspa), and successively Padina boryana Thivy in Corsica (VERLAQUE, 1989, as P. sphae- The records of this species from Amor- rocarpa), the Balearic Islands (BALLE- gos, Attica and Crete in Greece (see STEROS, 1992), and Milos Island in Greece NIZAMUDDIN, 1981, as Padina tenuis Bo- (LAZARIDOU, 1994). CORMACI et al. ry de Saint-Vincent) remain unconfirmed. (2004) considered N. sphaerocarpa to be in- Pending new observations, the occurrence troduced. However, the species is distrib- of P. boryana in Greece should be consid- uted worldwide and molecular data are need- ered as debatable. ed to locate its place of origin. Moreover, the identification of the species is not easy. Pylaiella littoralis (Linnaeus) Kjellman Consequently the absence of N. sphaero- There are only two records of this alga carpa in the Mediterranean prior to 1970 in Greece: one in the Aegean Sea could simply mean that it was overlooked (ANAGNOSTIDIS, 1968, with reservation), or mistaken for another Polysiphonia species. which probably corresponds to a misiden- Pending further investigations, we provi- tification of an Ectocarpus species (A. sionally consider this species as native in the ATHANASIADIS, pers. com.), and one in Mediterranean Sea. the Ionian Sea (SCHNETTER & SCHNETTER, 1981). Moreover, accord- Discussion ing to VERLAQUE et al. (in press) sever- al populations of this species could be na- Distribution and dynamics tive in the Mediterranean Sea. Therefore, pending new observations in Greece, P. lit- Since TSIAMIS et al. (2008), a large toralis should be considered as debatable. expansion has been observed for most es- tablished alien species, reaching 68 new dis- Excluded species tribution records on various Greek coasts (Table 3). The most rapid and broadest ex- Chorda filum (Linnaeus) Stackhouse pansion was observed for Asparagopsis tax- According to GERLOFF & GEISSLER iformis, a red alga with an heteromorphic (1974) and ATHANASIADIS (1987), the diplohaplontic life cycle, including a ga- single record of Chorda filum from Lesvos metophytic and a tetrasporophytic phase,

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | the latter known as ‘Falkenbergia’ phase. vealing a possible recent westward expan- A. taxiformis was found recently in Greece sion of the taxon during recent years. (TSIAMIS & PANAYOTIDIS, 2007), but Populations of Codium fragile subsp. since then it has exhibited rapid spread, fragile, originating from the N. Pacific Ocean especially in the South Aegean Sea. It is of- (CARLTON & SCANLON, 1985), are rel- ten found on shallow rocky bottoms, through atively restricted in Greece, found only at low to high abundance. As far as its ori- specific sites but usually in high abundance. gin is concerned, at least two cryptic taxa Most sites where it is observed are located of different origin (Atlantic and Indo-pa- near ports and polluted environments. It cific origin) coexist under the name of A. can be found on shallow rocky bottoms. The taxiformis in the Mediterranean Sea Indo-Pacific brown alga Colpomenia pere- (ANDREAKIS et al., 2004, 2007; N grina is known in Greece only through very CHUAL IN et al., 2004). On the basis few local records, although its distribution of their low survival rates in winter tem- could be wider since it can be easily con- peratures the Greek populations probably fused with the native species C. sinuosa belong to the invasive Indo-Pacific strain (Mertens ex Roth) Derbès & Solier. The (BARDAMASKOS et al., 2008). The sis- pantropical red alga Hypnea spinella also ter species, A. armata, may have a broader exhibits a restricted distribution with very distribution in Greece, but as the majority low abundance, found usually near ports as of Greek records refer to the Falkenbergia an epiphyte on other macrophytes. More- phase, which is nearly indistinguishable over, the Lessepsian immigrant Hypnea cor- from the Falkenbergia phase of A. taxiformis nuta is very rare in Greece, found only twice (N CHUAL IN et al., 2004), these records in the Aegean Sea: in 1894 in Rhodes Isl. could actually either correspond to (Reinbold 1898, as H. valentiae) and after A. taxiformis or A. armata. about one century (in 2007) in the Saronikos Caulerpa racemosa var. cylindracea is by Gulf by the authors. far the most common alien marine macro- The Lessepsian seagrass Halophila stip- phyte in Greece, presenting an extensive ulacea shows a wide distribution in Greece, spread along the majority of the Greek coasts. however, it is absent from the far N. Aegean This warm-temperate Caulerpa, originating Sea, probably because of the lower seawa- from SW Australia (VERLAQUE et al., ter temperature. It can be found on sandy 2003), can be found in various types of bot- bottoms at various depths, usually with the tom, from surface down to a depth of 40 m, native seagrass Cymodocea nodosa (Ucria) in both polluted and pristine sites. It often Ascherson. Occasionally, it can be abun- exhibits high abundances near ports and dant. The Lessepsian red alga Lophocladia fishing harbors. Conversely, the sister tax- lallemandii, has spread extensively during on C. racemosa var. lamourouxii f. requienii recent years in Greece, mainly in the S. has a very restricted distribution, found on- Aegean Sea. It is usually found as an epi- ly four times in Greece. Its distribution phyte during summer, especially at a depth requires further consideration since this tax- of several meters. Similarly, Stypopodium on can be easily confused with C. racemosa schimperi, another Lessepsian immigrant, var. cylindracea. Its discovery in the Ion- has exhibited a rapid expansion in the Aegean ian Sea represents the westernmost distri- Sea in recent years (see also bution record in the Mediterranean Sea, re- KATSANEVAKIS & TSIAMIS, in press),

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | revealing a gradual immigration from the the same location after about 30 years, in Levantive Sea towards the Western Mediter- 2008 (see also TSIAMIS et al., 2009); thus, ranean basin. It grows on rocky bottoms, revealing an established population in the from surface to several meters depth. area which still survives after 3 decades. Fi- Although it had been found in Greece nally, the Indo-Pacific red alga Womersleyella only once (SCHNETTER & SCHNETTER, setacea, found in various locations in both 1981) up till now, the red alga Antitham- the Aegean and the Ionian Seas, seems to nionella elegans was detected three times have been stable in Greece since 2007. The during the last year: once in the Ionian Sea taxon is usually found at a depth of sever- and twice in the Aegean Sea, all cases ster- al meters. ile and near shipping traffic, reflecting a In total, all established alien macrophytes possible spread of the species during recent (14 taxa) have been found in the S. Aegean years by vegetative propagation. In addi- Sea, most of them being present in the Sa- tion, the Lessepsian red alga Sarconema sci- ronikos Gulf. Similarly, most records of es- naioides, which was reported from Greece tablished alien species come from the South only in the Saronikos Gulf in the early 1980’s Aegean Sea, while the fewest come from the (DIAPOULIS et al., 1985), was re-found at North Aegean Sea (Fig. 3). These findings

Fig. 3: Records of established alien marine macrophytes in Greece until 2009.

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | reflect (i) the more extensive surveys on alien (OCCHIPINTI-AMBROGI & SAVINI, marine macrophytes in the S. Aegean Sea 2003; DUNSTAN & JOHNSON, 2007; (the Saronikos Gulf, Rhodes Island); (ii) the VALENTINE et al., 2007). role of Piraeus port, which is the largest port An invasive behavior was also observed in Greece, as a major site for introduction for Womersleyella setacea, but only once, in via shipping; and (iii) the role of the South the N. Aegean Sea in the middle 1990’s Aegean Sea as migration route for the Lessep- (ATHANASIADIS, 1997); however, simi- sian species coming from the Levantine coast lar cases could have been overlooked due (see also PANCUCCI-PAPADOPOULOU to its rather deep habitat. In addition, there et al., 2005). are also indications of invasive behavior Finally, we note an interesting increase of the seagrass Halophila stipulacea on sandy in established species, from 9 taxa in 2008 bottoms due to its high abundances; how- (TSIAMIS et al., 2008) to 14 taxa in 2009. ever, no displacement of native species due This is because of new records and findings, to H. stipulacea has hitherto been reported leading to the inclusion of originally casu- in the Mediterranean Sea. al (Caulerpa racemosa var. lamourouxii f. re- Finally, invasive behavior of the brown quienii, Hypnea cornuta, Sarconema sci- alga Stypopodium schimperi is known along naioides) and debatable species (Antitham- the Levantine coasts (BOUDOURESQUE nionella elegans, Asparagopsis taxiformis) in & VERLAQUE, 2002), although such be- the established category. Thus, established havior was not observed for any Greek site, alien marine macrophytes in Greece exhibit at least not until July 2009, when extreme- higher abundance than originally thought. ly high abundances and occasionally mo- nopolization of the substrate were detect- Impact on benthic assemblages ed in Rhodes Island. Until now, in Greece, there is not any The direct impact has not been docu- scientific proof of a negative impact of the mented yet for any alien marine macrophyte invasive marine macrophytes on human ac- in Greece. However, there are several cas- tivities, such as fisheries and tourism, al- es where losses of biodiversity and alter- though a few local cases have been report- ations of communities are evident, espe- ed where fishermen blamed invasive species, cially in the case of the invasive alien taxa. such as Codium fragile subsp. fragile, for the In such cases there is displacement of the reduction of fish abundance. native flora and monopolization of sub- strates by the alien invaders. Caulerpa race- Conclusion mosa var. cylindracea often exhibits such be- havior, especially in stressed ecosystems, as In conclusion, there are at least 14 alien well as in rather deep sandy bottoms. Codi- marine macrophytes that are established in um fragile subsp. fragile and Asparagopsis Greece. Most of them are still in progress taxiformis display invasive behavior exclu- along the Greek coasts, and further expan- sively in cases of ports and harbors, where sion, as well as new introductions, can be nitrophilous Chlorobionta, such as Ulva expected in the near future. Taking into ac- spp., also thrive. This supports the view that count that the arrival of invasive alien species invasive species are favored in ecosystems has brought about severe changes in the ma- already disturbed by human activities rine communities in other parts of the

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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 29/09/2021 09:09:02 | Mediterranean Sea (BOUDOURESQUE ANDREAKIS, N., PROCACCINI, G. & & VERLAQUE, 2002; SCHAFFELKE & KOOISTRA, W., 2004. Asparagopsis HEWITT, 2007), the spread of these aliens taxiformis and Asparagopsis armata (Bon- along the Greek coasts, especially from the nemaisoniales, Rhodophyta): genetic South Aegean Sea towards both the Ionian and morphological identification of and the North Aegean Sea, should be mon- Mediterranean populations. European itored. Moreover, more knowledge about Journal of Phycology, 39: 273-283. their biology, ecology and impact on the na- ANDREAKIS, N., PROCACCINI, G., tive communities would be helpful to elu- MAGGS, C. & KOOISTRA, W.H.C.F., cidate more aspects of alien introductions. 2007. Phylogeography of the invasive Asparagopsis (Bonnemaiso- Ackowledgements niales, Rhodophyta) reveals cryptic di- versity. Molecular Ecology, 16 (11): 2285- The authors are very grateful to the 2299. Greek State Scholarships Foundation for ATHANASIADIS, A., 1987. A survey of the the support of this study. They are also in- of the Aegean Sea with taxo- debted to Michèle Perret-Boudouresque nomic studies on species of the tribe An- for bibliographic assistance, as well as to tithamniae (Rhodophyta). PhD thesis, Athanasios Athanasiadis, Stelios Katsanevakis, University of Göterbog, Göterbog, Swe- Ioannis Issaris, Maria Salomidi, Giorgos den, 174 pp. Bardamaskos, Vasilis Roussis, Vasilis Ger- ATHANASIADIS, A., 1997. North Aegean akaris and Sotiris Orfanidis for their valu- Marine IV. Womersleyella setacea able contributions and comments. We would (Hollenberg) R.E. Norris (Rhodophy- also like to thank the anonymous reviewers ta, Ceramiales). Botanica Marina, 40 (1- and the editor for their valuable remarks. 6): 473-476. BAILEY, J.W., 1848. Continuation of the References list of localities of algae in the United States. American Journal of Sciences, AGARDH, J.G., 1842. Algae maris Mediter- Series 2, 6: 37-42. ranei et Adriatici, observationes in diag- BALLESTEROS, E., 1992. Contribucifi al nosin specierum et dispositionem generum. coneixement algolfigic de la Mediter- Paris: Fortin, Masson et Cie, X + 164 rània Espanyola. IX. Espècies interes- pp. sants de les Illes Balears. Folia Botani- ALONGI, G., CORMACI, M. & PIZZUTO, ca Miscelanea, 8: 77-102. F., 1993. La macroflora epifita delle BARDAMASKOS, G., TSIAMIS, K., foglie di Halophila stipulacea (Forssk.) PANAYOTIDIS, P. & MEGALO- Aschers. del Porto di Catania. Biologia FONOU, P., 2008. New records and range Marina Mediterranea, Suplemento al No- expansion of alien fishes and macroal- tiziario SIBM, 1: 287-288. gae in Greek waters (SE Ionian Sea). ANAGNOSTIDIS, K., 1968. Untersuchun- JMBA2 - Biodiversity Records, 2: 1-9. gen über die Salz- und Sü‚wasser- Thio- BEN ALAYA, H., 1970. Flore marine de bioconosen (Sulpuretum) Griechenlands. Tunisie. I. Liste préliminaire des algues PhD thesis, Aristotle University of Thes- du golfe de Tunis. Bulletin de l'Institut saloniki, Thessaloniki, Greece, 457 pp. National Scientifique et Technique

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