Alien macrophytes in the

Research Signpost 37/661 (2), Fort P.O., Trivandrum-695 023, Kerala, India

Recent Res. Devel. Environ. Biol., l(2004): 153-202 ISBN: 81-7736-217-8

I

Alien macrophytes in the l Mediterranean Sea: A review Cormacl Mario, Fumari Giovanni, Giaccone Giuseppe and Serio Donatdla Dipanimento di Botanica dell'universita di Catania, via Antonino Longo 19 95 125 Catania, Italy

Abstract A lis, of accepted alien macrophytes occurring in the Mediterranean Sea is presented. Of each species, a brief description, both world and Mediterranean distribwions, the chorology and /he possible vector of introduction are given. The list consists of 52 Rhodophyra, 15 Ochrophyta, 13 Chlorophyta and I ~Monocotyledonesfor a total of81 taxo at spec~jlcand infrospecrjc level and 3 t a w at generic level. The present list does not include several tara currently considered as introduced in literature. Such taxa are separately reported in hvo distinct lists: "tma not introduced': consisting of 12 Rhodophyta, I Ochrophyta, I Chlorophyta and " f m a exchrdenda" consisting of 6 Rhodophyta, 3 Ochrophyta and I Chlorophyta. Of each laxon included in the last hvo lists, reasonsjustrj$ing such a placement are given too. Correspo~~de~ice~eprinl requesl: Dr. Cormaci Mario, Dipartimsnto di Bot~nicadell'Universitl di Calania. via Antonino Longo 19, 95125 Catania, Italy. E-mail: [email protected]

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Cormaci Mario er al.

Introduction In recent years, the problem of the occurrence of "alien" species in the Mediterranean Sea came into the limelight mostly because of the invasive behaviour of some introduced green algae (Cawlerpa larifolia, C. racemosa) interacting with Posidonia oceanica (Linnaeus) Delile meadows [43, 44, 75, 100, 101, 127, 128, 1451 and the occurrence of several alien species in the Thau Lagoon, France mainly due to mollusc culture [189]. Other areas particularly affected by the occurrence of alien species were the Lagoon of Venice, ltaly [67, 68, 70, 941, the Gulf of Taranto, ltaly [46,48, 1431 and the Tuscany coast, ltaly [148, 1461. As a consequence, in several papers are reported lists of alien species to the Mediterranean Sea [35, 37, 38, 97, 98, 156, 188, 1891. Criteria for recognition of an alien species were proposed by several authors like [80, 156, 1881. They can summarized as follows: in order to be considered as introduced, a species should be new to the area studied; the area should be near to harbours, aquaculture farm, laboratory, aquarium; the new stations should be very localized and disjunct each from other; the kinetics of extension of the range should consist in a progressive extension in other neighbouring stations. In particular, a fast and massive extension is typical of "invasive7' species. However, in our opinion, several species reported in the above lists don't fit the above criteria so that they shouldn't be considered as alien. Since problems dealing with mechanisms and factors of successful introduction of alien species as well of their both negative and positive effects on local ecosystems and attempts to control of introduced species are deeply treated in literature [37, 4 1, 136, 156, 18 l], the present review is focused on species that in our opinion are actually alien to the Mediterranean Sea. Within each division, taxa are listed in alphabetical order. Of each alien species, the first record from the Mediterranean Sea is reported; as concerns records fiom other Mediterranean States only the first record per each State is reported. Of each taxon by us considered not introduced as well of those excludenda (separately listed) reasons for such a treatment are given.

Rhodophyta Acrothamnion preissii (Sonder) E.M. Wollaston (Ceramiales, Ceramiaceae) Type locality: Rottnest Island, Australia Thallus pale red in colour, filamentous, consisting of a creeping prostrate portion attached by digitate rhizoids, and erect branches to 3-4 cm high, bearing generally two opposite distichously-pinnate whorl-branchlets distichously arranged on each cell of the branch axis and 2 reduced whorl-branchlets side by

Medilerranean alien macrophytes

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side between the opposite branchlets. Each whorl branchlet ends with a terminal gland cell. World distribution - Indian Ocean: Australia, South Africa. Pacific Ocean: Japan, Philippines Chorology: Indo-Pacific Mediterranean distribution - Leghorn, Italy, first record from that sea [5 l]; France [28]; Balearic Islands [85]; Monaco [ l o l l . Vector of introduction: by fouling.

Agardhiella subulata (C. Agardh) Kraft et M.J. Wynne (Gigartinales, Solieriaceae) Type locality: probably eastern coast of Canada Thallus deep red in colour, cartilaginous, 5-20 cm long, terete, with apices acuminate, arising from discoid holdfast. Main axes alternate and distichously branched to 3-4 orders, with branches tapered at their bases. In transverse section the thallus shows an outermost cortical cells small and rounded; an inner cortex composed of four layers of cells decreasing in size towards the inside; a medulla consisting of primary filaments (10-12 pm in diameter) interspersed among interconnecting and rhizoidal filaments (1 5 2 pm in diameter). Tetrasporangia forming in branches strongly constricted at their bases. World distribution - Atlantic Ocean: from Canada to Brazil. Indian Ocean: India, Sri Lanka, Mauritius. Chorology: Circumboreal Mediterranean distribution - Italy, first record from that sea [45]; France [189]. Vector of introduction: by aquaculture (mollusc culture). NOTE - Even though the first published record of this species from the Mediterranean Sea is that by Cecere [45], the occurrence of the species in that Sea dates back to 1984, as reported in the paper by Verlaque [I89]. Only tetrasporic specimens are known in the Mediterranean Sea, where the species mostly reproduces vegetatively. According to Perrone & Cecere [l431 it should represent a Tethyan relict.

Aglaothamnionfeldmanniae Halos (Ceramiales, Ce ram iaceae) Type locality: Baie de Morlaix, France Thallus pink in colour, filamentous, consisting of both prostrate and erect axes, 2-5 cm high. Main axes distinct, with apex easily identifiable, bearing first order branches, arranged in a regularly alternate-distichous series. The basal cell of such branches bears an abaxial branchlet. First order branches bear second order branches with a similar branching pattern. World distribution - Atlantic Ocean: Great Britain, France, Spain

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Connaci Mario et a/,

Chorology: Boreo-Atlantic Mediterranean distribution - Leghorn, Italy, first record from that Sea [I 681; France [l 991. Vector of introduction: by fouling andfor ballast waters. NOTE - Only the tetrasporophyte is known from the Mediterranean Sea.

Ahnfeltiopsis jlabelliformis (Hawey) Masuda (Gigartinales, Phy Ilophoraceae) Type locality: Shimoda, Japan Numerous erect gametophytic axes, dark red in colour, arise from a basal crust (tetrasporophytic). Erect axes, fan shaped, are flattened, 5-6 cm high, branched 8-14 times dichotomously. In transverse section, erect axes show an outer cortex of small rectangular cells in radial rows and an inner compact medulla consisting of elliptic medullary cells increasing in size towards the centre. World distribution - Pacific Ocean: from Russia to Queensland (Australia); Hawaiian Islands; Peru. Chorology: Pacific Mediterranean distribution - Lagoon of Thau, France, first record from that Sea and Europe [l 861. Vector of introduction: By aquaculture (mollusc culture). NOTE - Unfortunately, Verlaque [l861 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

Antithamnion Ceramiaceae)

amphigeneum

A.

Millar

(Ceramiales,

Type locality: Coffs Harbour, Australia Thallus pale pink in colour, filamentous, ecorticate, consisting of creeping and erect axes to 8 mm long, with opposite and distichous whorl branchlets. Whorl branchlets, with the basal cell isodiametric, bear opposite distichous simple or branched branchlets with only abaxial branchlets near the distal region. Lateral indeterminate branches arise replacing a branchlet and with the suppression of the opposite branchlet. Gland cells are abundant on the adaxial side of both normal or special branchlets touching 2-3 ceIls. World distribution - Pacific Ocean: SE Australia Chorology: Pacific Mediterranean distribu/ion - Algeria, as Antithamnion algeriense M. Verlaque et Seridi, first record from that Sea [198]; Spain, as A. algeriense [ I S ] ; Morocco, as A. algeriense [106]; Italy, as A. algeriense [159]; Monaco [ l 921. Vector of introduction: by fouling.

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Mediterranean alien macrophytes

NOTE - Only male gametophytic and tetrasporic specimens were found in the Mediterranean Sea.

Antithamnion pectinatum (Montagne) Brauner ex Athanasiadis et Tittley (Ceramiales, Ceramiaceae) Type locality: Auckland, New Zealand Thallus red in colour, filamentous, 1-1.5 cm high, consisting of erect axes bearing whorl branches oppositely and distichously arranged. Whorl branches, 9-13 celled, with the basal cell isodiametric, show a distichous and opposite ramification with branchlets mostly abaxially arranged. Secondary axes replace a whorl branch in a pair without suppression of the opposite one. Gland cells on special 2-5 celled branchlets, touching 2-3 cells. World dis~ribu~ion - Atlantic Ocean: Connecticut, Azores. Pacific Ocean: New Zealand, Tasmania, Hawaiian Islands, SE Australia, Korea, S Japan. Chorology: Pacific Medilerranean dislribulion - Thau Lagoon, France, as A. nipponicum Yamada el Inagaki, first record from that Sea [197]; Italy [69]. Vec~orof inlroduclion: by aquaculture (mollusc culture).

Antithamnionella ternifolia (J.D. Hooker et Hawey) Lyle (Ceramiales, Ceramiaceae) Lectotype locality: Cape Horn. Thallus red in colour, filamentous, 2-4 cm high, consisting of prostrate axes becoming erect towards the apex, giving origin to secondary erect axes bearing primary whorled branches. Such branches, 8- 19 celled, are simple or poorly ramified, straight or slightly curved. When a secondary axis forms the suppression of the opposite lateral occurs. Generally, each whorled branch bears one elliptic gland cell on the second (rarely on the third or the fourth) cell of the branch. World distribu~ion - Atlantic Ocean: from Great Britain to Spain, Argentina. Pacific Ocean: Japan, Australia, Tasmania, New Zealand, Macquarie Islands, Ecuador, Chile. Indian Ocean: Chagos Islands. Chorology: Subcosmopolitan Medilerranean dis~ribu~ion - Toulon, France, as A. sarniensis Lyle [l 991. Vec~orofin~roduc~ion: by fouling.

Apoglossum gregarium (E.Y. (Ceramiales, Delesseriaceae)

Dawson)

Type locality: lsla S. Lorenzo del Norte, Mexico

M.J.

Wynne


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Thallus pink in colour, consisting of dense groups of small spathulate blades to 7 mm high and to 5 mm wide, arising from creeping terete axes. Blades show a consvicuous central ecorticate midriband indistinctlateral veins. The cells in secondary and higher cell rows have an irregular shape, often sigmoid. World disrriburion - Atlantic Ocean: Puerto Rico, Bermuda. Indian Ocean: South Africa. Pacific Ocean: California, Mexico, Hawaii. Chorology: Indo-Pacific Mediterranean distribution - Tuscan Archipelago, Italy, first record from that Sea [l 661; Spain [53]; Monaco [ l 921. Vector ofinfroducfion: unknown NOTE - According to Sartoni & Boddi [166], the occurrence in the Mediterranean Sea of both sexual and asexual plants of this species living in deep waters lets to doubt that it is a recently introduced species.

Asparagopsis armata Bonnemaisoniaceae)

Harvey

(Bonnemaisoniales,

Syntype localities: Garden Island and King George Sound, Western Australia; Tasmania Gametophytic thallus purplish-red in colour, consisting of a cylindrical axis to 1 mm wide and 20 cm long, arising from creeping stolons, irregularly branched, with branches densely and irregularly radially branched branchlets to 10 mm long. Scattered bare, but retrorsely spinous branches also occur. Tetrasporic thalli (known as "Falkenbergia-phase") consisting of densely tufted filaments 40-60 pm in diameter, with 3 pericentral cells around each axial cell. In the Mediterranean Sea, tetrasporophyte occurs throughout the year, while gametophytes only in late spring occur. World distribution - Atlantic Ocean: from Ireland to Canary Islands, Azores. Indian Ocean: Burma, South Africa, India, Australia. Pacific Ocean: Australia, Tasmania, New Zealand, Chile. Chorology: Cosmopolitan Mediterranean distribution - Algeria, first record from that Sea [169]; Italy [89]; Turkey [107]; Greece [171]; Tunisia [130]; Croatia [180]; Morocco [54]; Malta [62]; France and Spain [147]; Monaco [l 0 l]. Vecfor ofintroduction: by fouling.

Audouinella codicola (Bergesen) Garbary (Acrochaetiales, Acrochaetiaceae) Type locality: Canary Islands Thallus pale red in colour, filamentous, to 7.5 mm, consisting of creeping filaments bending round along the walls of the utricles of the host (Codium spp.) and erect filaments richly and irregularly branched. Filaments are cylindrical,


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not constricted at the cross walls. Cells with several lobate chloroplasts each provided with a pyrenoid. Monosporangia oval to obovate, sessile or pedicellate, generally in series adaxial along branches of 2"d or 3'd order. World dislribution - Atlantic Ocean: Canary Islands, Morocco, Azores, Madeira, Brazil. Indian Ocean: South Africa. Pacific Ocean: Japan, Korea. Chorology: Subcosmopolitan Medi~erraneandislribution - France, first ascertained record from that Sea [27]. Vector of in~roduc~ion: via the Straits of Gibraltar by ship transport. NOTE - According to Bidoux & Magne [27], it is possible that records by Boudouresque, Perret & Knoepffler [34] as Rhodotharnniella codii (P. et H. Crouan) Feldmann from Banyuls-sur-mer (France), Ballesteros & Romero [l41 as Audouinella codii (P. er H . Crouan) Garbary from Spain, Giaccone et al. [99] as Acrochae~iurncodii Bornet from Sicily (Italy), Meiiez & Mathieson [l301 as Acrochaetiurn codicolurn (sic) Bmgesen from Tunisia, should be referred to as this species.

Audouinella robusta (Bergesen) Garbary Type locality: St.Thomas, Virgin Islands Thallus pale red in colour, filamentous, to 2 mm high, originating from a basal cell constricted in the median part, generally merged into host's cells. Erect filaments end in a roundish cell and are unilaterally andlor irregularly branched. Monosporangia ovate, generally sessile or on a 1-2 celled pedicel. World distribulion - Atlantic Ocean: North Carolina, Caribbean Sea; Madeira. Indian Ocean: widely distributed in that Ocean. Pacific Ocean: China, Vietnam, Philippines, Hawaiian Islands. Chorology: Subcosmopolitan Medi~erraneandistribution - Egypt, as Acrochuetium sargassicola Bmgesen [3]. Veclor of introduc~ion:by fouling.

Audouinella subseriata (Bergesen) Garbary Type locality: Tuticorin, India Thallus pale red in colour, filamentous, to 1 mm high, caespitose, arising from a basal system of entangled filaments irregularly branched. Erect filaments are branched mainly in apical parts bearing branches irregularly or unilaterally arranged. Monosporangia ovate, sessile, borne in unilateral series in the apical parts of branches. World dis~ribu~ion - Indian Ocean: India, Pakistan, Mauritius, Seychelles. Pacific Ocean: Vietnam. Chorology: Indo-Pacific Mediterranean dislribution - Egypt, as Acrochaetiurn subseriaturn Bwgesen [3]. Vector of inlroduc~ion:by fouling.

Cormaci Mario et al.

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Bonnemaisonia hamifera Bonnemaisoniaceae)

Hariot

(Bonnemaisoniales,

Type locality: Yokosuka, Japan Gametophytic thallus dark red in colour, to 20 cm high, consisting of main axes 1-2 mm wide, bearing opposite branches, lacking in lower portions. Branchlets with conspicuous tendril-like, hooked ends. Tetrasporic thalli (known as "Trailiella-phase") consisting of uniseriate filaments, irregularly ramified. Cells of filaments often separate in their apical part one (rarely two) small gland cell. World distribution - Atlantic Ocean: from Faeroes to Canary Islands; Azores; South Africa (doubtful record). Pacific Ocean: Japan, Russia, California, Mexico. Chorology: Circumboreal Mediterranean distribution - Galite Island, Tunisia, first record from that Sea [144]; France [32]; Italy [50]; Spain [56]; Morocco [l06]. Vector of introduction: by fouling andlor by natural way through the Straits of Gibraltar. NOTE - Only "Trailiella-phase" is known from the Mediterranean Sea.

Botryocladia madagascariensis Feld mann-Mazoyer (Rhody meniales, Rhodymeniaceae) Type locality: Irakoka, Madagascar Thallus dark red in colour, to 10 cm high, with an axis pseudodichotomously branched. Ellipsoidal vesicles occur all around the axis; they have 4-6 layers of cells and show a continuous outer cortex consisting of small cells 5ym in diameter. Cells increase in size towards the inner side where they reach 70-80 pm in diameter. Gland cells 2-4 (rarely 5-6) are borne both on medullary cells and on special cells protruding into the vesicular cavity. World distribution - Indian Ocean: Madagascar, South Africa. Chorology: Indian Mediterranean distribution - Lampedusa Island, ItaIy, first record from that Sea [60]; Malta [62]; Turkey [183]. Vector of introduction: via the Suez Canal by natural way andlor by fouling. NOTE - Probably, the species has a wider distribution in the Mediterranean Sea since it could have been conhsed with B. botryoides [60].

Ceramium strobilifrme G.W. Lawson et D.M. John (Ceramiales, Ceram iaceae) Type locality: Vernon Bank, Ghana.


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Thallus red in colour, filamentous, simple, to 4 mm high, fixed by means of short basal rhizoids. Cortical cells form nodal cortical bands throughout the thallus except near the apex where the cortication is continuous. World distribution - Atlantic Ocean: Ghana. Chorology: Atlantic boreal Mediterranean distribution - Salina Island, Italy [60]. Vector of introduction: by fouling. NOTE - In the Mediterranean Sea both tetrasporophyte and female gametophyte epiphytic on Dicryota dichotoma (Hudson) J.V. Lamouroux and on Cystoseira spp. were found.

Chondria collinsiana M.A. Howe (Ceramiales, Rhodomelaceae) Type locality: Mayaguana, Bahamas Thallus pink in colour, terete, soft, 1-3 cm high, consisting of an erect terete axis arising from a crustose holdfast, irregularly branched with lateral simple branches with concave apices and attenuate base. In transverse section the axial cell surrounded by five periaxial cells is clearly observable. World distribrrtion - Atlantic Ocean: Caribbean Sea, Florida. Indian Ocean: Red Sea, Qatar, Kenya, Tanzania. Chorology: Pantropical Mediterranean distribution - Sithonia Peninsula, Greece, first record from that Sea [7]; Turkey [9]. Vector ofintroduction: probably by natural way through the Suez Canal NOTE - According to note 89 in Gomez Garreta et al. [l 041, the presence of this species in the Mediterranean Sea requires confirmation. According to Athanasiadis [7] it is uncertain whether it represents a recent introduction inthe Mediterranean Sea. On the basis of its distribution area it could be considered as a Tethyan relict.

Chondria curvilineata Collins et Hervey Type locality: Heron Bay, Bermuda Thallus red to yellowish in colour, soft in texture, 1-3 cm high, very slender (the main axes to 0.5 mm in diameter), bearing a few alternate branches of two orders generally contracted at base and apex. Growing point at bottom of an apical pit; swollen ends of the pericentral cells show through the cortication lines curved toward the apex of the branch. World distribution - Atlantic Ocean: North Carolina, Caribbean Sea, Salvage Islands. Indian Ocean: Maldives. Pacific Ocean: Philippines. Chorology: Pantropical Medirerranean dishibution - Egypt, first record 6om that Sea [36]; France [l 881. Vector of introduction: by fouling.

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Chondria polyrhiza Collins et Hervey Type locality: Shelly Bay, Bermuda. Thallus pale rose in colour consisting of creeping axes from which short erect axes arise. Erect axes ending with tapering apices, are cylindrical and bear simple branches irregularly arranged. The thallus is provided throughout with bunched rhizoids which are free and multicellular. World distribution - Atlantic Ocean: Bermuda, Caribbean Sea, Brazil. Indian Ocean: Haldabra Islands, RCunion. Pacific Ocean: Japan, Philippines, Hawaiian Islands. Chorology: Pantropical Mediterranean distribution - Sithonia, Greece, first record from that Sea [7]; Italy [47]. Vector of introduction:by fouling. NOTE - In the Mediterranean Sea only the tetrsporophyte was found.

Chondria pygmaea Garbary et Vandermeulen Type locality: Elat, Israel, Red Sea. Thallus red in colour, 10-15 cm high, fixed to the substratum by a large basal disc from which two or more erect axes arise. Axes are cylindrical, simple or bearing branches simple and strongly constricted at base. The cortex is unistratose with a typical brick-like arrangement of cortical cells. World distribution - Indian Ocean: Red Sea, Seychelles. Chorology: Indian A4editerranean distribution - Catania, Italy, first record from that Sea [60]; Albania [I 121; Malta [G2]. Vector of introduction: via the Suez Canal together with Halophila stipulacea on leaves of which it lives. NOTE - Even though the first published record of this species fiom the Mediterranean Sea is that by Connaci et al. [GO], the occurrence of the species in that Sea dates back to 1974, when some specimens were found on leaves of Halophila stipulacea collected in Albania, as published by Kashta & Pizzuto [l 121.

Chondrus giganteus Yendo f.fIabellatus Mikami (Gigartinales, Gigartinaceae) Type locality: Omazaki, Japan Thallus dark red in colour, cartilaginous, flabellate, flattened, 10-20 cm high, consisting of several erect axes arising from a discoid holdfast. Axes 4-8 times subdichotomously ramified with apices rounded to truncate. At margins of axes numerous proliferations occur. In longitudinal section the thallus consists of a cortex of 7-9 rows of oblong cells, of a sub-cortical layer of stellate cells and of a medulla of elongate cells arranged in longitudinal rows provided with


163

secondary pit connections. Tetrasporangia cruciate in chains in the medulla. In the Mediterranean Sea tetrasporophyte as well both male an female gametophytes were found. World distribtttion - Pacific Ocean: Japan Chorology: Pacific Mediterranean distribution - Thau Lagoon, France [l 961. Vector of introduction: By aquaculture (mollusc culture).

Chrysymenia wrightii (Harvey) Yamada (Rhodymeniales, Rhody meniaceae) Type locality: Hokkaido, Japan Thallus brownish in colour, terete, gelatinous, 20-60 cm high, attaching to substratum by means of a discoid holdfast, irregularly branched 3-6 times. Branches and branchlets gradually or abruptly constricted at base, tapering at apex. Axes in transverse section consisting of cortical layer of 2-3 rows of cells and a medullary layer of 3-5 rows of cells surrounding a central cavity with the innermost cells producing gland cells and hyphae-like filaments protruding into the central cavity. In the Mediterranean Sea both tetrasporophyte and female gametophyte were found. World distribution - Pacific Ocean: Russia, Japan, China. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France [25]. Vector of introduction: By aquaculture (mollusc culture).

Dasya sessilis Yamada (Ceramiales, Dasyaceae) Type locality: Ooshima, Japan Thallus dark red in colour, 10-25 cm high, consisting of erect axes polysiphonous, irregularly pinnately branched, presenting a pyramidal outline. The main branches, naked below, are covered with filiform branchlets above emerging from every articulation. Stichidia, ovate to conical, produced near the base of branchlets, sessile or with a short pedicel monosiphonous. World distribution - Pacific Ocean: Japan, Korea, Philippines. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France [l 901. Vector of introduction: By aquaculture (mollusc culture).

Dasysiphonia sp. (Ceramiales, Rhodomelaceae) To this genus, distributed in the Atlantic Ocean (N and S Carolina) and in the Pacific Ocean (Korea, Japan), were attributed with doubt some specimens collected at the Thau Lagoon (France) in May 1998 by Verlaque [l 891, probably introduced by mollusc culture. Later, Boudouresque & Verlaque [38] listed


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among Rhodophyta introduced into the Mediterranean Sea Heterosiphonia japonica Yendo with a note in which they put forward the hypothesis that it is probably a Dasysiphonia species. On this basis, no conclusions on the occurrence either of a species of Dasysiphonia or of Heterosiphonia japonica in the Tahu Lagoon can be reached.

Grateloupiafilicina (J.V. Lamouroux) C . Agard h v. luxurians A. et E.S. Gepp (Gigartinales, Halymeniaceae) Type locality: Farm Cove, Sydney, Australia Thallus dark red-brown in colour, 10-50 cm high, mucilaginous when young and cartilaginous when old, arising from a discoid holdfast, branched pinnately for several orders. Axes compressed provided with occasional proliferous branchlets on their surfaces. In transverse section axes consist of a cortex 5-8 layered with the innermost cells slightly stellate, a medulla lax (denser near the cortex) of widely spacely filaments. World distribution - Atlantic Ocean: Great Britain and France (introduced). Pacific Ocean: Australia and Tasmania. Indian Ocean: Australia. Chorology: Indo-Pacific Mediterranean distribution - Thau Lagoon, France [l 891. Vector of introduction: By aquaculture (mollusc culture). NOTE Unfortunately, Verlaque [l891 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

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Grateloupia lanceolata (Okamura) Kawaguchi Type locality: Kii, Japan Thallus purplish-red to yellowish-red in colour, lanceolate, gelatinous when young to leathery in old plants, arising from a small discoid holdfast generally through a short stipe. Blades, to 60 cm high and 20 cm broad, are lanceolate or irregularly lobed. In transverse section blades consist of a cortex of dichotomously branched anticlinal filaments of 10-20 cells, the inner cells rounded to stellate the outermost rounded and of a medulla of filaments periclinally aligned. World distribution - Pacific Ocean: Japan, Korea Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, first record from that Sea and Europe [I 891. Vector of introduction: By aquaculture (mollusc culture). NOTE Unfortunately, Verlaque [l891 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

-


165

Grateloupiapatens (Okamura) Kawaguchi et H.W. Wang Type locality: Shinoshima, Japan Thallus red brownish in colour, 10-15 cm high, flattened, with an irregularly pinnate branching pattern with branches distichous inserted at nearly a right angle narrow at the base and spathulate at the apex. Tetrasporangia in sori linear forming in ultimate branches. World distribution - Pacific Ocean: Japan, Korea. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, as Prionitis patens Okamura, first record from that Sea and Europe [189]. Vector of introduction: By aquaculture (mollusc culture). NOTE Unfortunately, Verlaque [l891 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

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Grateloupia turuturu Yamada Type locality: Hokkaido, Japan Thallus red in colour, narrowly to broadly lanceolate with several blades, 20-60 cm high, arising from a holdfast. In transverse section blades are characterized by a thin cortex of roundish cells and an abrupt transition between cortex and medulla. Medullary filaments show an anticlinal arrangement. World distribution - Atlantic Ocean: both Eastern and Western coasts (introduced); Pacific Ocean: coast of NE Asia. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, as G. doryphora (Montagne) M. Howe, first record fiom that Sea [162]; Lagoon of Venice, Italy, as G. doryphora [l 821. Vector of introduction: by aquaculture (mollusc culture). NOTE - This species was often quoted with the misapplied name G. doryphora. But, the Mediterranean record of G. doryphora fiom the Straits of Messina [72]corresponds to a distinct taxon (Gargiulo, personal communication).

Griffithsia corallinoides (Linnaeus) Trevisan (Ceramiales, Ceramiaceae) Type locality: in Mari Europaeo Thallus dark red to orange red in colour, consisting of numerous much branched erect filaments, attached by tangled rhizoidal filaments. Erect axes with apical cells of 50-100 pm in diameter increasing towards the base to 500 pm in diameter. Cells initially inflated at both ends, becoming markedly pyriform constricted near cross-walls. Branching pseudodichotomous every 1-3 cells. Spermatangial heads form dense bands around the nodes of sub apical

166

Cormaci Mar10er al.

cells. Cystocarps surrounded by an involucre of incurved 1-2 celled filaments. Tetrasporangia are borne in a ring around nodes, enclosed by 2-celled incurved involucral filaments. World distribution - Atlantic Ocean: from Noway to Canary Islands. Indian Ocean: India, Sri Lanka, Mauritius, Australia. Pacific Ocean: Japan. Chorology: Indo-Atlantic Mediterranean distribution - Palermo, Italy, first record from that Sea [102]; France, with doubt [22]; Turkey [9]; Tunisia [24]; Algeria, with doubt [142]. Vector of introduction : via the Straits of Gibraltar probably by natural way.

Herposiphonia parca Setchell (Ceramiales, Rhodomelaceae) Type locality: Arue Reef, Tahiti Thallus dark red in colour, filamentous, polysiphonous (8-10 pericentral cells), consisting of a prostrate axis with regular sequence of three erect determinate branches alternating with erect indeterminate branch. Single segments not bearing indeterminate branches often occurring between two determinate branches. The prostrate axis is attached by rhizoids with radiating, branched, often multicellular tips; determinate branches to 20 segments strongly curved toward apex when young, nearly straight at maturity. World distribution - Atlantic Ocean: Belize. Indian Ocean: Tanzania, Kenya, India, Maldives, Seychelles, RCunion. Pacific Ocean: from Japan to Philippines, Hawaiian Islands, Fiji, Tahiti. Chorology: Indo-Pacific Mediterranean distribution - Thau Lagoon, France, first record from that Sea and Europe [l 891. Vector of introduction: By aquaculture (mollusc culture). NOTE - Unfortunately, Verlaque [l891 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

Hypnea cornuta Hypneaceae)

(Kiitzing)

J.

Agardh

(Gigartinales,

Type locality: Guinea Thallus red in colour, terete, 6-20 cm high, repeatedly alternately branched with branches, tapering towards the apices, provided with numerous simple spinulose branchlets 1-2 mm long as well as with sessile or briefly pedicellate stellate branchlets of 3-5 points. In transverse section the axial cell is well distinguishable from bigger medullary cells. World distribution - Atlantic Ocean: Gulf of Guinea, eastern tropical and subtropical coasts of Americas. Indian Ocean: widely distributed in that Ocean. Pacific Ocean: Japan, Queensland (Australia); Fiji.


167

Chorology: Pantropical Mediterranean distribution - Harbour of Alexandria, Egypt, first record from that Sea 121; Israel [132]; Italy (Cecere & Petrocelli, personal communication). Vector of introduction: by fouling andor by natural way via the Suez Canal NOTE - Only sterile specimens were found in the Mediterranean Sea

Hypnea spicifea (Suh r) Harvey Type locality: Algoa Bay, South Africa (Indian Ocean) Thallus dark red to green in colour, terete, filiform, 1-2 mm in diameter, arising from branched interwoven, thinner prostrate axes attached to the substratum by means of several haptera. Branching irregular, sometimes proliferous, with branches tapering towards the apex; fertile specimens with numerous simple or branched spine-like branchlets bearing reproductive structures. In transverse section the axial cell is not distinguishable. World distribution - Atlantic Ocean: South Africa, Namibia. Indian Ocean: widely distributed in that Ocean. Chorology: Indo-Atlantic Mediterranean distribution - Cleopatra, Egypt, as Hypnea hurveyi Kiitzing [S]. Vector of introduction: unknown. NOTE The Aleem's record is based on specimens collected in 1956.

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Hypnea spinella (C. Agardh) Kii tzing Type locality: West lndies Thalli dark red in colour, terete, filiform, forming densely entangled mats 23 cm wide. Branching in all directions; axes and branches provided with spine like tapering branchlets. In transverse section the axial cells is much smaller than the medullary cells in older axes and branches; such a distinction is less evident in younger branches. World distribution - Atlantic Ocean: eastern tropical and subtropical coasts of Americas; from Morocco to Gabon, Canary Islands. Indian Ocean: widely distributed in that Ocean. Pacific Ocean: from Japan to Queensland (Australia); Hawaiian Islands and Polynesia. Chorology: Pantropical Mediterranean distribution - Balearic Islands, Spain, first record from that Sea, as Hypnea cervicornis J. Agardh [105]; Greece [7]; Italy, as H. cervicornis [59]; Turkey [203]; Tunisia [77]; Algeria [37]; Lebanon, as H. cervicornis [I 141. Vector of introduction: by fouling.

Hypnea valentiae (Turner) Montagne Type locality: Red Sea.

168


Thallus from red to yellow-greenish in colour, terete, filiform, consisting of erect axes in tufts or clumps arising from a basal disc, 10-30 cm high. Axes irregularly branched; branches with apices occasionally curved, more often straight, acute, with branchlets simple, forked or spine-like to 3 mm long. In transverse section the axial cell is small and easily distinguishable. World distribution - Atlantic Ocean: Caribbean Sea, Bermuda; Ghana, Canary Islands. Indian Ocean: widely distributed in that Ocean. Pacific Ocean: widely distributed in that Ocean. Chorology: Pantropical Mediterranean distribution - Rhodes Island, Greece, first record £tom that Sea [154]; Syria, as Hypnea hamulosa J.V. Lamouroux [124]; Sidi Bishre, Egypt [5]; Thau Lagoon, France [198]. Vector of introduction: via the Suez Canal by natural way. In the Thau Lagoon probably by aquaculture (mollusc culture).

Laurencia okamurae Yamada (Ceramiales, Rhodomelaceae) Type locality: Japan Thallus purplish green in colour, terete, 15-20 cm high, consisting of erect axes percurrent, arising from a discoid holdfast becoming stoloniferous. Branches patent, oppositely, alternately or subverticillately arranged. In fresh material one corp en cerise is detectable in both epidermal cells and trichoblasts. Four pericentral cells per axial segment. In transverse section cortical cells without palisade-like arrangement; lenticular thickenings abundant in the walk of medullary cells. Tetrasporangia parallel arranged. World distribution - Pacific Ocean: Japan, Korea, China, Taiwan, Philippines. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, first record from that Sea and Europe [l 891. Vector of introduction: By aquaculture (mollusc culture). NOTE - According to Verlaque [189], this species was introduced into the Lagoon probably in the 1970s with the massive importations of Japanese oysters; unfortunately, he didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

Lithophyllum yessoense Foslie (Corallinales, Corallinaceae) Type locality: Yezo, Japan Thallus pink to whitish in colour, entirely calcified, strongly adherent to the substratum, forming crusts smooth with irregular margins. The structure is dimerous consisting of primigeneous and postigeneous filaments. Secondary pit connections between cells of contiguous filaments present. Hypothallus


169

unistratose with subquadrate cells. Old asexual conceptacles buried in the thallus. World distribution - Pacific Ocean: Japan Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, first record from that Sea and Europe [189]. Vector of introduction: by aquaculture (mollusc culture). NOTE - Unfortunately, Verlaque [l891 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

Lomentaria hakodatensis Lomen tariaceae)

Yendo

(Rhodymeniales,

Type locality: Hakodate, Japan Thalli brownish-purple in colour, 5-10 cm high, attached to the substratum by a stoloniferous discoid holdfast. Axes and branches hollow, terete or slightly flattened. Axes branched to five orders, with branches oppositely arranged (rarely alternate or three whorled). Fusions among branches of the same thallus are frequent. Septa 3-5 layered occur throughout the thallus. In young branches and branchlets constrictions in correspondence of septa are very obvious. World distribution - Atlantic Ocean: France (introduced); Spain (introduced). Pacific Ocean: Japan, China, Korea, Russia, Philippines, Hawaiian Islands, Queensland (Australia, probably introduced); from British Columbia to Gulf of California. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, first record from that Sea [189]; Italy (Curiel, unpublished data). Vector of introduction: by aquaculture (mollusc culture).

Lophocladia lallemandii (Montagne) F . Schmitz (Ceramiales, Rhodomelaceae) Type locality: Red Sea. Thallus red dark in colour, filamentous, to 12 cm high, attached to the substratum by rhizoids. Axis dichotomously ramified, lightly corticated below, with a monopodial structure with each axial cell developing 4 pericentral cells, clothed with spirally arranged trichoblasts. Tetrasporangial stichidia slightly spirally twisted. World distribution - Indian Ocean: widely distributed in that Ocean. Pacific Ocean: Japan, Philippines, Queensland (Australia). Chorology: Indo-Pacific

170

Connaci Mario er al.

Mediterranean distribution - Saronikos Gulf, Greece and Tripoli, Libya, first records from that Sea [144]; Algeria and Tunisia [84]; Italy [93]; Syria [124]; Spain [179]; Egypt [S]; Turkey [203]; Lebanon [l 141. Vector of introduction:via the Suez Canal by natural way. NOTE - In the Mediterranean Sea, in summer and autumn, due to its high capability of vegetative reproduction [63], this species often covers bottom biocoenoses, causing severe damages.

Neosiphonia harveyi (J.W. Bailey) M.S. Kim, H.G. Choi, Guiry et G.W. Saunders (Ceramiales, Rhodomelaceae) Type locality: Stonington, Connecticut Thallus dark red in colour, filamentous, polysiphonous with 4 pericentral cells, 4-10 cm high, with one to few main axes arising from a discoidal holdfast. Erect axes alternately to irregularly branched to several orders becoming bushy and pyramidal in outline, the ultimate branches tapering to acute apices. Branches replacing trichoblasts at irregular intervals. Tetrasporangia formed in last three orders of branching, in long spiral series, spermatangial axes borne at the first dichotomy of fertile trichoblasts, cystocarps globose to ovoid. Rhizoids cut off from periaxial cells by cross-walls. World distribution - Atlantic Ocean: from Georgia to Connecticut; from Scandinavia to Canary Islands. Pacific Ocean: from the Bering Sea to Korea (as Polysiphonia japonica Harvey). Chorology: Atlanto-Pacific ~editerranekndistribution - Thau Lagoon, France, as Polysiphonia mottei Lauret, first record from that Sea [l 161; Italy, as P. mottei [49]; Syria, as P. mottei [124]; Egypt, as P. mottei [26]; Algeria, as P. mottei [172]; Spain, as P. mottei [l 21. Vector of introduction:By aquaculture (mollusc culture). NOTE - In the Mediterranean Sea the species was also quoted as both Polysiphonia insidiosa (J. Agardh) P.L. et H.M. Crouan and P. harveyi J.W. Bailey [92].

Neosiphonia sphaerocarpa (Bergesen) M . S. Kim et I. K. Lee Type locality: St. Thomas, Virgin Islands Thallus dark brown in colour, filamentous, polysiphonous (with four pericentral cells); ecorticate, to 2 cm high, consisting of a prostrate axis, attached to the substratum by unicellular rhizoids separated from pericentral cells by a cross wall, from which erect axes arise. Erect axes are subdichotomously branched. Tetrasporangia spirally arranged. Spermatangial branches arise from a branch of the trichoblasts. Cystocarps globular.


171

World distribution - Atlantic Ocean: Caribbean Sea, Georgia, North and South Carolina; Canary Islands and Salvage Islands. Indian Ocean: Maldives and Seychelles. Pacific Ocean: Philippines; Hawaiian Islands, Samoan Archipelago, Fiji, Australia. Chorology: Pantropical Mediterranean distribution - Tunisia, first record from that Sea, as Polysiphonia sphaerocarpa Brargesen [18]; Spain, as P. sphaerocarpa [13]; France, as P. sphaerocarpa [187]; Greece, as P. sphaerocarpa [l 171. Vector of introduction: by fouling.

Pleonosporium caribaeum (Bsrgesen) R.E. Norris (Ceramiales, Ceramiaceae) Type locality: Annaberg, St. John (Virgin Islands) Thallus pink in colour, filamentous, erect, 1-2 cm high, arising from a fibrous base. Axes with nearly isodiametric axial cells, branched several times with branches issuing multilaterally to all sides; branchlets on successive axial cells arranged in spiral, rarely distichously, incurved around branches apices. Tetrasporangia sessile in adaxial series arranged. Cystocarps globular loosely involucrate. World distribution - Atlantic Ocean: Caribbean Sea, Canary Islands, South Africa. Indian Ocean: India and Africa (from Kenya to South Africa). Pacific Ocean: Japan, Hawaiian Islands, Samoa, Micronesia. Chorology: Pantropical Mediterranean distribution - Spain, as Mesothamnion caribaeum Brargesen, first record from that Sea [178]; France [188]. Vector of introduction: via the Straits of Gibraltar by natural way or by fouling.

Plocamium secundatum (Kutzing) Kutzing (Gigartinales, Plocamiaceae) Type locality: Hermite Island, Chile Thallus erect, to 4 cm high, irregularly ramified with flattened main axes. Four-six branches in alternating series up to median parts of the axes, ten-fifteen (rarely thirty to forty) unilaterally arranged in the apical parts. The lowennost branches are generally simple, while the others are alternately ramified near the base, unilaterally near the apex. Tetrasporangial stichidia are simple or branched 2-3 times. Tetrasporangia are arranged in two rows. World distribution - Antarctic and the sub Antarctic islands. Chorology: Subantarctic Mediterranean distribution - Catania, Italy [6 l]. Vector of introduction: unknown, probably by fouling.


172

Polysiphonia morrowii Harvey (Ceramiales, Rhodomelaceae) Type locality: Hakodate, Japan Thallus red to brown in colour, to 50 cm high, polysiphonous, with 4 pericentral cells, consisting of creeping axes fixed to the substratum by unicellular rhizoids in open connection with the pericentral cell and by erect axes spirally branched. Axes branched to four orders. Basal first order branches are simple and strongly curved. Terminal branchlets are tapering. Cystocarps are urceolate, spermatangial branches replacing trichoblasts, tetrasporangia in straight series (up to 9). World distribution - Atlantic Ocean: The Netherlands (introduced). Pacific Ocean: from the Commander Islands to China; Australia and New Zealand (introduced) Chorology: Pacific Mediterranean distribution - Lagoon of Venice, Italy, first record from that Sea [68]; Thau Lagoon, France 11891. Vector of introduction:by aquaculture (mollusc culture). NOTE - Even though both the papers by Curiel et al. [68] and by Verlaque [l891 were published in the same year, we consider as the first record from the Mediterranean Sea that by Curiel et al. [68] since they refer to a collection made in 1999, see also Curiel et al. 1661.

Porphyra yezoensis Ueda (Bangiales, Bangiaceae) Type locality: Miyagi, Hokkaido, Japan Gametangial thallus red in colour, monostromatic, consisting of lanceolate blades with entire margins somewhat undulate and cordate or cuneate at the base. Cells of blades, elliptical to rectangular in surface view, rectangular in transverse section. A single stellate chloroplast per cell. Spermatangia and zygotosporangia in sori, intermixed. World distribution - Atlantic Ocean: Helgoland (probably introduced). Pacific Ocean: Russia, Japan, Korea, China; Alaska. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France [l 881. Vector of introduction: By aquaculture (mollusc culture). NOTE - According to Verlaque [l881 this species was introduced into the Thau Lagoon in the 1982, but it was not found in the recent study on that Lagoon [l 891.

Pterosiphonia tanakae Rhodomelaceae)

Uwai

Type locality: Motoujina, Japan

et

Masuda

(Ceramiales,


173

Thallus bright red in colour, filamentous, polysiphonous (with 7-9 pericentral cells); consisting of a prostrate axis from which 1-3 erect axes arise, 2-7 cm high, terete below, becoming compressed upward. Erect axes bear numerous first order branches from every two segments alternate-distichously. First order branches form progressively shorter and more slender branchlets of up to five orders from every two segments, producing a fan shaped appearance. Branches of any order are coalesced with their parental axes or branches for 1.52.5 segments. Ultimate branchlets terete throughout, acute at the apices, recurved with age. World distribution - Pacific Ocean: Japan Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, as Pterosiphonia sp., first record from that Sea and Europe [l 891. Vector of introduction: By aquaculture (mollusc culture). NOTE - Boudouresque & Verlaque [38] attributed to P. tanakae Verlaque's [l891 Pterosiphonia sp. giving no reasons to their taxonomic conclusion.

Rhodophysema georgei Batters (Palmariales, Rhodophysemataceae) Type locality: Scilly Islands, Great Britain. Thallus epiphytic, dark purplish red in colour, crustose or cushion like and then irregularly globose, somewhat mucilaginous. The thallus consists of large medullary cells up to 200 pm in diameter, surrounded by cortical filaments with cells about 10 pm in diameter. Tetrasporangia in sori terminal with spores cruciately divided, interspersed with colourless curved 3-4 celled sterile filaments. Worlddistribution - Atlantic Ocean: British Isles, from Norway to northern Spain, Newfoundland, Long Island. Pacific Ocean: British Columbia to Oregon, Russia, Japan. Chorology: Circumboreal Mediterranean distribution - Thau Lagoon, France [l 861. Vector of introduction: By aquaculture (mollusc culture). NOTE - According to Verlaque [l 861 this species was introduced into the Thau Lagoon in the 1978, but it was not found in a recent study on this Lagoon [189].

Rhodymenia erythraea Zanardini (Rhodymeniales, Rhodymeniaceae) Type locality: Hodeida, Yemen Thallus dark red in colour, coriaceous in texture, consisting of blades 6-7 cm high arising from a small basal disc. Blades cuneate, with smooth margins

174


towards the base and provided with proliferations in apical parts, are branched subdichotomously. In transverse section the thallus shows a medulla of 2-3 layers of large oblong cells and a cortical layer of small cells. Tetrasporangia cruciately divided, form in the cortical layer and are scattered over the whole thallus. Cystocarps protrudent, few in the basal parts, densely aggregated in upper parts also in marginal proliferations. World distribution - Indian Ocean: Yemen, Somalia Chorology: Indian Mediterranean distribution - Port Said Harbour, Egypt [2]. Vector of introdz~ction:via the Suez Canal by natural way or by fouling.

Sarconemafiliforme (Sonder) Kylin (Gigartinales, Solieriaceae) Type locality: Western Australia Thallus dark red in colour, cartilaginous, 10-20 cm high, consisting of a discoid holdfast from which several erect axes 0.5-1 mm in diameter, becoming basally stoloniferous, arise. Axes terete regularly subdichotomously branched. The thallus with a multiaxial structure consisting of a medullary core of compact longitudinal filaments and a broad cortex 10-12 layered with inner cells 20-60 pm in diameter, decreasing to 2-3 pm in diameter in the outer layer. Cystocarps sunken but distinctly bulging the branch. Tetrasporangia scattered in outer cortex, zonately divided. World distribution - Indian Ocean: widely distributed in that Ocean. Pacific Ocean: from China to Southern Australia and French Polynesia. Chorology: Indo-Pacific Mediterranean distribution - Port Said, Egypt, as Sarconema furcellatum Zanardini, first record from that Sea [2]; Israel 11521; Syria [124]. Vector of introduction: via the Suez Canal by natural way.

Sarconema scinaioides Bsrgesen Type locality: Karachi, Pakistan Thallus dark red in colour, fleshy, 10-20 cm high, consisting of a discoid holdfast from which several erect axes to 2 mm in diameter arise. Axes terete, pseudodichotomously branched, irregularly and regularly in tetrasporophytes and female gametophytes, respectively. The thallus shows a multiaxial structure consisting of a medullary core of longitudinal loose filaments and a broad cortex 8-10 layered with inner cells 50-60 pm in diameter, decreasing to 12 pm in diameter in the outer layer. Cystocarps and tetrasporangia as in the genus. World distribution - Indian Ocean: Pakistan, India, Yemen, Oman, Kenya, Tanzania. Chorology: Indian Mediterranean distribution - Gulf of Saronikos, Greece [74].

Mediterranean alien rnacrophytes

175

Vector of introduction: via the Suez Canal by natural way.

Solieria dura Solieriaceae)

(Zanardini)

F.

Schmitz

(Gigartinales,

Syntype localities: Hodeida and Mokha, Yemen Thallus pink to dark red in colour, cartilaginous in texture, to 15 cm high, terete, consisting of erect axes arising from a callous discoid holdfast. Axes irregularly branched to three orders with ultimate branchlets crowded near the apex. In transverse section they show an outermost cortex of small oblongate cells and an inner four-layered cortex of large rounded cells the innermost reaching 120 pm of diameter; the medulla consists of medullary filaments and 1-2 celled interconnecting filaments. Tetrasporangia, zonately divided, are scattered in the outer cortex. World distribution - Indian Ocean: India, Yemen, Somalia, Tanzania. Pacific Ocean: Philippines. Chorology: Indo-Pacific Mediterranean distribution - Port Said, Egypt, first record from that Sea [3]; Israel [152]; Syria [124]. Vector of introduction: via the Suez Canal by natural way.

Solieriafiliformis (Kiitzing) P. W . Gabrielson Type locality: Antigua Island, Antilles Thalli pink to dark red in colour, 5-10 cm high, terete, consisting of axes with tapering apices arising from a stoloniferous base. Axes sub-dichotomousIy ramified to three orders; in transverse section they show an outermost cortex of small rectangular cells and an inner three-layered cortex of spherical cells; primary medullary filaments conspicuous and straight; in the inner part of the section characteristic spur-shaped interconnecting cells and 1-2 celled interconnecting filaments occur. Worlddistribution - Atlantic Ocean: from North Carolina to Brazil; Ghana, Gabon, Southern British coasts and Canary Islands. Chorology: Boreo-Atlantic Mediterranean distribution - Taranto, Italy [143]. Vector of introduction: unknown. NOTE: According to Perrone & Cecere [l431 it should represent a Tethyan relict.

Symphyocladia marchantioides (Ceramiales, Rhodomelaceae)

(Harvey)

Falkenberg

Syntype localities: Cape Kidnapper & Hawke's Bay, New Zealand

176


Thallus brownish red in colour, foliaceous, with prostrate blades attached to the substratum by pluricellular rhizoids. Blades consist of coalescent polysiphonous axes. Tetrasporangia are produced in laterally hsed stichidia and occur singly in each fertile segment. World distribution - Atlantic Ocean: Azores, South Africa. Indian Ocean: South Africa. Pacific Ocean: from Russia to Japan; Hawaiian Islands; West coasts of Australia; New Zealand. Chorology: Indo-Pacific Mediterranean distribution - Leghorn, Italy, as Symphyocladia sp. [165]. Vector of introduction: by fouling. NOTE - The attribution of Symphyocladia sp. from Leghorn (Mediterranean Sea) [l651 to S. marchantioides was made by Rindi et al. [l601 on the basis of fertile specimens recently collected at that locality.

Womersleyella setacea (Hollenberg) R.E. Norris (Ceramiales, Rhodomelaceae) Type locality: Koko Head, O'ahu, Hawaiian Islands Thalli brown red in colour, forming densely entangled tufts consisting of prostrate axes polysiphonous, with 4 pericentral cells, ecorticate, fiom which erect axes to 1 cm high mostly unbranched arise. Rhizoids are cut off fiom the pericentral cell; at first unicellular, at maturity end in a multicellular pad. Tetrasporangia in slightly spiral series, up to 30 in a series, in terminal parts of erect axes. World distribution - Atlantic Ocean: Caribbean Sea, Canary Islands. Indian Ocean: Indonesia; Maldive. Pacific Ocean: Hawaiian Islands, Samoan Archipelago, Fiji Islands, Caroline Islands, Philippines, Columbia. Chorology: Pantropical Mediterranean distribution Var, France, first record from that Sea, as Polysiphonia setacea Hollenberg [I87]; Alboran Sea, as P. setacea [l]; Spain [148]; Malta [62]; Greece [g]; Italy, as Polysiphonia sp. [19]. Vector of introduction: by fouling. NOTE - Even though the first published record of this species from the Mediterranean Sea is that by Verlaque [187], the occurrence of the species in that Sea dates back to 1986, when some specimens were found at Rosignano Solvay (near Leghorn, Italy) as published by Benedetti Cecchi & Cinelli, as Polysiphonia sp. [I9]. In the Mediterranean Sea the species is invasive.

Ochrophyta Acrothrrjcgracilis Kylin (Chordariales, Chordariaceae) Type locality: Sweden Sporophytic macrothallus brown-yellowish in colour, to 20 cm high, with up to 3 orders of branching, with branches irregularly alternately arranged,

177


consisting of a single central axial filament with trichothallic growth, a medulla composed of 2-3 layers of hyaline cells (65-100 pm long and 50-90 pm wide in transverse section) and a cortical layer composed of roundish (8-50 pm in diameter in transverse section) cortical cells and uniseriate assimilatory filaments 7-12 celled of two types: nearly isodiametric throughout the length and with the terminal cells swollen to one side. Only unilocular sporangia, obovoid, sessile on the basal cells of assimilatory filaments or on the cortical cells with a stalk cell are present. Plurilocular sporangia in gametophytic microthalli. World distribution Atlantic Ocean: North European coast (from Scandinavia to Ireland). Pacific Ocean: Japan and Alaska. Chorology: Circumboreal Mediterranean distribution Thau Lagoon, France [ l 891. Vector of introduction: by aquaculture (mollusc culture) NOTE - Unfortunately, Verlaque [l891 didn't give any descriptions of Thau Lagoon's specimens supporting his taxonomic conclusion.

-

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Chorda jilum Chordaceae)

(Linnaeus)

Stackhouse

(Larninariales,

Type locality : Atlantic Ocean Thallus cartilaginous, to 3-4 m long, pale brown in colour, simple, threadlike, gradually tapering toward both ends, arising from a small discoid holdfast composed of bundles of rhizoids. Axis tube-like, composed of an outer meristoderm (single-celled layer); a cortex (6-7 celled layer) and a medulla consisting of hyaline hyphae with a reticular arrangement delimiting the internal hollow interrupted at various intervals by diaphragms of lobed cells. World distribution - Atlantic Ocean: widely distributed along the northern part of both eastern and western coast of the Ocean, Canary Islands. Pacific Ocean: China, Japan and Alaska Chorology: Circumboreal Mediterranean distribution - Thau Lagoon, France [161]. Vector of introduction: by aquaculture (mollusc culture) NOTE - According to Hamel [l 101 in the Thuret Herbarium, held in PC (MusCum National &Histoire Naturelle, Paris, France) there is a specimen of this species collected at Nice (Mediterranean coast of France) by Sake around the half of the 19Ih century. According to Athanasiadis [7] the record of this species from Lesbos (Greece) by Candargy [40] is based on a misidentification.

Colpomenia peregrina Scytosiphonaceae)

Sauvageau

(Scytosiphonales,

Syntype localities: different stations in the European Atlantic coast


178

Thallus olive brown in colour, spherical or saccate in shape, membranous, hollow, with 1-2 layers of small cortical cells and 2-4 layers of large medullary cells delimiting the central cavity. Plurilocular sporangia grouped in irregular and extensive sori not covered by a cuticle; hairs frequent, unicellular and club-shaped. World distribution - Atlantic Ocean: From Scandinavia to Canary Islands, Azores, NW African coast. Pacific Ocean: from Alaska to California, Japan, Australia and New Zealand. Chorology: Subcosmopolitan Mediterranean distribution - Banyuls, France, first record from that Sea [129]; Algeria [33]; Italy (J. Feldmann in Boudouresque Ch.F. & Boudouresque E.) [33]; Spain [l 851; Morocco [2 l]. Vector of introduction:via the Straits of Gibraltar by natural way.

Fucus spiralis Linnaeus (Fucales, Fucaceae) Type locality: Atlantic Ocean Thallus dark brown to olive green, coriaceous, 15-30 cm high, arising from a conical discoid holdfast. Axis dichotomously branched with branches on a plane or sometimes a little twisted, 2 cm wide, with margin slightly undulate. Receptacles terminal, simple or forked, obovate and swollen. World distribution - Atlantic Ocean: from Scandinavia to Western Sahara in the east coast, from Newfoundland to New York in the west coast. Pacific Ocean: from Alaska to Washington. Chorology: Circumboreal Mediterranean distribution - Ceuta, Morocco, first record from that sea [30]; Spain [SS]; France [164]. Vector ofintroduction: via the Straits of Gibraltar by natural way.

Halothrix lumbricalis Elachistaceae)

(Kiitzing)

Rein ke

(Chorda riales,

Type locality: Baltic Sea Thallus yellow-greenish in colour, forming small brush-like tufts on host, consisting of a basal system of compacted rhizoids from which numerous erect filaments arise. Erect filaments simple or rarely branched towards base, with an intercalary growth zone near base. Plurilocular sporangia, solitary or in 1-6 cell chains, multiseriate, formed by subdivision of cells of erect filaments. World distribution - Atlantic Ocean: from Scandinavia to France, Maine, New Jersey. Pacific Ocean: Japan, China, British Columbia. Chorology: Circumboreal Mediterranean distribution - Acicastello, Italy, first record from that sea [58]; Turkey [10]; France [l 891. Vector ofintroduction: by fouling or ballast waters.


179

Laminaria japonica A reschoug (Laminariales, Laminariaceae) Type locality: Japan Thallus olive brown in colour, 2-6 m high, consisting of a coriaceous undivided lanceolate blade, with entire margins, tapering to the base in a short stipe attached to the substratum by hapteres. Worlddistribution - Pacific Ocean: Japan, China Chorology: Pacific Mediterranean distribution - Thau Lagoon, France [139]. Vector of introduction: by aquaculture (mollusc culture) NOTE - This species was not found in a recent study on the Thau Lagoon [l 891.

Leathesia dijjformis (Linnaeus) Areschoug (Chordariales, Corynophlaeaceae) Type locality: Sweden Thallus yellow brown in colour, fleshy and mucilaginous in texture, globose, hollow, to 5-8 cm in diameter. It consists of a basal layer from which erect filaments arise. Erect filaments di-trichotomously branched comprising large and almost stellate medullary cells becoming smaller towards the periphery. The outer layer consists of small spherical cells bearing paraphyses and hairs. Unilocular sporangia globose; plurilocular sporangia, simple and uniseriate. World distribution - Atlantic Ocean: from Faeroes to Canary Islands, Azores; from Mauritania to South Africa; from Labrador to North Carolina. Indian Ocean: South Africa. Pacific Ocean: from Alaska to California; Philippines, Australia, New Zealand; sub Antarctic Islands. Chorology: Cosmopolitan Mediterranean distribution - Thau Lagoon, France, first record from that Sea [l 861; Italy [I 71; Morocco [21]. Vector of introduction: by aquaculture (mollusc culture) NOTE - According to Ben Maiz et al. [24], the record from Tunisia reported by Ribera et al. [l571 should be referred to Leathesia mucosa Feldmann; while, in the Thau Lagoon it was not found in a recent study [189].

Padina boergesenii Dictyotaceae)

Allender

et

Kraft

(Dictyotales,

Type locality: Danish West lndies Thallus light brown in colour, with fafl-shaped fronds moderately calcified, 5-20 cm wide in a semicircle, arising from a stupose holdfast. Fronds mostly tristromatic; sporangial son non-indusiate, arranged in concentric rows and separated from each other by a sterile zone.


180

World distribution - Atlantic Ocean: Danish West Indies. Indian Ocean: India, Pakistan, Indonesia. Pacific Ocean: Micronesia, Fiji. Chorology: Pantropical Mediterranean distribution - Mikhmoret, Israel, as P. gymnospora sensu Vickers, first record from that sea [149]; Italy, as P. gymnospora sensu Vickers [177]; Libya as P. gymnospora sensu Vickers [133]. Vector of introduction: via the Suez Canal by natural way.

Padina boryana Thivy Type locality: Tonga Thallus yellow-brown in colour, with fan-shaped fronds moderately calcified, 8-10 cm wide in a semicircle, arising from a stupose holdfast. Fronds distromatic throughout except at the base where are tristromatic; sporangial sori non-indusiate, arranged in concentric rows bounded by hair lines. World distribution Atlantic Ocean: S l o TomC. Indian Ocean: widely distributed in the ocean. Pacific Ocean: from Japan to Philippines; Samoan Archipelago, Tonga, Fiji, Tahiti. Chorology: Indo-Pacific Mediterranean distribution - Abu Qir, Egypt [S]. Vecror of introduction: via the Suez Canal by natural way.

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Sargassum muticum (Yendo) Fensholt (Fucales, Sargassaceae) Type Locality: Hokkaido, Japan Thallus dark brown in colour to 3 m tall, consisting of a fibrous holdfast from which a terete main axis arises. Main branches repeatedly ,and alternately branched; all branches bear leaves linear-lanceolate in the basal portion of the thallus, to 10 cm long and with margins toothed and narrow, to 4 mm long with margins generally entire in the upper portions. Pneumatocysts borne in clusters or single in leaf axils. Receptacles on special branches arising from leaf axils, cylindrical (rarely forked); 1-2 mm in diameter. World distribution - Atlantic Ocean: from Norway to Spain. Pacific Ocean: Japan, China, Hawaii, from Alaska to California. Chorology: Indo-pacific Mediterranean distribution - Thau Lagoon, France (KnoepMer & PCrez in Critchley et al. 1641 first record from that sea); Lagoon of Venice, Italy 1941; Spain [ l 12al. Vector of introduction: by aquaculture (mollusc culture)

Scytosiphon dotyi Scytosiphonaceae)

M.J.

Wynne

(Scytosiphonales,

Type locality: Pillar Point, San Mateo County, California


181

Thallus greenish to dark brown in colour, consisting of cylindrical (sometimes flattened) filaments without colourless unicellular paraphyses, mostly uncostricted or with few constrictions, to 12 cm long, arising from a discoid holdfast. Plurilocular sporangia grouped in sori covering most of surface of thallus. World distribution - Atlantic Ocean: England, Spain, Canary Islands. Pacific Ocean: North America coast. Chorology:Circumboreal Mediterranean distribution Trieste, Adriatic sea, Italy, first record from that sea [96]; France [l 891. Vector of introd~~tion: by aquaculture (mollusc culture)

-

Sorocarpus sp. (Ectocarpales, Sorocarpaceae) To this genus, distributed in the Atlantic Ocean (both eastern and western coast of Northern Hemisphere) and in the Pacific Ocean [Japan and Southern Australia (introduced)], were attributed some specimens collected at Giudecca Island (Lagoon of Venice, Italy) by Curiel et al. [65]. According to Curiel et al. [65] the italian specimens, of unknown vector of introduction, seem to not correspond to any of the currently known species of Sorocarpus, so that it could represent a new endemic Mediterranean species.

Sphaerotrichia firma Chordariaceae)

(E. Gepp) Zinova (Chordariales,

Type locality: Shantung, Weihaiwei Boyden, China. Thallus dark brown in colour, to 20 cm high, consisting of single or tufted filaments filiform, hollow, gelatinous, provided with long colourless hairs, arising from a basal disc. The main axis is irregularly branched with branches, inserted at nearly a right angle, bearing short branchlets (2-3 cm long); often curved. It consists of: a cortex composed of 4-5 celled assimilatory filaments, the terminal cell of which is spherical; a subcortical zone 2-3 layered (4-8 in old parts) of short irregular cells; a medullary zone of large cells longitudinally elongate delimiting the central cavity. Unilocular sporangia generally single at the base of assimilatory filaments. World distribution - Pacific Ocean: Japan, China, Russia. Chorology: Pacific Mediterranean distribution Thau Lagoon, France, as S. divaricata (C. Agardh) Kylin [161]. Vector of introduction:by aquaculture (mollusc culture)

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Stypopodium schimperi (Buchinger ex Kiitzing) Verlaque et Boudouresque (Dictyotales, Dictyotaceae) Type locality: Nuweiba, Egypt.

182


Thallus yellowish brown in colour, coriaceous in texture, to 30 cm high, consisting of a disciform fibrous holdfast from which a stipe to 1 cm long, expanding upwards into palmate blades irregularly cleft, arises. The thallus is composed of a cortical layer of small squarish cells and 2-3 to 6-10 layers of medullary cells in the apical and in the basal parts of the thallus, respectively. Tetrasporangia, one cell stalked, developing from the outer cells, are scattered mostly on the upper part of blades. Hairs and paraphyses are lacking. World distribution - Atlantic Ocean: Canary Islands. Indian Ocean: Red Sea, Somalia. Chorology: Indian Mediterranean distribution - Syria, with the misapplied name S. zonale (J.V. Lamouroux) Papenfuss, first record from that sea [125]; Libya, as S. tubruqense Nizamuddin et Godeh [135]; Egypt, Cyprus and Turkey [193]; Lebanon [l 141. Vector of introduction: via the Suez Canal by natural way or by fouling. NOTE - According to Verlaque & Boudouresque [193], Mediterranean records of S. zonale should be referred to S. schimperi. Even though the first published record of this species from the Mediterranean Sea is that published in May 1991 by Mayhoub & Billard [125], with the misapplied name S. zonale (J.V. Lamouroux) Papenfuss, the occurrence of the species in that Sea dates back to 1982, as reported in the paper published in August 199 1 by Verlaque & Boudouresque [l 931.

.Undaria pinnatifida Alariaceae)

(Harvey) Suringar (Laminariales,

Type locality: Simoda, Japan. Sporophytic thallus yellowish brown in colour, to 90 cm high, attached to the substratum by hapteres fibrous and terminally forked, consisting of a stipe and a blade. The stipe, simple, continues upward as a percurrent prominent midrib through the blade. The blade, ovoid to lanceolate, is pinnately sectioned forming leaflets 5-15 cm long and 2-5 cm wide. Sporangia formed in sporophylls forming two opposite wings along the stipe. World distribution - Atlantic Ocean: from England to Spain, Patagonia. Pacific Ocean: Japan, Korea, Australia, New Zealand. Chorology: Atlanto-Pacific Mediterranean distribution Thau Lagoon, France, first record from that sea [140]; Italy [163]. Vector of introduction: by aquaculture (mollusc culture).

-


183

Chlorophyta Acetabularia calyculus Polyphysaceae)

J.V.

Lamouroux

(Dasycladales,

Type locality: Shark Bay (Western Australia) Thallus green in colour, 1-3 cm high, consisting of an unbranched, cylindrical stalk forming in succession at the growing tip whorls of colourless hairs distichously ramified to the fifth order. At maturity a whorl of 22-30 gametangial rays terminally truncate to slightly bilobed, forming a concave cuplike structure (4-7 mm in diameter), terminates the stalk. It grows in sheltered places on sandy bottoms. World distribution - Atlantic Ocean: Caribbean Sea, Brazil, Canary Islands. Indian Ocean: widely distributed in the Ocean. Pacific Ocean: Polynesia, Japan, E Australia, New Zealand, Philippines. Chorologv: Pantropical Mediterranean distribution - Majorca Island, The Balearic Islands (Valet, 1968, first record from that sea); Sabkhat el Bardawil, Egypt [ l 191; Morocco [106]. Vector of introduction: via the Suez Canal by natural way, or by fouling on the hulls of ships. NOTE - Lipkin [l 191 states that his record is the first from the Mediterranean Sea ignoring the previous record published by Valet [184].

Batophora sp. (Dasycladales, Dasycladaceae) To this genus, distributed in the Atlantic Ocean [the Caribbean Sea and the Canry Islands (probably introduced)] were attributed some sterile specimens, of unknown vector of introduction, collected at Taranto (Italy) by Bottalico et al. [31]. Although some specimens were cultured [31] to date no taxonomic conclusions at specific rank could be reached by authors since no reproductive structures were obtained.

Caulerpa mexkana Sonder ex Kiitzing Type locality: Mexico Thallus bright green.in colour, with a long horizontal stolon to 1 mm thick, from which rhizoidal outgrowths downwardly and erect axes upwardly arise. Erect axes, to I0 cm high, are compressed, with a midrib to 1.5 mm wide and bear pinnae to 3-4 mm long, compressed as well, contracted at the base and acuminate, opposite and distichously arranged, with the middle to sub apical part overlapping the middle part of the next pinna. World distribution - Atlantic Ocean: Caribbean Sea, Brazil, Canary Islands. Indian Ocean: widely distributed in the Ocean. Pacific Ocean: from Malaysia to Australia and New Zealand.


184

Chorology: Pantropical Mediterranean distribution - Israel, Lebanon and Syria, as C. crassifolia (C. Agardh) J. Agardh [l 501. Vector of introduction: via the Suez Canal by natural way, or by fouling or ballast waters.

Caulerpa racemosa (Forsskil) J . Agardh (Bryopsidales, Caulerpaceae) Type locality: Suez (Red Sea) Thallus bright green in colour, with a long horizontal stolon, to 2 mm in diameter, naked, attached by rhizoidal outgrowths and bearing erect axes (very variable according to varieties), terete, simple or occasionally branched, to 15 cm high, with short cylindrical to clavate pinnae (absent in$ requienii), distichously to radially arranged more or less crowded. According to Verlaque et al. [194], at least three distinct taxa of C. racemosa CO-existin the Mediterranean Sea: a variety intermediate between C. racemosa v. turbinata (J. Agardh) Eubank and C. racemosa v. uvifera (C. Agardh) J. Agardh [to which the taxon recorded by Hamel [l081 for the first time from the Mediterranean Sea, should be referred], C. racemosa v. lamourouxii (Turner) Weber-van Bosse E requienii (Montagne) Weber-van Bosse (to which Eastern Mediterranean records should be referred) and the invasive variety attributed to C. racemosa aff. v. occidentalis (J. Agardh) Bmgesen. Later, Verlaque et al. [l951 concluded that the invasive taxon should be referred to C. racemosa v. cylindracea (Sonder) Verlaque, Huisman et Boudouresque. It grows on both rocky and sandy bottoms. World distribution - Widely distributed with all of its varieties in the tropical zone of all Oceans. Chorology: Pantropical. Mediterranean distribution - Tunisia, first record from that sea [log]; Egypt [2]; Korallorizon Island, Greece [l 1l]; Syria [l 1l]; Israel [153]; Libya [134]; Turkey [52]; Italy [6]; Greece [137]; France [194]; Lebanon [l 141; Spain [194]. Vector of introduction: for all varieties, except the invasive v. cylindracea, two hypotheses were put forward: i. taxa entered the Mediterranean Sea via the Suez Canal by natural way, or by ships [188], ii. taxa are relicts in the Mediterranean Sea from the Tethys Sea flora [loo, 1011. As concerns v. cylindracea, Verlaque et al. [l951 put forward the hypothesis of an introduction from Western Australia into the Mediterranean Sea via shipping (ballast waters, anchors, etc.) or by an accidental escape from a Mediterranean aquarium or by an intentional act.

Caulerpa scalpelliformis (Brown ex Turner) C. Agardh Type locality: Southern coast of Australia


185

Thallus bright green in colour, with a horizontal stolon to 1.5 mm thick, from which rhizoidal outgrowths downwardly and erect axes upwardly arise. Erect axes, to 20 cm high, compressed, bear pinnae compressed as well, oppositely placed, not contracted at the base, up-curved, with rounded apex and upper and outer margins generally denticulate, with the apex of a pinna generally overlapping the base of the next pinna. The midrib of axes is wider than the length of pinnae. World distribution - Atlantic Ocean: Caribbean Sea, Canary Islands. Indian Ocean: widely distributed in the Ocean. Pacific Ocean: Japan, Indonesia, Australia, New Zealand. Chorology: Indo-Pacific Mediterranean distribution - Syria, first record from that sea [109]; Palestine [42]; Lebanon [150]; Israel [l51]; Turkey [79]. Vector of introduction: via the Suez Canal by natural way, or by fouling or ballast waters.

Caulerpa taxifolia (Vahl) C. Agardh Type locality: St. Croix, Virgin Islands Thallus bright green in colour, with stolons richly branched bearing downward growing branchlets with groups of rhizoids and erect axes. Erect axes, to 60 cm high, compressed, bear pinnae compressed as well, oppositely placed, not contracted at the base, up-curved, with acuminate apex, generally not overlapping. World distribution - Atlantic Ocean: Caribbean Sea, Gulf of Guinea, Canary Islands. Indian Ocean: widely distributed in the Ocean. Pacific Ocean: Western Pacific, from Japan to Australia and New Zealand. Chorology: Pantropical. Mediterranean distribution - Monaco, first record from that Sea [128]; Italy [155]; Croatia, France and Spain [126]. Vector of introduction: escaped from the aquarium of Monaco and dispersed by anchors, fishing tools. Invasive.

Cladophoropsis javanica (Kii tzing) P.C. Silva (Cladophorales, Siphonocladaceae) Type locality: Java lsland Thallus dark green in colour, consisting of entangled filaments forming mats (sometimes small cushions) provided with rhizoids establishing a connection with the surrounding plants in the mat. Filaments 150-200 pm in diameter, irregularly ramified bearing laterals generally unilateral arising in open connection with the lower cell. Chloroplast parietal, reticulate, with numerous pyrenoids, numerous nuclei per cell.

186


World distribution - Indian Ocean: widely distributed in the Ocean. Pacific Ocean: from Japan to Philippine Islands, Papua. Chorology: Indo-Pacific Mediterranean distribution - Alexandria, Egypt, as C. zollingeri (Kiitzing) Reinbold, first record from that Sea [2]; Israel, as C, zollingeri [l 5 l]. Vector of introduction: via the Suez Canal by natural way.

Codium fragile (Suringar) Hariot ssp. tomentosoides (Goor) P.C. Silva (Bryopsidales, Codiaceae) Type locality: the Netherlands Thallus dark green in colour, erect, terete, subdichotomously ramified attached by a broad basal disc. The thallus consists of an internal medulla of slender, branched, interwoven filaments and a peripheral cortex of swollen utricles. Utricles slightly clavate to cylindrical, frequently with broad constriction at or just below middle, ending with apices pointed and thickened forming a mucron to 70 pm long. Chloroplasts numerous, without pyrenoids. Garnetangia ovoid to cylindrical, borne laterally on the sub-apical parts of utricles. World distribution - Atlantic Ocean: both North America (from North Carolina to Maine) and European coast (from Great Britain to Spain, the Azores Islands). Pacific Ocean: California, South Australia, New Zealand. Chorology: East Atlantic (introduced in other areas) Mediterranean distribution - France, first record from that sea [83]; Italy [go]; Spain [141]; Tunisia [76]; Algeria [172]; Turkey [203]; Slovenia [15]; Morocco [20]. Vector of introduction: by aquaculture (mollusc culture). Invasive. NOTE - According to Silva [175], this subspecies should be considered as an introduced plant also in the area where it was originally described (the Netherlands). Its invasive behaviour is due to its broad physiologica1 tolerance and unusually great parthenogenetic reproductive capacity.

Codium taylorii P.C. Silva Type Iocality: Pass-a-Grille Beach, Pinellas County, Florida (USA) Thallus dark green in colour, erect, terete, spongy, bushy, dichotomously ramified, to 15 cm high attached by a basal crust. Branches usually flattened, especially at dichotomies. Utricles cylindrical to slightly clavate, 110-260 pm in diameter, 650-1 150 pm long, with rounded (sometimes truncate) apices. Gametangia, ovoid to fusiform, 1-2 per fertile utricle, pedicellate. World distribution - Atlantic Ocean: Caribbean Sea, Brazil, Uruguay, Ghana, Canary Islands. Indian Ocean: Thailand, India, Seychelles. Chorology: Indo-Atlantic


187

Mediterranean distribution - Israel, first record from that sea [174]; Egypt PI. Vector of introduction: by fouling, ballast waters, anchors, fishing tools. NOTE - Even though the first published record of this species from the Mediterranean Sea is that by Silva [174], the occurrence of the species in that Sea dates back to 20.ii.1958, when some specimens were dredgeg from Eastern harbour of Alexandria, as published by Aleem [5].

Derbesia boergesenii (Iyengar et Ramanathan) Mayhoub (Bryopsidales, Derbesiaceae) Type locality: Krusadi Island, India Thallus dark green in colour, with filamentous sporophyte, consisting of coenocytic, irregularly branched filaments, 15-36 pm in diameter, 1 cm long. Sporangia, cut off by a'wall plug, are pyriform to spherical, stalked and arranged unilaterally. Vesiculate gametophyte ("Halicystis boergesenii Iyengar et Ramanathan" stage) consists of a coenocytic utricle, pyriform, attached to the substratum by a stoloniferous system. World distribution - Indian Ocean: India Chorology: Indian Mediterranean distribution - Syria [124]. Vector of introduction: via the Suez Canal by unknown vector. NOTE - In the Mediterranean Sea only the sporophyte was found [124].

Derbesia rhizophora Yamada Type locality: Horie pier, Matsuyama (Ehime Prefecture, Japan) Thallus dark green in colour, consisting of creeping filaments irregularly ramified and erect filaments to 1.5 cm high, bearing branches laterally or sub-oppositely, slightly constricted at the base. Some lower branches are separated from the axis by a small septation at their bases from which downwardly a rhizoidal filament arises. Sporangia, shortly stalked, obovate to pyriform, often truncate at apices, are scattered on the upper parts of erect filaments. World distribution - Pacific Ocean: Japan. Chorology: Pacific Mediterranean distribution - Thau Lagoon, France, as Derbesia sp. first record from that Sea and Europe [23]. Vector of introduction: by aquaculture (mollusc culture) NOTE - Derbesia sp., recorded by Ben Maiz et al. [23], was referred to Derbesia rhizophora by Verlaque [l891 but giving no descriptions of his specimens supporting that taxonomic conclusion.


188

Monostroma obscurum (Kiitzing) J . Agardh (Ulotrichales, Monostromataceae) Type locality: Biarritz (France) Thallus pale green in colour, foliaceous, monostromatic, 30-40 cm high, with an irregular outline and wavy margin, tapering to a stipe, attached to the substratum by a basal disc. Cells of the blade mostly unordered. World distribution -,Atlantic Ocean: Iceland, from Norway to Morocco, coast of North America. Pacific Ocean: British Columbia, Oregon, Washington. Black Sea. Chorology:Circumboreal Mediterranean distribution - Thau Lagoon, France [l 891. Vector of introduction: by aquaculture (mollusc culture).

Ulva pertusa Kjellman (Ulvales, Ulvaceae) Type locality: Japan Thallus bright dark green in colour, foliaceous, distromatic, 15-20 cm high, without microscopic marginal teeth, 80-400 pm thick in the basal regions and 40-130 pm thick in the middle regions, provided or not with a stipe, perforated with numerous scattered roundish openings, cells in surface view irregularly arranged. World distribution - Pacific Ocean: Japan, China, Korea, Philippines. Indian Ocean: Yemen, Kenya, Tanzania, Mauritius. Chorology: Indo-Pacific Mediterranean distribution - Thau Lagoon, France [l 891. Vector of introduction: by aquaculture (mollusc culture). NOTE - According to Verlaque et al. [l911 U. pertusa would appear to have the capacity to colonize large areas in temperate European regions both in the Mediterranean Sea and Atlantic ocean.

Monocotyledones Halophila stipulacea Hydrocharitaceae)

(Forsskiil)

Ascherson

(Helobiae,

Type locality: Red Sea Rhizome 0.5-2 mm wide, erect lateral shoots with 2 scales only at the base and at most 5 pairs of petiolate leaves at the top. Leaves linear, with ascending cross-veins. Leaf blades 3-6 cm long, 2.5-8 mm wide with margin serrulate and apex obtuse. Petiole 0.5-1.5 cm long, sheathing obliquely at base. Worlddistribution - Indian Ocean: widely distributed in the western part of the Ocean. Chorology: Indian

189


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Mediterranean distribution Rhodes Island, Greece, first record from that sea [88]; Crete [l 381; Egypt [4]; Lebanon and Cyprus [120]; Malta [l 151; Albania [l 121; Italy [200]. Vector of introduction: via the Suez Canal by natural way.

Taxa to not be considered as introduced Rhodophyta Acanthophora nayadiformis (Delile) Papenfuss (Ceramiales, Rhodomelaceae) This Indo-Pacific species (syntype localities: Alexandria and Suez) was reported as introduced by Boudouresque & Ribera [35], Ribera & Boudouresque [l 561, Boudouresque & Verlaque [38], Giaccone [97, 981. But, for this species described by Delile, as Fucus nayadiformis [71], on the basis of specimens collected also in the Mediterranean Sea (Alexandria, Egypt), Ribera & Boudouresque [l561 don't exclude the possibility that the species is a pre-lessepsian immigrant entered the Mediterranean through the canals dug by the ancient Egyptians as hypothesized by Aleem [2]. Conversely, we agree with the opinion of Feldmann [82], Verlaque [l881 and Cormaci et al. [57], that the species is a Tethyan relict.

Antithamnionella spirographidis (Schiffner) E.M. Wollaston (Ceramiales, Ceramiaceae) This is a sub-cosmopolitan species described by Schiffner as Antithamnion spirographidis [l701 from the Adriatic Sea (type locality: Trieste, Italy) and widely distributed in the Mediterranean Sea [104]. Ribera & Boudouresque [156], Giaccone [97, 981 and Boudouresque & Verlaque [38] list it as a possible introduced species in the Mediterranean Sea. Boudouresque & Verlaque [37] report this species among those probably introduced into the Mediterranean Sea via ship transport, with the following note in their table 4: "The species has been described in a region were it was introduced, before the discovery of its region of origin". That is probably based on the statement of Lindstrom & Gabrielson [l 1 81 that "...Most reports ofA. spirographidis ...have been from harbors or dockyards, except in the north Pacific, where, ......, the taxon is widespread .... Antithamnionella spirographidis may have been introduced to localities outside the north Pacific by man....". That could be correct for southern Australia where the species was recently recorded only from harbours [189], but, in our opinion, it is not correct for the Mediterranean Sea, from where the species was described and where it occurs in several natural habitats.


190

Asparagopsis taxiformis (Delile) Trevisan (Bonnemaisoniales, Bonnemaisoniaceae) This is a Pantropical species, described by Delile (1813, as Fucus taxiformis) from Alexandria (Egypt). It was considered by Verlaque [188], Boudouresque & Verlaque [37, 381 and Giaccone [97] as a possible introduced species. However, Verlaque [l881 didn't exclude the hypothesis that the species could be considered as a Tethyan relict, while Boudouresque & Verlaque [37] in note b added to the above hypothesis that the species could be a pre-lessepsian migrant introduced via waterways built during Pharaonic and Roman times. We agree with the Tethyan relict hypothesis.

Ceramium bisporum Ceramiaceae)

D.L.

Ballantine

(Ceramiales,

This species described by Ballantine [l l ] from Grappler Bank, Puerto Rico, was first recorded from the Mediterranean Sea (Tuscany coast, Italy) by Sartoni & Boddi [167]. We agree with Sartoni & Boddi [167], in not considering this species as introduced since it is highly probable that it was misidentified with Ceramium codii (H. Richards) Feldmann-Mazoyer, a widely distributed species in the Mediterranean Sea [104].

Ganonema farinosum (J.V. Lamouroux) Fan et Y.C. Wang (Nemaliales, Liagoraceae) This species (type locality: Suez, Egypt) was listed by Verlaque, as Liagora farinosa J.V. Lamouroux [188], among introduced species via the Suez canal but at page 8 he didn't exclude that, since the species is distributed in both eastern and western Mediterranean Sea, could be considered as a Tethyan relict. We agree with the latter hypothesis.

Goniotrichopsis sublittoralis G.M. Smith (Porphyridiales, Porphy ridiaceae) This species, distributed in the Pacific Ocean from Alaska to California (type locality: Monterey, California) and in the Fiji, was first reported in the Mediterranean Sea by Magne [l221 from Majorca, Balearic Islands, Spain. It was considered as introduced by Verlaque [l 881 and Boudouresque & Verlaque [37, 381. But, we agree with Magne [l221 that it could have a wider world distribution since it can be easily misidentified with cosmopolitan species of Stylonema from which it differs only in the presence of numerous chloroplasts per cell without pyrenoids (only one chloroplast per cell provided with one pyrenoid in Stylonema).


Halymenia ulvoidea Zanardini (Gigartinales, Halymeniaceae)

191

[nomen

iffegitimum]

This taxon was reported for the first time in the Mediterranean Sea by Aleem [5] from Alexandria (Egypt). Later, it was listed among introduced species by Verlaque [l881 on the basis of Aleem's [5] record. H. ulvoidea Zanardini is an illegitimate name being a later homonym of H. ulvoidea (Sonder) Kiitzing (= Gelinaria ulvoidea Sonder). It was reduced to a variety of H. j7oresia (Clemente) C. Agardh by Codomier as H. m r e s i a v. ulvoidea Codomier, an endemic taxon of the Mediterranean Sea. Aleem [S] considered H, ulvoidea (Sonder) Kiitzing as synonym .of H. ulvoidea Zanardini. Moreover, he stated that the thallus he collected at Alexandria was similar to the Indo-Pacific species Halymenia dilatata Zanardini. Therefore, in absence of a study of Aleem's specimen, it is impossible to ascertain to which taxon it belongs.

Laurencia caduciramufosaMasuda et Kawag uchi (Ceramiales, Rhodomelaceae) This species, described by Masuda et Kawaguchi in Masuda et al. [l231 from Hon Tre Island (Vietnam), was first recorded in the Mediterranean Sea by Furnari et al. [91] from Lachea Island (Sicily, Italy) and Linosa Island (Pelagean Islands, Italy). We agree with Furnari et al. [91] that is early, for the moment, to consider this species as introduced in the Mediterranean Sea. In fact, while L. caduciramulosa was described on specimens collected in Vietnam in 1993, the species was present in the Mediterranean Sea since 1991 [91]. Moreover, it should be pointed out that due to the taxonomic complexity of the genus Laurencia it can not be excluded a wider distribution of the species, due to possible misidentifications with other Mediterranean Laurencia spp..

Laurencia intricata J.V. Lamouroux This species widely distributed in tropical to warm temperate regions in the world (type locality: The Antilles) was first recorded in the Mediterranean Sea by Godeh et al. [I031 from Libya. Later, it was recorded from both Pantelleria Island (the Straits of Sicily) and the Tremiti Islands (Adriatic Sea) by Furnari et al. [91]. In our opinion, it isn't an introduced species, since it is highly probable that it was misidentified with other Mediterranean Laurencia spp..

Laurencia chondrioides Bsrgesen This amphi-Atlantic species (type locality: San Jan, Puerto Rico), occurring also in the Philippines, was first recorded in the Mediterranean Sea by Boisset et al. [29] from Aeolian Islands (Italy), Balearic lslands (Spain), Isla Columbretes

Connaci Mario et al.

192

(Spain), Lachea Island (Italy). We agree with Boisset ef al. [29] in not considering this species as recently introduced into the Mediterranean Sea because it is high probable that it has not previously been there reported because of both its deep habitat and its similarity with Chondria spp. with which it can be misidentified.

Laurencia majuscula (Hawey) A.H.S. Lucas This species (type locality: Ronnest Island, Western Australia), widely distributed from tropical to warm temperate regions of the world, was first recorded in the Mediterranean Sea by Caccamese et al. [39] from Sicily (Italy). Afterwards, L. majuscula was recorded from different Mediterranean localities [173]. Agreeing with Serio et al. [l731 that the species can be easily confused with L. obfusa (Hudson) J.V. Lamouroux, we think that its distribution in the Mediterranean Sea is much wider than to date known and that it can not be considered as a recently introduced species.

Polysiphonia fucoides Rhodomelaceae)

(Hudson)

Greville

(Ceramiales,

This Indo-Atlantic species (type locality: York, England), was reported as introduced by Boudouresque & Ribera [35] as P. nigrescens (Hudson) Greville], by Verlaque, as P. nigrescens [188], by Ribera & Boudouresque as P. nigrescens [l561 and by Giaccone [97, 981. But, since it is present in ancient Mediterranean floras as P. violacea (Roth) Sprengel (one of its numerous synonyms), in our opinion it isn't an introduced species into the ~Mediterranean Sea.

Ochrophyta Desmarestia viridis (O.F. Miiller) (Desmarestiales, Desmarestiaceae)

J.V.

Lamouroux

This Circumboreal species (type locality: Drrabak, Norway) was listed by Verlaque [186], Verlaque [l881 and Verlaque [l 891 as an introduced species to the Thau Lagoon (France) by aquaculture. Also Bellemo et al. [l61 and Giaccone [97] considered this species as newly introduced into the Lagoon of Venice (Italy) by aquaculture. Even though it is possible that the species came in the above basins by aquaculture, it cannot be considered as an alien species to the Mediterranean Sea because it was recorded in the Adriatic Sea since 1849 by Kiitzing [l 131. Its occurrence in that Sea was confirmed by Ercegovic [78]who recorded it from Jabuka Island (Croatia) as Desmaresfia adria~ica Ercegovic.

Mediterranean alien rnacrophytes

193

Chlorophyta Parvocaulis parvula (Solms-Laubach) S. Berger, Fettweiss, Gleissberg, Liddle, Richter, Sawitzky et Zuccarello (Dasycladales, Polyphysaceae) This Indo-Pacific species (syntype localities: tropical India, Celebes) was reported as a lessepsian migrant by Aleern [2] as A. moebii Solrns-Laubach. The species was later recorded from Palestine by Rayss as A. moebii [l511 and reported in the paper by Verlaque [l881 as Acetabularia parvula SolmsLaubach. We agree with both Rayss and Verlaque in thinking more probable that such a species is a Tethyan relict, possibly passed unnoticed in floristic studies.

Taxa excludenda Rhodophyta Audouinella spathoglossi (Bsrgesen) Garbary (Acrochaetiales, Acrochaetiaceae) This species, known only from the Indian Ocean (type locality: Tuticorin, India) was listed by Boudouresque & Ribera [35], Boudouresque & Verlaque [38], Ribera & Boudouresque [l561 and Verlaque [l881 as a lessepsian migrant on the basis of the only record by Aleem [3]. But, Aleem's record [as Kylinia spathoglossi (Bsrgesen) Aleem] is based on some specimens found epiphytic on cast ashore thalli of Spatoglossum variabile (see below), found in the trait of coast between Port Said and El Arish (Egypt). Since no settled specimens were ever collected in the Mediterranean Sea, the species should be excluded from the Mediterranean algal flora.

Galaxaura rugosa (J. Ellis et Solander) J.V. Lamouroux (Nemaliales, Galaxauraceae) This Pantropical species (type locality: Jamaica) was first reported as introduced in the Mediterranean Sea by Boudouresque & Verlaque [38]. Unfortunately, in that paper neither taxonomic nor collection data were given.

Gracilaria arcuata Zanardini (Gracilariales, Gracilariaceae) This Indo-Pacific species (type locality: Aquaba, Jordan, Red Sea) was reported as int~oducedby Boudouresque & Ribera [35], Verlaque [188], Ribera & Boudouresque [l561 and Boudouresque & Verlaque [38] on the basis of records by Feldmann [81] from Tunisia and Aleem [2] from Alexandria and Port Said (Egypt). But, according to Gargiulo et al. [95] the occurrence of this species in the Mediterranean Sea is doubtful.

194


Gracilaria disticha (J. Agardh) J. Agardh This species, known only from the Indian Ocean (type locality: Coast of Eritrea), was reported as introduced by Verlaque [l881 and Ribera & Boudouresque [IS61 (both with doubt) and later with no doubts by Boudouresque & Verlaque [38]. All the above citations are based on the only Lyle's finding at Port Said published by Fox [87]. Due to the taxonomic complexity of this genus, the occurrence of this species in the Mediterranean Sea should be confirmed.

Hypnea esperi auctorum (Gigartinales, Hypneaceae) This Indo-Pacific species, with both nomenclatural and taxonomic complicated histories, see Silva et al. [176], was reported in the Mediterranean Sea by Lipkin [I 191 from Israel. Later, it was reported by Athanasiadis [7] from the Saronikos Gulf, on the basis of the record by Diapoulis & Haritonidis [73]. In that paper, Athanasiadis stated that Israel record needed to be confirmed. Since no information aiding the correct identification of the species was given either by Diapoulis & Haritonidis [73] or by Athanasiadis [7] and no other records of this species from the Mediterranean Sea do exist, this species, in our opinion, should be excluded from the Mediterranean flora.

Hypnea nidifica J. Agardh The only records of this species (type Iocality: Hawaiian Islands) from the Mediterranean Sea are those by Reinbold [l541 from Rhodes Island (Greece) and Forti [86] from Simi Island (Greece). Aleem [2], referring to the above two records, considered the species as a lessepsian migrant. But, since both Reinbold's and Forti's records are based on cast ashore specimens and no other records of this species fiom the Mediterranean Sea do exist, H. nidiJica, in our opinion, should be excluded from the Mediterranean flora.

Ochrophyta Cystoseira myrica (S.G. Gmelin) C. Agardh (Fucales, Cystoseiraceae) This Indo-Pacific species (lectotype locality: Kamchatka Peninsula), was listed with doubt by Verlaque [188]. The only Mediterranean record is that by Muschler [ l 3 l] from Alexandria (Egypt) reported as doubtful by both Aleem [S] and Lipkin [ l 191.

Spatoglossurn asperum J. Agard h (Dictyotales, Dicty otaceae) We agree with Verlaque [l 881 that the record from Mediterranean coast of Israel [121], of this species known only from the Indian Ocean (type locality: Sri Lanka), is due to a misidentification.


195

Spatoglossum variabile Figari et De Notaris (Dictyotales, Dictyotaceae) This Indo-Pacific species (syntype localities: Suez and Aqaba) was listed by Boudouresque & Ribera [35], Verlaque [188], Ribera & Boudouresque C1561 and Boudouresque & Verlaque [38] as a lessepsian migrant on the basis of the only record by Aleem [3]. But, Aleem's record is based on cast ashore thalli found in the trait of coast between Port Said and El Arish (Egypt). S'mce no settled specimens were ever collected in the Mediterranean Sea, the species should be excluded from the Mediterranean algal flora.

Chlorophyta Cladophora cf patentiramea (Cladophorales, Cladophoraceae)

(Montagne)

Kii tzing

This Indo-Pacific species (type locality: Tahiti), was listed by Verlaque [188], Ribera & Boudouresque [156], Boudouresque & Verlaque [37] as a

possible lessepsian migrant. We think that is hazardous to include among alien species a taxon not identified with certainty whose presence in the Mediterranean Sea wasn't later confirmed.

Acknowledgements This work was carried out in connection with the "non indigenous marine species of Italian coast project" of ICRAM (Jstituto Centrale per la Ricerca scient~jicae tecnologica Applicata a1 Mare). This work was also supported by a Grant of the University of Catania.

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