Mediterranean Marine Science
Vol. 12, 2011
One hundred years after Pinctada: an update on alien Mollusca in Tunisia
ANTIT M. Département de Biologie. Faculté des Sciences de Tunis, Université El Manar, 2092 Tunis GOFAS S. Departamento de Biologia Animal, Facultad de Ciencias, Universidad de Malaga, E-29071 Malaga SALAS C. Departamento de Biologia Animal, Facultad de Ciencias, Universidad de Malaga, E-29071 Malaga AZZOUNA A. Département de Biologie. Faculté des Sciences de Tunis, Université El Manar, 2092 Tunis http://dx.doi.org/10.12681/mms.53
Copyright © 2011
To cite this article:
ANTIT, M., GOFAS, S., SALAS, C., & AZZOUNA, A. (2011). One hundred years after Pinctada: an update on alien Mollusca in Tunisia. Mediterranean Marine Science, 12(1), 53-74. doi:http://dx.doi.org/10.12681/mms.53
http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:33 | Mediterranean Marine Science Research Article Indexed in WoS (Web of Science, ISI Thomson) The journal is available on line at http://www.medit-mar-sc.net
One hundred years after Pinctada: an update on alien Mollusca in Tunisia
M. ANTIT1, S. GOFAS2 C. SALAS2 and A. AZZOUNA1
1 Département de Biologie, Faculté des Sciences, Université de Tunis El Manar, 2092 Tunis, Tunisia 2 Departamento de Biolog a Animal, Facultad de Ciencias, Universidad de M laga, E-29071 M laga, Spain
Corresponding author: [email protected]
Received: 25 October 2010; Accepted: 21 December 2010; Published on line: 11 April 2011
Abstract
The occurrences of non-indigenous marine molluscs in Tunisia are reviewed, based both on a literature survey and on original material. Species are accepted as established if there are two inde- pendent reports, either geographically separate or at least one month apart in time. On these grounds, fourteen species are accepted (twelve alien and two expanding their range from elsewhere in the Mediterranean), three aliens need confirmation but are likely to meet the standards for accept- ance in the near future, and five records are rebutted or questioned. Two more species may be con- sidered as cryptogenic, the reports are reliable but there is no clear indication that they are not indigenous. Two of the alien species are reported for the first time in Tunisian waters: the nudi- branch Polycerella emertoni qualifies as established, and the bivalve Anadara transversa is tentatively identified from a live juvenile specimen, which awaits further confirmation. The occurrence of aliens in Tunisia is balanced between presumably Lessepsian species of trop- ical Indo-Pacific origin, and species from other sources including species from the tropical Atlantic introduced through shipping. Nevertheless there is a prevalence of Lessepsian species towards the Gulf of Gabès in the south, while the shipping activity in Tunis harbour may be the main pathway of introduction in the north.
Keywords: Alien mollusca; Polycerella emertoni; Anadara transversa; Tunisia.
Introduction this respect, the Mollusca are one of the most intensively studied groups (GOFAS The fauna of the Mediterranean Sea is & ZENETOS, 2003; ZENETOS et al., 2004) known to be undergoing important changes and may be taken as representative of more related to the establishment of many exot- general trends in the benthos. ic species as a consequence of several fac- The coastline of Tunisia spans the tran- tors, such as human intervention or climate sition between Eastern and Western basins warming (ZENETOS et al., 2008; 2010). In and therefore is a key area for understand-
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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:33 | ing the progression of alien species in the tion of records follows ZENETOS et al. Mediterranean as a whole. The knowledge (2004: 31) & ZENETOS et al. (2010). Records of the Tunisian molluscan fauna relies heav- are considered as established if the species ily on studies published in the late 19th and have permanent, self-maintaining popula- early 20th century (DAUTZENBERG, 1883; tions, and this must be documented by at 1895; PALLARY, 1904-1906, 1914). The least two reliable and independent records. rather intensive surveys conducted at that Where such records originate from the same time, although biased towards the Gulf of authors they ought to be based on separate Gabès, have the advantage of providing a samples, collected at least one month apart reasonably accurate baseline for the mala- if they originate from the same site. Records cofauna in the early 20th century. are therefore evaluated in one of these cat- This knowledge nevertheless badly needs egories ‘Established, Casual, Questionable, updating, particularly regarding small and Cryptogenic, or Excluded’ following inconspicuous species, recently recognized ZENETOS et al. (2010), but considering Mediterranean taxa, and introduced species. apart the species cultured in marine farm- At the time when the ‘CIESM Atlas of Ex- ing. Contrary to the CIESM Atlas, we al- otic Species’ (ZENETOS et al., 2004) went so discuss range extensions from within the to press, there were only five Indo-Pacific Mediterranean, therefore including species alien species of molluscs reliably reported that were not native to Tunisia but had a in the Tunisian fauna: the gastropods Cerithi- historical range elsewhere in the basin and um scabridum, Melibe fimbriata and Bur- therefore do not qualify as ‘alien’ species. satella leachii and the bivalves Pinctada ra- Records will be presented in chronological diata and Fulvia fragilis. To those the gas- order of discovery in Tunisia, established tropods Echinolittorina punctata and records first, those in need of confirmation Siphonaria pectinata may be added, possi- next. bly arrived from other parts of the Mediter- Introduction pathways are inferred as ranean by their own means, and the oys- Lessepsian for species of Indo-Pacific ori- ter Crassostrea gigas repeatedly introduced gin which have been recorded in the Red for farming. These numbers have now near- Sea and Suez Canal and first entered the ly doubled and it is the goal of this paper to Mediterranean in the Levantine Sea. In- analyze the progress of the more ancient troduction as a consequence of shipping ac- immigrants, to report on the input of addi- tivity is presumed when an alien species is tional species and to discuss the pathways first detected in the vicinity of a major har- which are now documented for Tunisia. bour. Material from around the island of Djer- Materials and Methods ba was examined in the Muséum National d’Histoire Naturelle, Paris. This material The records of exotic molluscs in Tu- was collected in several localities around nisia have been searched for in the pub- the island, during one month in 1982 (ex- lished literature and over the World Wide pedition leader Philippe Bouchet), using Web; a particular effort has been made to scuba diving and an airlift pump on all kinds screen the ‘gray’ literature, including con- of substrates. This systematic collecting, not gress reports and memoirs written for ac- biased towards particular species, is partic- ademic degrees. The rationale for evalua- ularly useful for assessing presence/absence
54 Medit. Mar. Sci., 12/1, 2011, 53-73
http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:33 | of the common species. (ZENETOS et al., 2004 and references there- The original records presented in this in) and first appeared in Gulf of Gabès pri- paper are based on our own ongoing work, or to 1892. DAUTZENBERG (1895) gave dedicated to a survey of selected coastal en- a wealth of details regarding early reports vironments in the Bay of Tunis. on the species and noted that it was inva- The malacofauna of the infralittoral al- sive by 1892, whereas it was not detected in gal mat was sampled monthly at 1-2 m wa- a previous survey in 1882. It is today one of ter depth, on an artificial rocky pier situ- the most common molluscs on the whole ated at La Goulette (36Æ49.15'N - 10Æ18.60’E). Tunisian coast and is locally a dominant from February 2009 to February 2010. Sam- species, particularly in Gulf of Gabès. How- ples were collected on five replicas of squares ever its distribution seems to end abruptly of 25 x 25 cm. Algae were washed to re- at Bizerte (ZAKHAMA-SRAIEB et al., moving all associated animals and dried for 2009; TLIG-ZOUARI et al., 2010) and the determination of their biomass. The asso- reports in the Western Mediterranean are ciated fauna proceeding from washing as scanty (e.g. BOUDOURESQUE, 1999). It sieved over a 0.5 mm sieve and preserved is uncommon in our material from Tunis in 70% ethanol for picking out specimens. Bay. Samples from soft bottoms at La Goulette The valid name for this species has been were sampled with a dredge at 3-4 m debated although there is a consensus among (36Æ49.1'N - 10Æ18.9’E) and 10-15 m recent authors on using the name radiata (36Æ49.1'N - 10Æ19.6’E) depth. The dredge for the Red Sea/Mediterranean species was towed for 5 minutes parallel to the shore, (WADA & T MKIN, 2008). The type lo- at a speed of approximately one knot, and cality was nevertheless stated, with a ques- samples were replicated three times. The tion mark, as Caribbean and the name is al- sample was sieved on a column of sieves of so used for the very similar, if not identical, 1 cm, 5 mm, 2 mm and 0.5 mm mesh and Western Atlantic species otherwise known preserved in 70% ethanol. as P. imbricata (Röding, 1798). All samples were later sorted at the lab, Echinolittorina punctata (Gmelin, 1791) each size fraction separately, to separate is a predominantly tropical eastern Atlantic the molluscs. All shelled species were then species with an extensive historical range in dried, whereas selected specimens includ- the southern Mediterranean, along the ing all opisthobranchs were kept in ethanol. Moroccan and Algerian and along the Egypt- This material was then sorted down to species ian and Levantine coasts (PALLARY, 1912). level and identified. ANTIT et al. (2007) supported the view that the species arrived in Tunisia and Sicily at Results some time in the first half of the 20th cen- tury since it is unlikely that (1) Species recorded prior to the completion MONTEROSATO (1878) and PALLARY of the CIESM Atlas (2002) and considered (1914) would have missed it in their re- as established (Table 1) spective areas if it had existed at their time. Pinctada radiata (Leach, 1814) is the Crassostrea gigas (Thunberg, 1793) has oldest reported Lessepsian migrant in the been repeatedly introduced for aquaculture Mediterranean. It was first found in the since at least 1932, first in the Ghar el Melh Mediterranean at Alexandria in 1874 lagoon, formerly known as Porto Farina
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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | Origin pathway occurrence status 2003] IP lessepsian T G established et al., 1990 [this paper] IP (combined) G established et al. Table 1 2005 TP (unknown) T G established - 2004 WA shipping T established - 2005 TP (unknown) T established et al., et al., et al. 2007] 2010 WA shipping T established et al., et al., [ANTIT First published report [Report of earliest occurrence] WIMART-ROUSSEAU, 2004 1996 BEN SOUISSI (N: North coast, T: Gulf of Tunis, H: Gulf of Hammamet, G: Gulf of Gabès). fourth column the geographic origin (ATL: Western Atlantic; IP: tropical Indo-Pacific, The last column indicates our statement of the status of the species, as discussed in the text. (Gmelin, 1791) <1950 ENZENROß & ENZENROß, 2001 MED own means N T established MED: other parts of the Mediterranean; TP: temperate North Pacific; WA: tropical West Africa); (Benson in 2004 BEN SOUISSI Philippi, 1848 1997 ENZENROß & ENZENROß, 2001 [this paper] IP (combined) T G established (Linné, 1758) <1998 ENZENROß & ENZENROß, 2001 MED own means N T established a Edmunds, 1968 2003 BEN SOUISSI Verrill, 1881 2009 this paper ATL shipping T established de Blainville, 1817 1982 ENZENROß & ENZENROß, 2001 [this paper] IP lessepsian T G established fifth column the suspected pathway of introduction; sixth column detail of current occurrence within Tunisia (Thunberg, 1793) <1932 DANTAN & HELDT, 1932 TP farming N farmed (Leach, 1814) <1890DAUTZENBERG, 1895 IP lessepsian N T H G invasive (Lamarck, 1810) 2003 WIMART-ROUSSEAU & IP lessepsian G established List of ascertained exotic molluscs in Tunisian waters, in chronological order of earliest record (second column); (Duclos, 1846) 2008 ANTIT (Forsk l 1775) 1990PASSAMONTI, 1996 [BEN SOUISSI third column indicates the first published report of the species and the reference to the earliest finding, if different; Kelaart, 1848 1982 CATTANEO-VIETTI Ruditapes philippinarum Melibe viridis Crassostrea gigas Bursatella leachii Fulvia fragilis (Adams & Reeve, 1850) Cerithium scabridum Siphonaria pectinata SpeciesPinctada radiata Echinolittorina punctata Year Favorinus ghanensis Erosaria turdus Cantor, 1842) Mitrella psilla Musculista senhousia Polycerella emertoni
56 Medit. Mar. Sci., 12/1, 2011, 53-73
http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | (DANTAN & HELDT, 1932, as Crassostrea (2008), a Tunisian specimen was represented angulata) and later in the Bizerte lagoon in the Atlas of Mediterranean Nudibranchs and nearby Lake Ichkeul (AZOUZ, 1966; of CATTANEO-VIETTI et al. (1990). The MEDHIOUB & ZAOUALI, 1988; DRIDI species was present in the Gulf of Gabès as et al., 2006). Populations were generally not early as 1982 as documented by photographs self-sustained and successful reproduction taken during the MNHN expedition (Fig. was only reported in 2002-2003 for the Biz- 1), that is more or less simultaneously with erte Lagoon (DRIDI et al., 2006). the discovery of the Greek populations. Melibe viridis Kelaart, 1858 (= M. fim- Bursatella leachii de Blainville, 1817 is briata Alder & Hancock, 1864) is an Indo- at the forefront of progression of Lessep- Pacific species whose occurrence in the sian immigrants towards the western Mediter- Mediterranean was first recorded in Greece ranean. Its existence in Gulf of Gabès was (THOMPSON & CRAMPTON, 1984). also documented by the MNHN expedition Although its occurrence in Tunisia was first in 1982 (Fig. 2), nearly two decades be- formally reported on by CECALUPO et al. fore the first published report for Tunisia
Fig. 1-2: Opisthobranch molluscs photographed during the expedition of Muséum National d’Histoire Naturelle to Djerba, in May 1982. 1. Melibe viridis. 2. Bursatella leachii (Photographs by Claude Huyghens and Françoise Danrigal).
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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | (ENZENROß & ENZENROß, 2001, col- et al., 2008 and references therein). This is lected in 1996). Bursatella leachii has made an invasive species which must be watched its way along the Italian coasts of the Tyrrhen- for and should by no means be introduced ian sea (FASULO et al., 1984), to the Balear- deliberately, since it poses a risk for local es (OLIVER & TERRASA, 2004), to the extinction of R. decussatus. Spanish eastern peninsular coast Cerithium scabridum Philippi, 1848 is a (WEITZMANN et al., 2009; RAMOS- very early settler in the Eastern Mediter- ESPL et al., 2010) and, within Tunisia, to ranean but its colonization history is more the north coast (ZAKHAMA-SRAIEB et haphazard. Documented on the Levantine al., 2009). In all cases it seems to favour la- coast in the late 19th - early 20th century goonal or lagoon-like environments and to (PALLARY, 1912), it did not reach Sicily have a discontinuous distribution. (Brucoli) until 1972 (DI NATALE, 1978). Fulvia fragilis was first reported in Tu- Nevertheless, this species was not seen dur- nisia by PASSAMONTI (1996), in Gulf of ing the collecting expedition of the Muséum Gabès, but actually has been there at least National d’Histoire Naturelle of Paris at since 1990 (BEN SOUISSI et al., 2003) and Djerba in 1982, therefore was probably not not in the 1982 material in MNHN. It was present at that time. It was present at Djer- common a few years later (ENZENROß & ba in 1997 (material in MNHN, Paris, col- ENZENROß, 2001, collected 1996-1998). lected by SG) and its occurrence in Tunisia It is still common on the whole coast and since 1998 was first reported by ENZENROß has, like Bursatella leachii, reached far into & ENZENROß (2001). Larval dispersal the Western basin; in eastern Spain since should not be a limiting factor for this species 2002 (ZENETOS et al., 2004; TAMAYO with planktotrophic larvae, but the note- GOYA, 2008) and in southern Italy worthy occurrence of several native species (CROCETTA, 2005; CROCETTA et al., of Cerithium in Tunisian waters, particularly 2008). the Gulf of Gabès (NORDSIECK, 1974), may have hampered its progression for some Ruditapes philippinarum (Adams & time. It is now common, and locally inva- Reeve, 1850) has been reported in the south- sive, along the whole coast and it has been ern section of the Lac de Tunis coastal la- recently reported in Greek waters, filling goon since 1996 (BEN SOUISSI et al., 2005). backwards the gap in its Mediterranean dis- By 2005 it is reported by these authors as tribution (ZENETOS et al., 2009). Never- still rare and restricted to ‘a restricted theless, its progression towards the Western biotope close to the Gulf of Tunis’. It was basin is shy, currently limited to the extreme further reported from the Gulf of Gabès south of peninsular Italy (CROCETTA et area by ZAMOURI-LANGAR et al. (2006). al., 2008). We have not seen it in our material from Siphonaria pectinata was reported as La Goulette, in which the native Ruditapes early as 1998 from Tabarka and Bizerte decussatus (Linné, 1758) still seems healthy. (ENZENROß & ENZENROß, 2001) and Nevertheless the two species are known to in 2007 from La Marsa in the Gulf of Tu- compete for the same poorly sorted sedi- nis (ANTIT et al., 2007) and currently per- ments in areas of low hydrodynamism and sists or progresses in these localities. Like Ruditapes philippinarum has displaced the Echinolittorina punctata, it has an historical native species elsewhere (e.g. PRANOVI range inside the Mediterranean, on the Span-
58 Medit. Mar. Sci., 12/1, 2011, 53-73
http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | ish, Moroccan and Algerian coasts multaneously by BEN SOUISSI et al. (2005) (MONTEROSATO, 1878), but contrary to based on trawling in late 2003 and 2004 in Echinolittorina punctata, its arrival on the the Gulf of Gabès. It now seems to be com- Tunisian coast is recent and has been wit- mon throughout the Gulf of Gabès area nessed in recent surveys. (ROBIN, 2007) and is easily found because of its habit of entering the clay pots used in (2) Species reported posterior to the comple- octopus fisheries. The species was also re- tion of CIESM Atlas (2002) and considered ported on the Libyan coast on the Internet as established forum ‘cowries.info’ of Felix Lorenz These are the species meeting the cri-
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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | followed throughout the sampling survey in Western Italy, Malta and Greece. Further- the infralittoral algae of La Goulette (Table more, it is associated with the Bryozoan 2) and the egg capsules were observed in Zoobotryon verticillatum (Delle Chiaje, 1822) many samples (Figs 5-8). The egg capsules which itself is recognized worldwide as an in- are of a semitransparent texture, barrel- vasive species in the fouling community shaped attached by their base to the sub- (AMAT & TEMPERA, 2009) and is listed strate and reinforced laterally by small in the Global Invasive Species Database (In- vertical ridges; the upper surface acts as vasive Species Specialist Group an operculum to release the juvenile snail.
Table 2 Abundance of Mitrella psilla in the samples studied in algal wash at La Goulette (36Æ49.15'N - 10Æ18.60’E). A, adults; M, medium-grown specimens, more than half the adult size but without formed peristome as on figure 8; J, juveniles; E: egg-cases.
R1 R2 R3 R4 R5 Date A M J E A M J E A M J E A M J E A M J E 27/02/2009 1 2 3 - 2 - 4 1 3 - - - - - 1 - - - 31/03/2009 29/04/2009 2 - 1 2 - 1 2 1 - 8 1 27/05/2009 11 12 8 3 3 2 2 1 1 - 20 7 4 - 29/06/2009 - 3 3 2 - 2 1 - - - 1 ------2 1 - 23/07/2009 2 2 ------1 - 2 - 3 - 18/08/2009 2 4 4 - - 1 0 - - 2 - - - 1 - - 1 - 29/09/2009 3 1 ------29/10/2009 - - - - 9 3 - 13 1 - 1 1 - - - 3 - 4 4 24/11/2009 2 - 3 1 2 - - - 1 - - - - 23/12/2009 5 9763- - 3 -19-213 - -2- 19/01/2010 - -1-115------111--1 23/02/2010 3 1 10 10 5 - 3 - 3 6 1 2 4 - 1 - - - 3 -
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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | Table 3 Abundance of Polycerella emertoni in the samples studied at La Goulette (36Æ49.1'N - 10Æ18.9’E, 3-4 m and 36Æ49.1'N - 10Æ19.6’E, 10-15 m).
3-4 m R1 3-4 m R2 3-4 m R3 10-15 m R1 10-15 m R2 10-15 m R3 29/09/2009 29/10/2009 24/11/2009 23/12/2009 0 2 7 19/01/2010 127 68 66 0 57 25 22/02/2010 16 23 48 135 48 38 31/03/2010 77 29 180 120 33 23
(3) Species reported as alien or possibly alien imen of the native Crepidula moulinsii on the Tunisian coast, in need of confirma- Michaud, 1829. tion (Table 4) Acteocina knockeri (Smith, 1872) is a This section includes (1) ‘promising’ tropical West African species recorded by casual records based on only one recent ob- HOENSELAAR & GULDEN (1991) from servation, where the context allows the a single shell collected in March 1989 in the expectation of a confirmation in the near Kerkennah islands at Sidi Youssef future and (2) records where some elements (34Æ39’26"N, 15Æ58’13"E, plotted from in- allow doubt as to their validity. dications in the text). It was never mentioned Tornus jullieni Adam & Knudsen, 1969 again but in this case a confusion is possi- is a West African species reported by PIANI ble with the very similar Indo-Pacific Acteoci- & BRINI (1984) based on a single shell col- na mucronata (Philippi, 1849) which is def- lected on the East coast of Djerba initely established along the Levantine coast (33Æ46’50"N, 11Æ03’28"E, plotted from in- (ZENETOS et al., 2004). Further records dications in the text) in August 1973. It has are nevertheless needed to substantiate the never been reported again anywhere in the report from Tunisia. Mediterranean or NW Africa and thus, more Saccostrea cucullata (Born, 1778) was than a quarter of a century later, this record reported, without details of exact localities should be left as questionable rather than and material examined, by ROMDHANE venturing possible explanations. & TRITAR (2002) from the north coast [of Crepidula fornicata (Linné, 1758) was Tunisia] at 25-30 m and the Bizerte chan- reported from trawl samples collected in nel. These authors explain the finding as a the Gulf of Gabès (FEHRI-BÉDOUI, 1986) possible range extension from the Adriat- and the species has never been seen again ic. However, the Adriatic records refer to a in Tunisia. The most likely pathway for this different species Saccostrea commercialis species, which is invasive in Western Europe, (Iredale & Roughley, 1933) and were nev- would be the importation of oyster spat for er ascertained after 1990 nor outside the marine farming. There is no such activity in Venice Lagoon (see ZENETOS et al. 2004 the Gulf of Gabès and we can suspect that and references therein). Saccostrea cucul- the record was based on a very large spec- lata is a prevalently intertidal species in its
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http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | native area (LAM & MORTON, 2004) and we consider that this record must be considered as unwarranted unless
Table 1. further confirmed. Chromodoris quadricolor (Rüp- pell & Leuckart, 1830) is cited from the Bibane Lagoon near the Libyan border by BEN SOUISSI et al. (2004), collected in May 2003 at 5 m depth with abundant sponges. This species is a good candidate for becoming an established species in the Mediter- ranean. There is probably no con- nection between this and the first
Origin pathway occurrence status Mediterranean report in northern Italy in 1986 (see ZENETOS et al., 2004) but the species has been recently re- ported on the Turkish coast (ÖZTÜRK & CAN, 2006). Discodoris lilacina (Gould, 1852) was reported from the southern sec- tion of the Lac de Tunis coastal la- goon by BEN SOUISSI et al. (2005), Table 4 2005 ? cryptogenic T cryptogenic 2004and IP this record lessepsian is supported G casual by a pho- 2007tograph. Illustrations IP lessepsian in colour, of G casual spec- et al., et al., imens from the same source collect- et al., ed in January and March 2003, are al- so given on the Sea Slug Forum of the Internet First published report [Report of earliest occurrence] rum.net/find/discmacu> and there reidentified as the native species Dis- codoris maculosa (Bergh, 1884). The first report of Discodoris lilacina in the Mediterranean was that of VALDÉS & TEMPLADO (2002) and is based on specimens collected on the Le- (Rüppel & 2003 BEN SOUISSI vantine coast, but earlier BARASH & DANIN (1977) used the name Dis- (Gmelin, 1791) <2002 ROMDHANE & TRITAR, 2002 IP (unknown) N questionable (Linné, 1758) 1984 FEHRI-BÉDOUI, 1986 ATL (unknown) G questionable (Say, 1822) 2010 this paper ATL (combined) T casual (Smith, 1872) 1989 HOENSELAAR & GULDEN, 1991 WA (unknown) G questionable (Linné, 1758) 2007 BEN SOUISSI & ZAOUALI, 2007 IP (unknown) G questionable (Gould, 1852) 2003 BEN SOUISSI codoris concinna (Alder & Hancock, (Audouin, 1826) 2007 CAMPANI, 2008 ? cryptogenic N H cryptogenic Adam & Knudsen 1969 1973 PIANI & BRINI, 1984 WA1864) (unknown) for this Gspecies, questionable a species which (Gmelin, 1791) <2007also ZAOUALI has an Indo-Pacific type locality (in Madras, India). There is consid- erable taxonomic confusion regard- List of cited exotic or cryptogenic molluscs in Tunisian waters, in need of confirmation. Conventions and abbreviations as for ing this species which belongs to a Chromodoris quadricolor Saccostrea cucullata Discodoris lilacina SpeciesTornus jullieni Crepidula fornicata Acteocina knockeri Year Leuckart, 1830) Cellana rota Monetaria annulus Alvania dorbignyi Anadara transversa 62 Medit. Mar. Sci., 12/1, 2011, 53-73 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | complex of morphologically similar forms way as Erosaria turdus, before the species occurring in many tropical and temperate can be taken as an element of the Mediter- areas of the world. DAYRAT (2010) at- ranean fauna. tempted a revision of this complex, using Cellana rota (Gmelin, 1791) was re- for it the genus Tayuva Marcus & Marcus, ported by ZAOUALI et al. (2007) from the 1967 and taking the option to subsume sev- piers of the commercial harbour of Zarzis, eral species worldwide into Tayuva lilacina near the Libyan border, and also from Tripoli, ( ‘Tayuva lilacina of the Indo-West Pacific’ Libya. This occurrence seems straightfor- for Doris lilacina Gould, 1852 with a type ward since this species is in clear progress, locality in Hawaii, ‘Tayuva lilacina of the or even invasive, on the Levantine coast Panamic Eastern Pacific’ for Tayuva ketos (MIENIS, 2002). Further expansion is pre- Marcus & Marcus, 1967, ‘Tayuva lilacina of dictable and the species may soon prove to the Caribbean’ for Peltodoris crucis Bergh, be established in Tunisia. The question of 1880 and ‘Tayuva lilacina of the Eastern whether this is the valid name depends on Atlantic and Mediterranean’ for Doris mac- the resolution of the taxonomic controver- ulosa Bergh, 1884). Dayrat writes that the sy over whether this is a synonym of Cellana real Tayuva lilacina (i.e. his ‘Tayuva lilacina radiata (Born, 1778) or a separate species of the Indo-West Pacific’) is rare in the In- (e.g. DEKKER & ORLIN, 2000). dian Ocean and that all records under the Alvania dorbignyi (Audouin, 1826) was names lilacina or concinna in the Red Sea recorded from Tabarka by CAMPANI (2008) are mostly based on another species Se- and our own shore collections from Salak- badoris fragilis (Alder & Hancock, 1864) ta, in the Gulf of Hammamet in 2007 also (see discussion in DAYRAT, 2010: 126- yield many specimens of this species. There 127). Therefore, the most likely outcome is is some uncertainty regarding whether this that the Tunisian specimens are not Red species is an immigrant in the Mediter- Sea immigrants but belong to the native ranean, or just a native species which was member of the complex, Discodoris macu- overlooked due to its overall similarity to losa, as stated on the Sea Slug Forum. the well-known Alvania montagui Monetaria annulus (Linné, 1758) was (Payraudeau, 1826). The only grounds to reported by BEN SOUISSI & ZAOUALI consider this species as possibly Indo-Pa- (2007) as collected in spring 2006 from sev- cific are that most of Savigny’s material, eral points of the infralittoral of the Kerken- from which the taxon was described, origi- nah Islands, and off Tripoli, Libya. This is nated from the Red Sea, but actually it al- far from being the first report of this species so contains a few Mediterranean shells in the Mediterranean (see e.g. LOCARD, (BOUCHET & DANRIGAL, 1982). Even 1886: 94). There are holdings in the 19th if a native Mediterranean species, one can- century collection of Muséum National not rule out a recent progression within the d’Histoire Naturelle, Paris, of ornaments basin, but its absence in the 19th century labelled as coming from Tunisia and made checklists is far less conclusive than in the with shells of this species, which was wide- case of Echinolittorina punctata. spread throughout Africa as a currency and Anadara transversa (Say, 1822) is one a material for handicraft. Therefore, we be- of the three introduced species of Anadara lieve that such records must await repeat- currently recorded in the Mediterranean. ed observations of living animals in the same A juvenile living specimen (Figs 11-12) of Medit. Mar. Sci., 12/1, 2011, 53-73 63 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | Fig. 3-12: Alien species of molluscs collected since year 2008 in Gulf of Tunis. 3-4: Specimen of Mus- culista senhousia collected in algal wash at La Goulette (actual size 10 mm). 5-7: Egg capsules of Mitrel- la psilla, collected 23.01.2010 at La Goulette in infralittoral algae; scale bars are 1 mm. 8: Adult speci- men of M. psilla, same locality. 9-10: Polycerella emertoni, specimens from off La Goulette, 10-15 m col- lected 19.01.2010. The specimen on the right has just produced an egg mass (actual size ca. 3 mm). 11- 12: Juvenile specimen of Anadara sp. tentatively identified as Anadara transversa (Say, 1822), off La Goulette, 10-15 m collected 19.01.2010. an Anadara species collected off La Goulette know the juvenile stage of the native Anadara on a soft bottom at 10-15 m is at present the corbuloides (Monterosato, 1880). There- only clue we have to the arrival of one of fore, we prefer to await the collection of these species on the Tunisian coast. The na- somewhat larger individuals before ven- tive Anadara polii (Mayer, 1868) is definitely turing a species-level identification. ALBANO shorter at this size and lacks the long bris- et al. (2009) considered from morphologi- tles in the intercostal spaces, but we do not cal and molecular data that this species, 64 Medit. Mar. Sci., 12/1, 2011, 53-73 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | hitherto known in the Mediterranean as A. species (20 molluscs) added between 2005 demiri (Piani, 1982) was very similar, if not and 2008, the increase is 10,2% (9,2% for identical, to the Western Atlantic A. trans- molluscs) and averages one species every 9 versa (Say, 1822). days (one mollusc every 42 days). Within Tunisia, the numbers are far from being so Discussion impressive but the acceleration in the last decade is comparable or, if anything, is sharp- Chronology of first arrival and rate of influx er, with three species (Mitrella psilla, Cel- Almost one century separates the ar- lana rota, Monetaria annulus) added to rival of the first alien species in Tunisian the previously recorded 17 (cryptogenic and seas and the next batch, which would in- intra-Mediterranean not counted), making clude Bursatella leachii, Melibe viridis and an increase by 17% between 2005 and 2008. Fulvia fragilis in the late 20th century (Fig. 13). Leaving aside the cupped oyster Cras- Origin of the newcomers sostrea gigas voluntarily introduced for farm- In the Mediterranean as a whole most ing, this represents only three additions of of the alien species are thermophilous species extra-Mediterranean species in this long in- of Indo-Pacific origin and their establish- terval of time. The rather intensive collect- ment in the Mediterranean is seen by POR ing expedition of MNHN in 1982, although (2009, 2010) as the result of congruence be- mostly unpublished, provides an archive of tween the present Climate Optimum (which specimens indicating that this is not an arte- is expressed in the warming of the sea), the fact and that, at least in the Gulf of Gabès, opening of the Canal of Suez as a pathway the numbers were really low and the alleged from the Indo-Pacific area, and the renewed newcomers are really new to the fauna. entry of species through the Straits of Gi- According to ZENETOS et al. (2008) braltar. POR therefore suggested that the a total of 903 alien species, among which migration of these tropical species to the are 216 molluscs, have been incorporated Mediterranean basin represents a certain into the Mediterranean waters. With 94 restoration of the ancient Tethyan biota. Fig. 13: Chronogram of arrival of the alien species of molluscs on the Tunisian coasts, distributed ac- cording to establishment status. Mediterranean and farmed species not included. Medit. Mar. Sci., 12/1, 2011, 53-73 65 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | This view should be considerably qual- southern area (ENZENROß & ified when coming to the Tunisian coast. ENZENROß, 2001; CECALUPO et al., Among the twelve established alien species 2008). in the littoral of Tunis, three have an At- When considering Tunisia as a whole, lantic origin (Favorinus ghanensis, Mitrella the sources of alien species are much more psilla, Polycerella emertoni) and three more balanced than in the Eastern Mediterranean, (Crassostrea gigas, Ruditapes philippinarum reflecting the position of this coastline at and Musculista senhousia) have a temper- the transition between the Eastern and ate North Pacific origin (Table 1; Fig. 14). Western basins. In the detail, the invasion These species appear in the Gulf of Tunis history of the Gulf of Gabès is definitely and on the northern coast and, with the ex- separate from that of the Gulf of Tunis and ception of Ruditapes, they have not reached the north coast. the southern coast and the Gulf of Gabès, which has the warmer sea surface temper- Pathways of introduction ature. Up to now, they have relatively low The complicated history of Cerithium density of population and rate of spread, scabridum indicates that it started its jour- and they cannot be considered as ‘invasives’, ney into the Mediterranean as a plain Lessep- although the impact on native popula- sian species, then its arrival in Sicily around tions should be studied. a major harbour has generally been inter- Of the six established alien species com- preted as helped by shipping and finally, the ing from the Indo-Pacific region, five first distance across the Sicily Straits to Tu- arrived in the Mediterranean through the nisia can be easily bridged by its plankton- Suez Canal and one (Melibe viridis) via ship- ic larvae. Melibe viridis first appeared in the ping. Two of these (Melibe viridis and Erosaria Mediterranean in Greece, not on the Le- turdus) are restricted to the southern coast vantine coast and therefore is not held as a of Gabès, probably in relation to the warm Lessepsian despite its Indo-Pacific origin. climate of this area. The other four (Cerithi- Again, the leg from the Ionian coast of um scabridum, Bursatella leachii, Pinctada Greece to Tunisia can best be explained by radiata and Fulvia fragilis) are present in its own means of dispersal, as such light- nearly the whole coast, from Gabès to Biz- bodied nudibranchs can not only disperse erte, but still they are more abundant in the as larvae but also drift as adults. Fig. 14: Origin of the non-indigenous species of molluscs established on the Tunisian coasts. 66 Medit. Mar. Sci., 12/1, 2011, 53-73 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | From these examples it is clear that Tu- majority of the established species are gen- nisia is certainly not the first foothold in the eralists. The diet of Mitrella psilla is not Mediterranean for most of the species known; whereas some species of Mitrella discussed herein, and that the pathway of are known as specialists feeding on the egg introduction from the area of origin may in- masses of other molluscs (RUEDA et al., volve a combination of various means. Of 2009), other columbellids seem to be also the 12 established aliens, only Mitrella generalist detritivores or omnivores. Cowries psilla and Favorinus ghanensis had their first may be algal grazers or sponge feeders, and Mediterranean report in Tunisia. Both are the exact diet of Erosaria turdus is not known. tropical West African species and their in- Only the two nudibranchs Favorinus gha- troduction is considered mediated by ship- nensis and Polycerella emertoni are definitely ping in the large commercial harbour of Tu- specialist grazers, but their prey are foul- nis. This may not be straightforward from ing bryozoans like Zoobotryon, so that the current shipping routes, where ship move- vector of their introduction will carry the ments from West Africa into the Mediter- supporting host as well. On the whole, ranean account for only 4,1% of total move- the successfully installed species are pre- ments, while more than half originate from dominantly generalist feeders, meeting one NW Europe and Morocco (DOBLER, 2002). of the conditions for being a prospective Even if routes are occasional, there may be invader. a ‘West African connection’ from which Among the 18 molluscan species stat- these species will start their movements ed as invasive somewhere in the Mediter- through the Mediterranean. They did not raean (ZENETOS et al., 2010), five or six enter gradually through the Straits of Gi- (Cerithium scabridum, Bursatella leachi, Pinc- braltar, where the fauna is well known and tada radiata, Musculista senhousia, Rudi- where recent surveys would have detected tapes philippinarum, and maybe Anadara them if they were present. transversa) are found in Tunisia, although The Western Atlantic Polycerella emer- not necessarily invasive there. This supports toni, and Anadara transversa if confirmed, the now well-known statement of ‘intro- have certainly been conveyed through ship- duced here, introduced elsewhere’. Two of ping at some stage (see GALIL et al., 2008) them are of Indo-Pacific origin, therefore but their arrival in Tunisian waters is best not so much supporting POR’s (2010) view explained as a secondary introduction by that thermophilic aliens of Tethyan origin their own means of dispersal with pelagic should be more ‘at home’ and less invasive larvae from other parts of the Mediterranean than those of other origins. (southern Italy, Malta ...) where they are al- ready established. Conclusions and Perspectives Is Tunisia prone to marine invasions? Composition of the invaders Factors that have been considered to pro- Of the fourteen molluscs (twelve alien mote the influx of alien species include and two Mediterranean) recently incorpo- (1) a local fauna with relatively low species rated in the Tunisian fauna, five bivalves diversity (‘oligospecific’) and (2) the area are filter-feeders, five gastropods are gen- being repeatedly furnished with introduced eralist grazers scrubbing off detritus or mi- individuals. Both factors are relevant in Tu- crolgae from the substrate. Therefore the nisia. In the Gulf of Gabès, the number of Medit. Mar. Sci., 12/1, 2011, 53-73 67 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | species is relatively low even if abundance to become established until the environ- of particular species may be extreme, and ment met their own requirements. In this in this respect the Gulf very much resem- scenario, the littoral of Tunis could be a bles the large Mediterranean lagoons which small scale model for the entire Mediter- are a haven for alien species. On the oth- ranean basin, because of its transient role. er hand, the highly impacted area of Tu- The first event would be the arrival of new nis harbour, with important international tropical species, most of them from the In- shipping activity, may be a major promot- do-Pacific area, on the southern coast and er for new and unexpected introductions in- some from the Atlantic area on the north- to the Mediterranean. ern coast. The second consequence would Two major pathways must therefore be be the spread of the Indo-pacific immigrants watched for in the future. One of them is towards the northern coast. The rate of im- the progress of Lessepsian migrants out migration and the speed of spread along of the ca. 1800 km long blackout of the the Tunisian littoral would indicate the de- Libyan coast, with the casual Cellana rota gree of ‘tropicalization’ of the Mediter- and Chromodoris quadricolor as forerun- ranean as a whole. ners. The other one is the occurrence of The next item on the agenda is not on- alien species in the Tunis harbour and la- ly to make counts of newcomers. We now goon. Introduction through aquaculture need an appraisal of the share that the nat- seems less important than in other parts of uralized species take of the marine envi- the Mediterranean. ronment, and of the extent to which there The role of the increase of sea surface is competition with local species. This will temperatures in the acceleration of the require baseline studies to know which species process should not be underestimated. In- are dominant or otherwise important in tra-Mediterranean movements were not communities unaffected by alien biota, and considered in the CIESM Atlas but there time series in communities where the alien is a growing body of evidence that some species have become incorporated. tropical species with a historical Mediter- ranean range, including molluscs, are ex- Acknowledgements panding their range inside the Mediter- ranean. The cases of Echinolittorina punc- Photographs of living animals from Djer- tata and Siphonaria pectinata were discussed ba are part of the collection in the Muséum at length by ANTIT et al. (2007) but an- National d’Histoire Naturelle, Paris and other species in progress is Eastonia rugosa, were taken by Claude Huyghens and a native element of the Tunisian fauna since Françoise Danrigal during the 1982 expe- PALLARY (1914) now found as far north dition. The authors are grateful to Mr. as the coast of Latium in Italy (ALBANO, Pasquale Micali for sending juveniles of the 2006), and Valencia in Spain (TAMAYO different Mediterranean species of Anadara GOYA, 2008). The role of temperature for comparison to Dr. Rudo von Cosel for change is obvious when it results in range advice on some West African bivalves, to extensions of Mediterranean species, but Dr. Argyro Zenetos for help with some ‘gray’ it may be a factor in the success of many literature references and thoughtful sug- alien thermophilic species, which may have gestions on the draft and to Fabio Crocetta had the opportunity in the past and failed for some corrections and for calling our at- 68 Medit. Mar. Sci., 12/1, 2011, 53-73 http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 03/08/2019 23:28:34 | tention on an overlooked reference. The cal Atlantic becomes an established collecting in Bay of Tunis would not have alien in the Mediterranean. 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