Cah. Biol. Mar. (2007) 48 : 327-337

The planktonic copepod communities from the southern (, ) with a re-description of Paracalanus indicus Wolfenden 1905 (Copepoda: Calanoida)

Meriem KHELIFI-TOUHAMI1*, Rachid SEMROUD2, El Hadi HAMDI3, Makhlouf OUNISSI1, Ahcène HARIDI1, Mohamed NÉJIB DALY3 and Patricia AISSA3 (1) Faculty of Sciences, University of Annaba, BP 12 , Annaba, Algeria * Corresponding author: Fax: 213 38 871 062. E-mail: [email protected] (2) ISMAL, Dely Brahim, Alger, Algeria (3) Faculty of Sciences, University of Carthage 7 novembre, , Tunisia

Abstract: The composition, diversity and ecological affinities of pelagic copepods from the eastern Algerian coast and the contiguous Tunisian littoral (Bizerte shelf) were studied. The synthesis of the results showed peculiarities of the communi- ties inhabiting these shelf waters. One hundred forty three copepod species were identified from which Copilia lata Giesbrecht 1891 is recorded for the first time in the Mediterranean Sea. The copepods of the continental shelf of eastern Algeria seemed to be abundant and highly diversified. Ecological affinities revealed the influence of local and regional oceanographic conditions. Paracalanus indicus Wolfenden 1905 was recorded for the first time in the Algerian and Tunisian coastal waters. Because of the close morphological characteristics of P. indicus and P. quasimodo Bowman 1971 inhabiting the Atlantic Ocean, a compared study to distinguish between both species was undertaken. Despite its abundance in the Algero-Tunisian coast, representing 46% of total copepods, P. indicus was not found in the Mediterranean oriental basin or the north occidental part of the Mediterranean Sea. Although with careful morphological study, it is very difficult to distinguish between the two species.

Résumé : Les peuplements de copépodes planctoniques du sud méditerranéen (Algérie, Tunisie) avec une redescription de Paracalanus indicus Wolfenden 1905 (Copepoda: Calanoida). La composition, la diversité et les affinités écologiques des copépodes pélagiques de la côte est algérienne et du littoral tunisien (Bizerte) voisin ont été explorées. La synthèse des résultats a montré que ces eaux côtières étaient peuplées de communautés caractéristiques. Cent quarante trois espèces de copépodes ont été identifiées dont Copilia lata Giesbrecht 1891 est observée pour la première fois en Méditerranée. Les peuplements de copépodes du plateau continental de l’est algérien apparaissent riches et très diversifiés. Leurs affinités écologiques révèlent l’influence des conditions océanographiques locales et régionales. Paracalanus indicus Wolfenden 1905 est pour la première fois observé dans les eaux côtières algériennes et tunisiennes. En raison des caractères morphologiques peu distinctifs entre P. indicus et Paracalanus quasimodo Bowman 1971 peuplant l’Atlantique, une étude comparative pour différencier les deux espèces a été réalisée. Malgré sa forte abondance dans les eaux algéro-tunisiennes, représentant jusqu’à 46% des copépodes totaux, P. indicus n’a été rencontré ni dans le bassin oriental ni dans la partie nord du bassin occidental de la Méditerranée. Une étude morphologique minutieuse ne suffit pas à distinguer les deux espèces.

Keywords: Copepods l Paracalanus indicus l Zooplankton l Coastal Waters l Mediterranean Sea

Reçu le 19 février 2007 ; accepté après révision le 15 octobre 2007. Received 19 February 2007; accepted in revised form 15 October 2007. 328 COPEPODS FROM COASTAL SOUTHERN MEDITERRANEAN SEA

Introduction l : 18 and 27 May 1996 l Annaba: February, March and August 2002; March Research on marine zooplankton and pelagic copepods has and April 2003 become rare in Algerian coasts, since the intensive and l Bizerte: January and August 2002 (only Paracalanus famous works of Rose in the beginning of the last century indicus population have been considered here). and those of Bernard in 1950-55. Since that time, only three Thirty five vertical tows were realized from bottom to articles had considered copepods (Seguin, 1972; Ounissi et surface between 12 and 110 m depth in 10 stations (Fig. 1) al., 1998; Ounissi & Khélifi-Touhami, 1999). Therefore, the using a WP2 net in Annaba, a Japanese net (30 cm opening Algerian coasts are subject to interesting oceanographic diameter, 100 µm mesh size) in El Kala and a plankton net phenomena (penetration, crossing and accumulation of (20 cm opening, 200 µm mesh size) in Bizerte. Using so Atlantic water) and the Mediterranean is changing in many different mesh sizes introduces in fact some errors in ways (Béthoux et al., 2002). As indicators of environment, comparing regional copepod densities. This should be the study of copepods may allow to explain some local or taken into account for any plankton quantitative studies. regional oceanographic situations. Tropicalisation of the Simultaneously, temperature and salinity measures were Mediterranean is demonstrated by transportation of taken at the surface and at 50 m depth (stations 4 to 7) using copepods by the Atlantic water (Razouls & Durand, 1991) a thermosalinometer Kent EIL 5005. In stations 1 to 7, and particularly from the and Indian Ocean (Lakkis, current velocity and direction were taken at surface and at 1984 & 1990; Lakkis et al., 2002). The synthesis works from 50 m depth using a currentmeter T.D.CM2. the 25 past years on the pelagic copepod classification from All the samples were preserved in a 4% formaldehyde the Mediterranean (Kovalev & Shmeleva, 1982; Gaudy, solution and zooplankton analysis was realized on four 1985, Estrada et al., 1987; Razouls & Durand, 1991, fractions each one representing 1/20 to 1/2 of the sampled Razouls, 1995; Razouls et al., 2005-2007) revealed the volume, according to the plankton density. The species allochtonous intrusions of Atlantic and Indian species. In the identification however concerned the whole sample. Mediterranean Sea, the paracalanidae family which counts 7 Abundance was expressed in individual per m3. species has been revised many times since the works of Rose The description of Paracalanus indicus was established (1930). In fact, several authors agree that some difficulties in considering the morphological differences existing between the identification arise within the parvus group (Bowman, the Algerian and Tunisian specimens (more than 10,000 1971; Bradford, 1978; Kang, 1996). Bowman (1971) has individuals from Algerian and Tunisian samples) and the demonstrated that some specimens from several regions literature data mentioned above. The measure of identified as Paracalanus parvus (Claus, 1863), were Paracalanus indicus length was made on more than 30 actually Paracalanus indicus Wolfenden 1905 female and 27 male specimens. (Mediterranean Sea, Atlantic and Indian oceans) or Paracalanus quasimodo Bowman 1971 (S-W Atlantic). The confusion within the parvus group still exists and is exten- Results sively treated at a genetic level (Bucklin et al., in prep.). Moreover, the species Paracalanus indicus has been observed in eastern Algerian coasts in high densities Hydrology (Ounissi & Khelifi-Touhami, 1999). The confusion In the Gulf of Annaba, surface salinity values ranged from previously existing between the species Paracalanus 36.5 to 37.6 and temperature varied from 14.0°C in winter quasimodo and P. indicus has been removed by F. Ferrari, (February) to 28.0°C in summer (August). In May 1996, C. Razouls (pers. com. 1999; see also Ounissi et al., 1998). salinity values in El Kala varied between 36.7 at the surface In this paper, we give a broad synthesis on the diversity and 37.2 in deep waters. The temperature reached 21.4°C at and the main ecological characteristics of copepods from the surface and 15.0°C in the depths. The Bay of Bizerte the eastern Algerian coast and we give a short re- had a salinity ranging between 31.0 and 36.3 during winter description of Paracalanus indicus, focusing on the (January) and 37.7 during summer (August). The tempera- distinction between this species and Paracalanus ture varied from 13.0 in winter to 28.5°C in summer. quasimodo inhabiting the Atlantic Ocean. External waters moved eastward (Fig. 1), as a branch of modified Atlantic water, with velocities varying between 10 Material and Methods cm.s-1 and 25 cm.s-1. The residual current changed direction becoming southward in inner waters. This circulation Coastal zooplankton samples were taken during several allows some renewing of bays and inner waters particular- cruises as follow: ly in winter and spring. M. KHELIFI-TOUHAMI, R. SEMROUD, E.H. HAMDI, M. OUNISSI, A. HARIDI, M. NÉJIB DALY, P. AISSA 329

Figure 1. Map of the sampled areas: Gulf of Annaba, El Kala shelf and the bay of Bizerte, with the depth of the sampled stations. Arrows indicate the major current directions. Figure 1. Carte représentant les zones échantillonnées : Golfe d’Annaba, plateau d’El Kala et baie de Bizerte, avec la profondeur des stations étudiées. Les flèches représentent la direction du courant.

Abundance and richness of copepods major (Dahl, 1894), Oncaea conifera Giesbrecht, 1891, Copilia mediterranea (Claus, 1863), C. mirabilis Dana, Among 400 zooplankton taxa, 143 copepods were recorded 1849, C. quadrata Dana, 1849, C. vitrea (Haeckel, 1864), in eastern Algeria. The major fraction of copepods Lubbockia aculeate Giesbrecht, 1891. comprised rare species and only 25 were regularly present Moreover, it is interesting to point out the presence of in the samples (Table 1). However a big taxonomic richness Atlantic water indicator species, which occur in this geo- was observed in the whole region: 123 species in Annaba, graphical area: Acartia danae Giesbrecht, 1889, 58 species off El Kala. The main components of copepods Centropages violaceus (Claus, 1863), C. bradyi Wheeler, showed neritic affinity: Clausocalanus spp, Paracalanus 1901, C. gracilis (Dana, 1849), Ctenocalanus vanus indicus, Acartia spp, Centropages kroyeri Giesbrecht, Giesbrecht, 1888, Paracalanus indicus, Pleuromamma 1892, Centropages typicus Kröyer, 1849, Corycaeus abdominalis, P. xiphias, Isias clavipes Boeck, 1864 and typicus (Kröyer, 1849), Euterpina acutifrons (Dana, 1847), Copilia lata. Farranula rostrata (Claus, 1863), Oithona nana In addition, Copilia lata was recorded for the first time Giesbrecht, 1892, Oithona helgolandica Claus, 1863, in the Mediterranean. Only females have been collected; Oithona setigera (Dana, 1849), Oncaea venusta Philippi, the males have never been observed. Low densities were 1843, Temora stylifera (Dana, 1849) (Table 1). recorded in the gulf of Annaba: 0.1 to 0.4 ind.m-3 while in Distribution, affinities and abundance of copepods are El Kala continental shelf (Table 1) Copilia lata was not rare given in Table 1 in which it can be seen that the prospected presenting densities ranging from 5 to 9 ind.m–3. area shelters dense and much diversified copepod popula- tions of various affinities: 103 oceanic, 34 deep, 20 Atlantic Abundance and re-description of Paracalanus indicus affinity species, 17 neritic and over 15 coastal taxa. Typical inner water species were found few in the prospected zone High densities were recorded in El Kala continental shelf: but develop dense populations: Acartia clausi Giesbrecht, up to 899 ind.m-3 while Annaba and Bizerte coasts showed 1889, Acartia discaudata mediterranea Steuer, 1929, P. lower and similar densities: 1.17 to 72.7 ind.m-3 (Table 2). indicus, E. acutifrons, O. nana, C. kröyeri. Oceanic affini- In Tunisian waters where the species was recorded for ty populations were quite frequent: Clausocalanus spp., the first time, densities decreased from the inner waters to Calocalanus spp, Calanus helgolandicus (Claus, 1863), the lagoon. Even though Paracalanus indicus is known as Mecynocera clausi Thompson, 1888, Corycaeus spp. In a neritic species, it was absent in the polluted inner waters. addition, the bathypelagic and mesopelagic species were As described in Bowman (1971), Björenberg (1981) and very numerous but weakly abundant: Pleuromamma Bradford et al. (1999) keys and as it was observed in our abdominalis (Lubbock, 1856), P. xiphias (Giesbrecht, study, the genus Paracalanus is characterized for the 1889), Eucalanus hyalinus (Claus, 1886), Candacia spp, female by: Heterorhabdus papilliger (Claus, 1863), Heterostylites (1) First leg basipod with inner marginal seta, 330 COPEPODS FROM COASTAL SOUTHERN MEDITERRANEAN SEA 1 . 9 1 1 2 2 1 - s ...... 0 2 7 2 8 5 s . . . . e f 2 3 1 4 6 4 i D 2 l 4 u r 5 7 7 4 4 1 6 1 5 8 e S 3 e q t n 6 i 8 1 4 h . . . t 7 5 2 2 c 9 a s . . . . s 5 3 3 4 2 2 7 7 9 7 a e i ...... 9 6 9 8 1 r r 0 3 8 2 9 a 2 7 6 4 9 2 0 1 a l 3 9 4 9 . . 8 é 5 2 4 M . t 2 h 9 8 6 6 a y . 4 1 2 9 c . 1 c . . . 4 4 0 5 3 3 a a 2 9 K 3 . . l 2 5 8 4 9 e r 9 1 9 8 7 0 a l a 1 1 1 2 M g - - 1 1 - c c 1 - - - . 9 - E n i - - - . - 6 2 5 5 - + + + + s a g . . 0 3 5 4 2 . 9 3 1 2 . . e . o 4 5 . . . 7 7 6 3 R l . l 5 1 1 2 1 1 4 3 7 a 4 1 o 6 6 p 2 4 c i e c 7 . 2 3 6 8 5 8 r n ...... i D 3 2 o r 2 0 3 0 6 7 j S 2 3 p a n 0 9 s a . m 3 6 5 4 3 3 3 3 9 4 4 9 0 r ...... e 3 8 2 r u 0 1 0 0 3 0 0 0 0 3 8 3 7 3 i 7 3 1 e l M . 7 7 0 5 e l i 6 3 . . . . 2 . 3 r . h t . 6 3 e t 1 8 1 p 2 e 4 8 1 5 g 7 7 d - – – e - A – 1 1 n – - - 1 0 . . n n . . 2 + a 7 7 9 3 a 1 1 3 2 n 2 8 1 1 . . 6 . . 6 3 R . t 3 e . . 5 . 1 i 1 0 s 0 0 0 r 2 a é o g 5 3 6 3 8 1 c l D ...... a 2 0 3 0 4 3 S n 3 t n a 0 s i a . r 9 7 8 3 5 4 1 4 9 4 0 4 0 e ...... e e e 2 c e 8 0 1 0 1 0 3 1 1 7 8 0 g t M 6 n l 9 h ô . e 8 6 3 . c A c . 8 3 s 7 7 4 r . . 2 4 2 é 4 e 1 . a a n . 1 2 2 r . – l 1 r 7 7 g 4 5 - - p 1 – - e - M n e 7 1 1 3 2 t : . . + + – 5 7 a s d – 8 3 9 - . 7 . 1 1 . . a . 1 + s 0 0 R . 0 1 2 1 e 0 3 e , 1 1 e d e u o h a t 4 4 6 q p D . . . i b é t 2 1 7 1 S m a p n n 0 o 0 o a n a 0 r . l 3 3 1 4 7 5 3 4 0 3 4 1 1 c ...... e t f n 2 0 s 6 0 2 1 0 0 0 0 9 0 0 0 2 s t 1 A M e A 2 s d 6 é d . f 4 u 6 t o . . i ) o 4 g p e 0 3 n . f 1 e u i D g 5 5 7 8 5 1 5 1 e l f p . – 5 2 2 3 n S c f – 7 2 1 0 7 7 A u . . . . - ( . . . . . 7 . o a + a n 1 1 1 0 7 c 0 6 4 2 0 2 0 2 . 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KHELIFI-TOUHAMI, R. SEMROUD, E.H. HAMDI, M. OUNISSI, A. HARIDI, M. NÉJIB DALY, P. AISSA 331 5 7 7 . . . 8 5 1 9 7 2 2 ...... 0 2 0 2 0 3 0 8 7 5 2 4 3 1 8 2 6 4 2 . . . . . 4 6 2 6 2 . . . . . 7 6 4 9 1 1 9 5 0 2 6 9 2 7 4 1 1 1 7 . . 1 9 8 7 6 . . . . 6 5 4 5 0 9 . . . . 7 1 9 1 3 1 2 8 4 9 1 3 1 1 4 - 1 - - - - 4 1 - - - - - . 7 + + + + + + + + 1 6 . - 2 5 9 3 . . 4 7 2 5 . . . 7 . . . . 3 5 8 1 4 4 1 1 5 8 6 2 1 4 5 1 8 1 . . . . . 0 0 0 5 1 5 7 1 3 3 6 1 8 0 7 ...... 0 0 0 0 0 0 1 6 1 0 8 7 . . 2 3 2 . . . 2 2 1 1 0 1 7 - - - - 2 3 3 7 - 0 . . . . . 7 1 2 4 2 7 4 1 1 7 1 . . . . . 0 0 0 0 1 2 8 . . 1 7 8 . 4 4 4 4 3 6 3 2 3 4 3 ...... 1 0 0 0 0 1 0 0 1 0 0 0 1 8 8 . . 8 2 1 7 7 7 9 2 9 9 5 2 9 – - 1 1 1 ...... + 0 1 0 3 0 1 0 2 1 1 1 5 . . 1 3 6 2 . . 0 1 1 1 9 1 2 2 . , . . . . 0 0 0 1 2 0 1 2 . . 2 4 5 5 - - 7 5 2 2 . . . , + 2 0 2 5 0 0 . . 1 2 9 7 9 7 5 7 8 6 ...... 0 0 0 0 0 1 2 2 6 4 5 5 3 8 9 2 8 9 3 8 2 1 . . . , ...... 0 0 1 0 0 0 0 0 0 1 6 2 0 2 8 . 3 . 7 3 3 0 9 0 3 ...... 1 2 2 2 2 6 6 1 1 ------3 0 7 5 7 0 . . . . . , 2 + 3 2 1 0 7 0 3 0 3 3 0 3 2 . 1 ...... 1 1 1 1 1 1 2 0 1 6 7 . . 9 2 5 6 3 1 9 1 7 9 1 0 ...... 0 2 1 0 1 5 9 1 0 1 6 . . 8 3 1 1 5 0 2 5 6 5 5 - - ...... + + + + + + + + + + + + + + 2 9 0 4 3 4 4 6 3 . . 4 7 ) ) S S S S S S S 2 ) S D D D D D D D D ) 5 c ) S S S S S S S S S S S S S S S S S S S S S S D D D D D D 9 7 o 3 A S S S S A A A S S S A S A A 9 0 ) c o o o o o o o o o o o o o o o o o o o n A n o o o o o o 1 9 8 8 5 6 n ) o o o o o o o o o o o o o o o 8 4 0 9 ) ) 4 9 6 4 8 1 8 3 8 1 1 6 ) 9 4 3 4 & 8 8 ) 2 9 3 8 1 , 1 ) 3 ) , 9 t 8 1 8 8 9 1 8 t 1 t 9 8 ) 1 , 3 8 ) ) , 6 7 ) ) 8 1 h 8 , 1 8 8 s , 4 9 1 9 h 1 s , 6 ) 5 6 3 1 k , 6 3 t 8 4 c 1 s , 8 t , 1 o a c 1 8 4 6 4 7 , c 0 ) u , 8 8 r 4 , 6 3 6 r 1 8 h k e h l 8 a ) , , n 1 8 a , e 1 n 8 a a o 8 4 9 1 8 9 r t 9 6 8 c l 8 , 1 c F h r 4 3 l 1 1 n c 9 y n 9 , 1 4 1 1 9 1 8 b 1 h b e , 1 8 a 4 s S ) 1 e 8 1 h , b e e 6 6 a a 9 n r 9 s d r 4 ( a C s , s 4 9 9 , b 4 , 1 , r , , 1 r c s 8 9 a t u r , 1 o , 4 ( a a 8 d r 8 8 e n a 8 4 s n 8 D 8 1 b e s u 8 , u a s s e b y D a a h 1 s , i 2 r n , 6 e r 1 n r 1 i e 1 s l B 8 r a 1 l D . 1 u a 1 n a s D 1 , d s u u u F a , L c 9 ( a s a 8 y l b t e e , g B , a h , , a 1 . e G , n ( , n a F , t C a a e a l i G t D u 1 i ( s ( l s 1 ö u r s c a a g S t C r r ( l , ( D l s ( a F a a a l i a e a t n i a t ( D h e e ( u ( , i r , a ( C t G n b e n r i G n a c l l a l C n 8 d u C n o k B s s c n a D a s ( a i p r l D t l k e s r s ( a i s h ( s a s K p n l i r i c 8 e c ( l a C o c . a a n l e s e G ( l i e s g e c a g a u s c t a u P n a n i s r e o p i i 8 u m S D t a u n D r e n ( a r C e t F o r F e r D s u j r a s D u s e c s u s t a b t t t o t a w ( 1 o r D n a e n a a e r v r i i o a s o a . G o e a r s s n s u f s u i a a a u u e f a a s i a p r l r o s , i s c g i a s i e i p p f i t b t 8 H T c e i u r l s r a c c l r s B i n r a l l t c c S s o l m c b ( l t i o n i t t u i v s n t t o i l c n 6 s s a i c u s s g i ( i o r s r e a a e u u e . a s r u e g n r s r t r a r o p o u c t s b u u s 9 o v m a u u G a g i v s y b o l c c p s p l i r s o u l u a a i n a a a y w a c d o e d d c u 1 a a e e e n n a a o s s t e h l n a a f f l l l a g c t a r r c r d p c u r o a l a r r l l i l a i b b t t n a , p m h t a a f l l l i r t e s s s s s s s s s s s s s d s l s s i 1 e l l s s u r i a a r a a a a a m e e e a a t m l y i l l u u u u u u u u u u u u u u l u u u e v t t a a a a e a a e e l m m q v l h h t c o e e t t l i i n k e e e e e e e e e e e e e u u a c c c g e e a n n n n n r r s l i i e t t a o h s e e c l t a a a a a s s i a a a a a a a a a a a a a c n n r o n a a o o b u u s i i i i i o o o p o c n m m l l a a i o i c p T c c c c c c c c c c c c c s s o a a r r r o o l l l l l d r r a c c y u e e i i i i i r r a a h h h y y y y y y y y y y y y y i b a o s i i r r . i e e e u u n t t e m c c l T p p p p p r r r r r r r r r r r r r l t t t c c c c c c c t r b b r r a c c e a a y y e a e i i i C e a e e e i o o o o o o o o o o o o o o o o o o u u u u u u a a t l l l l a u u u l . s C C C C C C C C C C C C C C C C C C C C C C C D E E E E E E F F H H H H I L L L L M M M M M I F 332 COPEPODS FROM COASTAL SOUTHERN MEDITERRANEAN SEA 6 1 8 . . . 9 6 . . 4 1 2 5 7 . 3 6 3 7 3 6 2 0 6 3 1 1 3 6 3 5 . . . 1 9 4 . . 1 4 9 4 5 7 . . . 1 3 8 1 4 1 7 4 2 2 6 4 9 5 4 4 9 4 3 . . . 9 3 5 3 7 3 . 9 1 5 2 9 5 0 8 . 6 6 6 1 5 8 6 - - 6 - - - - - . - 2 9 . 8 6 . 5 6 2 + + + 8 5 . . . . 4 7 . . 1 8 . 1 5 8 9 0 1 9 2 5 2 9 6 2 1 3 2 2 7 1 . . 6 1 3 5 . . . . 1 6 8 0 5 1 2 1 6 4 8 . . . 3 6 1 6 . . . . 3 7 2 0 3 2 0 2 2 1 3 3 5 3 . . 1 . . 3 . 2 7 . 0 4 7 , , 1 1 3 5 - 1 - 1 - - . - - 7 1 2 2 8 1 . 4 9 . . . . . 1 1 0 1 0 1 2 4 7 5 9 . . . . 1 . 2 0 4 9 0 5 1 1 3 1 7 7 . 7 5 6 2 . . . . 1 3 6 3 . . . 0 8 0 8 4 6 0 1 7 5 1 4 1 2 . 3 9 7 1 . . . 8 4 3 . 2 4 1 . 7 2 3 2 9 7 2 1 1 3 - - 9 7 - - . . - - 5 8 0 4 2 . . 6 . 3 5 . 2 6 . . 6 5 5 9 3 1 1 3 . 2 8 . . 5 2 2 0 3 8 . 2 6 6 1 4 . . . . . 7 4 0 3 0 0 5 2 7 . 5 2 3 6 . . . 5 6 6 8 - - 2 7 2 – – . . . + 8 1 0 1 2 2 4 . . . . 6 7 1 2 1 5 7 . . 6 5 5 2 7 . . . . . 8 2 6 2 0 6 7 1 4 2 8 5 7 . . . 2 6 2 9 6 9 3 3 2 3 5 5 ...... 1 7 6 1 2 2 0 3 0 0 5 0 0 1 0 1 1 4 9 . 4 3 1 6 . . . 9 0 3 0 . . . 8 1 1 4 4 6 6 6 1 9 1 - 4 1 - - - - 7 8 3 5 7 - - - 0 0 . . . . . 9 . . + 2 . 3 1 2 0 0 8 0 4 . 2 5 8 3 0 . . . 3 . . . 0 2 5 3 1 3 2 6 3 3 . 4 9 6 . . . 3 3 6 4 4 8 8 . . 3 4 5 . . . 8 5 6 2 4 1 4 5 7 . . 9 6 . . 6 6 0 3 0 2 1 1 1 - 1 - - . - + + + + + + + + + + + + + + + + + + + + + + + + 4 9 ) . 5 6 5 . . 6 . 2 4 4 5 4 1 ) ) 8 9 3 8 1 ) 8 6 S S 8 , D D D S S D S 3 5 8 8 S S S S S S S S S S S S S S S S D D D D D D D D D D 8 c k 1 o n S S S A A S o o o o o o S ) 9 D D 1 0 9 c c c c c 1 o o o o o o n n n n n ) o o o o o o o o o o c 1 o c c 9 o o o o o n 3 ) 8 9 ) 2 3 , , o , ) 8 ) t 3 t 6 ) t 1 9 9 1 6 9 1 b 8 3 3 ) 1 h 3 1 1 7 6 1 h 8 9 h , , 4 1 ) 8 2 2 9 b 0 6 6 l 2 9 ) 6 9 9 c c 7 , 9 8 1 c 4 8 8 , n 1 1 t 9 9 3 8 u 9 8 8 e h 9 4 8 l 8 8 9 3 2 1 e 8 , 8 e 8 e r 1 ) 9 3 l 3 , h 3 1 8 r 6 1 a 1 1 1 7 8 8 r s 1 1 1 L 1 1 8 9 , 1 d s 8 e , b 9 c 8 6 4 8 b 4 ( 1 7 , 8 b , , s , 0 1 1 , , , u , 8 8 , , , D s 1 n s t u 0 4 a t 1 s e 8 8 8 s s 8 n t 6 l , 7 n 1 s t 9 a , s , u w 1 1 e 8 r l e t e . a a h , n e 8 t 9 h 1 0 h 1 u u i l h i 1 6 , u 9 , e i a a f 1 a a l i l b a s , 1 n , t 6 a h F c c l r 1 h 1 a a l r 9 , , s c t a , 8 c C 1 s n , S r ( a h a c h r 9 u y C l l e G G 1 i s c l ( G o 6 , e , e 1 u d e h C 1 a e , D ( r a ( s s ( a r n n d r c 1 r e 9 r p i s s ( ( C C 1 e u r a n a C c D , i s s , b i n r x F i e b a b F ( r s a , b ( d l u ( D a 1 a p x W a i l n 9 s a n r e s r a b G s o u s l a ( t o s i b i a l s m a e s r e , a r e t C C S i 1 s s l r e a a b u a e u i s r r l e r a s c e k e i s B i ( ( B o s i s u b l C i B n s r c i t a e e o i a i s , e a i a e l h u e i d n s D r r s i p h u i d u r i i l n u a l e e a d s ( n s G c s a S n a G a a u p G l u F a i H e i i a G v i . r r i P v o i b c m r o m t n i t C a r i G r n r G r n c S i r a r F i u n r s p a s r a c u a b s d a x g p a G a e a r o e n g a i i a m s s c a a e r s e a s t t a r r G f i l S f h n o l u n t a r a i e c i d r a o t i r e e a a a a a a n a d s e t p s i s l a t g l i i g s o p c u a a a p t l a f i c a a u n l i i n a g i u . g i u i u v i i t l d . c e u g m t i t s s s s s o c s e s i m m m m m e a a d d n g u c l b b e e p n t t n n r u g s r e t b s p n l 1 u u u u u i p n m u u i n n n a l e e e l r a m m m m m p i o a a e b o o o i s o x e n u u p n n n n n u u o e m l n n l e o l n p r s s s b h e a a a a a d h c o r s b e m m n e s t n t v c m a a a a a l p a a a a h n c l l l l l a a a e l l a a a a a a a a a t b c m m m m m l l a a a a a a a a a a a a a a a n s i l s a a a a a a u a a l l n n n n n n n n n a o e e e e e e e e e e e e e e e o o o o o a n o c c c c c o c e o c c e e r r r r r o o o o o o o o o T l a a a a a a a a a a a a a a a n e c t t h a a a a a a a a n n u u u u u h h h h h h h h h c c c c c c c c c c c c c c c n u o i n n i c a r r r r r r r r t t t t t t t t t e e e e e c i i i i i i i i i n n n n n n n n n n n n n n n a e a a a a a a a a a h o o h h l l l l l N N O O O O O O O O O O O O O O O O O O O O O O O O O P P P P P P P P P P P P P P P P P R R 2

M. KHELIFI-TOUHAMI, R. SEMROUD, E.H.2 HAMDI, M. OUNISSI, A. HARIDI, M. NÉJIB DALY, P. AISSA 333 . 6 1 8 . . 7 3 7 9 , 2 4 0 2 5 6 9 8 7 6 6 3 3 . . . - 3 , - 1 0 0 0 2 4 2 3 . 9 2 6 6 9 3 . . . . 1 . 4 8 2 4 2 2 , 5 8 1 . 3 0 9 3 . . 7 2 6 0 2 7 3 . 2 – – 1 1 2 7 . 7 . . 0 9 4 0 . 2 8 5 9 . 4 . 2 4 . 3 5 6 7 - 8 . 9 3 3 – 9 . 4 8 . 4 ) 6 . 2 8 7 ) 3 0 8 . 9 8 2 ) 5 . 6 6 0 . 9 8 8 . 9 1 1 2 1 8 8 1 - - 8 2 3 2 9 ) , 8 8 1 t 8 . 6 9 2 1 9 8 1 1 . , . h 9 8 t 8 8 8 9 ) 9 1 9 9 , 1 c 3 . 8 t h 1 8 1 6 8 2 1 8 3 , a 6 . . 0 e t c . 8 h , 5 , 1 8 , 3 1 r d 2 t 6 t s c 8 . h 8 1 e

8 Figure 2. Paracalanus indicus. Male: lateral view. 3 3 , b , u - r . . 2 h c t 3 h t 1 e u , s 9 1 Figure 2. Paracalanus indicus. Mâle : vue latérale. b t a c r 0 0 c ) a h 1 e h 2 e 4 – 4 , s c b r . i h l c e 9 c e i 3 0 8 e k s 2 r b c r . . 6 i 2 C 4 e . e u

G (2) Third to forth leg exopod 3 external edge at least c s e . 1 b + b r 1 1 r e 7 9 ( i 9 0 8 s n e s o

r partly serrated and , G 9 a b . b i 1 i 4 4 7 1 t 8 ( e 6 e s b . a b s s G 2

. (3) Both leg 5 developed, symmetrical and 2-3 segmented. . i i s 8 2 s , a ( e u G e n b 2 6

1 The male (Fig. 2) has: 3 a l i 3 8 - i e 1 a G - t G h . . u a i s 0 s u 1 (1) A cephalic hump on the lateral view and a , n c a G 0 8 G t u s c n 2 L a D 2 t - a

a (2) The right leg 5 is 2-3 segmented. G ( i a s . ( a s e a 2 l n n a Paracalanus indicus female has: i t . s a n 8 t i 0 D o a i n e . e a y + + + + + + ( l

i (1) First antenna shorter than body and extending a 1 o 1 g n m a s t S n D d a r i 9 l d u beyond anal segment (Fig. 3), o D 1 l a o s a 9 o c m r n S S S S S S S S S S S S r D c A a A l a

e (2) Outer distal of exopod 3 of legs 2, 3 serrate and 4 r a 2 s r s a r a . . t l o o o o o o o o o o n n g o u a o i o e

i p naked (Fig. 4), u l s s 9 u c b d n r f n e i a m n o s i i u u 1 (3) Basipod 1 of legs 1-4 covered with developed orna- n l a c x x n n

a mentation (Fig. 4), , i i i y r a a a a a a a l r t a a r r r (4) A prosome three times longer than the urosome (Fig. g n n n n n n n l l e a s i i i i i i i h h h d 3), f e c a a t t t a r r r r r r r u i i i n a o i i i i i i i i

i (5) A dorsal hump slightly developed (Fig. 3) and c c o t r n c c c r h h h h h h h h o u u (6) Leg 5 symmetrical and 3 segmented (Fig. 5). h l c o e e e o t n p p p p p p p e e 1 l l l Paracalanus indicus male has: a t e n t l t h p p p p p b b p p m o o o b (1) Outer distal of exopod 3 of legs 2-4 serrate, a e a o e e a a a a a c c c u u a a T (2) The right leg 5 is 2 segmented and the left 5 S S S S S S S S S S S S T V X T L segmented (Fig. 5). 334 COPEPODS FROM COASTAL SOUTHERN MEDITERRANEAN SEA

Table 2. Abundance (ind m-3) of Paracalanus indicus in the sampled areas. Tableau 2. Abondance (ind m-3) de Paracalanus indicus dans la zone étudiée.

Station May February March August March April January August 1996 2002 2002 2002 2003 2003 2002 2002

1 (Annaba) 4.6 3.8 66.7 36.4 17.1 2 (Annaba) 9.9 6.0 12.8 21 3 (Annaba) 10.9 4.0 12 4 (El Kala) 899 5 (El Kala) 864.6 6 (El Kala) 493.6 7 (El Kala) 198.9 8 (Bizerte) 135.8 57.2 9 (Bizerte) 86.5 10 (Bizerte) 170.8 37.2

between the females of Paracalanus quasimodo and Paracalanus indicus in the legs 2-4 armature (Fig. 3) in spite of the serrated outer distal exopod 3 of leg 4 in P. quasimodo. Moreover there is no difference in the males as can be shown in Table 3.

Discussion

The pelagic copepods of the eastern Algerian coast were abundant and showed a big taxonomic richness in the sampling period. They showed large cohabitation of various affinities and origins (Atlantic, oceanic, bathy- pelagic and coastal water) indicating thus the opening of the region to the external water influences. From a quantitative point of view, copepod populations were largely dominated by neritic species. The majority of the 143 copepods collected were rare, comprising deep and Atlantic affinities species. In this context, the synthesis works on Mediterranean plankton biogeography (Furnestin, 1979; Gaudy, 1985; Moraitou-Apostolopoulou, 1985; Estrada et al., 1987; Scotto di Carlo et al., 1984 & 1991) demonstrated the occurrence of such contingents. Copilia lata, collected for the first time in our region, is also a new species for the Mediterranean. It is known to be an oceanic species, but it was largely present in neritic waters of the studied area and may be considered as a common component of the south-western Mediterranean Figure 3. Paracalanus indicus. Female: lateral view. a: dorsal pelagic copepods. This occurrence may indicate the hump. influence of external oceanic waters in the region, Figure 3. Paracalanus indicus. Femelle : vue latérale. a : bosse dorsale. advecting the modified Atlantic water (Ounissi & Khelifi- Touhami, 1999) to the coastal area. The presence of C. lata of south-western Mediterranean may also reflect the tropi- The female of Paracalanus indicus of the studied area calisation of the Sea. Many tropical copepod species had a body length of 660 to 990 µm (887 ± 98.1 µm, n = penetrate in the Mediterranean from the Red Sea or the 30). The male body length ranged from 880 to 1050 µm Gibraltar strait as a consequence of the general water (942.2 ± 43.3 µm, n = 27). Table 3 shows no differences warming (Lakkis et al., 2002). M. KHELIFI-TOUHAMI, R. SEMROUD, E.H. HAMDI, M. OUNISSI, A. HARIDI, M. NÉJIB DALY, P. AISSA 335

Figure 4. Paracalanus indicus. Female: L1-L4 leg 1 to 4. Figure 4. Paracalanus indicus. Femelle : L1-L4 pattes 1 à 4.

On the other hand, if the region is known to be largely the continental shelf of El Kala during late spring. In this influenced by the Atlantic water (Seguin, 1972; Furnestin, dry period temperature is above 21.0°C and salinity is up to 1979), the coastal copepod fauna is more similar to that of 36.7-37.2. Comparable distribution may be observed for the oriental Mediterranean basin than to the occidental tropical population in the Brazilian continental shelf off basin. This similarity may be expressed by the regular Sergipe and Alagoas (Araujo, 2006). In that area P. indicus occurrence of C. kroyeri, Acartia latisetosa, E. acutifrons was more abundant in the rainy months and even reached (Gaudy, 1985; Moraitou-Apostolopoulou, 1985; Lakkis, the maximum value of 2,433 ind.m-3. The author reported 1990). Being an integral part of the Sardinia channel, the that this species was recorded in salinity values from 25.8 studied area is subject to cyclonic eddies and modified to 39.3 and temperature values from 24.0 to 29.5ºC but Atlantic gyres (Astraldi et al., 1999; Millot, 1999; Testor et seems to avoid low salinity waters of the inner shelf. al., 2005) that allows supplying additional external and Despite its abundance in the Algero-Tunisian coast, deep species. In a local hydrodynamic context, the neritic representing up to 46% of total copepods, P. indicus was inner waters are influenced by eastward residual current not found in the Mediterranean oriental basin or the north (Ounissi et al., 1998) resulting from the general circulation occidental part of the Mediterranean Sea. Its absence in the of the modified Atlantic water stream. Consequently, the NE Atlantic (the contiguous potential supplying source, see high copepod richness found in the studied area is to be Greze et al., 1985 and Razouls et al., 2005-2007) is a sign correlated to such hydrodynamic situations. of the isolation in the Mediterranean. Besides, in view of Paracalanus indicus (Wolfenden, 1905) was recorded the taxonomic confusion in existing data, it would be for the first time in the Algerian and Tunisian coastal prudent to consider the similarities between P. indicus and waters. It can develop high densities (up to 899 ind.m-3) in P. quasimodo: genetic analyses may precise the state of 336 COPEPODS FROM COASTAL SOUTHERN MEDITERRANEAN SEA

Figure 5. Paracalanus indicus. a. & leg 5, lateral. b. ( leg 5, anterior. c. & posterior prosome and genital segment, lateral. Figure 5. Paracalanus indicus. a. & patte 5, vue latérale. b. ( patte 5, vue anterieure. c. & prosome posterieur et segment génital, latéral.

Table 3. Morphological comparison of Paracalanus parvus, P. indicus and P. quasimodo (Modified from Bradford, 1978). B1: basi- pod 1, Ex3: exopod 3, Gns: genital segment, L2-4: leg2-4. Authors 1: Bradford (1978), 2: Bowman (1971), 3: Kang (1996), 4: Sewell (1929), 5: Present study. Tableau 3. Comparaison morphologique de Paracalanus parvus, P. indicus et P. quasimodo (Modifié de Bradford, 1978). B1 : basipodite 1, Ex3 : exopodite 3, Gns : segment génital, L2-4 : leg2-4. Auteurs 1 : Bradford (1978), 2 : Bowman (1971), 3 : Kang (1996), 4 : Sewell (1929), 5 : Présente étude.

Species Outer distal edge of Ex3 serrate B1 of L2-L4 with many Gns with posterodorsal Authors L2 L3 L4 posterior surface spinules spinules in &

P. parvus & no no no no no 1, 2, 3 P. parvus ( ? ? ? no 2 P. indicus & yes yes no yes yes 2, 3, 4 yes yes no yes no 1, 5 Fig.4 P. indicus ( yes yes yes yes 2, 5 P. quasimodo & yes yes yes yes no 2 P. quasimodo ( yes yes yes yes 2

these species revealing possibly the resemblance of the two Acknowledgements populations. We also suggest a full reviewing of species from the parvus group (P. indicus, P. parvus, P. quasimodo) from This work was partly funded by the Ministry of the the Mediterranean focusing on the possible occurrence of Territory Management and Environment (CREAD 29.568). these species. The authors are very thankful for the strong effort spent by the two anonymous referees to improve the manuscript. M. KHELIFI-TOUHAMI, R. SEMROUD, E.H. HAMDI, M. OUNISSI, A. HARIDI, M. NÉJIB DALY, P. AISSA 337

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