ACTA ICHTHYOLOGICA ET PISCATORIA (2007) 37 (1): 7–15

DISTRIBUTION AND CHARACTERIZATION OF LESSEPSIAN MIGRANT FISHES ALONG THE COAST OF LIBYA

* Esmaile A. SHAKMAN and Ragnar KINZELBACH

Allgemeine und Spezielle Zoologie, Universität Rostock, Germany

Shakman E.A., Kinzelbach R. 2007. Distribution and characterization of Lessepsian migrant fishes along . the coast of Libya. Acta Ichthyol. Piscat. 37 (1): 7–15 Background. The Lessepsian migrant ichthyofauna along the approximately 2000 km of the Libyan coastline has been poorly known. A comprehensive study, addressing this issue, was needed and therefore the presently reported research project is intended to close the gap, through identifying the fish species and assessing their dis - tribution. Materials and Methods. The catches of commercial fisheries vessels were monitored and examined in a stan - dardized procedure between January 2005 and March 2006. The project covered a total of 4273 specimens, com - prising 1901 specimens of Siganus luridus , 1885 specimens of Siganus rivulatus , and 487 specimens of fourteen other Lessepsian fish species. Results. Sixteen Lessepsian fish species, representing 14 families, were recently found along the Libyan coast, two of which are considered to be first records for Libya: Herklotsichthys punctatus (Rüppell, 1837) and Liza carinata (Valenciennes, 1836). Approximately 50% of the immigrants were found along the entire stretch of the Libyan coast, 12.5% in the east and central regions, and 37.5% were restricted to the eastern part of the Libyan coast. All were found in the coastal area (1–50 m depth), 12.5% on the vegetation, 31.25% on sandy bottoms, 12.5% on rocks, while the majority of them (43.75%) were pelagic. Regarding the size, 75% were medium, 18.75% large, and 6.25% were categorized as small. Conclusions. The north African coast (west of the Nile delta)—compared with the Levantine- and Anatolian coast of the —was considered to be settled by immigrants from the Red Sea with some delay, due to sea currents. If so, this situation has changed meanwhile at least for fishes. More than 37% of the record - ed Lessepsian fish species are of commercial value, especially rabbitfish ( Siganus spp.). The future research should be focussed on monitoring the stocks, especially of the commercially valuable species, regarding the biol - ogy and ecology of the Lessepsian migrants. Also, possible further additions to the fish fauna, by new invaders, . should be recorded Keywords: biogeography, bioinvasions, Libya, Lessepsian migration, marine fish

INTRODUCTION of species, which have passed from the Mediterranean The term “Lessepsian migration” was coined by Por into the Red Sea; these are named anti-Lessepsian (1978) for the migration of organisms from the Red Sea migrants (Por 1978). into the eastern Mediterranean through the Suez Canal. The fish fauna of the Mediterranean has already Many species, which were able to adapt rapidly to the new undergone considerable change, as can be illustrated by environment, spread into the Mediterranean and estab - the following selected case studies. Papaconstantinou lished new populations. Information on the comparative (1990) reported that 11 species had reached the Aegean life histories of the immigrants is necessary (a) for an Islands (Dodecanese, Cyclades) through swimming along understanding of the selective mechanisms controlling the the coast of Anatolia. Twenty-two Lessepsian fish species passage through the Suez Canal, (b) for an assessment of live on the coasts of the eastern Mediterranean- and the adaptive changes of the newly established “neopopu - Aegean seas, with some of them becoming commercially lations”, and (c) for an evaluation of the extensive ecolog - important ( Torcu and Mater 2000). In 2002, thirty-three ical changes which invading species may produce in their Lessepsian fish species were documented on the new areas of distribution (Ben-Tuvia 1978). The number Anatolian coast (Bilecenoglu and Taşkavak 2002). For the of Lessepsian migrant species exceeds by far the number Egyptian coast, several authors have recorded Lessepsian

* Correspondence: , Mr. Esmaile A. Shakman, Allgemeine und Spezielle Zoologie, Universität Rostock, Universitätsplatz 2, D-18055 Rostock, Germany e-mail: [email protected] 8 and Shakman Kinzelbach fish species ( Ben -T uvia 1976, El-Sayed 1994). Two forsskali (Fourmanoir et Guézé, 1976); and Indian scad, Lessepsian fish species were recorded in Italy: Siganus Decapterus russelli (Rüppell, 1830) (cf. Bilecenoglu and luridus (Rüppell, 1829) appeared along the shallow Kaya 2006, Çinar et al. 2006, Corsini et al. 2006, Golani waters of the Pelagic Islands (Azzurro and Andaloro 2006, Golani and Sonin 2006). 2004), whilst Fistularia commersonii Rüppell, 1838 was Although it is clear that migrant Lessepsian fish recorded on the eastern coast of Lampedusa ( Azzurro et species have had an enormous impact on the eastern al. 2004) . Ktari and Boualal (1971) reported S. luridus for Mediterranean ecosystem, there has been no thorough the first time on the Tunisian coast. In 1974, S. luridus and study to assess this impact (Golani 2002). Many S. rivulatus Forsskål 1775 were recorded for the first time Lessepsian fish species have been recorded in Libyan in the Gulf of Gabes ( Ktari and Ktari 1974). After that, six waters (Stirn 1970, Zupanovic and El-Buni 1982, Indo-Pacific fish species were recorded in Tunisian waters Al-Hassan and El-Silini 1999, Ben-Abdallah et al. 2005, as newcomers ( Parexocoetus mento (Valenciennes, 1847); Shakman and Kinzelbach 2006, Shakman and Kinzelbach Pempheris vanicolensis Cuvier, 1831; Stephanolepis 2007). There is no comprehensive study of the Lessepsian diaspros Fraser-Brunner, 1940 ; S. luridus ; S. rivulatus ; marine species in this area, particularly of the fish species, and Priacanthus hamrur (Forsskål 1775) ) ( Bradai et al. and so the objectives of this paper are to present the dis - 2004). The bluespotted cornetfish, F. commersonii , was tribution and characterization of fish species along the also recorded along the Tunisian coast ( Ben-Souissi et al. almost 2000 km of Libyan coast and a general contribu - 2004). In the Adriatic Sea, Sphyraena pinguis Günther, tion to knowledge of exotic marine fish species in the 1874 was recorded in 2001 (Pallaoro and Dulčić 2001), Mediterranean Sea. and afterwards S. rivulatus was recorded for the first time in the same area ( Dulcic and Pallaoro 2004). The silver - MATERIALS AND METHODS stripe blaasop, Lagocephalus sceleratus (Gmelin, 1789), This study was carried out on samples collected has become an abundant species in the eastern between January 2005 and March 2006 along the Libyan Mediterranean, immediately after its first record (Akyol et coast at the depths of 1–50 m. The pelagic and benthic sam - al. 2005, Bilecenoglu et al. 2006). More recently, the fol - ples were collected with a trammel net (inner mesh 26 mm, lowing alien species were included to Mediterranean outer mesh 120 mm). The study area was divided to three ichthyofauna; Japanese threadfin bream, Nemipterus main regions according to topography and environment japonicus (Bloch, 1791); teira batfish, teira (east region, Sirt Gulf, west region). Two sites were (Forsskål, 1775); peacock wrasse, Iniistius pavo selected in the east region (Tubruk, Benghazi), one site in (Valenciennes, 1840); Red Sea goatfish, Parupeneus the Sirt Gulf (Musrata), and two sites in the west region

Fig. 1. Map of the Libyan coast, showing cities adjacent to the sampling sites 9 Lessepsian migrant fishes in Libya

80 70 60

e 50 g a t

n 40 e c r 30 e P 20 10 0 l l l l n c e e y y m a n i a a i r i o g k d u o g c r i i a c c m n r t r a r m a d l o s l n a e a e r e t e m s o m e p m o m n g c m m e o o v c C

Abundance Size Habitat Commercial value

Characterization

Fig. 2. Characterization of Lessepsian fish species along the Libyan coast

60

50

40 e g a t n

e 30 c r e

P 20

10

0 Along Libyan coast East and Sirt gulf East region

Distribution

Fig. 3. Distribution of Lessepsian fish species along the Libyan coast

East region 100 90 Sirt region 80 West region

e 70 g

a 60 t

n 50 e c

r 40 e

P 30 20 10 0 r i s s s a r u u ta i ii a n s s s s f o s d t a u n b o ro i u tu . p n ri la s g o a s p m s a H . e u u in s d r s a o t id lu v t p r je e a u n c U n . ri b . e d i q u n e S . o m . m d s c u r . S m . o la p c S S m A o S d . . . o c n H C c . u A . S . F S

Fig. 4. The distribution in percentage of each Lessepsian fish species along the Libyan coast 10 and Shakman Kinzelbach 1 e l b a T e e a y a e d l e e i e d i a a a i a e e h e d e d d h m a i a a d i i p t e a e t i a e e d d d n n a r n a e d i a a n i i e m F i e e e r a r a d d a d d o a n i a a i g c a i i h b i d r l d r r d l e i i r i a n i n n p l y y o u e r m p a l g n a a t m r h h n a h m o u u u o g g s e t l a i i p p y i p e c S S S S S C H F A C M S P M S M y l a l k a i u r h e s e p b o a m m b g a I a r r n A h a g u f s f a a n o f s l o i a r l l t M a A h y u a i e f a l l b S n i i a i a H a S , s a z z z r , L a , a y h h l d a m r w l r g g a r G h a a a h a a s s O h a , f f t a Y l a e , s d h s a K h r M M a a a e s a n t A e o , , M s a e K b o e o a a d b u y u s t t r c a a a l m o f a t t a o u m l h h k - n t r r i n o a a s s a e k o d r e m l t t l a a u m e o o i a b a a a a a u a r r a p a y b B B M M A S M G N S T S S B B H i L e h t f o ) s h e s ) i t fi ) c l ) e l h e e h e l r s p s d u e s i fi fi s k ) e m e r r l h c e i s e e a fi y i m ( g v p f e a l r m r e t i n o n e g e s h ( p a k h h s i w ( g i s s c k s e s h i n a a n ( l s fi i p d j m t i e a t g e r r v fi a e s b p s o e e n a o d r e g f n s t r r o S p h l r e d r E n ( e f t o a e a i b v u o d e a t z o r l L e a h h a c c i i e f r e n e s l e d a l r f i l s e w e d d r l s d r fi u o s p a n r h t h e d e d a u c i s t r c t n t a a e a o b a t r k p s o t s e b m o r - r d e i a s c o a o e r g o e p h l t t e a b p d w L l a y k s y u h s n s b k e o i b b m ’ k t l s c u e d e r r c i r n i s t r r e r o r u a a u d t a o u b l r l a a e a p h e e d m o r b s b b h s P k V k n r ) 0 0 0 4 8 ) 9 ) 1 7 ) 1 9 8 , 3 1 , 8 4 e r 8 0 9 8 8 e d 1 8 1 ) 1 è 3 1 n , 1 3 5 l , p 8 , n ) , l i e 8 7 1 n r u ) 6 e n r c 4 9 1 7 e , o 5 3 p a t l a ) 7 s 2 , 1 5 ) l B 7 l s 8 p r 9 - 8 d L 8 r l 7 5 o e 7 1 r e ü r ( 2 1 å 1 o i 7 7 p 1 e , a G 8 R , k , v 1 7 s p F s n l h r ( r t s 1 ( a e u 1 l å ü o c e e e s r s , i s å i n s k l r l h C R r u F a l s t k v n å u o R t i i e s s e s s ( e n u k s i i F a r v i s p s o t ü ( s o m n s r o c C u s i r c p n n o s o r n G F m T p n ü a s u e ( m n e - o F t l o s s r c u l e R a i n F c a r o e a s a a p ( ( i d e u s u l c a u s i i V m f s s d t q u g e B a ( s u n t n y s u a n s s m r b i e u l b i m a h i o d a u r r r o o t a i t u a o p v p d c h c o o r h d v a n e m n s i a a p p c l u e s i n a m o i l r u j c i n n i o r n s r s m o r d r e e s fi s t e e n a e u a i n a a a a u u o s h d t i a d r e b l l c i r r i i r e n n p n h n u k y y r n m e p a e a t a e p r m m e i h h u o s h e z g g e e e p r i t l e c i i i p p a c t S S S S S H H S F A A U C P L S S 11 Lessepsian migrant fishes in Libya

(Tripoli, Zwara); these were considered to be the most meristic counts were taken and documented. The abun - active sites for catches and were investigated monthly. Al- dance, habitat type, depth range, maximum size, and com - bardiah in the east region and Farwah in the west region mercial value were recorded. were also selected as border sites and were investigated The abundance is divided into two levels: rare and seasonally (Fig. 1) in order to standardize sampling common, according to Bilecenoglu and Taşkavak (2002). bench-marks and fishing effort, two fishing boats of the If the species is represented by less than 1% of the total same size and fishing gears were considered from each Lessepsian fish collected during the fieldwork, the species sampling site. A total of 4273 specimens were collected, is designated as rare. The general habitat type for each including 1901 specimens of S. luridus , 1885 specimens sample was classified as vegetated (including rocks with of S . rivulatus , and 487 specimens of fourteen other algae, sand with algae, and grass with algae), pelagic, Lessepsian fish species. The samples were immediately rocky, and sandy. The sizes (maximum total lengths) were washed with fresh water, and were identified using placed in three categories, as small (TL < 10 cm), medium Whitehead et al. (1984–1986) and Golani et al. (2002). (10 ≤ TL < 50 cm), and large (TL ≥ 50 cm). The commer - Specimens from each sample were kept in a solution of cial value was based on commercial information received formaldehyde and ethanol mixture. The samples were from the fishermen’s union and was roughly divided into subsequently washed with fresh water and stored in 5-% “commercial value” and “no commercial value”. formaldehyde. Standard morphometric measurements and

Table 2 Morphological and meristic characters for Herklotsichthys punctatus and Liza carinata

Total Average Scientific Depth Coordinate Number length Meristic Habitat Temperature name [m] [cm] [˚C]

32°03´50´´N H. punctatus 1 7.2 D15, A17, P16, V8 Pelagic 15 1–3 23°59´02´´E 32°04´43´´N D1, IV, D2 1 + 8, L. carinata 2 16.3–23.3 Pelagic 17 5–7 23°58´50´´E A III + 9P 15, V I + 5

Table 3 Abundance and habitat occupation for each species according to the main regions Regions Species Habitat West Region Sirt Region East Region % abundance % abundance % abundance S. luridus Vegetation 87.74 Common 81.27 Common 6.52 Common S. rivulatus Vegetation 9.91 Common 11.46 Common 75.18 Common S. obtusata Pelagic 0.21 Rare 0.77 Rare 2.43 Common S. pinguis Pelagic 0.36 Rare 3.41 Common 7.82 Common H. punctatus Pelagic — — — — 0.04 Rare S. undosquamis Sandy — — 0.31 Rare 0.99 Rare H. far Pelagic — — — — 3.24 Common F. commersonii Sandy 0.07 Rare 0.15 Rare 0.04 Rare A. lacunosus Sandy — — 0.46 Rare 0.76 Rare A. djedaba Pelagic 0.36 Rare 0.62 Rare 1.62 Common U. pori Sandy — — — — 0.09 Rare C. crenidens Sandy — — — — 0.09 Rare P. vanicolensis Rocky — — — — 0.09 Rare L. carinata Pelagic — — — — 0.09 Rare S. commerson Pelagic 1.07 Common 1.08 Common 0.54 Rare S. diaspros Rocky 0.29 Rare 0.46 Rare 0.45 Rare Total 100 100 100

— species absent. 12 and Shakman Kinzelbach

RESULTS DISCUSSION A total of 16 Lessepsian fish species were found Sixteen Lessepsian fish species were found, two of (Table 1). Two of them are recorded for the first time on them are additions to the Libyan fish fauna and are also the Libyan coast: H. punctatus (Rüppell, 1837) and additions to the list of Lessepsian fish migrants in Libya: L. carinata (Valenciennes, 1836). Their morphological H. punctatus (Rüppell, 1837) and L. carinata and meristic characters are given (Table 2). The species (Valenciennes, 1836) (Tables 1, 2). These species have names Sphyraena obtusata Cuvier, 1829 and S. pinguis been recorded in many areas of the eastern Mediterranean Günther, 1874 are consistent with those provided by Sea by Kosswig (1956), Mouneimné (1977), Whitehead Doiuchi and Nakabo (2005) in their recent revision. et al. (1984–1986) and El-Sayed (1994). When a species As regards the abundance, the majority (9) of the is found to be rare, or as single specimen only, this is con - Lessepsian fish species studied (56.25%) were rare, con - sidered to be the first step in establishing a successful stituting fewer than 1% of the total number of the population, as expressed by an increase in the population Lessepsian fish collected, while the remaining seven (Golani and Ben-Tuvia 1989). Three species have also species (43.75%) were common. In terms of the habitat been recorded from Libya by other authors: Parexocoetus occupation, the fish were found on vegetation (12.5%), in mento (Valenciennes, 1846) (cf. Ben-Tuvia 1966), the water column (=pelagic; 43.75%), on sandy bottom Sargocentron rubrum (Forsskål, 1775), and Upeneus (31.25%), and on rocky bottom (12.5%). moluccensis (Bleeker, 1855) (cf. Stirn 1970), but they According to size, 75% of the Lessepsian species were were not found during the present study. categorized as medium, followed by small (6.25%), and The abundance indicates that seven species can be large (18.75%). More than one-third (37.5%), were considered as common, namely Siganus luridus ; S. rivu - species known for their commercial value, whilst 62.50% latus ; Sphyraena obtusata Cuvier, 1829 ; Alepes djedaba were species with no commercial value (Fig. 2). (Forsskål 1775); Hemiramphus far (Forsskål 1775); Regarding the distribution, the 50% of the fishes surveyed Sphyraena pinguis ; Scomberomorus commerson (Lacepède, were caught along the entire Libyan coast, 37.5% in the east - 1800) (43.75%), while most of the Lessepsian fish species ern part, and 12.5% in the east and central part (Fig. 3). were rare, such as Fistularia commersonii ; Stephanolepis The abundance and the habitat occupation of the species diaspros Fraser-Brunner, 1940; Herklotsichthys punctatus in each of the main regions are presented in Table 3. The (Rüppell, 1837); Upeneus pori Ben-Tuvia et Golani, 1989; commercial value, size, and distribution of the species in Crenidens crenidens (Forsskål 1775); Pempheris vanicolen - each of the main regions are presented in Table 4. Fig. 4 sis ; Liza carinata (Valenciennes, 1836); Saurida illustrates the distribution of each species along the undosquamis (Richardson, 1848); and Atherinomorus Libyan coast. lacunosus (Forster, 1801) (56.25%) (Fig. 2). The abundance of these species differs between the main regions (Table 3), which may be due to a relation between the species’ early

Table 4 The size, commercial value, and distribution of each species according to the main regions

Regions West Region Sirt Region East Region Size Species Commercial Commercial Commercial [cm] Distribution Distribution Distribution value value value S. luridus Medium + Commercial + Commercial + Commercial S. rivulatus Medium + None + Commercial + Commercial S. obtusata Large + None + None + None S. pinguis Medium + None + None + Commercial H. punctatus Small — — — — + None S. undosquamis Medium — — + None + None H. far Medium — — — — + Commercial F. commersonii Large + None + None + None A. lacunosus Medium — — + None + None A. djedaba Medium + None + None + Commercial U. pori Medium — — — — + None C. crenidens Medium — — — — + None P. vanicolensis Medium — — — — + None L. carinata Medium — — — — + None S. commerson Large + Commercial + Commercial + Commercial S. diaspros Medium + None + None + None + present, — absent. 13 Lessepsian migrant fishes in Libya arrival and the species abundance. Golani (1998) showed As far as their commercial value is concerned, six that there is a correlation between species that arrived earli - species (37.5%) have become commercially valuable on er in the Mediterranean and their greater abundance. This the Libyan coast, and ten species (62.5%) are character - can be explained by (a) the longer they are in the ized as having no commercial value (Fig. 2). Of these six, Mediterranean, the greater the opportunity for them to build three ( S. pinguis , H. far , and A. djedaba ) were found in the up their pop ulations, or (b) the greater research effort, east part of the Libyan coast, two ( S. luridus , S. commer - which was much less intense in the past (Golani 2002). son ) all along the Libyan coast, and one ( S. rivulatus ) in On the Turkish coast, the abundance of the Lessepsian the east part and the Sirt Gulf only (Table 4). These species fish has the following proportions: 5 species (15%) are are now found regularly in the Libyan catches; most of categorized as rare and the remaining 28 species (84.8%) them have been recorded as commercially valuable in as common (Bilecenoglu and Taşkavak 2002). The pro - many areas in the eastern and central-south Mediterranean portions are different in the presently reported study. Sea ( Torcu and Mater 2000, Bilecenoglu and Taşkavak Regarding habitat occupation, the majority of the 2002, Golani 2002, Shakman and Kinzelbach 2006). Lessepsian migrant fish species were found in the coastal The distribution observed, implies that half of the area and usually at depths of 1–50 m (Fig. 2, Table 3), species (50%) are present all along the Libyan coast (Fig. 3). while only two species were found in the vegetation habi - There is a different concentration for the different species: tat, namely S. rivulatus and S. luridus (12.5%). S. rivula - S. luridus is concentrated more in the west part and the tus was found in several different overgrown habitats Sirt Gulf rather than in the east part of the Libyan coast, (rocks with algae, sand with algae, and grass with algae) whilst S. rivulatus is concentrated in the east part and whereas S. luridus was found in one specific vegetation decreases in the Sirt Gulf and the west part (Fig. 4). There habitat (rocks with algae). Both Siganus species are con - may be competition between S. luridus and S. rivulatus in sidered to be strictly herbivorous, and in the the east region, as these species together are less concen - Mediterranean only two fish species belong to a similar trated in the area from Zwara up to the Tunisian border, trophic guild— Sarpa salpa (L.) (Sparidae) and although there is an appropriate habitat for herbivorous Sparisoma cretense (L.) (Scaridae)—both in the eastern species in this area especially on the Farwah coast. Two and central basins (Azzurro and Andaloro 2004). Bariche species are distributed in the east part and Sirt Gulf ( S. et al. (2004) showed that S. rivulatus is able to settle on undosquamis , A. lacunosus ) and six species are distrib - a large range of substrates and habitats, including rock uted only in the east part ( H. punctatus , H. far , U. pori , C. pools, muddy harbours, and sea-grass beds. In the eastern crenidens , P. vanicolensis , L. carinata ) (Fig. 4). For Mediterranean, S. rivulatus has a wider settlement range a better understanding of Lessepsian immigration, addi - than that of S. luridus , probably due that S. rivulatus has tional taxonomic and biological investigations are benefited from the low diversity of native herbivorous required (Ben-Tuvia, 1978). It is expected that in some species (Bariche et al. 2004). Five species were found in cases the exchange of fauna and flora may have taken the sand habitat: S. undosquamis , F. commersonii, place before the opening of the Suez Canal, as a result of A. lacunosus, U. pori , and C. crenidens (31.25%), and two the elevation of sea levels and undulations of the Isthmus species were on rocks: P. vanicolensis and S. diaspros during the Pleistocene (Ben-Tuvia 1978). (12.5%). The many potential rock habitat site-related The presently reported study has shown that some of species from the Red Sea would not succeed, or would the Lessepsian migrants have successfully adapted to the only rarely succeed, in reaching that habitat in the different topography and environments of Libyan coast Mediterranean, since they would need to cross the Suez and two fish species are also recorded for the first time. Canal, the northern Gulf of Suez and the south-eastern Many species have become widespread along this coast, Mediterranean, all of which lack a continuous rocky habi - which means that they are contributing to the commercial tat (Golani 2002). The pelagic species are the largest cat - fish catch in Libya. egory (seven species): S. obtusata, S. pinguis, H. puncta - tus, H. far, A. djedaba, L. carinata, and S. commerson ACKNOWLEDGEMENTS (43.8%). The majority of these species tend to spread We would like to thank the Administration of the within a depth range of 20 to 40 m (Por 1978). Marine Biology Research Center (MBRC) in Libya (Mr. As regards of their size, twelve species (75%) were Noureddin Essarbout, Head of the Center, and Abdulbaset classified as medium, followed by three species considered Abuissa, Head of Studies and Consultation Director) for large (18.75%) and one species—small (6.25%) (Fig. 2 , their support of this study. We would also like to thank the Table 4). This result is similar to the results in the Turkish fishermen and the fishermen’s union for their collabora - seas, where 78.8% of species were medium-sized, fol - tion with us. We would like to express our thanks to Dr. lowed by large (12.5%) and small (9.1%) (Bilecenoglu H. Winkler (Zoology Institute, Rostock University, and Taşkavak 2002). However, these figures differ from Germany) for his advice and to Dr. A. Pont, London, for the results in the eastern Mediterranean where more than the linguistic revision of the text. This article constitutes half of the Lessepsian migrants were of medium size; a part of a PhD project. small- and large species were more or less equal in num - ber (13 and 12, respectively) (Golani 2 002). 14 and Shakman Kinzelbach

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