Anali za istrske in mediteranske študijeAnnali di Studi istriani e mediterranei Annals for Istrian and Mediterranean Studies 

UDK 5                       Annales, Ser. hist. nat., 29, 2019, 1, pp. 1-158, Koper 2019 ISSN1408-533X 
UDK 5 ISSN 1408-533X (Print) ISSN 2591-1783 (Online) 

Anali za istrske in mediteranske študije Annali di Studi istriani e mediterranei Annals for Istrian and Mediterranean Studies 
Series Historia Naturalis, 29, 2019, 1 
KOPER 2019 
Anali za istrske in mediteranske študije - Annali di Studi istriani e mediterranei - Annals for Istrian and Mediterranean Studies 
ISSN 1408-533X (Tiskana izd.)    UDK 5 Letnik 29, leto 2019, številka 1 ISSN 2591-1783 (Spletna izd.) 
Nicola Bettoso (IT), Christian Capapé (FR), Darko Darovec, Dušan UREDNIŠKI ODBOR/ Devetak, Jakov Dulčić (HR), Serena Fonda Umani (IT), Andrej COMITATO DI REDAZIONE/ Gogala, Daniel Golani (IL), Danijel Ivajnšič, Mitja Kaligarič, 
BOARD OF EDITORS: Marcelo Kovačič (HR), Andrej Kranjc, Lovrenc Lipej, Vesna Mačić (ME), Alenka Malej, Patricija Mozetič, Martina Orlando-Bonaca, Michael Stachowitsch (AT), Tom Turk, Al Vrezec 
Glavni urednik/Redattore capo/ Editor in chief: Darko Darovec 
Odgovorni urednik naravoslovja/ 

Redattore responsabile per le scienze naturali/Natural Science Editor: Lovrenc Lipej 
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Anali za istrske in mediteranske študije - Annali di Studi istriani e mediterranei - Annals for Istrian and Mediterranean Studies 
UDK 5                                                      Letnik 29, Koper 2019, številka 1                                       ISSN 1408-533X (Print) ISSN 2591-1783 (Online) 


VSEBINA / INDICE GENERALE / CONTENTS 2019(1) 
SREDOZEMSKI MORSKI PSI SQUALI MEDITERRANEI MEDITERRANEAN SHARKS  
Hakan KABASAKAL A Review of Shark Research in Turkish Waters ...... Pregled raziskav o morskih psih v turških vodah  1  
Hakan KABASAKAL Finally Under Protection! Status of the Angel Shark, Squatina squatina (Linnaeus, 1758) in Turkish Seas, with Notes on a Recent Sighting and Incidental Captures ........................... Končno zavarovan! Status navadnega sklata, Squatina squatina (Linnaeus, 1758) v turških morjih s posebnim poudarkom na novejša opazovanja in naključne ulove  17  
Sihem RAFRAFI-NOUIRA, Youssouph DIATTA, Almamy DIABY & Christian CAPAPÉ Additional Records of Rare Sharks from Northern Tunisia (Central Mediterranean Sea) ....... Dodatni zapisi o pojavljanju redkih morskih psov iz severne Tunizije (osrednje Sredozemsko morje)  25  
RECENTNE SPREMEMBE V SREDOZEMSKI BIODIVERZITETI CAMBIAMENTI RECENTI NELLA BIODIVERSITA MEDITERRANEA RECENT CHANGES IN MEDITERRANEAN BIODIVERSITY  
Thodoros E. KAMPOURIS, Costas CONSTANTINOU & Ioannis E. BATJAKAS Establishment of the Non-Indigenous Prawn Penaeus pulchricaudatus Stebbing, 1914 in the Marine Area of Cyprus ................................ Tujerodna kozica Penaeus pulchricaudatus Stebbing, 1914, ustaljena vrsta v vodah Cipra  37  

Aytaç ÖZGÜL & Okan AKYOL  
Occurrence of a Lessepsian Swimming Crab,  
Portunus segnis (Crustacea: Decapoda),  
in Southern Aegean Sea, Turkey ............................  43  
Pojavljanje lesepske plavajoče rakovice,  
Portunus segnis (Crustacea: Decapoda),  
v južnem Egejskem morju, Turčija  
Moez SHAIEK, Sihem RAFRAFI-NOUIRA  
& Christian CAPAPÉ  
Occurrence and Unusual Abundance  
of Reticulated Leatherjack Stephanolepis  
diaspros (Osteichthyes: Monacanthidae)  
from the Lagoon of Bizerte (Northern Tunisia,  
Central Mediterranean Sea) ..................................  49  
Pojavljanje in nenavadna številčnost afriškega  
kostoroga, Stephanolepis diaspros (Osteichthyes:  
Monacanthidae) iz lagune pri Bizerti (severna  
Tunizija, osrednje Sredozemsko morje)  
IHTIOLOGIJA  
ITTIOLOGIA  
ICHTHYOLOGY  
Domen TRKOV, Borut MAVRIČ,  
Martina ORLANDO-BONACA & Lovrenc LIPEJ  
Marine Cryptobenthic Fish Fauna  
of Slovenia (Northern Adriatic Sea) .......................  59  
Morska kriptobentoška ribja favna Slovenije  
(severni Jadran)  
Polona PENGAL, Eva HORVAT  
& Mihael J. TOMAN  
Spatial and Temporal Distribution of Anchovy  
(Engraulis encrasicolus) and Sardine  
(Sardina pilchardus) Eggs in Slovenian  
Territorial Waters (Northern Adriatic) ....................  73  
Prostorsko in časovno pojavljanje iker sardele  
in sardona v slovenskem teritorialnem morju  

Sihem RAFRAFI-NOUIRA, Youssouph DIATTA & Christian CAPAPÉ Overview of Trophic Levels Recorded in Teleost Species from Northern Tunisian Waters (Central Mediterranean Sea) ...................... Pregled trofičnih nivojev pri ribah kostnicah vzdolž severnotunizijskih voda (osrednje Sredozemsko morje)  85  
FLORA FLORA FLORA  
Nenad JASPRICA, Marija PANDŽA & Milenko MILOVIĆ Spontaneous Vegetation on Slag Heaps in Southern Croatia .............................................. Spontana vegetacija na jalovinah v južni Hrvaški  93  
Amelio PEZZETTA Le Lamiaceae della Flora Italiana: Distribuzione Regionale e Considerazioni Fitogeografiche .......... Ustnatice (Lamiaceae) v italijanski flori: regionalna razširjenost in fitogeografski pomisleki  103  
FAVNA FAUNA FAUNA  
Manja ROGELJA & Lovrenc LIPEJ Occurrence of Giant Tun, Tonna galea (Linnaeus, 1758) (Gastropoda: Tonnidae) in the Marine Waters Off Slovenia (Northern Adriatic Sea) .......... Pojavljanje velikega sodca, Tonna galea (Linnaeus, 1758) (Gastropoda: Tonnidae) v morskih vodah Slovenije (severni Jadran)  121  
Jure JUGOVIC & Živa MUHIČ Spatial Distribution of Three Species of Palaemon Shrimp (Crustacea: Decapoda: Caridea) in Badaševica River (SW Slovenia) .......... Prostorska razporeditev treh vrst kozic iz rodu Palaemon (Crustacea: Decapoda: Caridea) v Badaševici (JZ Slovenija)  125  

MISCELLANEA 
Khadija OUNIFI BEN AMOR, Mohamed Mourad BEN AMOR & Jamila BEN SOUISSI 
Abiotic Parameters in Tunis Southern Lagoon After an Environmental Restoration and Status of Macrobenthic Biocenosis (Northern Tunisia, Central Mediterranean Sea) ...... 135 
Abiotski parametri v tuniški južni laguni po okoljski obnovi in status makrobentoških biocenoz (severna Tunizija, osrednje Sredozemsko morje) 
DELO NAŠIH ZAVODOV IN DRUŠTEV 
ATTIVITA DEI NOSTRI ISTITUTI E SOCIETA ACTIVITIES BY OUR INSTITUTIONS AND ASSOCIATIONS 


Ana FORTIČ 
Protecting Adriatic Biodiversity in Kotor: The Congress AdriBioPro 2019 ............................. 143 
IN MEMORIAM 
In Memoriam of Fabio Perco (1946–2019) (Lovrenc Lipej) ..................................................... 147 
Navodila avtorjem ................................................ 149 Istruzioni per gli autori .......................................... 151 Instruction to Authors ............................................ 153 
Kazalo k slikam na ovitku ..................................... 156 Index to images on the cover ................................ 156 

received: 2019-01-24 DOI 10.19233/ASHN.2019.13 

SPATIAL DISTRIBUTION OF THREE SPECIES OF PALAEMON SHRIMP (CRUSTACEA: DECAPODA: CARIDEA) IN BADAŠEVICA RIVER (SW SLOVENIA) 
Jure JUGOVIC & Živa MUHIČ 
Department of Biodiversity, Faculty of Mathematics, Natural Sciences and Information Technologies, University of Primorska, Glagoljaška 8, 6000 Koper, Slovenia e-mail: jure.jugovic@upr.si 
ABSTRACT 
We investigated the presence, spatial distribution and possible co-existence of Palaemon species from Badaševica river and its draining channels. We used inverted bottles with a bait to trap the shrimp and recorded three species 
(P. 
elegans, P. adspersus, P. antennarius) from six out of seven surveyed localities. The localities mostly differed in distance from the coastline (0–3.6 km) and salinity (3.9–36.2), while high variation was recorded in other hydrologi­cal, physio-chemical and biochemical parameters among the localities and species. P. elegans was present only at the river mouth at salinity of 33.2, while P. antennarius was predominantly found at localities in a distance more than 2 kilometres inland. P. adspersus was most common and prevailed at localities with denser vegetation. Although 

P. 
adspersus and P. elegans were both found at a single locality, their microhabitat clearly differed (P. elegans was recorded exclusively at the river mouth, P. adspersus ca. 50 metres inland). A single specimen of P. antennarius was trapped together with P. adspersus but in general their spatial and ecological segregation was well expressed. Presence of ovigerous females in brackish environment was confirmed for P. adspersus and P. antennarius. Males outnumbered females in all three species. 


Key words: brackish environment, spatial segregation, co-occurrence, Palaemonidae, salinity 
DISTRIBUZIONE SPAZIALE DI TRE SPECIE DI GAMBERI DEL GENERE PALAEMON (CRUSTACEA: DECAPODA: CARIDEA) NEL FIUME CORNALUNGA (SLOVENIA SUD-OCCIDENTALE) 
SINTESI 
Gli autori hanno studiato la presenza, la distribuzione spaziale e la possibile coesistenza di specie del genere Palaemon nel fiume Cornalunga e nei suoi canali di drenaggio. Con l’uso di bottiglie rovesce con un’esca, hanno intrappolato tre specie di gamberetti (P. elegans, P. adspersus, P. antennarius) in sei delle sette localita campionate. Le localita sono posizionate a distanze differenti dalla costa (0-3,6 km), con diverse salinita (3,9-36,2) e ampie differenze pure negli altri parametri idrologici, fisico-chimici e biochimici tra localita e specie. P. elegans e presente solo alla foce del fiume (salinita pari a 33.2), mentre P. antennarius e presente prevalentemente in localita a una distanza maggiore ai 2 km verso l’entroterra. P. adspersus e piu comune, trovato prevalentemente in localita con vegetazione piu densa. Sebbene P. adspersus e P. elegans sono stati trovati in una sola localita, il loro microhabitat differisce chiaramente (P. elegans presente esclusivamente alla foce del fiume, metre P. adspersus a circa 50 metri di distanza verso l’entroterra). Un singolo esemplare di P. antennarius e stato trovato intrappolato insieme a P. adspersus, ma in generale la loro segregazione spaziale ed ecologica e ben espressa. La presenza di femmine ovigere in ambiente salmastro e stata confermata per P. adspersus e P. antennarius. Per tutte le specie i maschi campionati superavano in numero le femmine. 
Parole chiave: ambiente salmastro, segregazione spaziale, co-occorrenza, Palaemonidae, salinita 
125 
INTRODUCTION 
Palaemonidae are diagnosed by the size of their second chelipeds that are always larger (in some species slightly and in other extremely) than the first chelipeds (Bauer, 2004; De Grave et al., 2008; Christodoulou et al., 2016). The genus Palaemon consists of 87 species (De Grave & Ashelby, 2013; Carvalho et al., 2014; Tzomos & Koukouras, 2015), however there are only two genera and 14 species in the Palearctic region (De Grave et al., 2008; Tzomos & Koukouras, 2015). In Slovenia, five species of Palaemonidae shrimps are re­ported (Manning & Stevčić, 1982; Christodoulou et al., 2016), although this family is one of the most speciose within the Caridea (de Grave et al., 2008; Christodoulou et al., 2016) following the Atyidae (43 genera) with 14 genera in total (Christodoulou et al., 2016). 
Among them, three species were known to penetrate rivers and channels, but only Palaemon anntenarius H. Milne Edwards, 1837 is treated as a predominantly fresh­water species inhabiting lakes and rivers. Nevertheless, it can sometimes be found also in coastal brackish waters such as lagoons and estuaries in the Mediterranean ba­sin (Holthuis, 1961; d’Udekem d’Acoz, 1999; Falciai & Palmerini, 2002; Gottstein-Matočec & Kerovec, 2002). This species is on the list of protected species in Slovenia (Decree on protected wild animal species: Anonymous, 2004). Palaemon adspersus Rathke, 1837 and Palaemon elegans Rathke, 1837 are predominantly marine spe­cies but can be found in anchialine waters, and they can sometimes co-occur (Manning & Stevčić, 1982). In Slovenia, they were found together in few localities in the areas of tidal flats, where another Palaemon spe­cies, P. xiphias (Risso, 1816) can also occur (Manning & Stevčić, 1982). Palaemon serratus (Pennant, 1777), a fifth palaemonid species from Slovenia, is frequently reported from the coastal marine environments (Turk & Richter, 2007). 
In total, there are only three small rivers with direct outflow into the Adriatic Sea in Slovenia: Dragonja at the border with Croatia in the south, Rižana outflowing into the port of Koper in the north, and Badaševica that is flowing through the artificial channel throughout its lower flow, while in its upper part it flows through agricultural landscape. Spring of Badaševica that was our focal river is at 196 m above sea level and reach the Bay of Koper after 
9.5 kilometres with a gradient of only 2.07 %. In its lower part, gradient is even lower, only 0.14 % during the last 
2.2 kilometres before the outflow. Owing to its torrential nature, floods were frequent in its lower part, therefore the river channel was highly changed and redirected; the river banks were walled or covered with stone blocks and many draining channels were constructed since 1950s (Plut, 1979). Badaševica had its old channel outflowing in the San Canziano bay, however an artificial new main and some side (draining) channels were constructed south of it and the river has now its main outflow in the Bay of Koper (Plut, 1979). All three main rivers of the Slovenian part of Adriatic basin potentially contribute to the pollution of the gulf of Trieste (Turk, 2016). At their estuaries, freshwater and marine systems are interacting, and anchialinae water is present. According to salinity, habitat categories can be divided in freshwater (i.e. lim­netic: 0–0.5) and saline (oligohaline: 0.5–5; mesohaline: 5–15; polyhaline: 18–30; mixohaline: 30–40) environ­ments (Jones & Hallin, 2010). 
In the present study we provide the information on spatial distribution of three species of Palaemon within 


Tab. 1: Sampling localities (S) with hydrological, biological, physio-chemical and biochemical parameters (min–max value) in Badaševica river.Tab. 1: Vzorčna mesta (S) z razponom hidroloških, bioloških, fizikalno-kemijskih in biokemijskih parametrov v reki Badaševici.

S  Geographic coordinates  Distance of traps from the sea [m]  Channel type  Substrate  Vegetation  Tidal influence  T [°C]  Oxygen [mg/L]  Oxygen [%]  pH  Conduc­tivity [mS/cm]  Salinity  Nitrates [mg/L]  Chl a [µg/L]  
1  45°32'32.8”N 13°43'11.4”E  0–80  new artificial channel  silt, mud, rocks, walled  exposed, some different algae  strong  11.1–25.9  7.08–11.75  86.3–116.8  8.05–8.09  16.1–50.2  13.2–33.2  42.3–195.5  0.40–8.10  
2  45°32'25.5”N 13°43'14.9”E  250–300  new artificial channel  silt, mud, rocks, walled  some filamentous algae  strong  11.2–25.7  7.16–12.19  87.0–124.0  8.05–8.14  13.1–50.5  10.5–32.6  35.4–196.5  0.61–25.33  
3  45°32'09.4”N 13°43'45.8”E  1130–1150  new artificial channel  rocks (artificial), walled (incl. bottom)  some filamentous algae  medium  12.1–25.6  7.09–12.46  86.0–127.4  8.06–8.33  4.16–50.4  3.0–36.2  14.2–193.0  1.90–33.66  
4  45°32'23.1”N 13°43'45.3”E  780–820  draining channel  silt, mud  some phylamentous algae, Phragmites  low  11.4–25.7  9.31–11.34  102.1–130.6  7.62–8.06  16.46–42.0  6.3–12.7  40.3–130.5  1.41–33.91  
5  45°32'16.2”N 13°44'12.1”E  2110–2160 (B*)3520–3570 (SC*)  old channel  silt, mud  dense filamentous algae  low  13.5–27.1  7.81–15.34  85.3–145.8  8.15–8.20  25.20–47.4  9.5–13.7  59.1–173.5  3.14–34.61  
6  45°32'28.7”N 13°44'20.3”E  2500–2550 (B)3050–3100 (SC)  old channel  silt, mud  filamentous and other algae  low  17.8–28.9  15.34–21.72  175.4–227.2  8.33–8.70  27.1–45.6  15.7–33.9  68.6–151.0  4.41–16.78  
7  45°31'48.0”N 13°44'40.0”E  2550–2610  old channel  silt, mud, concrete, small rocks, walled  algae in periphyton  none  10.0–28.4  6.16–11.80  78.5–125.3  7.80–8.30  0.63–38.4  3.9–22.7  0.94–138.5  3.66–41.71  

*B – distance measured through new Badaševica channel to open sea; SC – distance measured through San Canziano bay to open sea 
a single river system (Badaševica in SW Slovenia) and discuss possible environmental factors influencing their spatial distribution. 
MATERIAL AND METHODS 
Specimens were collected with trapping at seven localities (Fig. 1) of Badaševica river and its draining channels from the outflow (locality 1) to artificial barrier 
2.6 km inland (locality 7) (Fig. 1, Tab. 1). Locality 7 was situated below a two metres high artificial dam, hence no traps were set upstream from the dam. Three plastic bottles (volume 1.5 l) with inverted opening were used per locality and the distance from the sea of each trap was measured (Tab. 1). Crushed mussels (Mytillus gal-loprovincialis) and chicken meat were used as bait and trapping lasted for four days during four sampling ses­sions in spring 2017. Samples were collected once per month in middle March, April, May and June. Sampling localities were georeferenced using a GPS and phys-icochemical water quality parameters (dissolved oxygen (mg*L-1), oxygen saturation (%), water temperature (°C), conductivity (mS*cm-1), and pH) were measured using a Portable multiparameter Aquaprobe AP-200 with a GPS Aquameter (Aquaread AP 2000). At each locality, sediment type and tidal influence (strong, medium, low, none) were assessed by naked eye (Tab. 1). Samples of water were transferred into the laboratory and chloro­phyll a (µg*L-1) was measured shortly afterwards. Salinity was calculated from water temperature and conductiv­ity. Distance from the sea was measured from orto-photo maps for each sampling locality as a range of distances between the three traps. 
Specimens were identified using the Olympus SZX7 stereomicroscope with a built-in camera. The identifica­tion key of González-Ortegón & Cuesta (2006) was used for species identification. Presence of ovigerous females and male-female ratio were checked. Nomenclature follows De Grave & Ashelby (2013). 
RESULTS 
Main observations and measurements on the hy­drological and physio-chemical parameters at seven sampling localities are compiled in Tab. 1. Altogether, three species were sampled during the study from locali­ties 1–6, while no animals were recorded at locality 7 (Fig. 1, Tab. 2). Most animals were collected in April and May with 52.6 % and 31.6 % of the total sample size, respectively, while in March and June only 7.9 % of the total sample size in each of those months was collected. Palaemon adspersus was most commonly sampled (69.7 % of sampled animals) and was found at four localities (1–4; see Tab. 2). 
At locality 4, where only P. adspersus was recorded, animals counted for 50.0 % of the total sample size. The ranges of chemical parameters measured at each sam­pling day and locality overlap among the three species, however, oxygen levels (concentration and saturation) were bit higher at locality 6 (and sometimes at locality 5) than at the other localities. Moreover, the distance from the sea and salinity greatly differ among the localities (Tab. 1). With time (i.e. consecutive sampling periods), 
P. adspersus was recorded progressively further from the coastline (Tab. 2). While it was recorded from localities 1, 2 and 4 (50–820 metres from the coastline) in March, it was found at localities 2, 3 and 4 (250–1150 metres inland) in April, and only at localities 3 and 4 (780–1150 metres inland) during May and June. P. elegans (5.3 % of total sample) was found only in one (outermost) of the three traps at the river mouth (locality 1) where salinity at the sampling day was 33.2, which is almost as high as salinity reported for the northern Adriatic Sea (ap­prox. 38–39, e.g. Grbec et al., 2007). P. adspersus was recorded together with P. elegans at locality 1, however it was found there in another trap that was set 50 m from the river mouth, at salinity of 13.2. In total, P. adspersus was found 50–1150 m inland, at salinity range 3.2–36.2. 
P. antennarius (25.0 % of total sample size) was found at longest distance from the sea, i. e. at two localities (5, 6) that are situated more than 2 kilometres from the coastline, with salinity range 19.0–30.9. Nevertheless, a single specimen of P. antennarius was found also at locality 2 (ca. 300 metres from the coastline) together with P. adspersus at salinity 10.5. As in P. adspersus, re­cordings of P. antennarius progressed further inland with time (Tab. 2). While the species was recorded at locality 2 in March, it was later recorded only from localities 5 (April, May; ca. 2.1 kilometres from the coastline) and 6 (June; ca. 2.5 kilometres from the coastline). 
During April and May, presence of ovigerous females was confirmed for P. adspersus (2 and 5 females at locali­ties 2 and 4, respectively) and P. antennarius (4 females at locality 5) but not for P. elegans. Males outnumbered females in all three species (100 % of males in P. elegans, 
75.0 % in P. antennarius and 78.4 % in P. adspersus). 
DISCUSSION 
We report on a presence and spatial segregation of three Palaemon species (P. adspersus, P. elegans, P. antennarius) in Badaševica river and channels nearby. Nevertheless, P. adspersus and P. elegans are frequently reported from the same environments (Berglund & Bengtsson 1981; Łapínska & Szaniawska 2006; Janas 2013). In estuaries with a low tidal influence both can inhabit meadows of Zostera marina, however, Berglund (1980; 1982) reports that in a latter case P. adspersus is more abundant, possibly due to larger body size in com­parison to P. elegans. Moreover, a spatial co-occurence of three species of Palaemon, P. antennarius, P. adsper­sus and P. varians with an ecological niche separation within Phragmites australis was reported by Dolmen et al. (2004). In this case, P. adspersus and P. elegans occurred 

Tab. 2: A list of species per each sampling locality (S), sampling date (month, in roman numerals) and number of sampled shrimps together with physio-chemical and biochemical parameters measured at each successful sampling. Tab. 2:. Seznam vrst po vzorčnih mestih (S) s podatki o datumu vzorčenja (mesec, označeno z rimskimi številka-mi) in številu zbranih kozic ter vrednostmi fizikalno-kemijskih in biokemijskih parametrov ob vsakem uspešnem vzorčenju. 
S  Species  Month  No.  T [°C]  Oxygen [mg/L]  Oxygen [%]  pH  Conductivity [mS/cm]  Salinity  NO3 [mg/L]  Chlorophyll a [µg/L]  
1  P. adspersus  III  1  11.1  10.44  94  8.09  16.08  13.2  42.3  8.10  
P. elegans  V  4  20.5  9.17  101.3  8.08  45.7  33.2  144.5  3.67  
2  P. adspersus  III  1  11.2  10.23  92.1  8.14  13.1  10.5  35.4  25.33  
IV  2  16.2  12.19  124  8.13  35.9  27.8  104.5  4.42  
P. antennarius  III  1  11.2  10.23  92.1  8.14  13.1  10.5  35.4  25.33  
3  P. adspersus  IV  4  16.4  12.46  127.4  8.33  4.97  3.2  26.1  18.92  
V  5  20.4  10.50  111.2  8.13  49.8  36.2  123.5  1.90  
VI  1  25.6  7.09  86.0  8.06  50.4  32.6  193.0  2.88  
4  P. adspersus  III  3  11.4  11.34  102.9  8.06  16.5  13.4  40.3  18.03  
IV  26  16.8  9.91  102.1  7.95  27.3  20.3  77.1  33.91  
V  8  24.4  9.31  110.6  7.99  42.0  27.3  96.7  1.41  
VI  1  25.7  10.76  130.6  7.62  34.2  21.1  131.5  5.88  
5  P. antennarius  IV  8  19.6  7.82  85.3  8.17  42.5  30.9  120.5  34.61  
V  7  25.9  8.68  106.0  8.15  31.2  19.0  79.2  3.14  
6  P. antennarius  VI  4  28.9  15.72  202.2  8.33  39.8  23.3  151.0  15.11  

in a microhabitat with lower and P. varians with higher density of P. australis (Dolmen et al., 2004). On the other hand, where a tidal influence is strong, P. adspersus and 
P. elegans in most cases select different microhabitats (Dolmen et al., 2004). In our case, selection of different microhabitats of the two species would be in line with observations of a strong tidal influence at our localities; whereas P. elegans was found only at the final point of the riverine mouth, P. adspersus occupied brackish part of the river up to approx. one kilometre from the coastline. Both species usually prefer well aerated systems with sa­linity higher than 15 (mostly polyhaline and mixohaline environments) (Barnes, 1994; Dolmen et al., 2004). 
Despite that there are records of occurrence of 
P. elegans below 6.5 of salinity from the Baltic sea (mesohaline to oligohaline waters; Dolmen et al., 2004; Łapínska & Szaniawska, 2006), both species are euryha-line, however in Badaševica only P. adspersus was found in a wide range of salinity (oligohaline to mixohaline). During our study, P. elegans was found only in mixoha-line water. Berglund & Bengtsson (1981) reported that 
P. elegans is more prone to hypoxy than P. adspersus. In our case, oxygen levels were similar for a locality with 
P. elegans and for localities with P. adspersus. Therefore, we cannot see the oxygen as a factor influencing spatial segregation of the two species in our case. Both spe­cies are mainly nocturnal (Berglund 1980; Hagerman & Ostrup, 1980; Guerao & Abello, 1996; Janas & Baran-ska, 2008), however pronounced nocturnal activity is usually less stressed in P. adspersus (Berglund, 1980; Hagerman & Ostrup, 1980). Since P. elegans is not a habitat specialist, more prone to hypoxia, more active and quicker in feeding than P. adspersus, it can occupy sites that are not preferential for stronger and larger P. adspersus (Berglund & Bengtsson, 1981). Especially when found in marine environments, P. adspersus pre­fers meadows of marine flowering plants (e.g. Posidonia, Zostera, Cymodocea) (Manent & Abella-Gutiérrez, 2006), while P. elegans can be abundant also at sandy and unprotected bottom (Berglund & Bengtsson, 1981). This could be in line with our case as the environment at the site with P. elegans was open and unprotected. Moreover, we recorded P. adspersus only at localities where dense vegetation was present. It should be noted that P. adspersus was by far most abundant at the end of a draining channel at locality 4 (73.1 % of a P. adspersus sample) with densest vegetation. Since it has been re­ported that P. adspersus rather avoid open spaces since it can be quickly detected by the predators owing to its large size, this locality seemingly offers this species an optimal habitat and reproduction site, and is probably lacking shrimp predators. Moreover, at the localities closest to the coastline (1, 2), shrimps were found only at the beginning of the sampling period (at locality 1, a single specimen in March, and at locality 2, a single and two specimens in March and April, respectively). No additional animals were found there during the continu­ation of the sampling period (May, June), and a species reached a peak of abundance in April at locality 4 (50.0 % of a P. adspersus sample). 
P. anntenarius represented 25.0 % of the total sample size and was found in two localities where neither of the above mentioned species occurred. P. antennarius is reported to be oligohaline species that inhabits freshwa­ter to brackish environments with muddy bottom. Most commonly it is mentioned as a true freshwater species (Dalla Via, 1987; Gottstein-Matočec et al., 2006; Ana-stasiadou et al., 2009, 2014; Christodoulou et al., 2016). During our study it was found in mesohaline and poly­haline waters and always above 15 of salinity, however, this is not contradicting its euryhaline nature as Dalla Via (1987) reports that it can be found at a wide range of salinity from 5 to 30. This species can adapt to quick salinity changes by metabolic adjustment if the changes in salinity are short-termed, while at higher salinities (>19) oxygen uptake strongly increases. Populations that inhabit brackish environments with higher salinity can adapt up to approximately 25 of salinity but in the same time brackish populations cannot easily adapt to low sa­linities (Dalla Via, 1987). This is in line with our results as this species was found at the range of salinity above 
10.5 (but even higher than reported in literature, up to 
30.9) and most abundant at the two localities (5, 6) with highest oxygen levels. Gottstein-Matočec et al., 2006 reported that P. antennarius shows higher reproductive success in environments with stable salinity, however during the reproductive period brackish waters were preferred (Dalla Via, 1987; Gottstein-Matočec et al., 2006) over waters with lower salinity in delta of Neretva (SE Croatia). Sheltered bays with stable environment (low tidal influence, stable salinity, warm temperature, abundance of food) and muddy bottom can therefore be optimal for the reproduction of the species, such as probably at localities 5 and 6 in our case where it was recorded after the beginning of its breeding season in April. Nevertheless, a single specimen of P. antennarius was found before the breeding season in March also at locality 2 in the main channel of the Badaševica river, in this case together with P. adspersus. We failed to find the data on a sympatric occurrence of those two spe­cies in the literature, so we think this co-occurence was coincidetial. However, we cannot exclude the option that P. antennarius can occasionally be present in the main channel of the river, possibly due to drift or active spreading of the species. 
Sex ratio in our samples strongly deviates towards males and this contradicts the reports for all three species where females are usually more abundant (Gottstein­Matočec et al., 2006; Łapínska & Szaniawska, 2006; Manent & Abelle-Gutiérrez, 2006). P. adspersus for which a reproductive period lasts from March to August (Guerao & Ribera, 1995; Manent & Abella-Gutiérrez, 2006) females were frequently noticed to lay eggs in shallow coastal marine waters (Barnes, 1994; Guerao & Ribera, 1999; Glamuzina et al., 2014). Nevertheless, we noticed some ovigerous females in the river channel itself 
(28.6 
% of all ovigerous females at locality 2), however, it is possible that some of them move towards the sea for egg laying which could cause their deficit upstream. Since most of ovigerous P. adspersus females (71.4 %) were found at locality 4 it is also possible that ovigerous females seek for protected and well-hidden places where they remain for the most of time. Hence, their trapping probability could be lower than for males and estimated sex ratio is then biased towards males. In P. antennarius, reproductive period lasts from April until August with a maximum in June (Gottstein-Matočec et al., 2006). This is in line with our data for Slovenian mesohaline and poly­haline environments, while no conclusions can be made on unusual sex ratio in P. antennarius (but see above for 

P. 
adspersus) and P. elegans. For the latter, a small sample size may prevent a more accurate conclusion. 


CONCLUSIONS 
In this study we reported on a well expressed spatial and ecological segregation among three species of Pal-aemon in a brackish environment of a river Badaševica. The following conclusions can be made: 
1. 
P. elegans and P. adspersus select different habi­tats near the river outflow that is under a strong tidal influence (P. elegans occupies more exposed rocky bottom at higher salinity at the final point of the outflow, and P. adspersus is present at a microsite with lower salinity in a better protected river channel only few tens of meters inland); 

2. 
P. antennarius and P. adspersus seem to be eco­logically and spatially well separated as the vast majority of P. adspersus was found in a drainage channel and upper the river channel with dense vegetation (mostly macrophytes), and P. anten­narius was found predominantly at sites more than 2 kilometres inland with low tidal influence in a well aerated water; 

3. 
Presence of ovigerous females was confirmed for 


P.adspersus and P. antennarius. 
ACKNOWLEDGEMENTS 
Authors thank Felicita Urzi for help in a laboratory, Martina Jeklar for help on the field and Domen Trkov for useful information on distribution of Palaemon shrimps in the area of Koper. Authors also thank two anonymous referees for constructive comments on the manuscript. 

PROSTORSKA RAZPOREDITEV TREH VRST KOZIC IZ RODU PALAEMON (CRUSTACEA: 

DECAPODA: CARIDEA) V BADAŠEVICI (JZ SLOVENIJA) 
Jure JUGOVIC & Živa MUHIČ 
Department of Biodiversity, Faculty of Mathematics, Natural Sciences and Information Technologies, University of Primorska, Glagoljaška 8, 6000 Koper, Slovenia e-mail: jure.jugovic@upr.si 
POVZETEK 
Preiskovali smo prisotnost, prostorsko razporeditev in možnost sobivanja vrst iz rodu Palaemon v reki Badaševici in njenih odvodnih kanalih. Za pasti smo uporabili plastenke z navznoter obrnjenim ustjem. Zabeležili smo tri vrste 
(P. elegans, P. adspersus, P. antennarius) s šestih od sedem preiskovanih vzorčnih mest. Vzorčna mesta so se najbolj razlikovala po oddaljenosti od morja (0–3.6 km) in slanosti (3.9–36.2), pri ostalih beleženih hidroloških, fizikalno--kemijskih in biokemisjkih parametrih pa smo zabeležili veliko stopnjo variabilnosti tako med vzorčnimi mesti kot vrstami kozic. P. elegans smo zabeležili le na ustju reke pri slanosti 33.2, P. antennarius pa smo skoraj izključno našli le na vzorčnih mestih, ki so od morja oddaljena več kot 2 km. P. adspersus je bila najštevilnejše zastopana vrsta in se je pojavljala v velikem številu predvsem na lokacijah z gosto vegetacijo. Čeprav smo P. adspersus in P. elegans našli na istem vzorčnem mestu ob izlivu reke v morje, smo opazili, da se izbira njunega mikrohabitata razlikuje (P. elegans smo našli neposredno na izlivu reke v morje, P. adspersus pa smo zabeležili prb. 50 m po toku navzgor). P. antennarius smo le enkrat (en osebek) našli na eni od vzorčnih mest skupaj s P. adspersus, a je bila sicer prostorska in ekološka ločitev med vrstami jasno vidna. Prisotnost ovigerih samic smo potrdili za vrsti P. adspersus in P. anten­narius. Samci so bili pri vseh treh vrstah številnejši od samic. 
Ključne besede: somorno okolje, prostorska ločitev, sobivanje, Palaemonidae, slanost 
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