Living benthic foraminifera of southeastern Mediterranean ultra-oligotrophic shelf habitats: Implications for ecological studies

Journal Pre-proof Living benthic foraminifera of southeastern Mediterranean ultra-oligotrophic shelf habitats: Implications for ecological studies Simona Avnaim-Katav, Ahuva Almogi-Labin, Mor Kanari, Barak Herut PII:

S0272-7714(19)30724-3

DOI:

https://doi.org/10.1016/j.ecss.2020.106633

Reference:

YECSS 106633

To appear in:

Estuarine, Coastal and Shelf Science

Received Date: 20 July 2019 Revised Date:

27 January 2020

Accepted Date: 5 February 2020

Please cite this article as: Avnaim-Katav, S., Almogi-Labin, A., Kanari, M., Herut, B., Living benthic foraminifera of southeastern Mediterranean ultra-oligotrophic shelf habitats: Implications for ecological studies, Estuarine, Coastal and Shelf Science (2020), doi: https://doi.org/10.1016/j.ecss.2020.106633. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2020 Published by Elsevier Ltd.

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Living benthic foraminifera of southeastern Mediterranean ultraoligotrophic shelf habitats: implications for ecological studies

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Simona Avnaim-Katava*, Ahuva Almogi-Labinb, Mor Kanaria, Barak Heruta,

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a

Israel Oceanographic and Limnological Research, Haifa 3108001, Israel

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b

Geological Survey of Israel, Jerusalem 95501, Israel

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*Corresponding author: Telephone: +909-734-9661, Email: [email protected]

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Abstract

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The Levantine Basin, the saltiest, hottest and the most ultra-oligotrophic basin in the Mediterranean Sea, continues to be affected by recent anthropogenic changes. That includes the long-term influence of the opening of the Suez Canal and the enhanced oligotrophy in this region due to the damming of the Nile River. This study explores the spatial distribution and diversity patterns of living benthic foraminifera in this impacted SE Levantine shelf, between 40 -100 water depths at 59 sites, sampled in August 2011 off the Israeli coast.

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Multivariate statistical analyses resulted in the identification of four distinct benthic foraminiferal assemblages, reflecting their ecological preferences distributed within four coherent biotopes with different environmental settings. Two biotopes were identified along the 40 m depth interval: 1. the middle and the southern shelf in which Deuterammina rotaliformis accompanied by Eggerelloides scaber predominate, and their abundance is positively related to Chl-a concentrations and negatively related to total organic carbon (TOC) and fine-grained sediment contents, and 2. the northern middle sandier carbonate rich shelf in which Lessepsian taxa and others calcareous foraminifera such as Quinqueloculina schlumbergeri and Ammonia tepida dominate the assemblage. The other two biotopes that occur between 60 m and 100 m water depths consist of high concentrations of fine-grained sediments, relatively rich with TOC. Hanzawaia rhodiensis, Asterigerinata mamilla and Rosalina spp. reveal a positive relationship with the carbonate-rich sediments of the northern outer shelf biotope. Lagenammina sp, Reophax scorpiurus, Glomospira charoides, Valvulineria bradyana, and Bolivina striatula exhibit a more positive relationship with higher clayey-silty organic rich sediment of the central-southern outer-shelf biotope.

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A comparison between the living assemblages investigated in the current study and during a previous study in the late 90s, at the same sites, indicates a prominent foraminiferal response to the ongoing human activity in this region. That includes (I) the expansion of some Lessepsian species into ~40 m water depths habitats indicating the availability of suitable bottom water conditions for these species attributed to the

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increase in ultra-oligotrophy at this water depth. (II) the very recent introduction (either by shipping/aquaculture) of Deuterammina rotaliformis to the Israeli coast, sometime between the late 90s and 2011and its becoming the most dominant species in the southern middle shelf, a region most affected by the ongoing consequences of the damming of the Nile.

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Keywords: Southeastern Levantine shelf; Live foraminiferal ecology; Redundancy Analysis; Environmental relations; environmental changes; Anthropic impact

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1 Introduction In the context of recent climate change and increased anthropogenic activities in coastal and offshore areas there is a growing interest to better understand the mechanisms and evolution of such events and their impact on faunal communities. Such knowledge may serve as an important reference point in studies aimed at improving bio-monitoring methods (e.g., Alve et al., 2016; Jorissen et al., 2018). Studying this context is highly relevant specifically to the Levantine Basin (LB). Located at the easternmost part of the Mediterranean Sea, the LB is considered as the most ultra-oligotrophic region within the entire Mediterranean, severely affected by the damming of the Nile River (Said, 1993; Krom et al., 1999; Herut et al., 2000; Almogi-Labin et al., 2009, 2010; D’Ortenzio et al., 2009; Lazzari et al., 2012). Additionally, this saltiest and hottest basin within the Mediterranean has been experiencing a substantial warming trend over the past 30 years (Macias et al., 2013; Ozer et al., 2017). This along with other modern human activity such as the opening of the Suez Canal facilitate the recent Lessepsian invasion (Rilov and Galil, 2009; Zenetos et al., 2012). Living benthic foraminifera are being increasingly used as tools for environmental exploration particularly for biodiversity, biological, ecological and biomonitoring studies, due to their widespread distribution in diverse modern habitats and their adaptability to various environmental conditions (Murray 2006; Schönfeld et al., 2012). Additionally, these organisms are highly sensitive to changes in the environment, and abundantly preserved in the marine fossil record. Thus a comprehensive knowledge of the living foraminiferal faunas when compared to the recent dead counterparts (Murray, 2006) serves as a robust proxy for paleoenvironmental studies (e.g., Avnaim-Katav et al., 2012, 2013, 2016b, 2017a, 2019; Elyashiv et al., 2016; Sivan et al., 2016; Milker et al., 2010, 2019). Little is known so far on the distribution of living benthic foraminifera in the LB shelf region. Basso & Spezzaferri (2000) studied the distribution of living benthic foraminifera in Iskenderun Bay, Turkey, a eutrophic basin bordering the oligotrophic northeastern LB. Other studies have been focusing on the diversity, spatial distribution and composition of the dead foraminifera in the southeastern Mediterranean inner shallow shelf, 0-40 m water depth (Hyams-Kaphzan et al., 2008; Avnaim-Katav et al., 2013, 2015). Others focused on the alien-Lessepsian foraminifera species that migrated to the eastern Mediterranean from the Red Sea since the opening of the Suez Canal (Langer and Hottinger, 2000; Hyams et al., 2002; Hyams-Kaphzan et al., 2008; Zenetos et al., 2012). Living (Rose Bengal stained) foraminifera were studied in the outer Levantine Shelf by Jannink (2001). She studied the foraminiferal distribution patterns in a time series sampled during 1996 – 1998 in

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three stations (40, 120 and 700 m) along a depth transect off the central Israeli coast. In this study relatively high numbers of miliolids and agglutinated taxa occur in the 40 m station, being regulated by the seasonal increase in organic matter following the winter primary productivity peak. In another study, Hyams-Kaphzan et al. (2009) compared between an oligotrophic site representing the natural background conditions and a eutrophic site controlled by an activated sewage sludge discharge near Palmahim, at 36 m water depth. They documented major seasonal variations during 2003 – 2004 in the oligotrophic station characterized by abundant and diverse assemblage composition compared to the eutrophic site which showed restricted seasonality and sustained an impoverished assemblage with low species richness composed mainly by opportunist species. These localized effects of organic matter enrichment on the foraminiferal assemblage were reviewed by Tadir et al. (2017), who gathered new seasonal data during 4 sampling cruises in 2012, in the same stations and in an additional control site. The unique natural setting and human induced environmental changes in the LB put its shallow marine communities at a high risk. Due to the limited information on this vulnerable ecosystem, our study is designed to: (1) identify and describe the spatial distribution and diversity patterns of the present day living benthic foraminifera, along the southeastern Levantine shelf (Fig. 1); (2) determine the relationships between a range of environmental factors and the foraminiferal faunas using multivariate statistical analyses in order to understand their ecological preferences and thus improve their use as environmental bio-indicators; (3) investigate possible migration of aliens/invasive species and their association with the local ecosystem as part of the efforts to improve environmental conservation. By achieving these aims, the outcome of this research is to add new inclusive knowledge on the present-day ecology of living foraminifera compared to previous studies, thus offering the possibility of reconstructing the historical evolution of anthropogenic activity and environmental changes in this region.

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The southeastern (SE) Levantine shelf is part of the Nile littoral cell that extends ~ 700 km from the western part of the Nile Delta to Haifa Bay (Fig.1A). Consequently, the main nutrient-rich freshwater and most sediments of the southeastern Mediterranean shelf originated from the Nile River, until its damming in 1965. The sediments were distributed predominantly northward along the Nile littoral cell by prevailing counterclockwise wave- and wind-induced long-shore currents (Said, 1993; Zviely et al., 2007). Our study area, the Israeli continental shelf, is 25–30 km wide in its southern part, narrowing northward to less than 10 km (Fig. 1B) (Neev, 1965; Almagor and Hall, 1984). Most of the inner shelf sediments, between the foreshore and ~35 m water depth and between Ashqelon and Haifa Bay, are mainly Nile-derived siliciclasts with a minor contribution from local sources (e.g., Golik, 1993; Zviely et al., 2007). Haifa Bay is the most distal part of the Nile littoral cell and further to the north towards Akhziv, carbonate-rich sediments replace the siliciclastic quartz sediments, forming rocky, sandy and silty–clayey substrates (Almagor et al., 2000 and references therein; Almogi-Labin et al., 2012). Between 40 m and the shelf edge, at ~100 to 120 m, Nile-derived sand is progressively diluted by silt and clay that comprise ~85% of the sediment from 60 to 100 m water depths (Almagor et al., 2000;

2 Study Area

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Almogi-Labin et al., 2012). Total organic carbon (TOC) concentrations increase westward along with the decrease in grain size and range between 0.2 and 1.1 wt.% (Almogi-Labin et al., 2012). The LB is the warmest and saltiest basin in the entire Mediterranean Sea, with sea surface temperatures ranging between ~16 °C in winter and 30.8 °C in summer and sea surface salinity is as high as 39.7 psu during summer (Herut et al., 2000; Gertman and Hecht, 2002; Yogev et al., 2011; Ozer et al., 2017; Herut et al., 2018). Offshore, the upper ~150-250 m of the water column is well mixed in winter (December–March), enriched by nutrients and supports a modest phytoplankton bloom leading to the highest primary production observed for this region (Berman et al., 1984; Herut et al., 2000; Kress and Herut, 2001; D’Ortenzio et al., 2005). Whereas during summer, a sharp and well-developed halocline and thermocline appears below the mixed layer, restricted to the upper ~25 m and primary production decreases (Herut et al., 2000; Lazzari et al., 2012). At water depth of ~120 m, off the Israel coast, maximum Chlorophyll-a (Chl-a) concentration is measured during late fall to early winter (Herut et al., 2000; Jannink, 2001). The maximum in Chl-a was explained by cooling and consequent weakening and deepening of the thermocline, introducing nutrients into the mixed layer, and causing a seasonal increase in Chl-a concentration. A different pattern is observed in the seasonal Chl-a concentrations in the inner shelf shallow oligotrophic waters off the Israel coast. There, at 36 m water depth near Palmahim, Hyams-Kaphzan et al. (2009) and Tadir et al. (2017) documented maximum Chl-a concentrations in the top sediment layer during spring– summer and very low concentrations in the winter. The spring–summer maximum in shallow water is attributed to benthic primary producers (e.g., diatoms, living on the top sediment layer) that bloom and thrive with the rise of temperature during spring. The combined impacts of the damming of the Nile River (Said,1993; Krom et al.,1999) and the general W-E anti-estuarine circulation, causing intermediate water nutrients transported from the eastern Mediterranean into the north Atlantic, contributing to the ultra-oligotrophy of the LB (Bethoux, 1979).

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Fig. 1. (A) The study area, located in the southeastern Levantine Basin, eastern Mediterranean, is part of the Nile littoral cell that extends up to northern Israel and is influenced by the anticlockwise long-shore current (LSC); (B) The sampling sites located along the SE Levantine shelf at 40, 60, 80 and 100 m water depths are shown on the bathymetric map, following Sade et al. (2006). See also Table 1 for details. Overlaid colored areas designate the general abundance and species diversity per site. Legend of BF/g (large circles) and simple diversity (no. of species, small circles) determined in ArcGIS by using natural breaks method classification.

Station

1-60 1-80 1-100 2-40 2-60 2-80 2-100 3-40 3-60 3-80 3-100 4-40 4-60 4-80 4-100 5-40 5-60 5-80 5-100 6-40 6-60 6-80 6-100 7-40 7-60 7-80 7-100 8-40 8-60 8-80 8-100 9-40 9-60 9-80 9-100 10-40 10-60 10-80 10-100 11-40 11-60 11-80 11-100 12-40 12-60 12-80 12-100 13-40 13-60 13-80 13-100 14-40 14-60 14-80 14-100

Water depth (m)

Sand (63– 2000µm)

33.038592 33.046421 33.047784 32.972325 32.978308 32.980093 32.98096

60 80 100 40 60 80 100

3.6 29.5 9.2 70.1 12.1 14.8 8.5

67.8 58.9 66.9 26.5 52.6 70.7 73.8

34.9397

32.907809

40

75.8

34.920657 34.895023 34.89119 34.913058 34.888248 34.863608 34.852954 34.894507 34.85773 34.835216 34.825381 34.877273 34.832684 34.813291 34.805994 34.847674 34.804186 34.78611 34.775981 34.813572 34.768677 34.747376 34.732351 34.783483 34.720322 34.71395 34.685191 34.745962 34.675498 34.66446 34.658502 34.706618 34.654396 34.634354 34.614377 34.652779 34.609304 34.586636 34.572871 34.591095 34.549361 34.530686 34.51577 34.514974 34.502343 34.461477 34.422037

32.91071 32.914616 32.9152 32.809179 32.813286 32.817364 32.819128 32.696541 32.703665 32.708026 32.709931 32.616423 32.62642 32.630767 32.632403 32.523522 32.534064 32.538446 32.540901 32.416318 32.427856 32.433331 32.437193 32.28858 32.308607 32.310627 32.319743 32.186694 32.210102 32.213768 32.215748 32.090524 32.109135 32.116279 32.123398 31.979165 31.998254 32.008207 32.014251 31.879641 31.898202 31.906508 31.913142 31.794581 31.800071 31.817835 31.834979

60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100

65.6 5 9.1 nd 33.5 56.1 4.8 37.8 38 14.1 6.5 54.3 36.1 31.6 10.1 40.5 34.9 19 7.2 31.6 17.9 19.3 5 69.5 12.4 4.5 10.8 42.7 11.2 16.1 3.5 38.2 13.2 10.6 1.8 20 22.7 14.7 8.6 31.8 14.4 8.1 5.5 32.1 16.2 4.7 7.7

Water depth (m)

Longitude (E)

Akko

60 80 100 40 60 80 100

35.016829 34.969445 34.961196 34.98917 34.945097 34.931946 34.925559

Kiryat Yam

40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100 40 60 80 100

Location

Akhziv

Dado

Atlit

Dor

Cesarea

Michmoret

Poleg

Herzeliyya

Tel Aviv

Palmahim

Ashdod

Nizzanim

Latitude (N)

Silt (2– 63µm)

Clay (0.02– 2 µm)

CaCO3

TOC

Chl-a (mg/m3)

% 28.6 11.6 23.8 3.4 35.3 14.5 17.7

7.9 18 12.9 15.5 12.4 11.3 10.1

1.08 0.61 1.05 0.21 0.5 0.62 0.73

0.21 0.19 0.18 0.25 0.23 0.22 0.20

21.3

3

19.3

0.37

0.24

30.7 66.6 74.2 nd 54.5 37.4 73.7 55.1 50.3 69.1 73.7 39.2 52.4 56.1 73.3 52.2 52.7 68.4 66.8 56.3 68.1 67.2 73.3 25.7 73.7 74.9 74.9 48.9 75.2 70.8 75.8 50.6 72.6 75.5 80.1 65.3 66.2 72.6 77.3 57 73.8 79.4 74.7 59.4 71.7 73.9 77.5

3.8 28.4 16.7 nd 12 6.5 21.5 7.1 11.7 16.8 19.8 6.5 11.5 12.3 16.6 7.3 12.3 12.6 26 12.1 14 13.5 21.7 4.8 13.9 20.6 14.4 8.5 13.6 13.1 20.7 11.2 14.2 13.9 18.1 14.7 11.2 12.7 14.1 11.3 11.8 12.5 19.8 8.4 12.1 21.4 14.8

47.6 11.4 8.6 nd 32.6 37.2 5.9 34.6 20.3 11 9.5 14 15.8 14.8 11.2 16.5 11.2 10.8 9.6 14.9 9.8 9.3 9.2 15.2 9.2 8.8 7.8 12.7 9.3 8.6 8.5 12.1 9 8.7 8.2 13 9.6 9.3 8.2 12.9 9.4 8.6 7.2 10.1 9.8 9.1 7.3

1.03 0.95 1.09 nd 0.54 0.55 1.13 0.51 0.5 0.61 0.74 0.38 0.53 0.55 0.73 0.34 0.52 0.62 0.78 0.37 0.64 0.6 0.67 0.25 0.63 0.63 0.69 0.46 0.55 0.7 0.74 0.47 0.73 0.71 0.95 0.26 0.65 0.69 0.84 0.58 0.63 0.75 0.88 0.64 0.74 0.83 0.97

0.22 0.20 0.20 0.26 0.23 0.21 0.20 0.27 0.23 0.21 0.21 0.34 0.24 0.21 0.20 0.36 0.24 0.21 0.19 0.35 0.24 0.21 0.19 0.42 0.23 0.22 0.18 0.41 0.26 0.18 0.17 0.47 0.24 0.20 0.19 0.48 0.26 0.21 0.19 0.55 0.27 0.23 0.20 0.42 0.35 0.21 0.17

15-40 15-60 15-80 15-100

180 181 182 183 184 185 186 187 188 189 190 191

Ashqelon

40 60 80 100

34.459566 34.403691 34.390472 34.37214

31.695775 31.719286 31.724849 31.732563

40 60 80 100

16.7 13 6.8 3.7

70.8 69.4 81.4 77.1

12.4 17.5 11.8 19.1

8.3 9 9 5.5

0.59 0.77 0.85 0.97

Table 1. Station locations, water depths and coordinates of surface samples taken with a box corer by the R/V Shikmona during August 29-31, 2011 research cruise. The environmental dataset utilized in the current study, includes the following sediment properties: percentage of the clay, silt and sand fractions, TOC (wt.%) and CaCO3 content (wt.%), all from Almogi-Labin et al. (2012) and Chl-a (mg/m^3). (nd = no data).

3 Materials and Methods 3.1. Field sampling

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Fifty-nine box-corer (Ocean Instruments BX 700 AL) sediment samples were collected along fifteen east–west oriented transects, at 40, 60, 80 and 100 m water depths from Akhziv in the north to Ashqelon in the south, during August 2011 (Table 1, Fig. 1). Two sets of surface sediments (top 0–1 cm interval) were sampled for sedimentological and micropaleontological analyses from each box-core, at each station, using a 54 mm diameter mini corer. This interval was chosen for the micropaleontological analyses following earlier studies summarized in Schönfeld et al. (2012). The sediments samples of 5–44g dry weight were preserved in Rose Bengal solution (2 g l-1ethanol 95%) for two weeks to distinguish living from dead specimens. The sedimentological analyses of these samples that include grain size distribution, CaCO3 and TOC contents, published by Almogi-Labin et al. (2012), were incorporated in the environmental database of the present study along with other environmental data (Table 1).

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Grain size analysis was performed on sediment < 2 mm using a Malvern Mastersizer MS-2000. Bulk sediments were measured following the protocol of Crouvi et al. (2008). The clay/silt/sand content was calculated using thresholds of 2 µm and 63 µm, respectively. For TOC analysis, the modified Walkley–Black titration was used (Walkley and Black, 1934). The method is based on the oxidation of organic matter by potassium dichromate (K2Cr2O7)-sulfuric acid mixture followed by back titration of the excessive dichromate by ferrous ammonium sulfate. Relative error of the analysis is ±0.1%. The CaCO3 content was determined by gasometry.

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Chl-a concentrations were determined using analysis of Moderate Resolution Imaging Spectroradiometer, MODIS Aqua (1km2 resolution) (Herut et al., 2012). The annual averages of the monthly mean values calculated per each station (polygon with perimeter of 2km radius) over January to December 2011 are used in this study.

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3.3. Foraminiferal Analysis

3. 2. Analytical methods of environmental data The environmental dataset is composed of several parameters including percentage of the clay/silt/sand fractions, TOC (wt.%), CaCO3 (wt.%), and Chl-a (mg/m3) contents, all of which were determined in fifty nine samples (Table 1).

0.51 0.25 0.22 0.18

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Fifty nine samples were washed over a 63 µm sieve and dried at 50 °C and the >125 µm fraction was used for foraminiferal analyses. The living assemblage was studied in the entire samples following the protocol of Schönfeld et al. (2012) (Supplementary data No. 1). Specimens were identified to species level and counted to determine total number of benthic foraminiferal individuals per gram dry sediment (BF/g), species relative abundance, species diversity as raw diversity (species richness) and Fisher α-index and dominance (Fisher et al., 1943; Levin & Gage, 1998). Taxonomic identification to species level was based mainly on Loeblich and Tappan (1987, 1994), Cimerman and Langer (1991), Sgarrella and Moncharmont-Zei (1993), Hottinger et al. (1993), Jones (1994) and Hayward et al. (2017).) Living taxa with a relative abundance of >5% in at least one sample were photographed using a SMZ800 stereomicroscope (zoom range 1×–6.3×) equipped with a DS-Fi2 camera with a 5 million pixel resolution. The state of preservation of the tests was evaluated and is usually very good in all studied samples.

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The data set presented here, includes only samples with more than 50 shells in the numerical analyses (half of the samples contained 54 to 97 shells and the rest 106 to 290 shells) based on the commonly used minimum count number for analysis (Heiri and Lotter, 2001; Mischke et al., 2014; Avnaim-Katav et al., 2016a). We used only species with a relative abundance >5% in at least one sample in order to simplify the results and compare them with a similar approach used in previous studies in the area (Hyams-Kaphzan et al., 2008; Avnaim-Katav et al., 2013, 2015). The final data set consists of a total of 38 surface sediment samples which includes 27 living species with a relative abundance >5% in at least one sample. The dominant benthic species used in the final statistical analyses discussed in this paper accounts for 84% in average of the total census data (>71% in 35 samples and >52% in only three samples) (Table 2). Thus, a relatively large portion of the foraminiferal information is preserved making the statistical results more reliable. Cluster Analysis (CA) was used in order to classify the distribution of groups and subgroups in the foraminiferal community into homogeneous faunal zones (clusters). This method was processed by PRIMER version 6 software (Plymouth Routines In Multivariate Ecological Research, UK) following similar steps described in Avnaim-Katav et al. (2017b). A DCA was applied to the species dataset in order to provide further information about the patterns of variation in the faunal data and to determine the type of response (unimodal or linear) displayed by the species distribution to one or more environmental gradients (Lepš and Šmilauer, 2003). As DCA showed a linear species response, Redundancy Analysis (RDA) was applied to quantify the direct relationship between the distribution of the faunal structure and the available abiotic ecological variables including grain-sizes, TOC, CaCO3 content and Chl-a concentrations (Table 1). A summery on this ordination technique is given in Leyer and Wesche (2007). In order to further test the correlation between the species distribution and the environmental variables we used the parametric correlation coefficient Pearson's r. Both DCA and RDA were implemented using Canoco, version 5.10 software (Lepš and Šmilauer, 2003; Ter Braak and Šmilauer, 2002) following similar steps detailed in Avnaim-Katav et al. (2017b).

3.4. Multivariate statistical analysis

275 276

Sample name

A. glomeratum

A. planorbis

A. tepida

A. mamilla

A. stelligerum

B. subsphaerica

B. striatula

B. granulata

C. poeyanum

D. rotaliformis

D. compressa

E. scaber

F. subacuta

G. affinis

G. charoides

H. rhodiensis

P. calcariformata

P. mediterranensis

Polymorphina sp.

Q. bosciana

Q. schlumbergeri .

R. elongatastriata

Lagenammina sp.

R. scorpiurus

Rosalina sp.

T. bocki

V. bradyana

273 274

2-40 2-60 2-80 2-100 3-40 3-80 4-80 5-100 6-100 7-60 7-80 7-100 8-60 8-80 8-100 9-60 9-80 9-100 10-40 10-60 10-80 10-100 11-80 11-100 12-60 12-80 12-100 13-40 13-60 13-80 13-100 14-40 14-60 14-80 14-100 15-40 15-60 15-80

1 5 13 2 0 1 1 1 6 9 3 6 4 6 3 8 16 3 0 4 12 5 0 3 27 6 12 0 36 35 9 0 23 21 7 0 32 11

0 0 0 3 3 0 0 1 0 0 0 7 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 0 2 0 1 0 2

7 4 0 0 9 1 0 0 1 9 3 1 0 3 0 0 0 0 9 8 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0

2 1 1 0 0 2 7 0 1 0 0 0 3 3 0 0 1 0 0 0 0 0 0 0 0 0 1 2 1 0 0 0 0 0 0 0 0 0

5 2 0 0 3 3 4 0 2 9 3 1 8 1 1 2 4 6 0 1 8 1 1 1 0 1 0 0 7 0 2 2 16 1 0 0 3 2

0 0 0 0 0 0 5 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0

0 2 5 5 9 0 0 3 0 0 0 5 3 2 3 2 3 4 0 0 4 6 1 22 3 6 8 0 0 1 6 5 2 0 5 6 0 7

0 9 5 1 0 0 0 0 6 0 2 0 3 5 0 0 0 1 0 0 1 0 3 1 0 0 0 0 2 2 2 0 0 1 0 0 2 2

0 0 1 1 0 0 3 0 1 3 0 4 0 3 0 0 0 0 0 5 1 0 1 1 3 5 0 0 2 0 1 0 0 1 1 0 1 4

13 0 0 0 28 0 0 0 0 0 0 2 3 1 1 5 2 0 24 4 0 0 7 1 0 2 0 45 2 3 1 78 2 0 0 48 1 2

4 1 1 0 7 0 0 0 1 0 0 0 1 0 0 4 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 2 0 0 0 0

12 1 0 2 2 0 0 1 1 0 0 1 4 12 5 1 3 3 6 5 1 5 5 13 6 3 9 15 17 20 11 8 14 10 13 37 6 8

1 4 1 1 2 0 0 0 0 0 3 0 3 0 2 16 4 5 2 7 3 0 5 3 6 3 5 2 1 0 2 0 0 0 0 0 0 4

0 0 1 6 0 0 0 9 1 1 0 0 0 0 2 0 0 1 0 0 1 1 0 1 0 0 3 0 0 0 1 0 0 0 1 0 0 0

0 0 5 19 0 2 0 10 16 1 20 11 0 4 11 1 12 4 0 0 8 15 3 9 2 14 12 2 8 11 23 0 3 6 17 0 16 12

2 9 7 1 7 44 53 6 8 4 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

15 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 1 4 0 0 12 1 0 2 4 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0

0 0 0 3 0 1 0 0 1 3 0 2 0 5 0 0 0 1 0 4 1 1 1 1 0 5 0 0 1 0 0 0 6 1 0 1 2 1

5 1 0 0 0 0 0 0 5 0 2 0 0 0 0 1 0 0 6 1 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 1

15 2 0 1 5 1 0 0 2 3 2 2 0 2 2 1 1 1 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 2

0 18 18 5 5 10 5 13 8 10 23 14 31 2 13 27 15 20 4 32 15 10 42 0 29 20 9 3 8 1 1 3 5 5 0 1 3 9

0 2 17 6 0 11 1 12 2 6 8 10 18 26 23 7 9 27 0 8 22 23 17 23 6 8 10 0 7 8 17 3 12 22 15 0 17 8

0 0 5 4 0 4 0 10 13 4 8 19 7 7 13 8 12 10 0 1 13 11 7 4 4 12 6 0 1 11 1 0 5 10 3 0 2 17

1 1 1 0 0 1 5 3 0 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 4 1 3 0 1 0 0 3 3 0 0 3 4 0 9 0 0 1 2 4 1 0 0 1 0 0 0 2 1 0 0 6 0 0 4 1

0 0 0 1 0 0 0 0 0 0 2 0 0 2 0 0 2 0 0 0 0 5 0 1 0 0 14 0 0 0 2 0 0 2 10 0 0 0

Table 2. Relative abundances of the common (>5% in at least one sample) benthic foraminifera living in surface sediments in the southeastern Levantine Basin.

277 278 279

4 Results

280 281 282 283 284 285 286 287 288 289 290 291

4.1. Environmental properties Sediment and water properties vary along the middle-outer Israeli shelf (Fig. 2, Table 1). The sediments at the 40 m depth contour are sandier and the water contains higher Chl-a concentration than the 60 - 100 m water depths (43±18% vs. 16±13% on average, and 0.4±0.1 vs. 0.2±0.03 mg/m3 on average, respectively). Silts comprise the majority of the finer sediments (up to 82%) between 60 and 100 m water depths. These sediments are relatively rich in TOC content, increasing westward along with the decrease in grain size and ranging from 0.7±0.1 wt.% on average, at 60-80 m water depths to 0.9±0.2 wt.% on average, at 100 m water depth. Moreover, a coarsening trend accompanied by elevated CaCO3 concentrations is observed between the central part of the shelf (around Poleg) and further north mainly

292 293 294

295 296

at 40-60 m water depths. This trend is more pronounced in the middle shelf and is accompanied by a decrease in TOC content and Chl-a concentration.

297 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 321 322 323 324 325 326 327 328 329 330 331 332 333 334 335 336 337 338 339 340 341 342 343 344 345 346

Fig. 2. Distribution patterns of environmental variables including sand silt and clay contents (wt.%), CaCO3 (wt.%), TOC (wt.%) and Chl-a (mg/m3) plotted on a bathymetric chart of the SE Levantine shelf. See also Table 1. 4.2. Distribution patterns of live foraminiferal communities A total of 105 live taxa were identified during the current study, 73 of which identified at species level (Supplementary data No. 1). The relative abundances of the common species (>5% in at least one sample) are presented in Figure 3 and Table 2. The most abundant agglutinant species are Reophax scorpiurus, Lagenammina sp., Glomospira charoides, Eggerelloides scaber and Adercotryma glomeratum that live mostly in the outer shelf between 60 and 100 m, whereas Deuterammina rotaliformis occupy mostly the middle shelf sediments at 40 m water depth (Fig. 4). Among the common calcareous species Rectuvigerina elongatastriata and Hanzawaia rhodiensis, both, are living in the deeper water samples, however the latter one is more frequent in the northern outer shelf. Q-mode cluster analysis (CA) visually identified in the dendrogram four clusters of samples of the living benthic faunas encompassing specific geographic borderlines: clusters 1a, 1b, 2 and 3 (Fig. 5A, B). The most significant species, contributing to each cluster are shown with their relative contribution based on the ‘similarity percentages’ (SIMPER) routine in Fig. 5 (Supplementary data No. 2). Clusters 1a and 1b represent all the samples along the 40 m isobath from Ashqelon to Akhziv (Fig. 5A, B). The total abundances of the benthic foraminifera in these samples were below 3 BF/g. Species richness (per sample) in Cluster 1a, that includes all the southern sites didn’t exceed 12 species, whereas, in Cluster 1b that occurs north of Herzeliyya, it ranged between 17 and 23 species (Figs. 5A, B; Supplementary data No. 1 and 2). The samples of Cluster 1a are characterized by moderate amounts of sandy sediments with 17 - 32% quartz sands (27±9% on average) and the highest values of Chl-a (0.5±0.1 mg/m3 on average) (Table 3). This cluster was dominated by D. rotaliformis, E. scaber and R. elongatastriata (Figs. 4 and 5A; Supplementary data No. 2). The sediments of Cluster 1b samples are coarser and the sandy fraction that ranged between 43 and 76% (63±18% on average) was relatively enriched in carbonates (Tables 1 and 3). The most significant species in this cluster are D. rotaliformis (though with lower contribution than in cluster 1a), Ammonia tepida, E. scaber and Quinqueloculina schlumbergeri (Figs. 1, 4 and 5A, B; Supplementary data No. 1 and 2). Cluster 2 consists of clayey-silty samples from water depths between 60 and 100 m that occur in all sites located south of Michmoret and additional two samples from northern sites located at 100 m water depth. Silt concentrations were the highest in these sites ranging from 66 to 81% (74±4% on average) with 0.6–1 wt.% TOC (Tables 1 and 3). The total abundances of the benthic foraminifera in these samples was relatively higher ranging between 4 and 23 BF/g, and species richness per sample varied between 15 and 37 species (Figs. 1, 4 and 5A, B; Supplementary data No. 1). The most significant species contributing, in descending order, to the similarity between the samples of this cluster are: Lagenammina sp., R. elongatastriata, R. scorpiurus, A. glomeratum, G. charoides and E. scaber (Fig. 4; Supplementary data No. 1 and 2). Cluster 3 contains samples from water depth between 60 and 100 m from Cesarea and further to the north with lower amounts of silts compared with Cluster 2 and higher content of CaCO3, between 10 and 12% (11±1% on average). The total

347 348 349 350 351 352

353 354 355 356 357 358 359

abundances of the benthic foraminifera in all these samples were below 7 BF/g, while species richness usually fluctuated between 18 and 34 species per sample (Figs. 1 and 5A, B; Supplementary data No. 1). The most significant species contributing to this cluster are as follows: R. elongatastriata, Lagenammina sp. R. scorpiurus, H. rhodiensis and A. glomeratum (Fig. 4; Supplementary data No. 1 and 2).

Fig. 3. Relative abundances of the common (>5% in at least one sample) benthic foraminifera living in surface sediments in the southeastern Levantine Basin. The 59 studied stations were ordered along North-South oriented transects, at 40, 60, 80 and 100 m water depths. See also Table 2.

360 361 362 363 364

Fig. 4. Living (stained) benthic foraminifera collected from surface sediments from the Levantine shelf (SE Mediterranean), 40–100 m water depths. The scale bars of all specimens equal100 µm.

365 366 367 368 369 370 371 372 373 374 375 376 377 378

(1) Ammodiscus sp. lateral view. (2a) Ammonia tepida (Cushman, 1926) umbilical view. (2b) A. tepida spiral view. (3a) Asterigerinata mamilla (Williamson, 1858) umbilical view. (3b) A. mamilla spiral view. (4) Astrononion stelligerum (d'Orbigny, 1839), side view. (5) Biloculinella subsphaerica (Wiesner, 1923) side view. (6) Bolivina striatula Cushman, 1922, side view. (7a) Buccella granulata (di Napoli Alliata, 1952) umbilical view. (7b) B. granulata spiral view. (8) Cribroelphidium poeyanum (d'Orbigny, 1839) side view. (9) Adercotryma glomeratum (Cushman, 1910) side view. (10a) Deuterammina rotaliformis (Heron-Allen & Earland, 1911) umbilical view. (10b) D. rotaliformis, spiral view. (11) Discammina compressa (Goës, 1882) side view. (12) Eggerelloides scaber (Williamson, 1858), side view. (13) Fursenkoina subacuta (d'Orbigny, 1852), side view. (14) Globobulimina affinis (d'Orbigny, 1839) side view. (15a–b) Glomospira charoides (Jones & Parker, 1860). (16a) Hanzawaia rhodiensis (Terquem, 1878) umbilical view. (16b) H. rhodiensis spiral view. (17) Lagenammina sp. side view. (18a) Pararotalia

379 380 381 382 383 384

calcariformata McCulloch, 1977 umbilical view. (18b) P. calcariformata, spiral view. (19) Planorbulina mediterranensis d'Orbigny, 1826, the attached side. (20) Quinqueloculina schlumbergeri (Wiesner, 1923), side view. (21) Rectuvigerina elongatastriata (Colom, 1952), side view. (22) Reophax scorpiurus Montfort, 1808, side view. (23a) Valvulineria bradyana (Fornasini, 1900) umbilical view. (23b) V. bradyana spiral view.

385 386 387 388 389 390 391 392 393 394 395 396

Fig. 5. A. Dendrogram of Q-mode Cluster Analysis (CA) of the 38 samples from the southeastern Mediterranean shelf (40-100 m water depth), using group-average linking of Bray–Curtis similarities calculated on square root transformed count data of the common benthic foraminiferal species (samples with >5% relative abundance in at least one sample). Taxa that make significant contributions to the similarity within each cluster, based on SIMPER routine (Supplementary data No. 2), are shown, with their % contribution. B. Non-metric multi-dimensional scaling (MDS) illustration in two dimensions using the same similarity matrix of the hierarchical clustering (A). Notice the good separation in the MDS diagram of the four assemblages and one outlier distinguished in the dendrogram.

397 398 399

Cluster

Number of samples

Depth range [m]

Geographic borderlines

Sand (63– 2000µm)

Silt (2– 63µm)

Clay (0.02– 2 µm)

CaCO3

TOC

Chl-a (mg/m3)

% 1a

400 401 402 403 404 405 406 407 408 409 410 411 412 413 414 415 416 417 418 419 420 421 422 423 424 425 426 427 428 429 430 431 432 433 434 435 436 437 438 439 440

3

1b

3

2 3

24 7

40 40 60-100 60-100

Southern Middle Shelf Central-Northern Middle Shelf Central-Southern OuterShelf Northen Outer Shelf

27±9

62±7

11±2

10±2

0.6±0.0

0.5±0.1

63±18

32±15

5±3

16±3

0.3±0.1

0.3±0.1

10±5

74±4

16±4

9±1

0.7±0.1

0.2±0.0

15±10

65±9

20±9

11±1

0.7±0.2

0.2±0.0

Table 3. Clusters’ characteristics. 4.3. Relationships between foraminiferal composition and environmental variables via ordination technique Table 4 shows that the short length of the first DCA axis in turnover units is rather short (2.49 SD) suggesting that the foraminifera demonstrate a linear response to one or more environmental gradients and thus a linear ordination method (i.e., RDA) for the classification of foraminiferal species–environmental relationships is expected to perform well. The Q-mode CA results were reinforced by the RDA which constrained the relationships between the foraminiferal assemblages representing each sample cluster and the most important environmental drivers significantly influencing their distribution (Fig. 6; Supplementary data No. 1 and 2). Monte Carlo tests suggest a significant influence (p < 0.05; Table 5) of all environmental parameter (sand, silt, clay, TOC, CaCO3 and Chl-a content) on the species distribution in our data set. The silt, sand fractions and TOC, each explain 1617% of the variance in the data set. Chl-a, carbonate and clay fraction content justify 14.3%, 11.7% and 7% of the variance in the data set, respectively. These significant environmental variables are correlated to the first two axes, calculated with the RDA, which explain 54% of the cumulative variance of the species data and 83% of the species- environment relationship (Table 5, Fig. 6). In general, the results of the RDA (Fig. 6A) support the Q-mode CA (Figs. 5 A, B). The distance between the samples with relatively good separation of the four assemblages is distinguished in the dendrogram (Figs. 5 A, B and 6A ) and shows the dissimilarities between the sample clusters and the similarities within each sample cluster. The similarities within cluster 2 are clearly the greatest. In the RDA ordination diagram cluster 1a samples, representing the southernmost middle shelf at 40 m water depth, are plotting around Chl-a concentrations whereas the rest of the shallow water samples from similar water depths of cluster 1b from the central-northern shelf are plotting close to the sand fraction and to some extent to Chl-a concentrations and carbonate contents (Fig. 6A). Cluster 2 samples, representing the central-southern outer-shelf sediments, from deeper water depths (60-100 m) are plotting close to silt and clay fractions and TOC and mostly in the opposite direction to clusters 1a and 1b samples. Samples of cluster 3 are also from deeper water depths (60-100 m) however from the northern outershelf and are plotting mostly close to the clay fraction. The relationships between, the environmental parameters (Table 5) and abundance of the 27 foraminiferal species are shown in Fig. 6B. Strong correlations (positive or negative) are based on the arrows lengths and their directions calculated by the RDA and combined and tested with the parametric correlation coefficient

441 442 443 444 445 446 447 448 449 450 451 452 453 454 455 456 457 458 459 460

Pearson's r. Deuterammina rotaliformis is dominating the foraminiferal assemblage of cluster 1a samples and is positively and strongly (r = 0.8) correlated with Chl-a concentrations (Figs. 5A, 6A, B; Supplementary data No. 1). Ammonia tepida and Q. schlumbergeri (associated with cluster samples 1b) are positively correlated (r = 0.5) with sand fraction. These species are negatively correlated with TOC (r values range from -0.5 to -0.7). Hanzawaia rhodiensis (with the highest relative abundance in cluster 3), and Rosalina sp. are positively and relatively strongly (r values range from 0.6 to 0.8) correlated with carbonate content (Figs. 5A, 6A, B; Supplementary data No. 1). The relative abundance of species grouped in cluster 2 increases in the centralsouthern outer-shelf (60–100 m), indicating a positive relationship with silt, clay and TOC (Figs. 4, 5A, 6A, B). Lagenammina sp. indicates a close relationship with these three parameters demonstrating positive correlations (r values range from 0.4 to 0.5). Although the arrows of these environmental parameters point relatively to the same directions, several species seem to have stronger correlations primarily with one of them, for example R. scorpiurus has stronger correlation (r = 0.5) mainly with silt fraction. Whereas, G. charoides and Valvulineria bradyana are positively correlated with TOC (r = 0.5) and Buccella granulata is positively correlated with clay (r = 0.5). Other foraminiferal species shown in the RDA (Fig. 6B) such as R. elongatastriata seem to have very weak correlations with the investigated parameters.

Eigenvalues Explained variation (cumulative) Gradient length

461

Axis 1 0.2701

Axis 2 0.1956

Axis 3 0.057

Axis 4 0.0326

23.7

40.86

45.87

48.73

2.49

2.16

1.16

1.05

Table 4. Statistical results of Detrended Correspondence Analysis (DCA).

462 Capture variance (%)

pseudoF-value

P value

3.9

0.0005

% Silt

7.6

0.0005

17.5

% Sand

7.2

0.0005

16.6

TOC (wt.%)

6.9

0.0005

16.1

Chl-a (Annual Averege)

6.0

0.0005

14.3

CaCO3 (wt.%)

4.8

0.0005

11.7

% Clay

2.7

0.015

7.0

Eigenvalues Explained variation (cumulative percentage variance of species data ) Pseudo-canonical correlation (Speciesenvironment correlations) Explained fitted variation (cumulative percentage variance of speciesenvironment relation)

Axis 1

Axis 2

Axis 3

Axis 4

0.2241

0.0904

0.0399

0.0175

22.41

31.45

35.44

37.18

0.8311

0.7963

0.7139

0.6396

59.2

83.09

93.62

98.23

Permutation Test Results (On All Axes) Correlations

463

Table 5. Statistical results of Redundancy Analysis (RDA).

464 465 466 467

468 469 470 471 472 473 474 475 476

Fig. 6. A. Redundancy Analysis (RDA) for all 38 surface sediment samples showing retrospective projection of the sample—environmental variables in the upper diagram. Clusters sample distinguished in the Q-mode CA (Fig. 5) were incorporated into the RDA results. B. The species–environment relationships are shown on biplot diagram summarizing the effects of sand (% >63 µm size fraction), silt (% 2-63 µm size fraction), clay (% <2 µm size fraction), CaCO3 content (wt.%), Chl-a (mg/m3) and TOC (wt.%), upon the benthic foraminiferal distribution.

477 478 479 480 481

5 Discussion 5.1. Relationship between living species and abiotic environmental parameters as a base for spatial biotopes delineation

482 483 484 485 486 487 488 489 490 491 492 493 494 495 496 497 498 499 500 501 502 503 504 505 506 507 508 509 510 511 512 513 514 515 516 517 518 519 520 521 522 523 524 525 526 527 528 529 530

In general, the species richness values of the benthic foraminifera (BF) are lower than 37 and the total abundance is less than 23 BF/g per sample. These parameters were relatively variable in the middle and outer Levantine shelf. A patchy distribution of benthic foraminifera is a common characteristic in most shelf ecosystems and is accredited to highly variable environmental conditions (Murray, 2006). The LB foraminiferal species richness values are relatively lower than those in the siliciclastic shelf environments of the western Mediterranean Sea (Gulf of Lions, France) (Mojtahid et al., 2009; Goineau et al., 2011). The variability in living foraminiferal spatial distribution, composition and biodiversity in these environments is attributed mainly to eutrophic conditions controlled by the quality (fresh or degraded), quantity and origin (continental or marine) of the organic matter reaching the fine-grained sediment via varied hydro-sedimentary processes. Thus the overall low general foraminiferal abundances measured in the LB may largely be explained by the ultra-oligotrophy of this area compared to western Mediterranean shelf regions (de Rijk 1999, 2000). The living foraminiferal species exhibit a distinct lateral bathymetric zonation which follows changes in the environmental parameters (Figs. 5-8). The combined CA and RDA results, which were further confirmed by the parametric correlation coefficient Pearson's r, enabled the determination of four geographical coherent biotopes, with only two local exceptions (2 samples out of the 27 of biotope 2, which fall out of its geographic boarders) (Fig. 9). A general and clear faunal break is separating between the ~40 m water depth in which two biotopes were well defined, 1a and 1b, and the deeper part of the shelf, between 60 and 100 m, where biotopes 2 and 3 prevail (Fig. 9). The low species richness and the lowest foraminiferal densities which characterize the mid-shelf biotopes is explained by the coarser substrate which is accompanied by a decrease in organic matter content as shown also in previous studies (Hyams-Kaphzan et al., 2008, 2009; Avnaim-Katav et al., 2015: Tadir et al., 2017). Over the past 50 years, following the Nile River damming in 1965 and consequently cessation in its input of fines, sediments at ~40 m water depth within the Nile littoral cell became coarser and are composed at present by silty to muddy sand (Inman and Jenkins, 1984; Halim et al., 1995; Almogi-Labin et al., 2009, 2012). The comparison between the two mid-shelf biotopes reveals that the species which are dominating the foraminiferal assemblage of biotope 1a are E.scaber and especially D. rotaliformis. In this biotope, located proximal to the Nile Delta, Chl-a concentration is the highest along the entire Israeli shelf (Figs. 4-7). This observation is in agreement with Jannink, (2001) who showed maximal numbers of E. scaber following an increase in Chl-a concentrations in the central part of the Israeli mid shelf region. An opposite trend, however, was shown by Hyams-Kaphzan et al. (2009) and Tadir et al. (2017) who studied a highly disturbed eutrophic site controlled by an activated sewage sludge discharge. A broader range of ecological tolerance of Eggerelloides scaber is observed at the Ría de Vigo embayment located in the northwest of Spain (Diz and Francés, 2008) and from the French Mediterranean coasts regardless of the ecosystems quality (Barras et al., 2014; Jorissen et al., 2018). The central northern middle shelf biotope 1b is relatively richer in carbonate sandy sediments (Figs. 7, 9). Species positively correlated with this biotope based on the RDA results, include Lessepsian taxa such as Pararotalia calcariformata (Fig. 7C), and others such as Q. schlumbergeri and especially A. tepida already reported among the dominant recent species in the dead (Hyams-Kaphzan et al., 2008;

531 532 533 534 535 536 537 538 539 540 541 542 543 544 545 546 547 548 549 550 551 552 553 554 555 556 557 558 559 560 561 562 563 564 565 566 567 568 569 570 571 572 573 574 575 576 577 578 579 580

Avnaim-Katav et al., 2013, 2015) and the living assemblages (Hyams-Kaphzan et al., 2009). The northern outer shelf biotope 3 resembles biotope 1b by the relatively higher concentrations of carbonate compared to the other biotopes however it consists of much finer sediments (Figs. 7B, 9). Hanzawaia rhodiensis, A. mamilla and Rosalina spp. show a strong correlation with carbonate-rich sediments forming a distinct part of the assemblage in biotope 3 (Figs. 5A, 6A, B). Hanzawaia rhodiensis, known as a shallow infaunal specie, is capable of adapting to different environmental conditions (e.g., Jorissen, 1988; Jannink, 2001;Hyams-Kaphzan et al., 2008). Asterigerinata mamilla, occurs in the western Mediterranean between 40 and 80 m water depth where it is closely linked to the distribution of coarse-grained substrates rich in rhodolites (Milker et al., 2009). This species and Rosalina spp. as well, has an epiphytic mode of life and both are generally associated with carbonate-rich sediments (e.g., Jorissen, 1987; Kitazato, 1988; Milker et al., 2009). The central-southern outer-shelf biotope 2 has several similarities with biotope 3, in terms of relatively high TOC and clay contents and low Chl-a concentrations (Fig. 9). Disparities between these biotopes include higher silt concentrations in biotope 2 vs. biotope 3, accompanied by a relatively high foraminifera density and species richness (Figs. 1, 9). In general higher species richness is observed in biotopes 2 and 3, compared to the shallower biotopes 1a and 1b. This may be explained by the higher organic carbon contents associated to fine-grained sediments at these depths. The relationship between higher organic matter content in clayey sediment and water depth has been already documented while studying the distribution patterns of dead recent foraminiferal assemblages in the study area (Avnaim-Katav et al., 2015), and as also observed in other regions in the Mediterranean such as the Adriatic Sea (Jorissen, 1987). The most characteristic taxa, clearly related to biotope 2, are Lagenammina sp., R. scorpiurus, G. charoides, V. bradyana and B. striatula (Figs. 6, 8, 9). Lagenammina sp. is shown to be positively correlated with silt, clay and TOC while R. scorpiurus has specific stronger correlation with silt. These findings accord well with available records known from the deeper part of Iskenderun Bay (eastern Turkey) showing that these species are inhibiting fine grained sediments rich in terrigenous mud with low CaCO3 and relatively high organic matter contents (~615%) (Basso & Spezzaferri, 2000). Jorissen (1987) showed that V. bradyana has a minimum water depth of 40 m in the Adriatic Sea in the outer part of the organic carbon enriched clay belt. In the Rhône prodelta, where the organic carbon content is almost double (0.8–1.9%) than in our biotope 2, V. bradyana proliferate in a diverse assemblage, responding to a higher proportion of marine organic matter, at its deepest distal part, sheltered from the stressful river mouth influence (Mojtahid et al., 2009). Additionally, V. bradyana is reported to be opportunistic species able to respond to seasonal variability i.e., fresh organic phytodetritus input with increased reproduction rates (Goineau et al., 2015 and references therein). In the Rhône prodelta, higher relative contributions of this living species was reported at the end of the spring bloom than during the summer time when conditions were more oligotrophic (Goineau et al., 2011). Moreover, Frezza and Carboni (2009) found similar percentages of this species and at similar water depth of biotope 2 in front of the Ombrone River mouth (Tyrrhenian Sea, Italy). Thus, the relatively lower abundance (max of 14 %) of this species in our deepest sites (100 m water depths) within biotope 2, may be primarily attributed to the relatively elevated TOC values characterizing this biotope compared to the oligotrophic conditions prevailing mostly in the other

581 582 583 584 585 586 587 588 589 590 591 592

593 594 595 596 597 598

shallower sites and subordinately to the sampling period, used in this study (summer time). This finding agrees well with Barras el al. (2014) which designated it as a “stress-tolerant” species, when occurring in high percentages it is indicating stressed conditions, such as eutrophication or increasing contribution of fine-grained sediments. Glomospira charoides is shown to be positively correlated with silt, clay and TOC, occurring mainly at the 100 m water depth sediments. This species is reported from deeper muds, even below 1000 m depth in the Mediterranean Sea (Sgarrella and Moncharmont-Zei, 1993; Hyams-Kaphzan et al., 2018). Whereas in the Eastern Mediterranean it is usually reported at depths deeper than ~ 100 m with maximum abundances at ~600 m depth in clayey sediments consisting mostly kaolinite(Sgarrella and Moncharmont-Zei, 1993).

Fig. 7. Distribution patterns of specific species (%) against particular environmental variables plotted on a bathymetric chart of the SE Levantine shelf. The biotic and abiotic variables were chosen based on statistics applied in the current study including CA, RDA and the parametric correlation coefficient Pearson's r (full explanations –

599 600 601 602

within the text). A. Deuterammina rotaliformis vs. Chl-a concentrations (r = 0.9). B. Hanzawaia rhodiensis vs. CaCO3 contents (r = 0.6). C. Pararotalia calcariformata vs. sand concentrations (r = 0.5).

603 604 605 606 607 608 609 610 611 612

Fig. 8. Distribution patterns of specific species (%) against particular environmental variables plotted on a bathymetric chart of the SE Levantine shelf. The biotic and abiotic variables were chosen based on statistics applied in the current study including CA, RDA and the parametric correlation coefficient Pearson's r (full explanations – within the text). A. Reophax scorpiurus vs. silt concentrations (r = 0.6). B. Glomospira charoides vs. clay concentrations (r = 0.4). C. Bolivina striatula vs. TOC contents (r = 0.4).

613

614 615 616 617 618 619

Fig. 9. Spatial delineation of the four biotopes identified in this study on top of sampling site locations as resulted from ordination and classification analyses shown as colored shapes (Figs. 5 and 6) and black dots (analyzed samples excluded from the statistical methods) are shown on the bathymetric map, following Sade et al. (2006).

620 621 622 623 624 625 626 627

5.2. Decadal changes of the live foraminiferal community along the ~40 m water depth Since the damming of the Nile in 1965 the Israeli shelf sediment properties rapidly change, affecting to a large degree the 40 m depth interval located between the sandy and the fine-grain sediment belts (Almogi-Labin et al., 2009, 2010, 2012). The changes at 40 m include ongoing coarsening of grain size, increase in carbonate content and decrease in TOC (Almogi-Labin et al., 2009; Tadir et al., 2017; Tibor et

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al., 2019). Taking into account that these variables might affect the living assemblages, we compared the 40 m records of living foraminifera, sampled during August 2011 with those sampled in May 1997 (Hyams, 2000) and between 1996 and 1998 (Jannink, 2001). The northern most Akhziv station studied by Hyams (2000) was not included in this comparison because the seafloor at this station became lately rocky and therefore could not be sampled by a box-corer during the 2011 sampling campaign. In these studies the foraminifera were analyzed in the >150 µm size fraction and the >63 µm size fraction, respectively. It should be noted that comparing results determined by using different methodologies, for example the usage of different size fraction for the analyses, might introduce a possible bias of assemblage data in terms of abundance and species richness: both could be higher in the smaller fraction analyzed (e.g., Schönfeld et al., 2012). Despite this bias, the general trends for environmental interpretations remain the same for the 125 µm and 150 µm, regardless of the chosen size fraction (Weinkauf & Milker, 2018). Thus, we proceed with our comparison in order to evaluate the influence of decadal changes in the sedimentary regime on the foraminifera. The total abundance of the benthic foraminifera in the late 90th is quite similar to the current study being usually below 3 BF/g. Unlike the nearly stable values of the numerical abundance of the BF since the 90th, species richness had declined considerably. In the late 90th species richness varied between 23 and 41 species per sample, on average (Hyams, 2000, and Jannink, 2001, respectively) compared to 12 species per site, on average, in the current study indicating a considerable decline, by approximately 30 to 50%, towards 2011. This decrease might be related to the continuous impact of the damming of the Nile River, the main source of nutrients in this region until 1965 (Said, 1993; Krom et al., 1999; Herut et al., 2000). The lower species richness compared to earlier studies in the region (Hyams, 2000; Jannink, 2001; Hyams-Kaphzan et al., 2009; Tadir et al., 2017) seems to reflect a general ongoing trend of increasing oligotrophy in the studied area. Concerning the living foraminiferal assemblage composition, three major differences are observed while comparing between the late 90th and the current 2011 situation: 1. Deuterammina rotaliformis dominate the assemblage in 2011 along the entire 40 m belt (Figs. 3-5), comprising 33% on average, as compared to its absence in earlier studies (Hyams, 2000; Jannink, 2001; Hyams-Kaphzan et al., 2009). This specie is identified in the Mediterranean Sea for the first time in the western Pontine Archipelago, Tyrrhenian Sea, in a living foraminiferal assemblage constituted merely of agglutinated species, with low abundance and diversity in sandy sediments, at 127– 137 m water depth, a region influenced by hydrothermal emission. (Di Bella et al., 2016). Similarly, it was found as the most frequent species in Panarea Island, Aeolian Archipelago at north of Sicily in the southern sector of Tyrrhenian Sea (Panieri et al., 2005), at 12 m water depth, also an area influenced by hydrothermal emission (Di Bella, personal communication). Interestingly, this species does not occur in the dead assemblages of the current dataset (in preparation) and the Tyrrhenian Sea samples (Di Bella, personal communication) neither it occurs in the paleo (fossil) records from the Israeli region as well (e.g., Avnaim-Katav., 2013, 2016b, 2019). Consequently, its appearance, merely in the last decade in increased numbers may be attributed to two possibilities: a) it is a fragile organically-cemented agglutinated species subjected to taphonomic processes such as shell destruction after death (Schröder, 1988; Bender, 1995; Edelman-Furstenberg et al., 2001) and thus only present in the live assemblage; b) it is a new recent entry introduced to the East Mediterranean either by shipping or

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aquaculture pathways (Zenetos et al., 2012). Its success in inhabiting the 40 m depth biotope is mostly attributed to its ability to occupy this rapidly changing environment affected considerably by ongoing grain size coarsening (discussed in Chapter 5.1). 2. A remarkable drop in the numerical abundances of A. mamilla in the 2011 campaign compared to 1997 (3% versus 10% on average, respectively), and to some extent of A. tepida (3% versus 6% on average). Both of these species were mostly documented in the southern part of the shelf (Ashqelon to Hadera). The first species is an epifaunal suspension feeder and is permanently or temporarily attached to coarse substrates, such as bioclastic sands (Murray, 2006; Milker et al., 2009). However, in the study area it lives at the surface and deeper down to 5 cm in the sediment even in higher numbers (Hyams, 2000). The infaunal preference may be attributed to presence of more organic matter at depth as a food resource, although it may represent species response to environmental controls such as sediment disturbance, burrowing or predation (Jorissen, 1999 and references therein). Ammonia tepida, is an opportunistic species known to occur in maximal numbers at the sediment surface when food resources are abundant (Hyams-Kaphzan et al., 2009; Jorissen et al., 2018). It is associated with anthropogenic pollution (Burone et al., 2006) such as an activated sewage sludge discharge (Hyams-Kaphzan et al., 2009; Tadir et al., 2017). The decrease of this species shown in this study was already documented by Tadir et al. (2017) who compared foraminiferal records in three stations near Palmahim between the years 2003 and 2012. The ongoing far-reaching decreased numbers of this species probably represent continued reduction in TOC content in the oligotrophic eastern Mediterranean attributed to the damming of the Nile and its reduced nutrients input (Almogi-Labin et al., 2009). 3. The occurrence of the following symbiont bearing alien/ Lessepsian species that invaded the southeastern Mediterranean from the Red Sea through the Suez Canal (Hyams et al., 2002; Hyams-Kaphzan et al., 2008; Langer and Hottinger, 2000; Zenetos et al., 2012): Amphistegina lobifera, Hauerina diversa, Heterostegina depressa and P. calcariformata. These species usually live on rocky substrate in areas covered by macroalgae (Hyams-Kaphzan et al., 2008). Hyams-Kaphzan et al. (2014) reported that all these species were found alive at hard bottom/rocky habitats down to 18 m water depth off the northern Israeli coast (Carmel Head and Akhziv). Dead specimens of A. lobifera were recorded by Hyams et al. (2002) down to 25 m water depth. In the current study, these species (except H. depressa) are reported for the first time as expanding their depth habitat, to 40 m water depth along the central northern part of the shelf. Thus our findings indicate the ability of these alien/ Lessepsian species to live also in deeper water habitats of the Israeli shelf, because they became more suitable for them. 5.3. Classification of species for the Mediterranean FORAM-AMBI The current study might serve as a new additional independent dataset which adds new useful information for foraminiferal bio-monitoring of soft substrates in the Mediterranean Sea, which can be used in different ecological indices such as TSIMed, Foram-AMBI and similar methods (Barras et al., 2014; Jorissen et al., 2018). Among the 27 dominant (>5% in at least one sample) species living in the study area, four are classified as 3rd order opportunists including; B. striatula, Cribroelphidium poeyanum, E. scaber and Textularia bocki. Additional two species are considered as 2nd order opportunists including: A. tepida and V. bradyana. The relative proportions

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of opportunistic species assigned to both of these categories are lower than 20%, as expected in the study area considered as a natural oligotrophic environment, excluding 4 samples from the southern shelf proximal to the Nile Delta in which their contribution reached up to 43%. It is worth mentioning that this group has higher densities in the dead assemblages (paper under preparation), representing predamming Nile faunas with enhanced nutrients input (e.g., Almogi-Labin et al., 2009). Consequently, the comparison of live and dead faunas deserve to be further explored aiming at gathering potential information regarding the opportunistic specie response to ecosystem organic carbon impoverishment. Such information is required to test earlier assignments of common Mediterranean benthic foraminiferal taxa (i.e., stresstolerant and/or stress-sensitive taxa) to ecological categories serving as an essential tool for bio-monitoring (Barras et al., 2014; Jorissen et al., 2018).

New insights on the abundance, diversity, spatial distribution and ecological preferences of living benthic foraminifera in the southeastern Levantine shelf (40–100 m water depths) are presented and compared to previous records from this region. Quantifications of the environmental drivers controlling the distribution patterns of foraminiferal assemblages in an area which recently (last 150 years) has been severely affected by human activity (opening of the Suez Canal and damming of the Nile River) adds information on specific species habitat preferences on a wider scale and their potential use as environmental bio-indicators. The 40 m depth habitat represents the most vulnerable environment for the living benthic foraminifera of the southeastern Mediterranean shelf. At this depth ongoing sediment coarsening, continuous decrease in TOC and increase in CaCO3 content started several decades ago and still continues. This seems to affect mainly species richness which decreased significantly in 2011 compared to the late 90s living faunas records reflecting the ongoing increasing oligotrophy in the region, mainly due to the recent shutdown of the Nile River discharge to the southeastern Mediterranean Sea.

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Some symbiont bearing Lessepsian species that were distributed in the late90th mainly down to a water depth of 10-25 m start inhabiting the 40 m depth zone indicating availability of suitable bottom water conditions for these species.

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Deuterammina rotaliformis, that recently was found living in central Mediterranean (Di Bella et al., 2016) appeared recently in the southeastern Levantine coast becoming the dominant species at the southern and central parts of the 40 m depth zone.

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Finally, for the first time, this study illustrate how the benthic foraminifera living in the SE Levantine shelf respond to recent environmental changes. This study would benefit from additional research to further support these data; and moreover on the surface dead assemblages to be compared with the living assemblages in order to improve our understanding of the environmental processes acting on benthic foraminiferal assemblages in this area.

6 Conclusions

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Acknowledgments This study was carried out with the support of the Ministry of Energy Grant (no. 01406-211) within the project “Research and Monitoring the Israeli Mediterranean continental shelf as a base for sustainable decisions on marine infrastructures”. We thank the Israeli Ministries of Environmental Protection and Energy for partial support in frame of the National Monitoring Program of Israel's Mediterranean coastal waters. We gratefully acknowledge John Hall for the constructive suggestions regarding the sampling design and the crew of the R/V Shikmona (IOLR), for their dedicated work during the sampling campaign. Thanks are due to Yoav Ben-Dor and Gloria Lopez for technical assistance in sample collection. Michael Kitin, Ruth Binstock, Stephen Abrahams, Hadar Elyashiv, Gideon Tibor, Lana Ashkar from the GSI and IOLR, are highly appreciated for the different analyses carried on during this project. Editor Daniel Baird, reviewer Pia Nardelli and the other anonymous reviewer are deeply appreciated for their constructive comments, suggestions and helpful revision resulting in a significantly improved manuscript.

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Table captions Table 1. Station locations, water depths and coordinates of surface samples taken with a box corer by the R/V Shikmona during August 29-31, 2011 research cruise. The environmental dataset utilized in the current study, includes the following sediment properties: percentage of the clay, silt and sand fractions, TOC (wt.%) and CaCO3 content (wt.%), all from Almogi-Labin et al. (2012) and Chl-a (mg/m^3). (nd = no data).

1096 1097 1098

Table 2. Relative abundances of the common (>5% in at least one sample) benthic foraminifera living in surface sediments in the southeastern Levantine Basin.

1099 1100

Table 3. Clusters’ characteristics.

1101 1102

Table 4. Statistical results of Detrended Correspondence Analysis (DCA).

1103 1104

Table 5. Statistical results of Redundancy Analysis (RDA).

1105 1106 1107

Figure captions

1108 1109 1110 1111 1112 1113 1114 1115

Fig. 1. (A) The study area, located in the southeastern Levantine Basin, eastern Mediterranean, is part of the Nile littoral cell that extends up to northern Israel and is influenced by the anticlockwise long-shore current (LSC); (B) The sampling sites located along the SE Levantine shelf at 40, 60, 80 and 100 m water depths are shown on the bathymetric map, following Sade et al. (2006). See also Table 1 for details. Overlaid colored areas designate the general abundance and species diversity per site. Legend of BF/g (large circles) and simple diversity (no. of species, small circles) determined in ArcGIS by using natural breaks method classification.

1116 1117 1118 1119

Fig. 2. Distribution patterns of environmental variables including sand silt and clay contents (wt.%), CaCO3 (wt.%), TOC (wt.%) and Chl-a (mg/m3) plotted on a bathymetric chart of the SE Levantine shelf. See also Table 1.

1120 1121 1122 1123 1124

Fig. 3. Relative abundances of the common (>5% in at least one sample) benthic foraminifera living in surface sediments in the southeastern Levantine Basin. The 59 studied stations were ordered along North-South oriented transects, at 40, 60, 80 and 100 m water depths. See also Table 2.

1125 1126 1127 1128

Fig. 4. Living (stained) benthic foraminifera collected from surface sediments from the Levantine shelf (SE Mediterranean), 40–100 m water depths. The scale bars of all specimens equal100 µm.

1129 1130 1131 1132 1133 1134 1135 1136 1137 1138 1139 1140 1141 1142 1143 1144 1145 1146 1147 1148

(1) Ammodiscus sp. lateral view. (2a) Ammonia tepida (Cushman, 1926) umbilical view. (2b) A. tepida spiral view. (3a) Asterigerinata mamilla (Williamson, 1858) umbilical view. (3b) A. mamilla spiral view. (4) Astrononion stelligerum (d'Orbigny, 1839), side view. (5) Biloculinella subsphaerica (Wiesner, 1923) side view. (6) Bolivina striatula Cushman, 1922, side view. (7a) Buccella granulata (di Napoli Alliata, 1952) umbilical view. (7b) B. granulata spiral view. (8) Cribroelphidium poeyanum (d'Orbigny, 1839) side view. (9) Adercotryma glomeratum (Cushman, 1910) side view. (10a) Deuterammina rotaliformis (Heron-Allen & Earland, 1911) umbilical view. (10b) D. rotaliformis, spiral view. (11) Discammina compressa (Goës, 1882) side view. (12) Eggerelloides scaber (Williamson, 1858), side view. (13) Fursenkoina subacuta (d'Orbigny, 1852), side view. (14) Globobulimina affinis (d'Orbigny, 1839) side view. (15a–b) Glomospira charoides (Jones & Parker, 1860). (16a) Hanzawaia rhodiensis (Terquem, 1878) umbilical view. (16b) H. rhodiensis spiral view. (17) Lagenammina sp. side view. (18a) Pararotalia calcariformata McCulloch, 1977 umbilical view. (18b) P. calcariformata, spiral view. (19) Planorbulina mediterranensis d'Orbigny, 1826, the attached side. (20) Quinqueloculina schlumbergeri (Wiesner, 1923), side view. (21) Rectuvigerina elongatastriata (Colom, 1952), side view. (22) Reophax scorpiurus Montfort, 1808, side view. (23a) Valvulineria bradyana (Fornasini, 1900) umbilical view. (23b) V. bradyana spiral view.

1149 1150 1151 1152 1153 1154 1155 1156 1157 1158

Fig. 5. A. Dendrogram of Q-mode Cluster Analysis (CA) of the 38 samples from the southeastern Mediterranean shelf (40-100 m water depth), using group-average linking of Bray–Curtis similarities calculated on square root transformed count data of the common benthic foraminiferal species (samples with >5% relative abundance in at least one sample). Taxa that make significant contributions to the similarity within each cluster, based on SIMPER routine (Supplementary data No. 2), are shown, with their % contribution. B. Non-metric multi-dimensional scaling (MDS) illustration in two dimensions using the same similarity matrix of the hierarchical clustering (A). Notice the good separation in the MDS diagram of the four assemblages and one outlier distinguished in the dendrogram.

1159 1160 1161 1162 1163 1164 1165 1166

Fig. 6. A. Redundancy Analysis (RDA) for all 38 surface sediment samples showing retrospective projection of the sample—environmental variables in the upper diagram. Clusters sample distinguished in the Q-mode CA (Fig. 5) were incorporated into the RDA results. B. The species–environment relationships are shown on biplot diagram summarizing the effects of sand (% >63 µm size fraction), silt (% 2-63 µm size fraction), clay (% <2 µm size fraction), CaCO3 content (wt.%), Chl-a (mg/m3) and TOC (wt.%), upon the benthic foraminiferal distribution.

1167 1168 1169 1170 1171

Fig. 7. Distribution patterns of specific species (%) against particular environmental variables plotted on a bathymetric chart of the SE Levantine shelf. The biotic and abiotic variables were chosen based on statistics applied in the current study including CA, RDA and the parametric correlation coefficient Pearson's r (full explanations –

1172 1173 1174

within the text). A. Deuterammina rotaliformis vs. Chl-a concentrations (r = 0.8). B. Hanzawaia rhodiensis vs. CaCO3 contents (r = 0.6). C. Pararotalia calcariformata vs. sand concentrations (r = 0.5).

1175 1176 1177 1178 1179 1180 1181 1182

Fig. 8. Distribution patterns of specific species (%) against particular environmental variables plotted on a bathymetric chart of the SE Levantine shelf. The biotic and abiotic variables were chosen based on statistics applied in the current study including CA, RDA and the parametric correlation coefficient Pearson's r (full explanations – within the text). A. Reophax scorpiurus vs. silt concentrations (r = 0.5). B. Glomospira charoides vs. clay concentrations (r = 0.5). C. Bolivina striatula vs. TOC contents (r = 0.4).

1183 1184 1185 1186 1187

Fig. 9. Spatial delineation of the four biotopes identified in this study on top of sampling site locations as resulted from ordination and classification analyses shown as colored shapes (Figs. 5 and 6) and black dots (analyzed samples excluded from the statistical methods) are shown on the bathymetric map, following Sade et al. (2006).

1188 1189 1190 1191 1192 1193

Supplementary data captions

1194 1195

Supplementary data No. 1. Census data set of the live benthic foraminiferal dataset from 0-1cm >125µm size fraction.

1196

Supplementary data No. 2. A similarity percentages” (SIMPER) routine.

Sample name

A. glomeratum

A. planorbis

A. tepida

A. mamilla

A. stelligerum

B. subsphaerica

B. striatula

B. granulata

C. poeyanum

D. rotaliformis

D. compressa

E. scaber

F. subacuta

G. affinis

G. charoides

H. rhodiensis

P. calcariformata

P. mediterranensis

Polymorphina sp.

Q. bosciana

Q. schlumbergeri .

R. elongatastriata

Lagenammina sp.

R. scorpiurus

Rosalina sp.

T. bocki

V. bradyana

2-40 2-60 2-80 2-100 3-40 3-80 4-80 5-100 6-100 7-60 7-80 7-100 8-60 8-80 8-100 9-60 9-80 9-100 10-40 10-60 10-80 10-100 11-80 11-100 12-60 12-80 12-100 13-40 13-60 13-80 13-100 14-40 14-60 14-80

1 5 13 2 0 1 1 1 6 9 3 6 4 6 3 8 16 3 0 4 12 5 0 3 27 6 12 0 36 35 9 0 23 21

0 0 0 3 3 0 0 1 0 0 0 7 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 1 1 0 0 2

7 4 0 0 9 1 0 0 1 9 3 1 0 3 0 0 0 0 9 8 1 0 0 0 0 0 1 0 0 0 0 0 0 0

2 1 1 0 0 2 7 0 1 0 0 0 3 3 0 0 1 0 0 0 0 0 0 0 0 0 1 2 1 0 0 0 0 0

5 2 0 0 3 3 4 0 2 9 3 1 8 1 1 2 4 6 0 1 8 1 1 1 0 1 0 0 7 0 2 2 16 1

0 0 0 0 0 0 5 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0

0 2 5 5 9 0 0 3 0 0 0 5 3 2 3 2 3 4 0 0 4 6 1 22 3 6 8 0 0 1 6 5 2 0

0 9 5 1 0 0 0 0 6 0 2 0 3 5 0 0 0 1 0 0 1 0 3 1 0 0 0 0 2 2 2 0 0 1

0 0 1 1 0 0 3 0 1 3 0 4 0 3 0 0 0 0 0 5 1 0 1 1 3 5 0 0 2 0 1 0 0 1

13 0 0 0 28 0 0 0 0 0 0 2 3 1 1 5 2 0 24 4 0 0 7 1 0 2 0 45 2 3 1 78 2 0

4 1 1 0 7 0 0 0 1 0 0 0 1 0 0 4 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 0 0 2

12 1 0 2 2 0 0 1 1 0 0 1 4 12 5 1 3 3 6 5 1 5 5 13 6 3 9 15 17 20 11 8 14 10

1 4 1 1 2 0 0 0 0 0 3 0 3 0 2 16 4 5 2 7 3 0 5 3 6 3 5 2 1 0 2 0 0 0

0 0 1 6 0 0 0 9 1 1 0 0 0 0 2 0 0 1 0 0 1 1 0 1 0 0 3 0 0 0 1 0 0 0

0 0 5 19 0 2 0 10 16 1 20 11 0 4 11 1 12 4 0 0 8 15 3 9 2 14 12 2 8 11 23 0 3 6

2 9 7 1 7 44 53 6 8 4 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

15 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 1 4 0 0 12 1 0 2 4 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0

0 0 0 3 0 1 0 0 1 3 0 2 0 5 0 0 0 1 0 4 1 1 1 1 0 5 0 0 1 0 0 0 6 1

5 1 0 0 0 0 0 0 5 0 2 0 0 0 0 1 0 0 6 1 0 0 0 0 1 1 1 0 0 0 1 0 0 0

15 2 0 1 5 1 0 0 2 3 2 2 0 2 2 1 1 1 2 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0

0 18 18 5 5 10 5 13 8 10 23 14 31 2 13 27 15 20 4 32 15 10 42 0 29 20 9 3 8 1 1 3 5 5

0 2 17 6 0 11 1 12 2 6 8 10 18 26 23 7 9 27 0 8 22 23 17 23 6 8 10 0 7 8 17 3 12 22

0 0 5 4 0 4 0 10 13 4 8 19 7 7 13 8 12 10 0 1 13 11 7 4 4 12 6 0 1 11 1 0 5 10

1 1 1 0 0 1 5 3 0 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 4 1 3 0 1 0 0 3 3 0 0 3 4 0 9 0 0 1 2 4 1 0 0 1 0 0 0 2 1 0 0 6

0 0 0 1 0 0 0 0 0 0 2 0 0 2 0 0 2 0 0 0 0 5 0 1 0 0 14 0 0 0 2 0 0 2

14-100 15-40 15-60 15-80

7 0 32 11

0 1 0 2

0 0 0 0

0 0 0 0

0 0 3 2

0 0 0 0

5 6 0 7

0 0 2 2

1 0 1 4

0 48 1 2

0 0 0 0

13 37 6 8

0 0 0 4

1 0 0 0

17 0 16 12

0 0 0 0

0 0 0 0

0 0 0 0

0 1 2 1

0 0 0 1

0 0 0 2

0 1 3 9

15 0 17 8

3 0 2 17

0 0 0 0

0 0 4 1

10 0 0 0

Highlights 3 to 5 bullet points / maximum 85 characters, including spaces, per bullet point Living foraminiferal distribution in the SE Levantine shelf (40 -100 water depth)

Multivariate analyses were used for exploring species – environment relations

Benthic foraminiferal bioindicators responded to the ongoing recent human activity

We identified four biotopes with distinct depth zonation, sediment type and Chl-a

1

Conflict of Interest and Authorship Conformation Form On behalf of all listed authors I Simona Avnaim-Katav

declare that:

All authors have participated in (a) conception and design, or analysis and interpretation of the data; (b) drafting the article or revising it critically for important intellectual content; and (c) approval of the final version. This manuscript has not been submitted to, nor is under review at, another journal or other publishing venue. The authors have no affiliation with any organization with a direct or indirect financial interest in the subject matter discussed in the manuscript All authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Author’s name (Affiliation) Simona Avnaim-Katav (Israel Oceanographic & Limnological Research, Haifa, 31080, Israel)

Ahuva Almogi-Labin (Geological Survey of Israel, Jerusalem, 95501, Israel

Mor Kanari (Israel Oceanographic & Limnological Research, Haifa, 31080, Israel)

Barak Herut (Israel Oceanographic & Limnological Research, Haifa, 31080, Israel)