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Abstract

Tens of thousands of fish bones were recovered from the Final Natufian (Late Epipaleolithic) layer of the site of Eynan/Ain Mallaha (Northern Israel) dated to the end of the Pleistocene. Almost a hundred of them were attributed to a Salmoninae, most probably a trout Salmo cf. trutta. This is the southernmost attestation of a Salmo species in the Near East, past and present. It is suggested that trout were present locally as small populations at least at the end of the Pleistocene.

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... NEG II is located in the Upper Jordan Valley (UJV), 2 km east of the Sea of Galilee (Fig. 1), the largest body of freshwater in the southern Levant. The unique ecology created by the Jordan Valley river system provided local residents with special aquatic foraging opportunities that gave a local character to the transition to agriculture in this area (see also Borvon et al. 2018;Zohar in Valla et al. 2007). The Jordan Valley differs from the Mediterranean Hills region of the southern Levant in its significant permanent freshwater system comprised of rivers and lakes that buffered the region against water stress and provided unique hunting and gathering opportunities in aquatic and wetland habitats. ...
... The taxonomic data indicates that the hunter-fishers of NEG II were highly selective when it came to aquatic resources. This selectivity is even more pronounced when compared to the much more diverse fish and waterfowl assemblages from Ohalo II and Eynan which have eight and four times the respective number of aquatic species than NEG II (Table 2; Borvon et al. 2018;Bouchud 1987;Bridault et al. 2006, Desse 1987Pichon 1987;Simmons 2013;Simmons and Nadel 1998;Zohar et al. 2018). The same is true of the inverse of Simpson's Diversity Index for birds which is nearly five times greater at Eynan and 6 times greater at Ohalo II than at NEG II. ...
... Fish diversity is also 50% higher at Ohalo II (Table 2) than at NEG II. A detailed analysis of the fish has not yet been completed for the new excavations at Eynan, but in progress work on a single Final Natufian structure (228) revealed thousands of fish bones (NISP = 7475 and counting; Borvon et al. 2018). At least six species of fish representing a wide spectrum of body-sizes have been identified thus far. ...
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This paper investigates aquatic resource exploitation at the Late Natufian site (ca. 12,000 cal. BP) of Nahal Ein Gev II located 2 km east of the Sea of Galilee. Aquatic game, here fish and waterfowl, were an important component of the diverse small game resources that became important in the Late Epipaleolithic in Southwest Asia. We characterize local adaptations to the aquatic habitat and their economic and social implications at Nahal Ein Gev II. Taxonomic abundance and diversity, body-part representation, and fish body-size were investigated to evaluate the contribution of aquatic resources to human diets and butchery and transport strategies. Our results show that the residents of Nahal Ein Gev II were highly selective of the aquatic resources they captured and transported home. The hunters maximized foraging efficiency by nearly exclusively choosing the largest bodied species of fish and waterfowl and processing their carcasses to maximize meat utility before transporting them back to the site. The selectivity of these human foragers enables us to reconstruct rare details about the organization of forays for aquatic resources. When combined with evidence from other material classes from Nahal Ein Gev II and other sites, the results suggest that aquatic resource exploitation is only one of several specialized activities practiced at Nahal Ein Gev II. These along with other archaeological evidence provide evidence of task diversification that foreshadows the emergence of a more complex division of labor to come in the succeeding Neolithic period.
... Altogether the data at hand suggest at least repeated long stays of people. This is underlined by intensive activities, including the digging of semisubterranean oval buildings, burying the dead, tool making, artwork, and more [14], as well as intensive exploitation of every ecological niche available near the site, and game procurement throughout the year, including fish, birds and mammals [15][16][17][18][19][20][21][22]. Each building went through several phases of occupation, some of them involving changes in function (dwellings versus other activities), as shown by reorganizations of postholes and hearths inside the buildings. ...
... Snake meat contains approximately 93 calories per 100g of raw meat, varying according to the snake taxon [61]. However, at Eynan a variety of vegetation and animals were available, as attested by the floral and faunal remains recovered [18][19][20][21]62], and therefore two questions must be asked: Was the choice of eating specifically large "colubrine" snakes necessary at Eynan as a protein or fat source? Should the exploitation of supplementary species at the end of the Pleistocene be considered as associated merely with dietary requirements and calorie intake? ...
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During the Natufian period, more than 12,000 years ago, Eynan (Ain Mallaha) was an important human settlement in the Hula Valley, Israel. This study concentrates on the anuran and squamate assemblage from the ultimate stage of the Natufian period at the site, the Final Natufian. Over five thousand bones assigned to at least sixteen taxa were studied from a sampled segment of the excavated open-air site. Relative species abundance, spatial distribution, taphonomic observations and ecological considerations all pointed to the conclusion that the inhabitants of Eynan intensively exploited three large “colubrine” snakes species: the Large Whip Snake ( Dolichophis jugularis ), the Eastern Montpellier Snake ( Malpolon insignitus ) and an Eastern Four-lined Ratsnake ( Elaphe cf. sauromates ). These snakes were the most desired and were intensively gathered, while other snakes and lizards could have been opportunistically collected when encountered. We raise questions about whether the large “colubrines” exploitation should be interpreted as additional evidence of increasing diet breadth. We suggest challenging this line of reasoning and offer possible alternative motives.
... The species composition and the ecosystem structure of the Natufian site of Eynan (Ain Mallaha) are generally very similar to the recent Lake Hula ecosystem before its drainage (Bridault et al. 2008;Ashkenazi 2013;Biton et al. 2016). However, some evidence might hint at a somewhat cooler climate compared to the present (Rosen and Rivera-Collazo 2012;Borvon et al. 2018). Nevertheless, the existing spring near the site, with a constant year-round temperature of 21°-22°C, created an open pond and a short river about 3 km long in the past, before it entered the Hula Lake (Dimentman et al. 1992). ...
... During the Pleistocene in the Hula Valley, aquatic taxa such as ostracods, and gastropods were dominated by taxa of northern origin (Palaearctic or Holarctic) (Mienis and Ashkenazi 2011;Mischke et al. 2014). Moreover, recent studies from Pleistocene Hula Valley assemblages have identified the presence of hydrophilous fish and anuran genera of northern origin, previously undescribed from that area, that reached the Hula Valley through a water 'corridor' and support this scenario (Biton et al. 2013(Biton et al. , 2016Borvon et al. 2018). ...
Article
The (palaeo)biogeography of water voles is a puzzle that is not solved in detail yet. Extant species of the genus Arvicola cover a vast geographic area of the Palearctic. In this study, we collected morphometric data of extant and fossil Arvicola from Israel at the southern fringe of the water vole distribution area. The dental evolution of water voles is characterised by certain clear trends in the first lower molar (m1) related to crown height, tooth length, the proportion of the anteroconid-complex, and enamel differentiation. Trans-regional correlation of the latter trend shows that it did not develop completely synchronously in all the geographic areas of water voles' distribution, i.e., not at the same evolutionary rate. Based on the size and geographic distribution, we tentatively consider the Israeli material to belong to an A. persicus group, which facilitates the possibility of future taxonomic subdivision. Among the Israeli material, Late Pleistocene remains from Nahal Mahanayeem Outlet (NMO) show a very particular morphology of the first lower molar compared to samples from other localities in Israel and western Asia, which-in our opinion-justifies the establishment of a new species Arvicola nahalensis n. sp.
... The site is located in the Northern Jordan Valley (Israel), on a slope overlooking the Eynan spring (Fig. 1). The Eynan people settled in the vicinity of the Hula Lake and the vast adjacent wetland area, and exploited a wide array of resources and habitats, as reflected by the plant and faunal remains at the site (Bridault et al. 2008;Rosen 2010;Askhenazi 2013;Simmons 2013;Borvon et al. 2018;Bridault 2019). The local availability of water and grass made this area attractive to herbivore populations. ...
... Could the magnitude of these changes be observed in the archaeological record? Were human behavioural patterns, mode of prey procurement, seasonality and foraging methods influenced as a result? Some studies have pointed out the relative stability of the environmental conditions at Eynan (Askhenazi 2013;Biton et al. 2016), while others support the existence of a somewhat cooler climate compared to the present (Bar-Matthews et al. 1997Hartman et al. 2016;Valla et al. 2007;Borvon et al. 2018). ...
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The Final Natufian level Ib from Eynan/Ain Mallaha provided a large and diversified assemblage of ungulates used here to study the seasonality of game procurement. The objective of this study was to evaluate the advantage of combining two methods, dental eruption and wear, and tooth microwear. Tooth microwear results allowed us to confirm the seasonality of procurement of red and roe deer and to narrow down its duration for the gazelle and the fallow deer. Game procurement occurred throughout the year for the roe deer. Red deer were hunted in spring and summer, and gazelle in late autumn-early winter. This approach allowed us to improve the accuracy of seasonality estimates and to increase sample size by using the entire population (young and adult individuals) for each species. The few individuals that were analysed with the two methods also permitted us to characterize the diet of the population at the time of death.
... Teeth from the Cyprinid species Luciobarbus longiceps, Casiobarbus canis and Acanthobrama hulensis have been identified thus far. A few Salmonids (e.g., trout) may also be present (see also [121], but these identifications must be confirmed with an appropriate comparative assemblage. ...
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Nineteen broken and complete bone fish hooks and six grooved stones recovered from the Epipaleolithic site of Jordan River Dureijat in the Hula Valley of Israel represent the largest collection of fishing technology from the Epipaleolithic and Paleolithic periods. Although Jor-dan River Dureijat was occupied throughout the Epipaleolithic (~20-10 kya the fish hooks appear only at the later stage of this period (15,000-12,000 cal BP). This paper presents a multidimensional study of the hooks, grooved stones, site context, and the fish assemblage from macro and micro perspectives following technological, use wear, residue and zooarch-aeological approaches. The study of the fish hooks reveals significant variability in hook size, shape and feature type and provides the first evidence that several landmark innovations in fishing technology were already in use at this early date. These include inner and outer barbs, a variety of line attachment techniques including knobs, grooves and adhesives and some of the earliest evidence for artificial lures. Wear on the grooved stones is consistent with their use as sinkers while plant fibers recovered from the grooves of one hook shank and one stone suggest the use of fishing line. This together with associations between the grooved stones and hooks in the same archaeological layers, suggests the emergence of a sophisticated line and hook technology. The complexity of this technology is highlighted by the multiple steps required to manufacture each component and combine them into an integrated system. The appearance of such technology in the Levantine Epipa-leolithic record reflects a deep knowledge of fish behavior and ecology. This coincides with significant larger-scale patterns in subsistence evolution, namely broad spectrum foraging, which is an important first signal of the beginning of the transition to agriculture in this region. PLOS ONE PLOS ONE | https://doi.org/10.1371/journal.pone.
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Significance Decreases in hunter-gatherer mobility during the Late Pleistocene altered relationships with animal communities and led to domestication. Little is known, however, about how selection operated in settlements of varying duration. This study of mice in modern African mobile settlements and ancient Levantine sites demonstrates competitive advantages for commensal mice when human mobility is low and niche partitioning with noncommensal wild mice when mobility increases. Changing mice molar shapes in a 200,000-y-long sequence from the Levant reveal that mice first colonized settlements of relatively settled hunter-gatherers 15,000 y ago. The first long-term hunter-gatherer settlements transformed ecological interactions and food webs, allowing commensal house mice to outcompete wild mice and establish durable populations that expanded with human societies.
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Analysis of both uni-(two mtDNA gene sequences) and bi-parentally (seven microsatellite loci) inherited genetic markers, together with analysis of 40 morphological characters, described Salmo ohridanus as a highly divergent member of the genus Salmo. Based on comparative substitution rate differences at the cytochrome b gene, and a rough estimated age of the Salmo trutta complex (i.e. at least 2 million years), the S. ohridanus and Salmo obtusirostris clade probably split from a common ancestor of brown trout Salmo trutta > 4 million years ago, overlapping with minimum age estimates of the formation of Europe's oldest freshwater habitat, Lake Ohrid. Comparative analysis with Lake Ohrid brown trout (known regionally as Salmo letnica), supported the notion that these fish have more recently colonized the lake and phylogenetically belong to the Adriatic lineage of brown trout. It is further suggested that species-specific saturation in the mtDNA control region underestimated the divergence between S. ohridanus and S. trutta. Evidence of rare hybridization between S. ohridanus and Lake Ohrid brown trout was seen at both mtDNA and microsatellite markers, but there was no support for extensive introgression. (c) 2006 The Fisheries Society of the British Isles.
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Salmo tigridis, new species, from the Tigris River drainage, Turkey, is distinguished from the other species of Salmo in Turkey and adjacent basins by having a greater number of scale rows between the end of the adipose-fin base and lateral line (19–20, vs. 12–17); a greater number of scale rows between dorsal-fin origin and lateral line (32–35, vs. 23–32); and a deeper and stouter caudal peduncle (its depth 11.5–12.6 % SL, vs. 9.3–11.5, except in S. cf. macrostigma).
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Despite the fact that fish are a common component of coastal and other aquatic archaeological sites, cut marks are rarely reported on archaeological fish remains. To assess whether butchering practices leave cut marks on fish bones, we butchered 37 fish using stone tools and a metal knife following methods provided in ethnographic accounts and by modern fish processors. In contrast to archaeological analyses, our research demonstrates that butchering commonly produces cut marks on fish bones, with 4019 cut marks and 2167 cut mark clusters iden-tified on the bones of 30 fish. Cut marks occurred frequently on vertebral neural and haemal spines, vertebral transverse processes, pterygio-phores, ribs, and other bones not generally identified to low taxonomic categories by zooarchaeologists (e.g., family, genus, or species). To test our experimental data, we also analyzed 9391 archaeological fish remains from a Late Holocene shell midden on the California Coast, noting 33 previously undocumented cut marks. We hypothesize that the scarcity of cut marks reported on archaeological fish bones is the result of researchers overlooking cut marks because they occur primarily on undiagnostic bones, taphonomic factors such as root etching that may destroy or obscure cut marks, differences between fish, mammal, and bird anatomy, or ancient butchering strategies that relied on limited cutting of fishes.
Article
The spatial distribution of the European freshwater eel (Anguilla anguilla) was very different in historic and prehistoric times in comparison to the present. A database of the spatial and temporal distribution of eel remains in archaeological and palaeontological sites is presented and used to assess the spatial distribution of populations from the height of the last glacial maximum. The results show that the eel was absent from northern Europe until about 11 000 years ago. The reason was probably a southerly displacement of the Gulf Stream carrying the larval migration from the Sargasso Sea. However, additional factors preventing eel populations in northern Europe may have also been the colder temperatures in the Arctic tundra landscape that existed at the time and the extreme distance to the European Atlantic coast along the Channel River. The archaeological record shows that eels were absent from the Baltic Sea until about 6700 cal BC, but there is some indication of an earlier presence during the Yoldia Sea stage at the beginning of the Holocene. Only in southern Europe south of the Gironde river basin were eel populations maintained through the last glaciation. The species may have survived the last glaciation in a relatively restricted area in the Mediterranean and the Atlantic coast of western Europe. Published palaeontological and genetic information gives important insights into climatic, geologic, and tectonic events on longer time scales. The oldest subfossil remains from Pleistocene sediments in northern Europe are approaching the age of the estimated genetic divergence of the European and American eel populations, and hence the species identity of the oldest subfossil remains may be ambiguous.
Article
The eastern Mediterranean continental paleoclimate during the past 25,000 years was determined by a high-resolution petrographic, stable isotopic, and age study of speleothems from Soreq Cave, Israel. δ18O–δ13C trends indicate that all speleothems older than 7000 yr formed under conditions that differ from those of today. The period from 25,000 to 17,000 yr B.P. was characterized by the highest δ18O and δ13C values, which indicate deposition at temperatures of 12°–16°C, annual rainfall of 300–450 mm, and vegetation typical of a mixed C3–C4type. From 17,000 to 10,000 yr B.P. (deglaciation in northern Europe) δ18O values dropped progressively, correlative with warming (2°–3°C) and a gradual increase in precipitation. A simultaneous decrease in δ13C gives a range expected for C3-type vegetation. This period also shows significant δ18O and δ13C “spikes” which are correlatable with global events (e.g., Heinrich events and the Younger Dryas Stade). The speleothems that grew between 10,000 and 7000 yr B.P. have a unique petrography showing irregular thin laminae of various colors and much detritus. They have the lowest δ18O (corresponding to ∼1000 mm rain) coupled with the highest δ13C (caused by flooding events which stripped the soil cover). From 7000 to 1000 yr B.P. conditions became closer to those of today. This study demonstrates that global events which were recognized in Northern Europe and North Africa are also evident in the eastern Mediterranean and are reflected principally by large changes in the rainfall rate.
Article
The climate of the Eastern Mediterranean region of the last 60 ky was determined by a high resolution study of the oxygen and carbon isotopic composition (1500 measurement pairs) of speleothems from the Soreq cave, Israel, with chronology provided by 53 precise ^(230)Th–^(234)U (TIMS) ages. The high precision of the speleothem TIMS ages permits us to determine the timing of regional climatic events in the Eastern Mediterranean region and to see if they correlate with global events. During the period from 60 to 17 ky, the δ^(18)O and δ^(13)C values were generally 2–2.5‰ higher than during the period from 17 ky to present. This is consistent with the climatic transition from glacial to interglacial. Within the 60 to 17 ky period, the Soreq cave stable isotope profile includes four cold peaks (at 46, 35, 25 and 19 ky) and 2 warm peaks (at 54 and 36 ky). In addition, the period <17 ky has two more cold peaks at 16.5 and from 13.2 to 11.4 ky. The ages of four of the six cold peaks correlate well with the ages of three Heinrich events (H1, H2, H5) and with the age of the Younger Dryas. However, the other two Heinrich events are not reflected in the Soreq cave record. Several other isotope peaks which appear during the last 7 ky are contemporaneous with regional climatic events in the Middle East and North Africa. In addition to the drop in δ^(18)O and δ^(13)C observed between the last glacial and the Holocene, sharp simultaneous drops in (^(234)U/^(238)U)_0 ratios, Sr concentrations and in ^(87)Sr/^(86)Sr are also observed, suggesting that the latter are climate related. These variations are interpreted in terms of major changes in the temperature, the mean annual rainfall and its isotopic composition, the isotopic composition of the Mediterranean vapor source, the soil moisture conditions, and in the mixing proportions of sources with different ^(87)Sr/^(86)Sr ratios (sea spray, dust particles and dolomitic host rock).
Article
Phylogeographic, nested clade, and mismatch analyses of mitochondrial DNA (mtDNA) variation were used to infer the temporal dynamics of distributional and demographic history of brown trout (Salmo trutta). Both new and previously published data were analyzed for 1,794 trout from 174 populations. This combined analysis improved our knowledge of the complex evolutionary history of brown trout throughout its native Eurasian and North African range of distribution in many ways. It confirmed the existence of five major evolutionary lineages that evolved in geographic isolation during the Pleistocene and have remained largely allopatric since then. These should be recognized as the basic evolutionarily significant units within brown trout. Finer phylogeographic structuring was also resolved within major lineages. Contrasting temporal juxtaposition of different evolutionary factors and timing of major demographic expansions were observed among lineages. These unique evolutionary histories have been shaped both by the differential latitudinal impact of glaciations on habitat loss and potential for dispersal, as well as climatic impacts and landscape heterogeneity that translated in a longitudinal pattern of genetic diversity and population structuring at more southern latitudes. This study also provided evidence for the role of biological factors in addition to that of physical isolation in limiting introgressive hybridization among major trout lineages.
Article
Complete sequencing of the mitochondrial control region was used to describe phylogenetic relationships of brown trout populations (Salmo trutta) in the Mediterranean river basins of Iberia and to review the historical biogeography of trout from the Mediterranean regions. Phylogenetic relationships among trout lineages suggested that the Danubian one is the most ancestral, in accordance with the eastern origin of most of the European freshwater fish species. Nested-clade and mismatch analyses suggested that the present distribution of haplotypes of the Adriatic and Mediterranean lineages resulted from population expansions originated, respectively, from central and western Europe, which favoured extensive secondary contacts between lineages. Reduced diversity detected within 50% of the analysed populations and large intrabasin differentiation indicated restricted gene flow in post-glacial periods.
Reconstruction of the habitats in the ecosystem of the final Natufian site of Ain Mallaha (Eynan)
  • S Ashkenazi
Ashkenazi, S., 2013. Reconstruction of the habitats in the ecosystem of the final Natufian site of Ain Mallaha (Eynan). In: Bar-Yosef, O., Valla, F.R. (Eds.), Natufian Foragers in the Levant. Ann Arbor, International Monographs in Prehistory, International Series 19 Oxford, pp. 312-318.
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Les états de Syrie. Richesses marines et fluviales
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