This article was published in an Elsevier journal. The attached copyis furnished to the author for non-commercial research and

education use, including for instruction at the author’s institution,sharing with colleagues and providing to institution administration.

Other uses, including reproduction and distribution, or selling orlicensing copies, or posting to personal, institutional or third party

websites are prohibited.

In most cases authors are permitted to post their version of thearticle (e.g. in Word or Tex form) to their personal website orinstitutional repository. Authors requiring further information

regarding Elsevier’s archiving and manuscript policies areencouraged to visit:

http://www.elsevier.com/copyright

Author's personal copy

The living and the dead: How do taphonomic processesmodify relative abundance and skeletal

completeness of freshwater fish?

Irit Zohar a,b,⁎, Miriam Belmaker c, Dani Nadel d, Sarig Gafny e,Menachem Goren a, Israel Hershkovitz f, Tamar Dayan a

a Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv 69978, Israelb The Leon Recanati Institute for Maritime Studies, University of Haifa, Mount Carmel, Haifa, 31905, Israel

c Harvard University, Department of Anthropology, Peabody Museum, 11 Divinity Ave. Cambridge MA 02138 USAd Zinman Institute of Archaeology, University of Haifa, Mount Carmel, Haifa 31905, Israel

e School of Maritime and Marine Environment Sciences, Rupin Academic Centre 40250, Emek Hefer, Israelf Department of Anatomy and Anthropology, Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv, 69978, Israel

Received 30 November 2006; received in revised form 8 September 2007; accepted 2 November 2007

Abstract

This study is designed to determine the extent to which taphonomic processes alter the taxonomic composition of fish remainsin lacustrine sediments. We wish to explore information loss in a bone assemblage relative to the original, living community. Weexamined fish bone assemblages from lacustrine sediments along the southern shore of Lake Kinneret (Sea of Galilee) andcompared them to modern living communities. For this purpose we randomly selected 24 squares, each 0.5 m2 in size, andexcavated them to a depth of 30–50 cm. Three lithofacies were recovered, spanning the past 1500 years (unccorected for reservoirage). The fish remains include 5037 bones and 758 scales, of which 1566 bones were identified to taxonomic group. The list ofidentified species was compared with the list of indigenous species known to live in Lake Kinneret in general and in a similar sandyhabitat in particular. The proportion of skeletal elements found was compared with the proportion known in a complete fish. Ourstudy indicates that differences exist between the three lithofacies in species diversity and composition, skeletal element richness,completeness, and relative abundance. In addition, the bones exhibit a clumped distribution pattern, regardless of depositionaldepth. From a taphonomic and paleoecological perspective, our findings demonstrate that fish remains retrieved from lacustrinesediments do not represent the composition and diversity of species as in the recent fish community.© 2007 Elsevier B.V. All rights reserved.

Keywords: Faunal assemblage; Fish remains; Freshwater fish; Jordan Rift Valley; Lacustrine sediments; Lake Kinneret; Taphonomy

Available online at www.sciencedirect.com

Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316www.elsevier.com/locate/palaeo

⁎ Corresponding author. The Leon Recanati Institute for Maritime Studies, University of Haifa, Mount Carmel, Haifa, 31905, Israel. Fax: +972 36407304.

E-mail addresses: zoharir@post.tau.ac.il (I. Zohar), belmaker@fas.harvard.edu (M. Belmaker), dnadel@research.haifa.ac.il (D. Nadel),gafny@post.tau.ac.il (S. Gafny), gorenm@post.tau.ac.il (M. Goren), anatom2@post.tau.ac.il (I. Hershkovitz), dayant@post.tau.ac.il (T. Dayan).

0031-0182/$ - see front matter © 2007 Elsevier B.V. All rights reserved.doi:10.1016/j.palaeo.2007.11.004

Author's personal copy

1. Introduction

Unraveling the mystery of ancient aquatic life anddeath in changing environments is a huge challenge forpalaeoecology. While considerable information is lostdue to bone destruction (Farlow and Argast, 2006),skeletal elements that do survive are incorporated intothe sediment, where they become the record of past fishcommunities. This record is significant for paleoecolo-gical (i.e., taxonomic composition and biodiversity;O'connell and Tunnicliffe, 2001; Tunnicliffe et al., 2001)and paleoenvironmental reconstruction (e.g. salinity,temperature, water level; Casteel, 1976; Brett and Baird,1986; Wilson, 1988; Weigelt, 1989; Cutler et al., 1999;Chen, 2000). Recently, a growing body of taphonomicresearch demonstrated the numerous depositional pro-cesses that affect fish bone preservation (Trueman andMartill, 2002; Trueman et al., 2003; Farlow and Argast,2006). It has been realized that biases in skeletal elementpreservation alter species richness and abundance (Elderand Smith, 1988; Ferber and Wells, 1995; Cutler et al.,1999; Martin, 1999; O'connell and Tunnicliffe, 2001).Clearly, the ability to estimate what and how much oforiginal information was lost is crucial for paleoecolo-gical and paleoenvironmental reconstruction. Therefore,field tests that compare living communities to associatedbone assemblages are the primary means of estimatingthe biological information loss in the palaeontologicaland archaeological record (Behrensmeyer, 1982, 1983;Kidwell, 1986; Kidwell and Bosence, 1991; Kidwell,2001, 2002).

Direct live-dead comparisons have demonstrated thata wide variety of factors play a role in the processesof fish carcass decay, disintegration, articulation, andpreservation in the aquatic sediments, and that acomplete fish skeleton will be preserved only underspecial circumstances (Schäfer, 1972; Soutar and Isaacs,1974; Elder and Smith, 1988; O'connell and Tunnicliffe,2001; Trueman and Martill, 2002; Trueman et al., 2003;Farlow and Argast, 2006). These include: water tem-perature, water depth, pressure, salinity level, oxygenlevel, oxygen stability, type of sediments, rate of sedi-mentation, carcass buoyancy, scavenger activity, waveaction, and sea bottom currents (Schäfer, 1972; Elder andSmith, 1988; Cutler et al., 1999; Reiche et al., 2003). Ofthese, water temperature has been regarded as the mostsignificant factor in determining the fate of carcasses(Schäfer, 1972; Elder and Smith, 1988; Collins et al.,1995). In all, it has been shown that fish diversity andtaphonomic processes are habitat dependent, and we stillneed to seek for generalities in the taphonomic processesinvolved. In order to do that, we must examine the

differential loss between the live-dead assemblagesbearing in mind that in most cases a bone assemblageis not necessarily biologically equivalent to a livingcommunity (Kidwell and Bosence, 1991; Kidwell, 2001;Kidwell, 2002; Tomasovych, 2006).

Although live-dead comparisons are fairly commonin the taphonomic literature on marine shell assemblages(e.g., Kidwell, 2001; Kidwell, 2002; Alin and Cohen,2004; Tomasovych, 2006) and terrestrial mammals (e.g.,Behrensmeyer, 1982, 1983; Behrensmeyer et al., 1989;Behrensmeyer, 1991; Cutler et al., 1999; Behrensmeyerand Barry, 2005), they are sparse for fish remains (Soutarand Isaacs, 1974; Stewart, 1989; Butler, 1993; O'connelland Tunnicliffe, 2001). Recently, a growing literature isalso concerned with the paleoanthropological recon-struction of ancient fish exploitation patterns in coastalcommunities (Brewer, 1991; Arnold, 1992; Cerón-Carrasco, 1994; Enghoff, 1994; Leach and Boocock,1994; Wing, 1994; Zohar et al., 1994; Fraser, 1998;Butler, 2000; Cannon, 2000; Rick and Erlandson, 2000;Butler, 2001; Galili et al., 2004). Some of the archae-ological sites have become submerged due to local orglobal water level changes (Nadel, 1993; Zohar et al.,2001; Galili et al., 2004). Therefore, while studying fishremains from lacustrine sediments, onemust askwhetherthe bone assemblage reflects human fishing activity orwhether it is merely natural death assemblage? Thus, thestudy of fish remains from waterlogged and submergedsites must be based on sound taphonomic analyses ofnatural fish accumulations that may enable researchers totease apart the remnants of economic practices from theimprint of natural deaths (Butler, 1987; Stewart, 1989;Gifford-Gonzales et al., 1999).

In the southern Levant no research, to our knowledge,has yet compared the fish species found alive in aquatichabitat to those found as bones in an adjacent shoreline,although this region is rich in important archaeologicalsites (e.g., Bar-Yosef, 1980, 1987a, 1987b; Bar-Yosef andBelfer-Cohen, 1989; Goren-Inbar et al., 2000). In order togain insight into taphonomic biases that affect bothpaleoecological and paleoanthropological reconstruction,we performed a pioneer study of fish remains naturalaccumulations in lacustrine sediments along the southernshore of Lake Kinneret (Sea of Galilee; Fig. 1). Threelithofacies were identified, spanning the past 1500 years.This study was designed to determine information loss(taxonomic diversity, species composition and skeletoncompleteness) relative to the living community and todetermine biases expected in environmental interpreta-tions from fish remains deposited in a sandy littoralsouthern shore of Lake Kinneret. The focus of this studywas to compare one modern shoreline habitat and the

293I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

facies that occur within a small stratigraphic distancebeneath it with nearby living communities from differenthabitats. This was used as a test case. Further research inother lacustrine habitat is required to test the generality ofour results and conclusions.

2. Locality

2.1. Lake Kinneret

Lake Kinneret (Sea of Galilee), located in the northernpart of the rift valley, is the largest fresh-water lake inIsrael. During the Pleistocene (70–15 ka) it was part ofLake Lisan, which extended from the Arava Valley in thesouth to the present Kinneret basin in the north, when itreached a maximum level of 170 m below sea level at

26 ka (Bartov et al., 2002; Hazan et al., 2005). Thedramatic changes in the lake's size, salinity, temperature,sedimentation, depth, as well as other factors, might haveaffected the composition of primary freshwater fish.

At present (July 2006) the lake level is about 211.09 mbelow sea level (BSL) (http://www2.kinneret.ac.il/kinneret/miflas.asp). It is ca. 21 km long and 13 kmwide, covering an area of ca. 168 km2 with a maximumdepth of 43 m (Gophen and Gal, 1992). It is a warmmonomictic lake, withwinter homeothermal temperaturesbetween 12 °C and 14 °C and summer temperaturesapproaching 30 °C (Hambright et al., 1994). The lake isfed by a number of freshwater streams and springs. TheJordan River is the major inflow. Lake level varies fromrainy to drought years, and in recent years according touse for domestic and agricultural purposes, andmay reach

Fig. 1. Location map of Lake Kinneret, Plaleo lake Lisan (after Hazan et al., 2005), and the study site, exhibiting the position of 24 random squaresalong the southern shore.

294 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

up to 5 m (between −209 and −214 mBSL) (Gafny et al.,1992; Gophen and Gal, 1992).

2.2. Depositional environments surrounding LakeKinneret

The littoral zone of Lake Kinneret extends to a depthof 3 m (between 209 to 212 m BSL), and is char-acterized by strong wave action, relatively large waterlevel fluctuations (WLF), and unfavorable light condi-tions (Gasith and Gafny, 1990). Studies showed that thehigh rate of sedimentation at Lake Kinneret is seasonal(April–May) and is highly correlated with flow dis-charge of the Jordan River and periodical bloom ofPeridinium gatunense (Stiller et al., 1983–84; Korenand Klein, 2000). The rates of sedimentation decreasesouthward along the coast as a result of the increaseddistance from the Jordan River mouth.

Four different depositional environments have beenidentified around Lake Kinneret (Hazan et al., 2005 andreferences therein): deep-water, fluvial, fan deltas, andshore environments. Each of these depositional envir-onments is represented by several sedimentary lithofa-cies. Differences between lithofacies are attributable tovariations in water depth, energy, and the limnologicalconfiguration of the lake (Hazan et al., 2005).

The lithological composition of Lake Kinneret shoresvaries from boulders and stones of various sizes to finersubstrates of sand, silt, and clay (Gasith and Gafny,1990; Hazan et al., 2005). For example, at someshorelines, finer sediments are washed away by waveaction creating beach-ridges made of pebbles or coarsesand (Stiller et al., 1983–84; Gasith and Gafny, 1990;Gafny and Gasith, 2000; Koren and Klein, 2000; Hazanet al., 2005). Offshore, the influence of surface waves isreduced and sediments become finer. Sediments withgrain sizes varying from sand to silt are deposited at thelittoral zone, under a water depth of up to a few meters.At deeper waters, clay to silt-size laminated sediments aredeposited (Hazan et al., 2005). Due to fluctuations inwater level (±5 m), periodic exposures and inundation ofthe littoral areas occur, followed by changes in the natureof the littoral zone (Gasith and Gafny, 1990; Gafny, 1992;Gasith et al., 1996; Gafny and Gasith, 2000).

3. Materials and methods

3.1. The bone assemblages

We examined fish bone assemblages from a sandybeach located on the southern shore of Lake Kinneret.This area is usually inundated by the lake, except during

drought seasons, when it is exposed. The excavation tookplace during July 2001, when water levels dropped to214 m BSL, and archaeological sites were exposed alongthe shore (Nadel, 1993). Hence, in order to avoidexcavation in an area previously inhabited, the areaselected for this study was carefully chosen to be devoidof archaeological finds (e.g., Ohalo-II; Nadel et al., 2002).

Using an eTrek global positioning system (GPS) wemarked an area of 100 m in length and 50 m in width. Wecalculated the position of 24 random squares (Fig. 1),each 0.5m2 in size. The squares were excavated to a depthof 30–50 cm and wet sieved through fine mesh (0.5 mm)screens.

We identified three lithofacies (A, B, and C from topto bottom; Fig. 2) based on grain size, sediment colors,and texture. We are concerned with the potentialconfusion in the use of the term “facies”. This may bedefined either as the lithological product and its processor the environment in which it was formed (Reading,1996). Following Badgley (1982, 1986) we use the termqfaciesq (i.e., lithofacies) to mean the lithologicalproduct only. Lithofacies A included the surface sandstratum (3–10 cm thick) (Gafny, 1992), which issubjected to wave, wind, water and human activity(walking and driving). Lithofacies B is a mixed sand-siltstratum (5–10 cm thick) and lithofacies C is a darkbrown stratum of fine grain silt and clay (5–10 cmthick), dated at the bottom to 1515± 50 y BP(uncalibrated 14C and uncorrected for reservoir age,University of Arizona, Tucson sample #RTT4385).Following Hazan et al. (2005), these changes inlithofacies are interpreted as representing changes inwater depth and energy during the shift between theshoreline and deepwater region of Lake Kinneret, andoverall reflect the receding shoreline of the lake.

Fig. 2. An excavated square of the death assemblages from thesouthern shore of Lake Kinneret (observe the changes from uppersandy layer to a dark bottom layer).

295I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

3.2. The living fish community

The present ichthyofauna of Lake Kinneret com-prises 19 indigenous fish species (Table 1) that representsix families originating from Africa, Central Asia, andEurope (Ben-Tuvia, 1978; Krupp, 1987; Banarescu,1990; Goren and Ortal, 1999; Durand et al., 2002). Thedistribution of fish species in different habitats along theLake Kinneret littoral zone was examined diurnally byone of us (S.G.). During three consecutive years (1988–1990) of low lake level, seven sampling excursions wereperformed at different times of the year, using a shore-operated electro-shocker (Gafny, 1992). By using a1 mm seine net mesh, an area of 100 m2 was encircledprior to the operation of the elcetro-shocker and laterused for seining. A total of 5915 fish, representing 15species of various body sizes, were collected from 8localities that differ in substrate and habitat (Table 2).Station E-9 is a sandy littoral shore that resembles thebone assemblage habitat in geographic location and

substrate. Therefore it is used for comparison with thebone assemblage.

3.3. Bone assemblage taxonomic identification

Fish bones were identified to the lowest anatomic andtaxonomic level possible, using a large osteologicalreference collection prepared by one of us (I.Z.) andhoused at Tel-Aviv University (Zohar, 2003). Whilemost postcranial and cranial bones were identifiable tofamily level, only few bones (atlas, axis, pharyngealbones, and teeth) were species specific. The issue of fishbone identification has been widely discussed in theliterature (Wheeler, 1978; Colley, 1984; Nichol andWild, 1984; Owen, 1994; Nicholson, 1996; Gobalet,2001), and it has been suggested (Owen, 1994) thatsampling and analytical procedures should be modifiedto suit specific situations. Therefore, to overcome theseproblems and to enlarge sample sizes, we grouped thebones according to five taxonomic groups as follow:

Table 1List of recent native freshwater fishes inhabiting Lake Kinneret, Israel. Taxonomy follows (Goren et al., 1973; Goren and Ortal, 1999; Durand et al.,2002)

Family Species Origin Habitat

Cyprinidae Acanthobrama lissneri Endemic-Jordan River drainage basin and Qishon R. Bottom of lakes, ponds and slow-flowingwater.

Mirogrex terraesanctae Endemic-Lake Kinneret. PelagicCarasobarbus canis Jordan River and Lake Kinneret. Small and wide streams and in lakes

and ponds.Barbus longiceps Jordan River and Lake Kinneret. Bottom dweller.Capoeta damascina Jordan River drainage basin, the entire Levant,

Mesopotamia and parts of southern Turkey.Bottom fish, occurs in lakes, streams withfast and slow-moving water currents andin clear and muddy water.

Garra rufa Jordan, Orontes, and Tigris–Euphrates river basins.Also in some coastal rivers in southern Turkeyand northern Syria and Israel.

Bottom dweller, found in rivers, lakes,small ponds, and small muddy streams.Hides under and among stones.

Hemigrammocapoeta nana Damascus, Jordan river basins, and Qishon River. Occurs in lakes, ponds and slow-movingstreams, in stones and vegetation.

Pseudophoxinus kervillei Orontes River, Litany and the central and northernpart of Jordan River drainage.

In slow-moving or still water amidststones and vegetation.

Balitoridae Nemacheilus tigris Asia–Europe. Rivers and lakes.Clariidae Clarias gariepinus Africa, Asia: Jordan, Israel, Lebanon, Syria

and southern Turkey.In quiet waters, lakes and pools and infast flowing rivers and rapids. Tolerant ofextreme conditions.

Cyprinodontidae Aphanius mento Israel, Anatolia, Iran. Tethys relict. Found in shallow water among vegetation.Cichlidae Tilapia zillii Africa and Middle East. Prefers shallow, vegetated areas in

rivers and lakes.Oreochromis aureus Africa and Middle East. River and lakes.O. niloticus Africa and coastal rivers of Israel. Rivers, lakes, sewage canals and irrigation

channels.Sarotherodon galilaeus Africa and Middle East. Vegetated areas in rivers and lakes.Haplochromis flaviijosephi Endemic Jordan River system. Shallow water among stones and vegetation

in lakes, ponds and streams.Tristramella sacra Endemic-Lake Kinneret. Lake KinneretT. simonis simonis Endemic-Lake Kinneret. Lake Kinneret

Blennidae Salaria fluviatilis Lake Kinneret and Bet Netofa. Rivers and streams with shallow waters.

296 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Table 2Frequency (No.), relative abundance (%), species richness, Brillouin index and evenness of modern ichthyofauna collected from November 1988 to March 1990, from various stations along the shoresof Lake Kinneret (Gafny, 1992)

Family Species Total E-13 Stoneand Tree

E-21 stone E-26-sand E-9 sandy E-26-Tree W-36Boulders

W-8 Stone W33/34boulder

N % N % N % N % N % N % N % N % N %

Cichlidae A. flaviijosephi 482 8.1 9 1.5 71 9.1 0 0 0 0 22 3 71 5.4 223 24.5 86 9.4O. aureus 505 8.5 1 0.2 30 3.8 116 52 82 20 186 25.2 75 5.7 4 0.4 11 1.2S. galilaeus 1176 19.9 153 24.7 109 13.9 2 0.9 279 66 125 16.9 205 16.0 16 1.8 287 31.3T. simmonis 1 0.017 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0.1T. zilli 924 15.6 193 31.2 54 6.9 79 36 38 9 107 14.5 158 12.0 101 11.1 194 21.2

Cyprinidae B. canis 49 0.8 0 0 0 0 0 0 0 0 0 0 46 3.5 0 0 3 0.3B. longiceps 2 0.03 0 0 0 0 0 0 0 0 0 0 1 0.1 0 0 1 0.1C. damascina 31 0.5 0 0 0 0 3 1.4 0 0 0 0 20 1.5 0 0 8 0.9

Small rockycarps

G. ruffa 105 1.8 0 0 4 0.5 0 0 0 0 0 0 46 3.5 23 2.5 32 3.5H. nana 441 7.5 0 0 2 0.3 0 0 0 0 0 0 278 21 1 0.1 160 17.5P. kervillei 21 0.4 2 0.3 0 0 0 0 0 0 7 0.9 9 0.7 0 0 3 0.3

Blennidae B. fluviatilis 1886 31.9 249 40.2 416 53.1 21 9.5 21 5 247 33.4 395 30 441 48.5 96 10.5Clariidae C. gariepinus 6 0.1 0 0 0 0 0 0 0 0 1 0.1 1 0.1 0 0 4 0.4Poecillidae G. affinis a 59 1 0 0 0 0 1 0.5 0 0 27 3.7 0 0 1 0.1 30 3.3Balitoridae O. tigris 227 3.8 12 1.9 98 12.5 0 0 0 0 17 2.3 0 0 100 11.0 0 0Total 5915 100 619 100 784 100 222 100 420 100 739 100 1305 100 910 100 916 100Sp. Richness 15 7 8 6 4 9 12 9 14Brillouin index 0.833 0.530 0.615 0.440 0.408 0.709 0.818 0.593 0.792Eveness 0.708 0.627 0.681 0.566 0.677 0.743 0.758 0.621 0.691

E = East side of Lake Kinneret.W = West side of Lake Kinneret.a Gambusia affinis is an exotic species introduced to Lake Kinneret since 1920s'.

297I.Zohar

etal.

/Palaeogeography,

Palaeoclim

atology,Palaeoecology

258(2008)

292–316

Author's personal copy

Mirogrex terraesanctae, large cyprinids (e.g., Caraso-barbus canis, Barbus longiceps, Capoeta damascina),small cyprinids (small bones that can be identified onlyto family level and might include Mirogrex sp., small/juvenile B. longiceps, C. canis, and C. damascina),Tilapinii, and Clarias gariepinus. For each taxonomicgroup the data was summarized using the number ofidentified specimens (NISP) and minimum number ofindividuals (MNI) (Grayson, 1984; Klein and Cruz-Uribe, 1984; Reitz and Wing, 1999).

For each taxon we calculated bones relative abun-dance (NISP; Appendix A). Since only few bones werespecies specific, we group the bones according to nineanatomic regions (following the terminology used byWheeler and Jones (1989): neurocranium, branchialregion, hyoid region, oromandibular region, and opercu-lar series from the cranial region, and appendicularskeleton, median fins, Weberian apparatus, and vertebralcolumn from the postcranial region; Table 3), andcranial vs. postcranial regions (following Butler, 1990).

3.4. Data analysis

In order to identify the taphonomic agents respon-sible for fish bone accumulation and information loss,we examined several attributes, and calculated variousquantitative and qualitative indices as described below.

3.4.1. Taxonomic richness and diversityTaxonomic richness and diversity were calculated

from species-specific bones. Species richness (S) wascalculated as the number of genera identified in eachsample (Krebs, 1999). Species diversity was calculated

using the Brillouin Index (HB), since it is most sensitiveto the abundance of the rare species in the sample (Krebs,1999).

We compared the HB diversity index for the boneand the living assemblages. Due to the paucity of data offish communities prior to stocking and commercialfishing, we used fishing reports as they partially reflectfish composition at the pelagic habitat. However, we didnot include in our analysis data on introduced exotic fishsuch as common carp, silver carp and rainbow trout(Sarig, 1982; Goren, 1983; Gophen and Gal, 1992;Goren and Ortal, 1999).

Since species richness depends on sample size, andmany paleontological samples are of small sizes,sample-size effects should always be evaluated beforedifferences or similarities are assumed between assem-blages (Koch, 1987; Krebs, 1999; Zohar and Belmaker,2005; Bulinski, 2007). Several methods have beendeveloped to deal with samples that differ in size and areneither random nor normally distributed (Grayson,1984; Kintigh, 1984; Plog and Hegmon, 1993; Kauf-man, 1998; Baxter, 2001). To test the adequacy of thesmaller sample sizes with respect to the lithofacies, wecalculated species richness expected in larger samplesizes, using “the equivalent alpha diversity method forabundifaction” (Koch, 1987; Hayek and Buzas, 1997).By fitting logarithmic series to the data, we calculatedalpha diversity, and used the statistics to predict thenumber of species expected in a larger sample size(Hurlbert, 1971; Heck et al., 1975; Grayson, 1984;Krebs, 1999; Baxter, 2001). To compare between theexpected species richness generated for various NISP,by lithofacies, a point by point rarefaction was

Table 3Number (NISP) and percentage of bones, by anatomic regions, expected in complete skeleton of five taxa of freshwater fish

Anatomic region ⁎ M. terraesanctae Large cyprinids ⁎⁎ Tilapinii C. gariepinus

NISP % NISP % NISP % NISP %

CraniaNeurocranium 51 23.83 51 22.08 48 24.62 42 24.0Branchial region 50 23.36 58–60 25.11 36 18.46 22 12.6Hyoid region 16 7.48 16 6.93 16 8.21 14 8.0Oromandibular region 20 9.35 20 8.66 20 10.26 18 10.3Opercular series 8 3.74 8 3.46 8 4.10 6 3.4

PostcraniaAppendicular skeleton 16 7.48 16 6.93 16 8.21 10 5.7Median fins 4 1.87 7 3.03 21 10.77 – –Weberian apparatus 10 4.67 10 4.33 – – –Vertebral column 39 18.22 45 19.48 30 15.38 63 36.0Total 214 100% 231–233 100% 195 100% 175 100%

⁎Does not include the following elements: Scales, Ribs, intermuscular, pterygiophore (except for the 1st dorsal or anal pterygiophore), soft fin ray,epural, hypural, urostyle and radials.⁎⁎Large cyprinids include Barbus longiceps, Carasobarbus canis and Capoeta damascina.

298 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

employed by using the program Analytic RarefactionV.1 (Holland, 2001).

3.4.2. Skeletal element presentationPrevious studies demonstrated that in different

depositional environments different agents of destruc-tion may be involved leading to differences in skeletalelement representation, abundance and density (i.e.,water level, temperature, wave activity, scavengersactivity, sedimentology, etc.; Behrensmeyer, 1983;Elder and Smith, 1988; Stewart, 1989; Butler, 1993).The issue of bone survivorship has been much discussedin the literature and has resulted in a wide variety ofmethods (e.g.,Grayson, 1984; 1991; Klein and Cruz-Uribe, 1984; Lyman, 1994a,b; Nagaoka, 2005; Reitzand Wing, 1999; Schick et al., 1989; Wolverton, 2002).

Due to the high variability in body forms and bonesamong fishes (Casteel, 1976; Colley, 1986, 1990;Gobalet, 2001; Nicholson, 1998; Wheeler, 1978;Wheeler and Jones, 1989) we opted to apply a variationof the method developed by Reitz and Zierden (1991;see Reitz and Wing, 1999:211 for further discussion).

To evaluate the survivorship of each bone, by taxon,we calculated survival (S.I.) and fragmentation indices(F.I.). The SI represents the ratio between the number ofobserved bones (NISP) to the number of expected bones(per skeletal element in each anatomic region; seeTable 3) in a complete fish skeleton (per species). Theexpected value of each anatomic region, per taxon, wascalculated as its ratio in a complete fish multiplied by thetotal NISP observed at the site (Zohar et al., 2001). WhenSI = 1, the observed NISP = expected NISP. SI larger than1 implies over-representation, while SI smaller than 1indicates under-representation.

Since abundant bones (SIN1) may reflect a highoccurrence of a body region, or high state of fragmen-tation that increases the NISP, we calculated a fragmen-tation index (F.I.) (see Zohar et al., 2001). FI is calculatedfrom the relative proportion of the retrieved bonecompared with a complete bone (81–100%; 61–80%;41–60%; 21–40%; less than 20%). A bone with high SI(N1) and low FI (∼10–50%) may reflect a high state offragmentation rather than good survivorship; it attests tohigh levels of post-depositional destruction (Zohar et al.,2001).

The differences in the number of observed andexpected bones (NISP) were compared using the chi-square contingency test.When contingency tables containcells of nb5, similar rows were combined or acontingency table was constructed with probability valuescalculated using a randomization test (Manly, 1991).

Table 4Fish remains NISP a and standardized BSF b calculated in 24 squares,by lithofacies volume c

NISP by Lithofacies Standardized BSF byLithofacies

Square no. A B C Total A B C

10 17 32 9 58 13.6 25.6 7.211 16 9 0 25 12.8 3.6 0.012 14 13 18 45 28.0 8.7 10.313 3 9 66 78 2.4 7.2 52.814 7 187 0 194 5.6 74.8 0.015 78 9 82 169 44.6 4.5 65.616 3 22 61 86 1.2 17.6 48.817 0 12 7 19 0.0 9.6 4.018 3 5 0 8 2.4 4.0 0.019 34 328 2532 2894 27.2 262.4 2025.620 19 39 20 78 25.3 22.3 16.021 31 42 154 227 31.0 33.6 123.222 23 36 14 73 46.0 36.0 6.225 0 19 5 24 0.0 15.2 4.027 3 28 216 247 12.0 11.2 172.828 0 8 16 24 0.0 4.6 12.829 19 383 22 424 25.3 218.9 8.830 5 44 21 70 20.0 17.6 16.831 0 43 333 376 0.0 24.6 133.234 22 0 18 40 88.0 0.0 14.435 31 20 23 74 24.8 8.0 18.436 6 0 440 446 12.0 0.0 352.039 2 48 47 97 8.0 19.2 37.640 3 9 7 19 12.0 3.6 5.6Total NISP 339 1345 4111 5795% NISP 6.0% 23.0% 71.0% 100%

Lithofacies A = the surface sand stratum (3–10 cm thick).Lithofacies B = a median layer of mixed sand-silt stratum (5–10 cmthick).Lithofacies C = the bottom stratum of fine grain silt and clay.a NISP = Number of identified specimen.b Standardized BSF = Bone scatter frequency: NISP bones/

sediment volume.c Lithofacies volume = Lithofacies maximum depth×0.5 m2.

Table 5Morisita index of dispersion, standardized bone scatter frequency(BSF), taxon richness and diversity (HB) calculated for the boneassemblage, by three lithofacies (df=23)

Indices Lithofacies Totalsample

A B C

Morisita Id 2.4 4.0142 9.6617 6.5207Morisita Mu 0.9665 0.9916 0.9922 0.9980Morisita Mc 1.0446 1.0112 1.0037 1.0062Morisita Ip 0.5295 0.5653 0.6882 0.6200Mean BSF 22.1 37.8 150.0Min. BSF 1.2 3.6 4.0Max. BSF 88.0 262.4 2025.6Taxon richness 5 4 4HB 0.347 0.243 0.061

Id = Morisita Index, Mu = Uniform index, Mc = Clumped Index, Ip =Standardizes Morisita Index of dispersion, HB = Brillouin diversity index.

299I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

3.4.3. Bone spatial distributionDifferences in the pattering of bone spatial distribution

is used to reflect differences in the agents of depositionand scattering, particulary for identifying the processesresponsible for creating these concentrations of bones(e.g., Behrensmeyer, 1982, 1983; Kidwell, 1985, 1986;Bonnichsen and Sorg, 1989; Lyman, 1994b; Tappen,1995 and references therein). Ecologists recognize threebasic types of spatial distribution: uniform, random, orclumped as indicators of paleoenvironmental conditionsduring carcass disarticulation and deposition (Wilson andBarton, 1996; Martin, 1999).

To test the bone spatial distribution pattern in eachlithofacies (across 24 squares) we used the standardizedMorisita index of dispersion (Hurlbert, 1971; Hecket al., 1975; Krebs, 1999). This index is one of the bestmeasures for distribution because it is independent ofpopulation density and sample size (Hurlbert, 1971;Heck et al., 1975; Krebs, 1999). We also calculated thefrequency (density) of the bones in each depositionalunit (standardized bone scatter frequency – BSF). Thestandardized BSF is calculated from the number ofbones in each square and lithofacies, divided bylithofacies volume (=lithofacies depth×0.5 m2) (Kid-well, 1985, 1986; Stewart, 1989; Zohar et al., 2001).

4. Results

4.1. The bone assemblage

The faunal remains recovered from the 24 excavatedsquares include a large number of freshwater mollusks,two rodent bones, 5037 fish bones and 758 scales. Here

we focus on the taphonomic processes affecting fishremains.

4.2. Spatial distribution of fish remains

The average number of fish remains (standardizedBSF) per square was 241 (SD 580), ranging from 0 to2025.6 bones when standardized per sediment volumein the lithofacies (Tables 4 and 5). The spatialdistribution exhibited a clear clumped pattern (MorisitaIndex, I pN0, Table 5) for each lithofacies. Moreover,the relative abundance of the remains increased withsediment depth (lithofacies), and distance from shore,independently of sediment volume. Of the 5037 skeletalelements retrieved, 6% were from lithofacies A, 23%from lithofacies B and 71% from lithoacies C (Table 4).When standardized by sediment volume similar ratiosare observed: 10% for lithofacies A, 19% for lithofcaiesB, and 71% for lithofacies C.

4.3. Taxonomic composition

In all, 1566 fish bones (ca. 31%; Appendix A) wereidentified to three families of primary and secondaryfreshwater fish (Cyprinidae, Cichlidae, and Clariidae;Fig. 3). Taxonomic identification to the species levelwas possible only for a small sample of bones (Table 6;NISP=256; MNI=65), and included the following fourspecies: Mirogrex terraesanctae, Barbus longiceps,Capoeta damascina, and Clarias gariepinus. To enlargesample size (NISP=1566, MNI=115) and includebones identified to genus or family level, we groupedthe bones into five taxonomic groups, as follows:

Fig. 3. The relative abundance (%) of each family per lithofacies.

300 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

C. gariepinus, Tilapinii (Tilapia sp., Oreochromisand Tristamella sp.), Mirogrex sp., large cyprinids(B. longiceps, C. canis, and C. damascina), and smallcyprinids (small bones that can be identified only tofamily level and might include Mirogrex sp., small/juvenile B. longiceps, C. canis, and C. damascina;Fig. 4 and Table 6). Other fish species currently presentat Lake Kinneret (see Table 1 for the complete list) were

not included in the taxonomic groups listed above due toabsence of their distinctive bones as well as theirabsence from the sandy shore community (Gafny, 1992;Voskoboynik, 1995).

There is a consistency in taxonomic compositionbetween lithofacies (Figs. 3 and 4), except for theexclusive appearance of Clarias gariepinus in lithofa-cies A. As a result taxon richness was higher for

Table 6NISP, MNI (in parenthesis) and relative abundance (%) of fish remains from the bone assemblages, by taxon, compared to their frequency in livingassemblages from a modern littoral sampling of a southern sandy beach (E-9) (Gafny, 1992), and fishing reports (Ricardo-Bertram, 1947; Sarig, 1981;Gophen and Gal, 1992)

Taxonomicgroup

Bone assemblage bylithofacies

Total in boneassemblage

Littoral-live ass. Fishing reports

A B C NISP (MNI) % E-9 1935–36, 39 1940–1943 1980

M. terraesanctae 13 64 159 236 15.0 – 35.4 60.7 43.7(13) (14) (29) (56)

Small cyprinid a 89 363 697 1149 73.4 –(4) (14) (22) (40)

B. longiceps 0 4 1 5 0.32(4) (1) (5)

C. damascina 2 10 2 14 0.9(1) (1) (1) (3)

Barbus sp. b 0 1 2 3 0.2 – 33.1 22.1 7.9(1) (2) (3)

Tilapinii c 25 49 83 157 10.0 95.0 30.5 16.3 28.5(2) (2) (3) (7)

C. gariepinus 1 0 0 1 0.06 – 0.0 0.0 0.5(1) (1)

S. fluviatilis – – – – – 5.0Total NISP 130 492 944 1566 100% 420 fish 302 ton 417 ton 1950 tonTotal MNI 21 36 58 115

a Small cyprinids— small bones that can be identified only to family level and might includeMirogrex sp., small/ juvenile B. longiceps, C. canis,and C. damascina.b Large cyprinid = C. canis, B. longiceps. or C. damascina.c Tilapinii = Tilapia sp., Oreochromis and Tristamella sp.

Fig. 4. The relative abundance (%) of each taxon per lithofacies.

301I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

lithofacies A (S=5) and lower for lithofacies B and C(S=4). Due to differences in species richness and inthe relative abundance of each taxonomic group (r×crandomization test, χ2 =45.831, df=8, p=0.0001;Fig. 4), the Brillouin Diversity Index (HB) differedsignificantly between lithofacies (Table 5). The up-per lithofacies displays the highest diversity value

(HB=0.347) while the bottom lithofacies (C) displaysthe lowest value (HB=0.061).

4.4. Skeletal parts representation and preservation

Of the total 5037 fish elements retrieved, 2277 (45%)were identified to body elements (Appendix A) and

Table 7NISP of anatomic regions retrieved in the bone assemblages, by taxonomic group and lithofacies

Anatomic region Cyprinidae

M. terraesanctae Small Large Tilapinii Clarias Total

Lithofacies ACraniaNeurocranium 0 6 0 2 0 8Branchial region 2 6 2 1 0 11Hyoid region 1 8 0 1 0 10Oromandibular region 2 5 0 3 0 10Opercular series 0 4 0 3 0 7Cranial general 0 0 0 0 0 0

PostcraniaAppendicular skeleton 0 8 0 1 0 9Median fins 0 4 0 7 0 11Vertebral column 8 45 0 7 1 61Rib 0 3 0 0 0 3

Total Lithofacies A 13 89 2 25 1 130

Lithofacies BCraniaNeurocranium 1 20 0 6 0 27Branchial region 21 13 15 5 0 54Hyoid region 1 33 0 1 0 35Oromandibular region 1 45 0 1 0 47Opercular series 0 36 0 1 0 37Cranial general 0 0 0 0 0 0

PostcraniaAppendicular skeleton 1 54 0 3 0 58Median fins 0 18 0 13 0 31Vertebral column 39 137 1 15 0 192Rib 0 7 0 4 0 11

Total Lithofacies B 64 363 16 49 0 492

Lithofacies CCraniaNeurocranium 0 60 0 6 0 66Branchial region 47 14 4 5 0 70Hyoid region 12 66 0 0 0 78Oromandibular region 29 54 0 4 0 87Opercular series 2 74 0 3 0 79Cranial general 0 0 0 0 0 0

PostcraniaAppendicular skeleton 0 108 1 10 0 119Median fins 0 19 0 20 0 39Vertebral column 69 297 0 30 0 396Rib 0 5 0 5 0 10

Total Lithofacies C 159 697 5 83 0 944

Small cyprinids — most of them are probably skeletal remains of M. terraesanctae that are not species specific.Large cyprinid = C. canis or B. longiceps and C. damascina.Tilapinii = Tilapia sp., Oreochromis and Tristamella sp.

302 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

grouped by anatomic regions (Table 7). Most (84%) ofthe remains exhibited good preservation (low fragmen-tation—FI), i.e., more than 80% of the bone elementsurvived (Table 8). High fragmentation was observedonly for bones from the branchial region, appendicularskeleton, and ribs. As most of the bones were in agood state of preservation, the survival index (SI) wasexamined for each bone and by anatomic regions(Table 9). The bone survivorship (SI) by anatomicregions was found to vary with fish taxon and lithofacies(Table 9). For example, the oromandibular region wassignificantly over-represented for small cyprinid remains

from lithofacies B, and for M. terraesanctae remains inlithofacies C. The opercular region was over-representedfor small cyprinid remains from lithofacies B and C, andsignificantly differed from the expected in a complete fish.The median fin is significantly over-represented in alllithofacies. In lithofacies C the relative abundance of M.terraesanctae hyoid region was similar to a completeskeleton. A similar pattern was observed for smallcyprinids from lithofacies A and B. For small cyprinidsfrom lithofacies C, this region was significantly over-represented. Except for large cyprinids, the vertebralcolumn is over-represented (SIN1) in all samples.

The sole case inwhich bone over-representation can beattributed to high fragmentation is that of the appendicularskeleton which is over-represented for small cyprinidsfrom all lithofacies (Tables 8 and 9). An opposite trend isobserved for the branchial region, which despite highfragmentation is under-represented, in all samples, exceptfor M. terraesanctae remains from lithofacies C.

The relative frequencies of cranial and postcranialelements retrieved (Table 7), differ significantly fromthose expected in a complete skeletonized fish (Tables 3and 10). Except for large cyprinids, the postcranial regionis over-represented regardless of taxon and depositionalarea (Table 10). Fragmentation may have biased theperceived abundance of cranial bones since once they arebroken, only few bones can be identified. However, mostof the postcranial bones are well preserved (in 91.0% ofthe cases N80% of the bone is present; Table 8).

As the bones from the different anatomic regionsexhibit good preservation (Table 8), we cannot attributehigh SI to high rate of fragmentation, but rather totaphonomic processes. The improvement in bone pre-servation with stratigraphic depth (χ2=71.218, df=6,

Table 8Percentage (%) of skeletal elements state of preservation (FI) (alllithofacies combined), by anatomic region (NISP=3017)

State of preservation (%)

Anatomicregion

1–20%

21–40%

41–60%

61–80%

81–100%

Total%

Count

Crania regionNeurocranium 0.9 1.7 8.5 21.4 67.5 100.0 117Branchial region 3.7 27.9 34.6 19.9 14.0 100.0 136Hyoid region 1.6 2.4 8.7 19.8 67.5 100.0 126Oromandibular

region0.0 2.0 4.7 10.1 83.1 100.0 148

Opercular series 0.7 2.0 12.8 20.1 64.4 100.0 149

Postcrania regionAppendicular

skeleton3.3 14.1 20.2 9.4 53.1 100.0 213

Median fins 0.9 9.7 15.9 2.7 70.8 100.0 113Rib 0.0 16.7 29.2 4.2 50.0 100.0 24Vertebral column 0.1 0.0 0.3 4.0 95.5 100.0 1212unidentified 4.8 4.8 4.8 4.8 81.0 100.0 21Scales 0.0 0.0 0.0 0.1 99.9 100.0 758

FI = Fragmentation index.

Table 9Survival indices (SI) calculated for fish remains retrieved from the bone assemblages by taxonomic group, anatomic region and lithofacies

Anatomic region Small cyprinid Tilapinii M. terraesanctae

A B C A B C C

CraniaNeurocranium 0.30 ⁎ 0.25 ⁎ 0.39 ⁎ 0.32 0.49 0.29 ⁎ 0.00 ⁎

Branchial region 0.27 ⁎ 0.14 ⁎ 0.08 ⁎ 0.21 0.55 0.33 ⁎ 1.26Hyoid region 1.30 1.31 1.37 ⁎ 0.48 0.25 0.00 ⁎ 1.01Oromandibular region 0.65 1.43 ⁎ 0.89 1.17 0.20 0.47 1.95 ⁎

Opercular region 1.30 2.86 ⁎ 3.07 ⁎ 2.92 0.50 0.88 0.34

PostcraniaAppendicular skeleton 1.30 2.15 ⁎ 2.24 ⁎ 0.48 0.75 1.47 0.00 ⁎

Median fins 2.60 ⁎ 2.27 ⁎ 1.14 2.60 ⁎ 3.22 ⁎ 2.78 ⁎ 0.00 ⁎

Weberian apparatus 0.00 ⁎ 0.00 ⁎ 0.00 ⁎ – – – 0.00 ⁎

Vertebral column 2.60 ⁎ 1.94 ⁎ 2.19 ⁎ 1.82 1.99 ⁎ 2.35 ⁎ 2.38 ⁎

Due to small sample size, M. terraesanctae SI, by anatomic regions, is calculated only for facies C.⁎ Significantly different from the expected in a complete fish, pb0.05 (Chi-square test).

303I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

pb0.0001) suggests that the lithofacies are taphonomi-cally distinct in terms of fragmentation and that eachlithofacies assemblage has a unique taphonomic history.A similar conclusion can be drawn from the scales relativefrequencywithin the lithofacies. Although scaleswere notidentified to taxa, the 758 scales recovered (NISP bylitofacies: A=103; B=252; C=403) exhibited a decreasein their relative abundance with stratigraphic depth(A=30%, B=19%, C=10%).

4.5. The bone assemblages vs. the living assemblages

4.5.1. Fidelity measures among data setsComparison of the fish bone assemblages with the

living assemblages (Sarig, 1981, 1982; Gafny, 1992;Voskoboynik, 1995) exhibited the following discrepan-cies (Table 6):

1. Species richness: The taxon richness in the boneassemblages (S=4–5) is lower than that currentlyfound overall in Lake Kinneret (S=19; Table 1) and inrocky shores (S=14). However, low species richness(S=4) also characterizes the living community at thesouthern sandy shore (E-9; Table 2) (Gafny et al., 1992).

2. Diversity index: The diversity index (HB) of the boneassemblages is significantly lower (HB=0.061–0.347) than that of sandy shores living assemblages(HB=0.408; Table 2).

3. Family representation: Native representative from thefamilies Cyprinidae, Cichlidae, and Clariidae arefound in the bone assemblages. While these familiesalso appear in the fishing reports (pelagic region),Cyprinidae and Clariidae are absent from the littoral

sampling (E-9). The southern sandy shore (E-9) habitatis characterized solely by the presence ofCichlidae andBlennidae. Interestingly, contrary to the Cichlidpreponderance in the living community (95%), theyare scarce in the bone assemblages (10%; Table 6).

4. Taxon representation: Fidelity among live–deadassemblages can be examined by comparing thedominant taxa. In this study live–dead differenceswere high for all taxa identified, as follows: a). Themost abundant species in the sandy littoral sampling(E-9) is S. galilaeus (66%; Cichlidae) that alsoexhibits sharp periodical changes in its relativeabundance (Table 11). In the bone assemblagesTilapinii are less abundant and comprise only 10% ofthe identified bones. b). The littoral fish S. fluviatilis(Blennidae) is absent from the bone assemblages. c).M. terraesanctae is the most abundant taxon in thebone assemblages (15%; Table 6). This is surprisingas M. terraesanctae are pelagic fish that aggregate inschools as observed from their preponderance in thefishing reports (Sarig, 1981, 1982; Gafny et al.,1992). Moreover, although they do exploit the littoralzone during the wintertime for breeding, this has notyet been documented in a sandy shore (Voskoboynik,1995; Gasith et al., 1996). d). The absence of largecyprinids (C. canis, B. longiceps and C. damascina)from the sandy shore (E-9) possibly reflects habitatpreferences, as observed in their higher occurrence inthe fishery report (10%; Table 6). Interestingly,contrary to the littoral living assemblages, largecyprinids appear, although in small numbers, in thebone assemblages (Table 6). e). The catfish are rare inall samples (b1%). Their rarity in the fishing reports

Table 10Observed and expected percentage and SI of cranial and postcranial bones in the bone assemblages, by taxonomic group and lithofacies

Lithofacies Genus group Cranial Postcranial Total

Obs. % Exp. % SI Obs. % Exp. % SI NISP χ2

A M. terraesanctae 38.5 66.0 0.58 61.5 34.0 1.81 13 2.5 nsSmall cyprinids 32.6 63.0 0.52 67.4 37.0 1.82 89 16.4⁎

Large cyprinids 100.0 63.0 – 0.0 37.0 – 2 nsTilapinii 40.0 65.0 0.63 60.0 35.0 0.952 25 0.6 nsC. gariepinus 0.0 58.0 – 100.0 42.0 – 1 ns

B M. terraesanctae 37.5 66.0 0.57 62.5 34.0 1.84 64 10.1⁎

Small cyprinids 40.5 63.0 0.64 59.5 37.0 1.61 363 37.1⁎

Large cyprinids 93.7 63.0 1.50 6.2 37.0 0.169 16 4.6 nsTilapinii 28.6 65.0 0.45 71.4 35.0 1.13 49 5.2⁎

C M. terraesanctae 56.6 66.0 0.86 43.4 34.0 1.28 159 2.9 nsSmall cyprinids 38.4 63.0 0.61 61.5 37.0 1.66 697 83.9⁎

Large cyprinids 80.0 63.0 – 20.0 37.0 – 5 –Tilapinii 21.7 65.0 0.34 78.3 35.0 1.24 83 15.8⁎

⁎ Fisher exact test significant different pb0.05, df=1.SI was not calculated for large cyprinids and Clarias sp. due to small sample size.

304 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

can be attributed to their low economic value (thefishermen tend to throw the catfish back to the lake),rather than to actual abundance in this habitat.However, their appearance in the northern rockyshores and absence from the southern shores (Table 2)probably reflects habitat preferences and thereforemayexplain their rarity in the bone assemblages.

4.5.2. Taxonomic composition in responds to habitatsubstrate

To provide a visual representation of the patternof proximities (i.e., similarities or distances) amongspecies from the living communities according to theirrelative abundance at different habitat/substrate alongLake Kinneret, multidimentional scaling (MDS) wasused (Borg, 1981; Busing et al., 1996; Borg and Groene,1997). This analysis indicates that species compositiondiffers according to habitat substrate (Fig. 5). Therefore,the sandy shore communities (E-9Sa and E-26Sa) differfrom other habitat communities, and are close to thedeath assemblage community (Fig. 5; Stress=0.1911,Variance=0.98089). The presence ofM. terraesanctae atthe bone assemblage (natural) is unique since this speciesis absence from the living littoral communities sampled inthis study. This result demonstrates that there are notabledifferences between the expected taxonomic compositionby habitat and those observed at the bone assemblages, allprobably reflecting variations due to differential deposi-tion, dispersal, and preservation of bones within thelacustrine and coastal taphonomic processes.

4.5.3. Taxonomic composition and sampling biasDue to the differences in taxonomic composition

between the bone assemblages and living community,we examined the possibility that sample size biasedthe bone assemblage. Calculation of the rarefactioncurves (Holland, 2001) reveals that species richness oflithofacies C reached the asymptote at 300 bones, andrichness of lithofacies B reached the asymptote at 100bones. However, the sample size from lithofacies A didnot reach the asymptote (Fig. 6). For this lithofacies weexamined species richness estimated for larger samplesizes. Based on N1 of 130 bones and S1=5 we calculatedan alpha (α1) value of 1.032. We used the α1 to examinehowmany species (S2) we would expect to find in a largersample (N2) of 500 bones from lithofacies A. Speciesrichness (S2) increased to 6.4 for a sample size of 500bones (NISP; Fig. 6). This result exhibits that speciesrichness, diversity and composition in lithofacies A(surface) is potentially higher (SN6) in a larger sample,compared with other lithofacies (S=4) and with theliving assemblage. This result strengthens previous ob-servation that species composition among lithofaciesreflects variation between depositional microenviron-ments and taphonomic processes in lacustrine and coastalhabitats.

4.5.4. Fish body size distributionThe modern fish sampled from the littoral zone

exhibited an abundance of relatively small body sizedfish (Table 12). Of the eight potentially “large” species

Table 11Relative abundance (count and percent) of fish, by species, of modern ichthyofauna collected from November 1988 to March 1990, from variousstations along the shores of Lake Kinneret (Gafny, 1992)

Species Total Month

Nov. 88 Dec. 88 Jan. 89 March 90 April 89 July 89 Oct. 89

N % N % N % N % N % N % N %

H. flaviijosephi 482 39 8.1 5 1.0 23 4.8 30 6.2 9 1.9 167 34.6 209 43.4O. aureus 505 76 15.0 0 0.0 3 0.6 8 1.6 16 3.2 402 79.6 0 0.0S. galilaeus 1176 154 13.1 2 0.2 5 0.4 4 0.3 64 5.4 571 48.5 376 32.0T. simmonis 1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 100.0 0 0.0T. zillii 924 139 15.0 113 12.2 194 21.0 52 5.6 260 28.1 137 14.8 29 3.1C. canis 49 19 38.8 0 0.0 0 0.0 0 0.0 27 55.1 3 6.1 0 0.0B. longiceps 2 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 100.0C. damascina 31 11 35.5 0 0.0 9 29.0 5 16.1 3 9.7 2 6.4 1 3.2G. rufa 105 44 41.9 0 0.0 3 2.9 0 0.0 4 3.8 23 21.9 31 29.5H. nana 441 265 60.1 0 0.0 9 2.0 1 0.2 1 0.2 38 8.6 127 28.8P. kervillei 21 0 0.0 0 0.0 12 57.1 0 0.0 9 42.9 0 0.0 0 0.0S. fluviatilis 1886 34 1.8 167 8.8 55 2.9 317 16.8 265 14.0 459 24.3 589 31.2C. gariepinus 6 0 0.0 0 0.0 0 0.0 0 0.0 2 33.3 4 66.7 0 0.0G. affinis a 59 0 0.0 0 0.0 1 1.7 2 3.4 1 1.7 52 88.1 3 5.1N. tigris 227 5 2.2 3 1.3 0 0.0 2 0.9 15 6.6 121 53.3 81 35.7Total 5915 786 290 314 421 676 1980 1448

a Introduced species.

305I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

(N25 cm max. SL; Table 12) only three exhibitedsome larger sized fish: S. galilaeus, C. Canis, andC. Damascina. It could be argued that the preponder-ance of small body sized fish result from over-fishing ofthe larger one. However, considering that fishing reportsdo not suggest a dramatic reduction in fish body size andabundance over the last 50 years, this option wouldappear unlikely. Moreover while the benthic habitat ishighly exploited by the smaller sized species, larger fish

tend to exploit the pelagic zone during the day and arriveat the littoral zone exclusively at night or during theirbreeding season (Sarig, 1981; Gafny, 1992; Gophen andGal, 1992). Therefore, small sized fish characterizes thepopulation exploiting the littoral zone.

Small sized fish are abundant also at the boneassemblage as observed from the vertebrae widthdiameter (range: 1.1–3.7 mm; mean=1.7 mm±0.7;NISP=77). Due to low survival of atlas and axis in thebone assemblages, body size reconstruction was possibleonly on 19 atlas vertebrae of M. terraesanctae (Zohar,2003). Estimation of fish standard length exhibited arange between 65 to 127 mm. Since M. terraesanctaemaximum SL reach 220 mm, we assume that most of theremains represent juvenile fish at their second year(Ostrovsky and Walline, 1999).

5. Discussion

Many studies exhibited the significance of fishremains preserved in the sediments as proxies forpaleo-population dynamics (Soutar, 1966; Soutar andIsaacs, 1974; O'connell and Tunnicliffe, 2001; Wrightet al., 2005). They also demonstrated that the fidelity offossil assemblages and patterns of information lossshould be examined for each aquatic habitat (Kidwelland Bosence, 1991; Butler, 1996; Kidwell, 2001;O'connell and Tunnicliffe, 2001; Tunnicliffe et al.,2001; Kidwell, 2002). In this study several biases wereobserved for fish live-dead assemblages from thesouthern shore of Lake Kinneret (Table 13).

Fig. 6. Point by point rarefaction curves for species richness by lithofacies as function of NISP, and speies richness expecte in a larger sample size forfacies A (black dot).

Fig. 5. Multidimentional scaling for comparison of living species relativeabundance at different habitats at Lake Kinneret and the bone assemblage.

306 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

5.1. Presentation of life assemblages within the boneassemblages

Studies of sedimentary fish remains in differenthabitats show that if taxonomic biases occur then weexpect to uncover the most abundant species (Soutar,1966; Soutar and Isaacs, 1974; O'connell and Tunnicliffe,2001; Tunnicliffe et al., 2001). In this study an oppositepattern occurred (Fig. 6). While habitat preferences andpopulation dynamicsmay explain the low number of largecyprinids (C. canis, B. longiceps and C. damascina) andcatfish (C. gariepinus) they cannot explain the lowabundance of Tilapinii or the absence of Blennidae thatrepeatedly exploit the sampled habitat. Moreover, thepreponderance of small cyprinids, especially M. terrae-sanctae, is extremely surprising since they are not foundin the living assemblages of the sandy shore (Voskoboy-nik, 1995; Gasith et al., 1996).

Our findings demonstrate that in the southern shoreof Lake Kinneret the bone assemblages differ from thetaxonomic composition and diversity of the lifeassemblage, as observed from the presence of Cyprini-dae, absence of Blennidae and relatively low frequencyof Cichlidae.

Since all three lithofacies are similar in taxonomiccomposition (except for the single apperance ofC. gariepinus at litofacies A) and do not exhibit changesthrough time, we assume that the differences between thebone and living assemblages represent a respond todeposition and preservation processes under complex andfluctuating lacustrine and coastal environments (see nextsection).

5.2. Preservation and depositional processes

Consideration of bone preservation is necessary formeaningful ecological and taphonomic inferences espe-cially since studies showed that due to microbial attackmost buried bones do not survive into the fossil record(Berger, 1976; Trueman andMartill, 2002; Trueman et al.,2003). In general, the processes leading to bone pre-servation and fossilization are complex and difficult todefine (Lyman, 1984, 1994b; Trueman andMartill, 2002).

In this study we observed that clumps of disarticulatedfish remains characterize assemblages from the southernshore of Lake Kinneret. We assume that regardless ofdepositional area, the fish remains must have suffered fromphysical and chemical weathering since theywere subjected to repetitive cycles of lake level changesand vegetation growth (Hazan et al., 2005). However, sincethe bones' relative abundance (survival) tends to varyalong a taphonomic gradient (depth and distance fromshore), we attribute further mechanical and chem-ical abrasion to the bones embedded on the surface(lithofacies A). As a result, fishes are more common andwell preserved in collections made stratigraphically fartherfrom the recent shore and in deeper layers (lithofacies C).The occurrence of a taphonomic gradient is not exceptionalfor Lake Kinneret and has been described in severallacustrine deposits (Schäfer, 1972; Elder and Smith, 1988;Wilson, 1988; Wilson and Barton, 1996; Martin, 1999).These studies showed that although species represented inthe bone assemblages are, in many cases, not representa-tives of the long-term ecological, much less evolutionary,population dynamics, they do provide a “snapshot” into

Table 12Standard length (SL in mm) observed on modern ichthyofauna collected from November 1988 to March 1990, from various stations along the shoresof Lake Kinneret (Gafny, 1992) compared to data published on population body size a

Family Species Modern sample SL (Gafny, 1992) Population SL (mm) a

Count Mean Min. Max. Range Max.

Cichlidae A. flaviijosephi 482 63.2 10.0 115.0 20–100 128.0O. aureus 505 45.4 17.0 191.0 200–250 457.0S. galilaeus 1176 47.6 8.0 220.0 150–250 410.0T. zillii 924 86.7 30.0 183.0 100–200 400.0T. simmonis 1 – 41.0 – 150–200 250.0

Cyprinidae C. canis 49 92.7 60.0 271.0 200–400 450.0B. longiceps 2 123.0 108.0 138.0 250–500 500.0C. damascina 31 111.7 72.0 232.0 150–300 500.0G. rufa 105 66.7 27.0 151.0 50–120 140.0H. nana 441 72.0 38.0 108.0 40–80 120.0P. kervillei 21 47.5 39.0 59.0 20–60 100.0

Blennidae S. fluviatilis 1886 38.1 17.0 87.0 10–100 150.0Clariidae C. gariepinus 6 291.3 231.0 430.0 400–1000 1700.0Poecillidae G. affinis 59 27.7 20.0 51.0 20–40 40/70Balitoridae N. tigris 227 46.9 26.0 64.0 30–60 73.0

a Data from http://www.fishbase.org and Golani and Darom, 1997.

307I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

true biotic dynamics of short-term population fluctuations(Wilson and Barton, 1996; Martin, 1999). Which “snap-shots” can be recognized from the fish remains recovered atthe southern shore of Lake Kinneret?

The increase in the number of bones, as a function ofstratigraphic depth combined with their clumped distribu-tion and the presence of M. terraesanctae, conforms towinter-kill episodes. During this period water level ishigh, water temperature is low, bacteriological action islow and the small dead fish are trapped in the claysediments (Schäfer, 1972; Elder and Smith, 1988;Wilson,1988; Wilson and Barton, 1996; Martin, 1999). The lowpreservation of large cyprinids, cichlids, and catfish, mayreflect carcass flotation and wave energy (Schäfer 1972).Studies have shown that while small fish skeletons restinguncovered on the bottom of the lakewill remain depositedin the sediments, skeletons of large fish may be moved by

currents and scavengers (Schäfer, 1972; Elder and Smith,1988; Wilson and Barton, 1996). It has also been sug-gested that in relatively shallow water, warmer tempera-tures of the following spring may also cause decay andflotation of the largewinter-killed fish (Wilson andBarton1996). Thus the rarity of larger fish in the sampled areamay be explained either by habitat preferences or bydecay, flotation and wave energy during the followingspringwhenwater temperature rises (Schäfer, 1972; Elderand Smith, 1988; Wilson and Barton, 1996).

The rare and exclusive occurrence of catfish remains atthe surface lithofacies is surprising since clariids are one ofthe most commonly reported fish in African paleontologyand archaeology (Van Neer, 1986; Stewart, 1989; VanNeer, 1994; VanNeer, 2004). Clarias remains recovered atthe Nile Valley and Lake Turkana archaeological siteswere attributed to increased abundance due to the

Table 13Summary of fish bone assemblage taphonomic characteristics, in three lithofacies, from southern shore of Lake Kinneret

Characteristics Lithofacies A Lithofacies B Lithofacies C

Bone distribution Clumps of bones and scales

NISP range 2–78 bones 5–383 bones 5–2532 bonesNISP mean per 0.252 17 bones per square 61 bones per square 196 bones per squareBSF range 1.2–88 bones 3.6–262.4 bones 4–2025.6 bonesBSF mean 22 bones 38 bones 150 bones

Bone packing of accumulation Dispersed or in loosely packed clumps Densely packed clumps

Bone articulation status Disarticulated and sparse bones

Bone fragmentation (FI)FIb70% 14.2% 18.5% 8.7%FI 71–80% 4.2% 2.9% 3.5%FI 81–90% 17.2% 36.2% 42.1%FI 91–100% 64.6% 42.4% 45.7%

Bone sorting Poor sorting of size and shape. Poor sorting of size and shape. High degree of small sized bones

Taxon richness 5 4 4Evenness 0.577 0.510 0.127Brillouin diversity index 0.347 0.243 0.061

Taxonomic composition 3 families: Cyprinidae, Cichlidae, andClariidae M. terraesanctae, B. longiceps,C. canis, C. damascina., Tilapinii.,C. gariepinus.

2 families: Cyprinidae and Cichlidae 2 families: Cyprinidaeand Cichlidae

Mostly small cyprinids, andM. terraesanctae. Also B. longiceps,C. canis, C. damascina. Tilapinii,are less frequent.

Small cyprinids andM. terraesanctae arehighly abundant. B. longiceps,C. canis and C. damascina.are rare. Increase in Tilapinii.

Taxonomic composition vs.habitat substrate

No correlation

Skeletal remains Vertebral column is highly abundant

Crania vs. postcrania Postcrania region is over represented for all taxa except for large cyprinids.In large cyprinids the few remains recovered are pharyngeal bones.

308 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

combination of seasonal flooding and spawning activity(Van Neer, 1986; Stewart, 1989; Van Neer, 1989, 1993,2004). Boyer (1982) attributed catfish preservation toanaerobic environment and suggested that catfish that diein shallowwater are expected to buoy and further decay onthe shore, while only carcasses deposited in relativelydeep water would stay submerged despite the continuingdevelopment of gasses of decomposition. This scenariocombined with habitat preferences of catfish along LakeKinneret, may explain the rarity of catfish remains at thebone assemblage. Still, further research is needed toclarify which processes effect this species' preservation ingeneral and in the different lithofacies in particular.

5.3. Taxonomic composition and cultural bias

While discussing species composition of the boneassemblage, by lithofacies, we cannot disregard thepossibility that the upper lithofacies have been altereddue to fish stocking. Population dynamics observed inthe fisheries since 1939 exhibit fluctuations in largecarps (C. canis, B. longiceps and C. damascina) andTilapinii abundance's (Table 6) (Sarig, 1991, 1993).Large carps' populations sharply decreased since 1949,while since the 1980 s, an increase in the relativeabundance of introduced exotic species and Tilapinii isobserved (Table 6). While fish stocking may explain theabundance of Tilapinii in the modern littoral samplingassemblages, it is unclear in regard to their remains inlithofacies A of the bone assemblages. Moreover, inspite of stocking, large exotic fish such as the commonand silver carps are absent from the bone assemblages.

6. Conclusions

Live-dead assemblages of fish in the southern shore ofLake Kinneret present a unique case study to test differ-ences in species richness, abundance, and composition. Theliving assemblages represent fish communities that differbetween habitat/substrate type and that vary throughpopulation and seasonal dynamics. The bone assemblagediffers from the habitat specific community in speciescomposition and diversity. Although discrepancies in tax-onomic composition of living and bone assemblages havebeen described for other lacustrine deposits (e.g., Stewart,1991; O'connell and Tunnicliffe, 2001; Wright et al.,2005), in none were numerous pelagic fish found in littoralsampling. The absence/low frequency of littoral speciescontrary to the preponderance of pelagic species, especiallyM. terraesanctae, recovered at the southern shore of LakeKinneret, probably reflects seasonal mass death duringwintertime and small carcass deposition in clay.

From taphonomic and palaeoecological perspectives,the above findings suggest the following: 1) Bonesretrieved from lacustrine sediments may not represent thefull diversity of fish species and their relative abundancein the community. 2) Even when excavations are carriedout meticulously, erroneous conclusions regarding therelative abundance of fish can be deduced due to therandomly clumped nature of fish bone accumulations,regardless of depositional area and depth. 3) Fish remainsrecovered from coastal and inundated archaeologicalsites, may result also from natural death and do notnecessarily reflect human activity.

Our understanding of faunal preservation is stilllimited by the complexity of processes in time andspace. Patterns of fish preservation defy description bysimple rules based on readily observed environmentalgradients. Unique combinations of environmental andbiological conditions create unique taphonomic signa-tures. A comparative approach is required for furtherunderstanding of taphonomic processes and for paleoe-cological and anthropological reconstruction.

Acknowledgments

This study is based on dissertation research conductedby the first author at the Department of Zoology, Tel-AvivUniversity, the I. Meier Segals Garden for ZoologicalResearch, Israel, and the Natural History Museum,Brussels. The study was supported by the University ofHaifa, the Jacob Recanati fellowship from the Center ofMaritime Studies at the University of Haifa, the MariaRossiAscoli Fellowship, theMorrisM. Pulver Fellowship,and the Irene Levi Sala CAREArchaeological Foundation.

E. Geffen encouraged and supported this projectfrom initial to final phases; E. Boaretto performed theradiocarbon dating; D. Gifford-Gonzales providedwarm hospitality to the first author; S. Weiner providedaccess to labs and facilities housed at WeizmannInstitute; T. Goldman provided extremely helpful fieldassistance; G. Bar-Oz, S. Bar-David and the team ofOhalo-II project excavators helped during the field-work. We thank Wim Van-Neer and several anonymousreviewers for their helpful comments.

The bone assemblage project was partially funded bythe Ohalo II archaeological site project. Ohalo II wassupported by the Irene-Levi Sala CARE ArchaeologicalFoundation, The Israel Science Foundation (Grantno. 831/00), the Jerusalem Center for AnthropologicalStudies, the L.S.B. Leakey Foundation, the StekelisMuseum of Prehistory in Haifa, the MAFCAF Founda-tion, the National Geographic Society and the IsraelAntiquities Authority.

309I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Appendix A

NISP of identified bones (by alphabetic order), by taxonomic groups and lithofacies

Lithofacies A Cyprinids Tilapinii Clariassp.

Uniden. Total

Skeletal elements Mirogrex sp. Small Large

Angular/Articular 1 2 0 2 0 0 5Basapophysis 0 1 0 0 0 0 1Basioccipital 0 3 0 2 0 0 5Cranial bones 0 0 0 0 0 8 8Ceratohyal 0 7 0 0 0 0 7Cleithrum 0 1 0 0 0 2 3Coracoid 0 1 0 0 0 1 2Dentary 0 1 0 0 0 0 1Epihyal 0 1 0 0 0 0 1Fin ray 0 0 0 0 0 2 2Fin Spine 0 0 0 4 0 0 4First Dorsal Pterygiophore 0 2 0 1 0 0 3Frontal 0 2 0 0 0 0 2Hyomandibular 0 0 0 1 0 1 2Maxilla 0 2 0 0 0 0 2Neurocranium 0 0 0 0 0 1 1Opercle 0 4 0 2 0 0 6Opercle app. 0 0 0 0 0 1 1Pelvic Spine 0 0 0 1 0 0 1Pelvis 0 4 0 0 0 1 5Pharyngeal bone 2 0 0 1 0 0 3Pharyngeal teeth 0 6 2 0 0 0 8Preopercle 0 0 0 1 0 0 1Pterotic 0 1 0 0 0 0 1Pterygiophore 0 2 0 2 0 0 4Quadrate 1 0 0 1 0 1 3Rib 0 3 0 0 0 0 3Scapula 0 2 0 0 0 1 3Urohyal 1 0 0 0 0 0 1

Vertebral columnVertebrae general 0 0 0 0 0 14 14Atlas 3 0 0 0 0 0 3Atlas/Axis 1 0 0 0 0 0 1Axis 3 0 0 0 0 0 3Third vert. 1 0 0 0 0 0 1Fourth vert. 0 1 0 0 0 0 1Fifth vert. 0 1 0 0 0 0 1Ultimate vert. 0 1 0 1 0 0 2Thoracic vert. 0 15 0 3 0 0 18Precaudal vert. 0 0 0 2 0 0 2Precaudal/Caudal vert. 0 21 0 1 0 3 25Caudal vert. 0 3 0 0 0 0 3Penultimate vert. 0 2 0 0 0 0 2Tail vertebrae-elements 0 0 0 0 1 0 1Total 13 89 2 25 1 36 166

Lithofacies B Cyprinids Tilapinii Uniden. Total

Skeletal elements Mirogrex sp. Small Large

Angular/Articular 0 5 0 0 0 5Basapophysis 0 4 1 0 0 5Basioccipital 0 11 0 0 0 11Ceratohyal 0 12 0 0 0 12Cleithrum 0 13 0 1 5 19

310 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Coracoid 0 8 0 0 0 8Cranial bone 0 0 0 0 19 19Dentary 0 21 0 0 0 21Entopterygoid 0 1 0 0 0 1Epihyal 0 5 0 0 0 5Exoccipital 0 0 0 1 0 1Fin ray 0 0 0 0 16 16Fin Spine 0 0 0 9 0 9First Dorsal Pterygiophore 0 10 0 2 0 12Frontal 0 6 0 0 0 6Hyomandibular 0 16 0 1 1 18Interopercle 0 0 0 1 0 1Maxilla 0 6 0 0 0 6Mesethmoid 0 1 0 0 0 1Neurocranium 0 0 0 2 9 11Opercle 0 31 0 0 2 33Opercle app. 0 0 0 0 4 4Palatine 1 0 0 0 0 1Parasphenoid 0 0 0 0 1 1Pelvis 1 21 0 0 0 22Pharyngeal bone 15 0 0 5 0 20Molariform teeth 0 0 4 0 0 4Pharyngeal teeth 6 13 11 0 0 30Posttemporal 0 0 0 3 0 3Premaxilla 0 0 0 1 0 1Preopercle 0 5 0 0 3 8Pterotic 0 1 0 0 0 1Pterygiophore 0 8 0 2 0 10Quadrate 0 12 0 0 0 12Rib 0 7 0 4 0 11Scapula 0 12 0 0 0 12Spina neuralis 0 1 0 0 0 1Supracleithrum 0 0 0 2 0 2Supraoccipital 0 1 0 0 0 1Urohyal 1 0 0 0 0 1Vomer 1 0 0 0 0 1

Vertebral columnAtlas 14 0 0 1 0 15Atlas/Axis 0 2 0 0 0 2Axis 13 0 0 0 0 13Third vert. 12 0 0 0 0 12Fourth vert. 0 9 0 0 0 9Fifth vert. 0 10 0 0 0 10Thoracic vert. 0 93 0 10 0 103Precaudal vert. 0 1 0 1 5 7Precaudal/Caudal vert. 0 2 0 3 66 71Caudal vert. 0 4 0 0 0 4Penultimate vert. 0 2 0 0 0 2Ultimate vert. 0 3 0 0 0 3Vertebrae 0 1 0 0 112 113Os suspensorium 4th vert. 0 5 0 0 0 5Total 64 363 16 49 243 735

Lithofacies B Cyprinids Tilapinii Uniden.

Appendix A (continued )

Skeletal elements Mirogrex sp. Small Large

Total

(continued on next page)

311I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Lithofacies C Cyprinids Tilapinii Uniden. Total

Skeletal elements Mirogrex sp. Small Large

Angular/Articular 5 10 0 2 0 17Basioccipital 0 21 0 2 0 23Ceratohyal 0 27 0 0 0 27Cleithrum 0 30 1 1 14 46Coracoid 0 18 0 3 0 21Cranial bone 0 0 0 0 11 11Dentary 5 33 0 0 0 38Epihyal 0 17 0 0 0 17Epiotic 0 1 0 0 0 1Exoccipital 0 3 0 0 0 3Fin ray 0 2 0 0 14 16Fin Spine 0 0 0 18 0 181st Dorsal Pterygio. 0 10 0 0 0 10Frontal 0 9 0 0 0 9Hyomandibular 11 19 0 0 1 31Hypohyal 0 1 0 0 0 1Maxilla 10 2 0 1 2 15Neurocranium 0 0 0 1 4 5Opercle 2 31 0 3 5 41Opercle app. 0 9 0 0 3 12Palatine 4 2 0 0 0 6Parasphenoid 0 6 0 2 1 9Parietal 0 3 0 0 0 3Pelvis 0 32 0 2 3 37Pharyngeal bone 47 6 1 5 2 61Pharyngeal teeth 0 8 3 0 0 11Posttemporal 0 5 0 0 0 5Postcleithrum 0 1 0 0 0 1Preopercle 0 18 0 0 8 26Pterotic 0 10 0 0 0 10Pterygiophore 0 7 0 2 0 9Quadrate 5 7 0 1 1 14Rib 0 5 0 5 0 10Scapula 0 27 0 1 0 28Spina neuralis 0 2 0 0 0 2Subopercle 0 16 0 0 0 16Supracleithrum 0 0 0 1 0 1Supraoccipital 0 0 0 1 0 1Tripus 0 3 0 0 0 3Urohyal 1 2 0 0 0 3Ventral Postcleithrum 0 0 0 2 0 2Vomer 0 2 0 0 0 2

Vertebral columnAtlas 15 0 0 3 0 18Atlas/Axis 0 14 0 0 0 14Axis 28 0 0 2 0 30Third Thoracic vert. 22 0 0 0 0 22Fourth vert. 0 22 0 0 0 22Fifth vert. 4 5 0 0 0 9Thoracic vert. 0 213 0 9 0 222Precaudal vert. 0 0 0 3 0 3Precaudal/Caudal vert. 0 0 0 2 52 54Caudal vert. 0 20 0 3 0 23Penultimate vert. 0 3 0 1 0 4Ultimate vert. 0 6 0 1 0 7

Appendix A (continued )

312 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Vertebral columnVertebrae 0 2 0 6 309 317Tail vert.-Epural 0 0 0 0 1 1Tail vertebrae-elements 0 0 0 0 1 1Os suspensorium 4th vert. 0 7 0 0 0 7Total 159 697 5 83 432 1376

Lithofacies C Cyprinids Tilapinii Uniden. Total

Appendix A (continued )

Skeletal elements Mirogrex Small Large

References

Alin, S.R., Cohen, A.S., 2004. The live, the dead, and the very dead:taphonomic calibration of the recent record of paleoecologicalchanges in Lake Tanganyika, East Africa. Paleobiology 30 (1),44–81.

Arnold, J.E., 1992. Complex hunter-gatherer-fisher of prehistoricCalifornia: chiefs, specialists, and maritime adaptations of theChannel Islands. American Antiquity 57 (1), 60–84.

Badgley, C.E., 1982. Community Reconstruction of a SiwalikMammalian Assemblage. Unpublished Ph.D. dissertation Thesis,Yale University, New Haven, Connecticut.

Badgley, C.E., 1986. Taphonomy of mammalian fossil remains fromSiwalik rocks of Pakistan. Paleobiology 12 (2), 119–142.

Banarescu, P., 1990. Zoogeography of Fresh Waters, GeneralDistribution and Dispersal of Freshwater Animals. Fish andFisheries. AULA-Varlag GmbH, Wiesbaden, pp. 1–519.

Bar-Yosef, O., 1980. Prehistory of the Levant. Annual Review ofAnthropology 9, 101–133.

Bar-Yosef, O., 1987a. The Late Pleistocene adaptations in the Levant.In: Soffer, O. (Ed.), The Pleistocene Old World: RegionalPerspective. Plenum Press, New York, pp. 219–236.

Bar-Yosef, O., 1987b. Prehistory of the Jordan Rift. Israel Journal ofEarth Sciences 36, 107–119.

Bar-Yosef, O., Belfer-Cohen, A., 1989. The origins of sedentism andfarming communities in the Levant. Journal of World Prehistory 3 (4),447–498.

Bartov, Y., Stein, M., Enzel, Y., Agnon, A., Reches, Z., 2002. Lakelevels and sequence stratigraphy of Lake Lisan, the late Pleistoceneprecursor of the Dead Sea. Quaternary Research 57, 9–21.

Baxter, M.J., 2001. Methodological issues in the study of assemblagediversity. American Antiquity 66 (4), 715–725.

Behrensmeyer, A.K., 1982. Time resolution in fluvial vertebrateassemblages. Paleobiology 8, 211–228.

Behrensmeyer, A.K., 1983. Patterns of natural bone distribution onrecent land surfaces: implications for archaeological site formation.In: Clutton-Brock, J., Grigson, C. (Eds.), Animals and Archae-ology: 1. Hunters and their prey. British Archaeological ReportsInternational Series, Oxford, pp. 93–106.

Behrensmeyer, A.K., 1991. Terrestrial vertebrate accumulation. In:Allison, P.A., Briggs, D.E. (Eds.), Taphonomy: Releasing the DataLocked in the Fossil Record. Topics in Geobiology. Plenum Press,New York, pp. 291–337.

Behrensmeyer, A.K., Barry, J.C., 2005. Biostratigraphic surveys in theSiwaliks of Pakistan: A method for standardized surface samplingof the vertebrate fossil record. Palaeontologica Electronica 8 (1),15A, 24p.

Behrensmeyer, A.K., Gordon, K.D., Yanagi, G.T., 1989. Nonhumanbone modification inMiocene fossils from Pakistan. In: Bonnichsen,R., Sorg,M.H. (Eds.), BoneModification. Peopling of The Americas

Publications, Orono, Maine, pp. 99–120. Center for The Study ofThe First Americans, Institute of Quaternary Studies, University ofMaine.

Ben-Tuvia, A., 1978. Fishes. In: Serruya, C. (Ed.), MonographieBiologica: Lake Kinneret. Dr. W. Junk Publishers, Dordrecht, theNetherlands, pp. 407–430.

Berger, W.H., 1976. Biogenous Deep-Sea sediments: production,preservation and interpretation. In: Riley, J.P., Chester, R. (Eds.),Chemical Oceanography. Academic Press, London, p. 106.

Bonnichsen, R., Sorg, H.M. (Eds.), 1989. Bone Modification. Centerfor The Study of The First Americans, Institute for QuaternaryStudies, University of Maine, Orono, Maine, p. 535.

Borg, I. (Ed.), 1981. Multidimensional Data Representations: Whenand Why? Mathesis Press, Ann Arbor.

Borg, I., Groene, P. (Eds.), 1997. Modern Multidimensional Scaling.Theory and Applications. Springer, New York.

Boyer, B.W., 1982. Green River laminites: does the Playa-lake modelreally invalidate the stratified-lake model? Geology 10, 321–324.

Brett, C.E., Baird, G.C., 1986. Comparative taphonomy: a key topaleoenvironmental interpretation based on fossil preservation.Palaios 1, 207–227.

Brewer, D.J., 1991. Fishing in Prehistoric Egypt: Inferences fromfaunal remains. In: J.R. Purdue, W.E. Klippel and B.W. Styles(Editors), Beamers, Bobwhites, and Blue-Points. Illinois StateMuseum Scientific Papers, Vol. XXIII, and the University ofTennessee, Department of Anthropology Report of Investigationsno. 52, Springfield, pp. 333–340.

Bulinski, K.V., 2007. Analysis of sample-level properties along apaleoenvironmental gradient: the behavior of evenness as afunction of sample size. Palaeogeography, Palaeoclimatology,Palaeoecology 253, 490–508.

Busing, F.M.T.A., Commandeur, J.F., Heiser, W.J., 1996. PROXS-CAL: A Multidimensional Scaling Program for IndividualDifferences Scaling with Constraints. University of Leiden,Leiden, Netherlands.

Butler, V.L., 1987. Distinguishing natural from cultural Salmoniddeposits in the Pacific Northwest of North America. In: Nash, D.T.,Petraglia, M.D. (Eds.), Natural Formation Process and theArchaeological Record, vol. 352. B.A.R., Oxford, pp. 131–149.

Butler, V.L., 1990. Distinguishing natural from cultural Salmoniddeposits in the Pacific Northwest of North America. UnpublishedPh.D. Thesis, University ofWashington, Seattle, Washington, 218 pp.

Butler, V.L., 1993. Natural versus cultural Salmonid remains: origin ofthe Dalles Roadcut bones, Columbia River, Oregon, USA. Journalof Archaeological Science 20, 1–24.

Butler, V.L., 1996. Tui Chub taphonomy and the importance of marshresources in the western great basin of North America. AmericanAntiquity 61 (4), 699–717.

Butler, V.L., 2000. Resource depression on the Northwest coast ofNorth America. Antiquity 74, 649–661.

313I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Butler, V.L., 2001. Changing fish use on Mangaia, Southern CookIslands: resource depression and Prey Choice model. InternationalJournal of Osteoarchaeology 11, 88–100.

Cannon, A., 2000. Assessing variability in Northwest coast Salmon andHerring fisheries: Bucket–Auger sampling of shell midden sites onthe central coast of British Columbia. Journal of ArchaeologicalScience 27, 725–737.

Casteel, R.W., 1976. Fish Remains in Archaeology and Paleoenviron-mental Studies. Academic Press, London.

Cerón-Carrasco, R., 1994. The investigation of fish remains from anOrkney farm mound. In: Van Neer, W. (Ed.), Fish Exploitation inThe Past: Proceedings of the 7th meeting of the ICAZ FishRemains Working Group. Annales du Musee Royal de l'AfriqueCentrale, Sciences Zoologiques no. 274., Teruven, pp. 207–210.

Chen, P.F., 2000. Using fish taphonomy to reconstruct the environmentof ancient Shanwang Lake. Advances in Ecological Research 31,483–496.

Colley, S.M., 1984. Some methodological problems in the interprata-tion of fish remains from archaeological sites in Orkney. In: Desse-Berset, N. (Ed.), 2nd Fish Osteoarchaeology Meeting. Notes etMonographies Techniques. CNRS, pp. 117–131.

Colley, S.M., 1986. Site formation and archaeological fish remains. Anethnohistorical example from the northern Isles, Scotland. In:Brinkhuizen, D.C., Clason, A.T. (Eds.), Fish and Archaeology,vol. 294. BAR International Series, Oxford, England, pp. 34–41.

Colley, S.M., 1990. The analysis and interpretation of archaeologicalfish remains. In: Schiffer, M.B. (Ed.), Archaeological Method andTheory, vol. 2. University of Arizona Press, Tuscon, pp. 207–253.

Collins, M.J., Riley, M.S., Child, A.M., Turner-Walker, G., 1995. Abasic mathematical simulation of the chemical degradation ofancient collagen. Journal of Archaeological Science 22, 175–183.

Cutler, A.H., Behrensmeyer, A.K., Chapman, R.E., 1999. Environ-mental information in a recent bone assemblage: Roles oftaphonomic processes and ecological change. Palaeogeography,Palaeoclimatology, Palaeoecology 149, 359–372.

Durand, J.D., Tsigenopoulos, C.S., Unlu, E., Berrebi, P., 2002.Phylogeny and biogeography of the family Cyprinidae in theMiddle East inferred from cythochtome b DNA-Evolutionarysignificance of this region. Molecular Phylogenetics and Evolution22 (1), 91–100.

Elder, R.L., Smith, G.R., 1988. Fish taphonomy and environmentalinference in paleolimnology. Palaeogeography, Palaeoclimatology,Palaeoecology 62, 577–592.

Enghoff, I.B., 1994. Fishing from Medieval Holbæk/ Denmark, withnotes to reversed Platichthys flesus. Offa 51, 299–302.

Farlow, J.O., Argast, A., 2006. Preservation of fossil bone from the PipeCreek Sinkhole (Late Neogene, Grant County, Indiana, U.S.A.).Journal of the Paleontological Society of Korea 22 (1), 51–75.

Ferber, C.T., Wells, N.A., 1995. Paleolimnology and taphonomy ofsome fish deposits in Fossil and Uinta lakes of the Eocene GreenRiver Formation, Utah and Wyoming. Palaeogeography, Palaeo-climatology, Palaeoecology 117, 185–210.

Fraser, K., 1998. Fishing for tuna in Pacific prehistory. Paper Presentedto the 8th Congress of the International Council for Archaeozool-ogy, Victoria, B.C., Canada.

Gafny, S., 1992. The effect of substrate type on the structure andfunction of the littoral zone of Lake Kinneret. Unpublished Ph.D.Thesis, Tel-Aviv University, Tel-Aviv, 110 pp.

Gafny, S., Gasith, A., 2000. Spatial and temporal variation in thestanding biomass of emergent macrophytes: effect of water levelfluctuations. Archiv für Hydrobiologie— Advances in Limnology55, 301–316.

Gafny, S., Gasith, A., Goren, M., 1992. Effect of water level fluctuationon shore spawning of Mirogrex terraesanctae (Steinitz), (Cyprini-dae) in Lake Kinneret, Israel. Journal of Fish Biology 41, 863–871.

Galili, E., Lernau, O., Zohar, I., 2004. Fishing and coastal adaptationsat 'Atlit-Yam— A submerged PNC fishing village off the CarmelCoast, Israel. Atiqot 48, 1–34.

Gasith, A., Gafny, S., 1990. Effects of water level fluctuations on thestructure and function of the littoral zone. In: Tilzer, M.M.,Serruya, C. (Eds.), Large Lakes: Ecological Structure andFunction. Springer Series in Contemporary Bioscience. Springer,Berlin, pp. 156–171.

Gasith, A., Goren, M., Gafny, S., 1996. Ecological consequences oflowering Lake Kinneret water level: effect on breeding success ofthe Kinneret Sardine. Preservation of our World in the Wake ofChange. ISEEQS Pub., Jerusalem, pp. 569–573.

Gifford-Gonzales, D., Stewart, M.K., Rybczynski, N., 1999. Humanactivities and site formation at modern lake margin foraging campsin Kenya. Journal of Anthropological Archaeology 18, 397–440.

Gobalet, K.W., 2001. A critique of faunal analysis; Inconsistencyamong experts in blind tests. Journal of Archaeological Science 28,377–386.

Golani, D., Darom, D., 1997. Handbook of The Fishes of Israel. KeterPublishing House Ltd., Jerusalem. (in Hebrew).

Gophen, M., Gal, I., 1992. Lake Kinneret. Ministry of DefensePublishing House, Tel-Aviv, Israel. (in Hebrew).

Goren, M., Fishelson, L., Trewavas, E., 1973. The Cyprinid fishes ofAcanthobrama Heckel and related genera. Bulletin of The BritishMuseum (Natural History) Zoology 24 (6), 293–315.

Goren, M., 1983. Freshwater Fishes of Israel: Biology andTaxonomy. Hakibutz Hameuchad Publishing House Ltd., Tel Aviv.(in Hebrew).

Goren, M., Ortal, R., 1999. Biogeography, diversity and conservationof the inland water fish communities in Israel. BiologicalConservation 89 (1), 1–9.

Goren-Inbar, N., Feibel, C.S., Verosub, K.L., Melamed, Y., Kislev, M.E.,Tchernov, E., Saragusti, I., 2000. Pleistocene milestones on the out-of-Africa corridor at Gesher Benot Ya'aqov. Science 289 (5481),944–947.

Grayson, D.K., 1984. Quantitative Zooarchaeology: Topics in theAnalysis of Archaeological Faunas. Academic Press, New-York.

Grayson, D.K., 1991. Alpine faunas from the White Mountains,California: adaptive change in the late prehistoric great basin?Journal of Archaeological Science 18 (4), 483–506.

Hayek, L.C., Buzas, M.A., 1997. Surveying Natural Populations.Columbia University Press, New York. 563 pp.

Hambright, K.D., Gophen, M., Serruya, S., 1994. Influence of long-term climatic changes on the stratification of a subtropical, warmmonomictic lake. Limnology and Oceanography 39, 1233–1242.

Hazan, N., Stein,M., Agnon, A.,Marco, S., Nadel, D., Negendank, J.F.W.,Schwab, M.J., Neev, D., 2005. The late Quaternary limnologicalhistory of Lake Kinneret (Sea of Galilee), Israel. Quaternary Research63, 60–77.

Heck Jr., K.L., Van Belle, G., Simberloff, D., 1975. Explicitcalculation of the rarefaction diversity measurement and thedetermination of sufficient sample size. Ecology 56, 1459–1461.

Holland, S.M., 2001. Analytic Rarefaction. University of Georgia, Stra-tigraphic Lab. http://www.uga.edu/~strata/software/AnRareReadme.html.

Hurlbert, S.H., 1971. The nonconcept of species diversity: a critiqueand alternative parameters. Ecology 52, 577–586.

Kaufman, D., 1998. Measuring archaeological diversity: an applica-tion of the Jackknife technique. American Antiquity 63, 73–85.

314 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Kidwell, S.M., 1985. Paleobiological and sedimentological implica-tion of fossils concentrations. Nature 318, 457–460.

Kidwell, S.M., 1986. Models for fossil concentrations: paleobiologicimplications. Paleobiology 12, 6–24.

Kidwell, S.M., 2001. Preservation of species abundance in marinedeath assemblages. Science 294 (2 November), 1091–1093.

Kidwell, S.M., 2002. Time-averaged molluscan death assemblages:palimpsests of richness, snapshots of abundance. Geology 30 (9),803–806.

Kidwell, S.M., Bosence, D.W., 1991. Taphonomy and time averagingof marine shelly fauna. In: Allison, P.A., Briggs, D.E. (Eds.),Taphonomy: Releasing the Data Locked in the Fossil Record.Plenum, New York, pp. 116–209.

Kintigh, K.W., 1984. Measuring archaeological diversity by compar-ison with simulated assemblages. American Antiquity 49, 44–54.

Klein, R.G., Cruz-Uribe, K., 1984. The Analysis of Animal Bonesfrom Archaeological Sites. Prehistoric Archeology and EcologySeries. The University of Chicago Press, Chicago.

Koch, C.F., 1987. Prediction of sample size effects on the measuredtemporal and geographic distribution patterns of species. Paleo-biology 13, 100–107.

Koren, N., Klein, M., 2000. Rate of sedimentation in Lake Kinneret,Israel: spatial and temporal variations. Earth Surface Processes andLandforms 25, 895–904.

Krebs, C.J., 1999. Ecological Methodology. Harper Collins Publishers,University of British Columbia. 620 pp.

Krupp, F., 1987. Freshwater ichtyogeography of the Levant. In: Krupp,F., Schneider, W., Kinzelbach, R. (Eds.), Fauna and Zoogeographyof the Middle East. Beihefte Zum TAVO, Mainz, pp. 229–237.

Leach, F., Boocock, A., 1994. The impact of pre-European Maorifishermen on the New Zealand snapper, Pagrus auratus, in thevicinity of Rotokura, Tasman Bay. New Zealand Journal ofArchaeology 16, 69–84.

Lyman, R.L., 1984. Bone density and differential survivorship of fossilclasses. Journal of Anthropological Society 3, 259–299.

Lyman, R.L., 1994a. Quantitative units and terminology in zooarch-aeology. American Antiquity 59 (1), 36–71.

Lyman, R.L., 1994b. Vertebrate Taphonomy. Cambridge Manuals inArchaeology. Cambridge University Press, Cambridge.

Manly, B.F.J., 1991. Randomization and Monte Carlo Methods inBiology. Chapman and Hall, New York.

Martin, R.E., 1999. Taphonomy: A Process Approach. CambridgePaleobiology Series. Cambridge University Press, Cambridge.508 pp.

Nadel, D., 1993. Submerged archaeological sites on the shores of LakeKinneret. Atiqot XXII, 1–12.

Nadel, D., Tsatskin, A., Bar-Yosef Mayer, D.E., Belmaker, M.,Boaretto, E., Hershkovitz, I., Kislev, M.E., Rabinovich, R.,Simmons, T., Weiss, E., Zohar, I., Asfur, O., Emmer, G., Ghraieb,T., Grinberg, U., Halabi, H., Weissbrod, L., Zaidner, Y., 2002. TheOhalo II 1999–2000 Seasons of excavation: a preliminary report.Mitekufat Ha'even, Journal of the Israel Prehistoric Society 32,17–48.

Nagaoka, L., 2005. Declining foraging efficiency and moa carcassexploitation in southern New Zealand. Journal of ArchaeologicalScience 32, 1328–1338.

Nichol, R.K., Wild, C.J., 1984. Number of individuals in faunalanalysis: the decay of fish bone in archaeological sites. Journal ofArchaeological Science 11, 35–51.

Nicholson, R.A., 1996. Bone degradation, burial medium and speciesrepresentation: debunking the myths, an experiment-basedapproach. Journal of Archaeological Science 23, 513–533.

Nicholson, R.A., 1998. Bone degradation in a compost heap. Journalof Archaeological Science 25, 393–403.

O'connell, J.M., Tunnicliffe, V., 2001. The use of sedimentary fishremains for interpretation of long-term fish population fluctua-tions. Marine Geology 174, 177–195.

Ostrovsky, I., Walline, P., 1999. Growth and production of thedominant pelagic fish, Acanthobrama terraesanctae, in subtropicalLake Kinnerert, Israel. Journal of Fish Biology 54, 18–32.

Owen, J.F., Merrick, J.R., 1994. Analysis of coastal middens in South-Eastern Australia: sizing of fish remains in Holocene deposits.Journal of Archaeological Science 21, 3–10.

Plog, S., Hegmon, M., 1993. The sample size-richness relation: Therelevance of research questions, sampling strategies, and beha-vioral variation. American Antiquity 58 (3), 489–496.

Reading, H.G. (Ed.), 1996. Sedimentary Environments and Facies.Blackwell Scientific Publications, New York, p. 688.

Reiche, I., Favre-Quattropani, L., Vignaud, C., Bocherens, H., Charlet,L., Menu, M., 2003. A multi-analytical study of bone diagenesis:the Neolithic site of Bercy (Paris, France). Measurement Scienceand technology 14, 1608–1619.

Reitz, E.J., Zierden, M., 1991. Cattle bones and status fromCharleston, South Carolina. In: Purdue, J.R., Klippel, W.E.,Styles, B.W. (Eds.), Bemers, Bobwhites, and Blue-points: Tributesto the Career of Paul W. Parmalee. Illinois State Museum ScientificPapers, Springfield, pp. 395–407.

Reitz, E.J., Wing, E.S., 1999. Zooarchaeology. Cambridge UniversityPress, Cambridge. 455 pp.

Ricardo-Bertram, C.K., 1947. Third report on the fish and fisheries ofLake Tiberias. Department of Agriculture and Fisheries, Govern-ment of Palestine, Jerusalem.

Rick, T.C., Erlandson, J.M., 2000. Early Holocene fishing strategies onthe California coast: Evidence from CA-SBA-2057. Journal ofArchaeological Science 27, 621–633.

Sarig, S., 1981. The Israeli Fisheries in 1980 (in Hebrew). Fisheriesand Fish breeding in Israel 3 (16), 3–16.

Sarig, S., 1982. The Israeli Fisheries in 1981(in Hebrew). Fisheries andFish breeding in Israel 16 (4), 11–22.

Sarig, S., 1991. The Israeli Fisheries in 1986–1989 (in Hebrew).Fisheries and Fish breeding in Israel 3 (24), 123–131.

Sarig, S., 1993. The Israeli Fisheries in 1990–1991 (in Hebrew).Fisheries and Fish breeding in Israel 26 (1), 13–18.

Schäfer, W., 1972. Ecology and Paleoecology of Marine Environ-ments. Oliver and Boyd, Edinburgh.

Schick, K.D., Toth, N., Daeschler, E., 1989. An early paleontologicalassemblage as an archaeological test case. In: Bonnichsen, R.,Sorg, M.H. (Eds.), Bone Modification. Peopling of the AmericasPublications. Center for the study of the first Americans,University of Maine, Orono, Maine, pp. 121–137.

Soutar, A., 1966. The accumulation of fish debris in certain Californiacoastal sediments. California Cooperative Oceanic FisheriesInvestigations Reports 11, 136–139.

Soutar, A., Isaacs, J.D., 1974. Abundance of pelagic fish during the19th and 20th centuries as recorded in anaerobic sediment off theCalifornia. Fishery Bulletin 72, 257–273.

Stewart, K.M., 1989. Fishing Sites of North and East Africa in the LatePleistocene and Holocene, vol. 521. B.A.R., Oxford. 521.

Stewart, K.M., 1991. Modern fishbone assemblages at Lake Turkana,Kenya: a methodology to aid in recognition of Hominid fishutilization. Journal of Archaeological Science 18, 579–603.

Stiller,M., Ehrlich, A., Pollinger, U., Baruch, U., Kaufman, A., 1983–84.The late Holocene sediments of Lake Kinneret (Israel) — multi-disciplinary study of a fivemeter core. GSI, Current Research 83–88.

315I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316

Author's personal copy

Tappen, M., 1995. Savabba ecology and natural bone deposition:implications for early hominid site formation, hunting, andscavenging. Current Anthropology 36 (2), 223–260.

Tomasovych, A., 2006. Linking taphonomy to community-levelabundance: insights into compositional fidelity of the UpperTriassic shell concentrations (Eastern Alps). Palaeogeography,Palaeoclimatology, Palaeoecology 235, 355–381.

Trueman, C.N., Martill, D.M., 2002. The long-term survival of bone:the role of bioerosion. Archaeometry 44 (3), 371–382.

Trueman, C.N., Benton, M.J., Palmer, M.R., 2003. Geochemicaltaphonomy of shallow marine vertebrate assemblages. Palaeogeo-graphy, Palaeoclimatology, Palaeoecology 197, 151–169.

Tunnicliffe, V., O'connell, J.M., McQuoid, M.R., 2001. A Holocenerecord of marine fish remains from the Northeastern Pacific.Marine Geology 174, 197–210.

Van Neer, W., 1986. Some notes on fish remains fromWadi Kubbaniya(Upper Egypt; Late Pleistocene). In: Brinkhuizen, D., Clason, A.(Eds.), Fish and Archaeology. BAR International Series, Cam-bridge, pp. 103–113.

Van Neer, W., 1989. Fishing along the prehistoric Nile. In: Kryzaniak,L., Kobuseiwiciz, M. (Eds.), Late Prehistory of the Nile Basin andthe Sahara. Ponzan Archaeology Museum, Ponzan, pp. 49–56.

Van Neer, W., 1993. Fish remains from the last Interglacial at BirTarfawi (Eastern Sahara, Egypt). In: Wendorf, F., Schild, R., Close,A.E. (Eds.), Egypt During the Last Interglacial. Plenum Press,New-York, pp. 144–155.

Van Neer, W., 1994. Cenozoic fish fossils from the Albertine RiftValley in Uganda. Geology and Paleobiology of the Alertine RiftValley, Uganda–Zaire. CIFEG Occas. Publ., Orléans, pp. 89–127.

Van Neer, W., 2004. Evolution of prehistoric fishing in the Nile Valley.Journal of African Archaeology 2, 251–269.

Voskoboynik, A., 1995. Use of littoral resources by two CyprinidsHemigrammocapoeta nana and Garra rufa in Lake Kinneret,Israel. Unpublished MSc Thesis, Tel Aviv University, Tel Aviv,82 pp.

Weigelt, J., 1989. Recent Vertebrae Carcasses and Their Paleobiolo-gical Implications. University of Chicago Press, Chicago. 188 pp.

Wheeler, A., 1978. Problems of identification and interpretation ofarchaeological fish remains. In: Brothwell, D.R., Thomas, K.D.,

Clutton-Brock, J. (Eds.), Research Problems in Zooarchaeology.Occasional Papers of the Institute of Archaeology, London,pp. 69–75.

Wheeler, A., Jones, A.K.G., 1989. Fishes. Cambridge Manuals inArchaeology. Cambridge University Press, Cambridge.

Wilson, M.V.H., 1988. Reconstruction of ancient lake environmentusing both autochthonous and allochthonous fossils. Palaeogeo-graphy, Palaeoclimatology, Palaeoecology 62, 609–623.

Wilson, M.V.H., Barton, D.G., 1996. Seven centuries of taphonomicvariation in Eocene freshwater fishes preserved in varves:paleoenvironments and temporal averaging. Paleobiology 22 (4),535–542.

Wing, E.S., 1994. Patterns of prehistoric fishing in the West Indies.Archaeofauna 3, 99–107.

Wolverton, S., 2002. NISP:MNE and % whole in analysis ofprehistoric carcass exploitation. North American Archaeologist23 (2), 85–100.

Wright, C.A.,Dallimore,A., Thomson, R.E., Patterson, R.T.,Ware,D.M.,2005. LateHolocene paleofish populations in Effingham Inlte, BritishColumbia, Canada. Palaeogeography, Palaeoclimatology, Palaeoe-cology 224, 367–384.

Zohar, I., 2003. Fish Exploitation at the Sea of Galilee (Israel) by EarlyFisher-Hunter-Gatherers (23,000 B.P.): Ecological, Economical,and Cultural Implications. Unpublished Ph.D. Thesis, Tel AvivUniversity, Tel Aviv, 222 pp.

Zohar, I., Belmaker, M., 2005. Size does matter: methodologicalcomments on sieve size and species richness in fishbone assem-blages. Journal of Archaeological Science 32, 635–641.

Zohar, I., Dayan, T., Spanier, E., Galili, E., Lernau, O., 1994.Exploitation of gray triggerfish (Balistes carolinensis) by theprehistoric inhabitants of Atlit-Yam, Israel: a preliminary report.In: Van Neer, W. (Ed.), Fish Exploitation in The Past: Proceedingsof the 7th meeting of the ICAZ Fish Remains Working Group,vol. 274. Annales du Musee Royal de l'Afrique Centrale,Sciences Zoologiques, Teruven, pp. 231–237.

Zohar, I., Dayan, T., Galili, E., Spanier, E., 2001. Fish processingduring the early Holocene: a taphonomic case study from coastalIsrael. Journal of Archaeological Science 28, 1041–1053.

316 I. Zohar et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 258 (2008) 292–316