Palaeogeography, Palaeoclimatology, Palaeoecology
The living and the dead: How do taphonomic processes modify relative abundance and skeletal completeness of freshwater fish?
Introduction
Unraveling the mystery of ancient aquatic life and death in changing environments is a huge challenge for palaeoecology. While considerable information is lost due to bone destruction (Farlow and Argast, 2006), skeletal elements that do survive are incorporated into the sediment, where they become the record of past fish communities. This record is significant for paleoecological (i.e., taxonomic composition and biodiversity; Nicholson, 1998, Trueman et al., 2003) and paleoenvironmental reconstruction (e.g. salinity, temperature, water level; Casteel, 1976, Brett and Baird, 1986, Wheeler and Jones, 1989, Voskoboynik, 1995, Cutler et al., 1999, Chen, 2000). Recently, a growing body of taphonomic research demonstrated the numerous depositional processes that affect fish bone preservation (Tomasovych, 2006, Trueman and Martill, 2002, Farlow and Argast, 2006). It has been realized that biases in skeletal element preservation alter species richness and abundance (Elder and Smith, 1988, Ferber and Wells, 1995, Cutler et al., 1999, Manly, 1991, Nicholson, 1998). Clearly, the ability to estimate what and how much of original information was lost is crucial for paleoecological and paleoenvironmental reconstruction. Therefore, field tests that compare living communities to associated bone assemblages are the primary means of estimating the biological information loss in the palaeontological and archaeological record (Behrensmeyer, 1982, Behrensmeyer, 1983, Kidwell, 1985, Kidwell, 2002, Kidwell, 1986, Kidwell, 2001).
Direct live-dead comparisons have demonstrated that a wide variety of factors play a role in the processes of fish carcass decay, disintegration, articulation, and preservation in the aquatic sediments, and that a complete fish skeleton will be preserved only under special circumstances (Sarig, 1993, Soutar, 1966, Elder and Smith, 1988, Nicholson, 1998, Tomasovych, 2006, Trueman and Martill, 2002, Farlow and Argast, 2006). These include: water temperature, water depth, pressure, salinity level, oxygen level, oxygen stability, type of sediments, rate of sedimentation, carcass buoyancy, scavenger activity, wave action, and sea bottom currents (Sarig, 1993, Elder and Smith, 1988, Cutler et al., 1999, Reading, 1996). Of these, water temperature has been regarded as the most significant factor in determining the fate of carcasses (Sarig, 1993, Elder and Smith, 1988, Zohar et al., 2001). In all, it has been shown that fish diversity and taphonomic processes are habitat dependent, and we still need to seek for generalities in the taphonomic processes involved. In order to do that, we must examine the differential loss between the live-dead assemblages bearing in mind that in most cases a bone assemblage is not necessarily biologically equivalent to a living community (Kidwell, 2002, Kidwell, 1986, Kidwell, 2001, Tappen, 1995).
Although live-dead comparisons are fairly common in the taphonomic literature on marine shell assemblages (e.g., Kidwell, 1986, Kidwell, 2001, Alin and Cohen, 2004, Tappen, 1995) and terrestrial mammals (e.g., Behrensmeyer, 1982, Behrensmeyer, 1983, Behrensmeyer et al., 1989, Behrensmeyer, 1991, Cutler et al., 1999, Behrensmeyer and Barry, 2005), they are sparse for fish remains (Soutar, 1966, Soutar and Isaacs, 1974, Butler, 1993, Nicholson, 1998). Recently, a growing literature is also concerned with the paleoanthropological reconstruction of ancient fish exploitation patterns in coastal communities (Brewer, 1991, Arnold, 1992, Cerón-Carrasco, 1994, Enghoff, 1994, Krupp, 1987, Wilson and Barton, 1996, Zohar and Belmaker, 2005, Fraser, 1998, Butler, 2000, Cannon, 2000, Reitz and Wing, 1999, Butler, 2001, Galili et al., 2004). Some of the archaeological sites have become submerged due to local or global water level changes (Martin, 1999, Zohar et al., 1994, Galili et al., 2004). Therefore, while studying fish remains from lacustrine sediments, one must ask whether the bone assemblage reflects human fishing activity or whether it is merely natural death assemblage? Thus, the study of fish remains from waterlogged and submerged sites must be based on sound taphonomic analyses of natural fish accumulations that may enable researchers to tease apart the remnants of economic practices from the imprint of natural deaths (Butler, 1987, Soutar and Isaacs, 1974, Gifford-Gonzales et al., 1999).
In the southern Levant no research, to our knowledge, has yet compared the fish species found alive in aquatic habitat to those found as bones in an adjacent shoreline, although this region is rich in important archaeological sites (e.g., Bar-Yosef, 1980, 1987aBar-Yosef, 1987b, Bar-Yosef and Belfer-Cohen, 1989, Goren-Inbar et al., 2000). In order to gain insight into taphonomic biases that affect both paleoecological and paleoanthropological reconstruction, we performed a pioneer study of fish remains natural accumulations in lacustrine sediments along the southern shore of Lake Kinneret (Sea of Galilee; Fig. 1). Three lithofacies were identified, spanning the past 1500 years. This study was designed to determine information loss (taxonomic diversity, species composition and skeleton completeness) relative to the living community and to determine biases expected in environmental interpretations from fish remains deposited in a sandy littoral southern shore of Lake Kinneret. The focus of this study was to compare one modern shoreline habitat and the facies that occur within a small stratigraphic distance beneath it with nearby living communities from different habitats. This was used as a test case. Further research in other lacustrine habitat is required to test the generality of our results and conclusions.
Section snippets
Lake Kinneret
Lake Kinneret (Sea of Galilee), located in the northern part of the rift valley, is the largest fresh-water lake in Israel. During the Pleistocene (70–15 ka) it was part of Lake Lisan, which extended from the Arava Valley in the south to the present Kinneret basin in the north, when it reached a maximum level of 170 m below sea level at 26 ka (Bartov et al., 2002, Hazan et al., 2005). The dramatic changes in the lake's size, salinity, temperature, sedimentation, depth, as well as other factors,
The bone assemblages
We examined fish bone assemblages from a sandy beach 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 took place during July 2001, when water levels dropped to 214 m BSL, and archaeological sites were exposed along the shore (Nadel, 1993). Hence, in order to avoid excavation in an area previously inhabited, the area selected for this study was carefully chosen to be devoid of
The bone assemblage
The faunal remains recovered from the 24 excavated squares include a large number of freshwater mollusks, two rodent bones, 5037 fish bones and 758 scales. Here we focus on the taphonomic processes affecting fish remains.
Spatial distribution of fish remains
The average number of fish remains (standardized BSF) per square was 241 (SD 580), ranging from 0 to 2025.6 bones when standardized per sediment volume in the lithofacies (Table 4, Table 5). The spatial distribution exhibited a clear clumped pattern (Morisita Index, I p > 0,
Discussion
Many studies exhibited the significance of fish remains preserved in the sediments as proxies for paleo-population dynamics (Schick et al., 1989, Soutar, 1966, Nicholson, 1998, Wolverton, 2002). They also demonstrated that the fidelity of fossil assemblages and patterns of information loss should be examined for each aquatic habitat (Kidwell, 2002, Butler, 1996, Kidwell, 1986, Nicholson, 1998, Trueman et al., 2003, Kidwell, 2001). In this study several biases were observed for fish live-dead
Conclusions
Live-dead assemblages of fish in the southern shore of Lake Kinneret present a unique case study to test differences in species richness, abundance, and composition. The living assemblages represent fish communities that differ between habitat/substrate type and that vary through population and seasonal dynamics. The bone assemblage differs from the habitat specific community in species composition and diversity. Although discrepancies in taxonomic composition of living and bone assemblages
Acknowledgments
This study is based on dissertation research conducted by the first author at the Department of Zoology, Tel-Aviv University, the I. Meier Segals Garden for Zoological Research, Israel, and the Natural History Museum, Brussels. The study was supported by the University of Haifa, the Jacob Recanati fellowship from the Center of Maritime Studies at the University of Haifa, the Maria Rossi Ascoli Fellowship, the Morris M. Pulver Fellowship, and the Irene Levi Sala CARE Archaeological Foundation.
E.
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