66 Chapter 6 RESULTS This chapter presents results of the quantitative analyses applied to the faunal samples selected for study. First, the taxonomic composition of each component is presented and statistical measures used in the subsequent analyses are explained. Second, samples are divided between habitats and analyzed in terms of site catchment. Both general subsistence patterns and taxonomic diversity are examined as part of the site catchment analysis. Finally, the resulting subsistence economies for each habitat are discussed in terms of foraging efficiency tempered by paleoenvironmental and technological changes. The total number of identified vertebrate specimens examined in this study is 15,319 (Tables 6 through 8). Freshwater mussel weights are presented in Table 9. The Middle Period assemblage is comprised of 6,219 specimens, while the Late Period sample contains 9,100 specimens. Table 2. Diachronic Taxonomic Richness. Taxa Middle Period Late Period Difference Mammals 16 21 31% increase Birds 6 8 33% increase Fish 11 11 No Change Herpetofauna 3 7 133% increase Total 36 47 31% increase Richness was calculated for each site using the lowest taxonomic level of each taxon present. For example, if no specimens could be identified to species, such as Canis latrans, then the sum of the specimens identified to genus level, Canis, was used; if none could be identified to the genus level, then the sum of the specimens at the level of family, Canidae, was used, etc. As a result, 36 taxa were identified in the Middle Period and 47 taxa were identified in the Late Period, indicating a total increase of 31 percent in taxonomic richness over time. Common Name Artiodactyls Tule Elk Black-tailed Deer Pronghorn Antelope Carnivores American Black Bear Canids Raccoon River Otter Weasels and Minks Mink Spotted Skunk Striped Skunk Lagomorphs Black-tailed Jack Rabbit Cottontails Squirrels California Ground Squirrel Grey Squirrel Wood Rats Botta's Pocket Gopher Meadow Voles California Meadow Vole Deer Mice Kangaroo Rats Heermann's Kangaroo Rat North American Beaver Rodents Badger Mammal Richness Table 3. Mammals NISP. Scientific Name SAC-67 SAC-133 Artiodactyla 34 42 Cervus elaphus nannodes 1 Odocoileus hemionus columbianus 7 Antilocapra americana 3 Carnivora 7 Ursus americanus Canis sp. 9 Procyon lotor 2 Lutra canadensis Mustela Mustela vison 1 Spilogale putorius Mephitis mephitis Lagomorpha Lepus californicus 7 45 Sylvilagus sp. 2 25 Sciuridae 12 Spermophilus beecheyi 10 17 Sciurus griseus 2 Neotoma sp. Thomomys bottae 14 131 Microtus sp. Microtus californicus 4 Peromyscus sp. Dipodomys sp. 1 Dipodomys heermanni 1 Castor canadensis Rodentia 26 Taxidea taxus Totals 112 291 8 10 SAC-42 167 61 26 26 32 18 10 1 2 2 1 15 14 1 3 379 SAC-329 126 106 1 3 19 149 1 3 1 4 2 10 3 3 6 574 8 85 1104 SAC-267 36 29 2 1 2 1 1 1 2 7 50 39 32 25 7 15 3 4 204 461 SAC-29 207 201 41 22 1 24 42 1 1 3 4 40 3 6 596 Total 486 389 209 54 43 1 70 203 3 3 3 2 7 2 66 93 51 31 5 32 225 13 592 23 1 4 93 230 9 2943 13 16 15 12 21 67 67 8 1 Table 4. Birds NISP. Scientific Name SAC-67 SAC-133 Anseriformes 2 8 Cygnus columbianus Anserinae 4 Branta canadensis Anatinae/Aythyinae/Dendrocygninae/ 24 21 Merginae/Oxyurinae Mallard Anas platyrhynchos 9 Northern Pintail Anas acuta Gadwall Anas strepera Ruddy Duck Oxyura jamaicensis American Coot Fulica americana 1 Pie-billed Grebe Podilymbus podiceps Sandhill Crane Grus canadensis American Crow Crovus brachyrhynchos 1 California Quail Callipepla californica 1 Other (doves, perching birds, hawks, grouse, shorebirds) 2 Totals 39 34 Common Name Waterfowl Whistling Swan Geese Canada Goose Ducks Bird Richness 2 4 SAC-42 26 101 SAC-329 17 2 16 18 SAC-267 1 1 13 38 SAC-29 94 223 Total 27 1 127 29 425 45 19 1 2 6 200 4 15 16 8 96 2 4 8 67 76 4 35 432 11 4 15 16 122 19 8 2 3 59 868 6 4 4 4 11 Table 5. Herpetofauna NISP. Common Name Reptiles Western Pond Turtle Snakes Scientific Name Squamata Actinemys marmorata Serpentes 2 48 27 4 11 - 29 - 7 - 3 20 7 - 9 115 34 Garter Snake Thamnophis sirtalis 2 - - 1 - - 3 Gopher Snake Pituophis melanoleucus - - - 1 - - 1 Pit Vipers Crotalinae - - - - 1 - 1 Frogs North American Bullfrog Western Toad Salamander Anura Rana catesbeiana Bufo boreas Caudata 1 80 15 29 1 10 1 10 42 0 1 1 1 10 176 3 1 1 4 4 0 7 Totals Herpetofauna Richness 68 8 1 Common Name Sturgeon Salmon Chinook Salmon Minnows & Suckers Sacramento Sucker Minnows Hardhead Sacramento Blackfish Hitch Sacramento Pikeminnow Sacramento Splittail Thicktail Chub Sacramento Perch Tule Perch Scientific Name Acipenser sp. Salmonidae Oncorhynchus tshawytscha Cyprinidae/Catostomidae Catostomus occidentalis Cyprinidae Mylopharodon conocephalus Orthodon microlepidotus Lavinia exilicauda Ptychocheilus grandis Pogonichthys macrolepidotus Gila crassicauda Archoplites interruptus Hysterocarpus trask ii Table 6. Fish NISP. SAC-67 SAC-133 4 2 2 8 43 349 40 216 418 66 2 16 2 13 9 22 7 1 11 299 24 8 Totals Fish Richness SAC-42 20 19 286 1489 3 41 40 15 10 145 1390 20 SAC-329 13 4 103 57 50 100 79 50 221 865 78 SAC-267 8 34 1513 247 130 14 40 16 8 19 81 10 SAC-29 160 427 1444 6 154 5 62 22 29 6 128 109 - Total 207 427 67 3349 898 2257 97 195 191 162 74 524 2768 116 856 706 3478 1620 2120 2552 11,332 10 8 11 10 11 10 11 Table 7. Invertebrates (weights in grams). Common Name Scientific Name Freshwater Mussel Unionoidea - - - 4723.5 1634 163 6520.5 Western Pearlshell Mussel Margaritifera falcata Gonidea angulata - 132.5 - - - 30 162.5 - - 151.3 - - 320 471.3 Anodonta nuttalliana - - 0.1 - - 262 262.1 - 132.5 151.4 4723.5 1634 775 7416.4 Rocky Mtn Ridged Mussel Nuttall's Anadon Totals 69 8 1 70 However, taxonomic richness (i.e., the number of taxa present in an assemblage) is frequently a function of sample size (Lyman 2008:180). Sample size can also affect taxonomic evenness, or how well identified specimens are distributed among the assemblage. Given differences in sample sizes between the sites analyzed, the Simpson’s Diversity Index was used to assess whether sample size influenced taxonomic diversity. Simpson’s Diversity Index evaluates both the richness and evenness of a sample. As species richness and evenness increase, so does diversity. The formula used to calculate Simpson’s D for a finite population is: ∑ ni (ni -1) D= N(N-1) ni = the total number of specimens of a particular species (NISPi) N = the total number of specimens of all species (NISP total) D= Simpson’s Index of dominance (adapted from Lyman 2008) When the reciprocal of D (1/D) is computed, the larger the value, the more evenly specimens are distributed. Conversely, the lower the value of 1/D, the more an assemblage is dominated by one or a few taxa (Lyman 2008:197). The reciprocal of Simpson’s Diversity Index was applied to each of the site samples. The results indicate an increase in diversity in all habitats over time. Table 8. Diachronic Taxonomic Diversity. Habitat Richness NISP Site Time 1/D SAC-67 Middle Period Freshwater Marsh 23 499 2.65 SAC-329 Late Period Freshwater Marsh 34 2205 5.43 SAC-133 Middle Period Grassland 25 570 4.75 SAC-267 Late Period Grassland 38 762 7.39 SAC-42 Middle Period Riparian Woodland 31 2374 2.75 SAC-29 Late Period Riparian Woodland 26 1739 8.62 71 When a scatter plot is created comparing diversity with NISP (Figure 14), the best fit line shows there is no relationship between NISP and diversity, indicating that the 31 percent increase in diversity/richness seen between Middle Period and Late Period assemblages is not being driven by sample size. 10.00 CA-SAC-29 8.62 9.00 CA-SAC-267 7.39 8.00 7.00 CA-SAC-329 5.43 6.00 1/D R² = 0.0006 5.00 CA-SAC-133 4.75 4.00 CA-SAC-42 2.75 3.00 2.00 CA-SAC-67 2.65 1.00 0.00 20 520 1020 1520 2020 2520 NISP Figure 14. Scatter Plot of Diachronic Taxonomic Diversity (1/D) and NISP. One of the problems in using NISP for the analysis of prehistoric diets is that it can differentially exaggerate taxonomic abundance. For example, NISP does not control for the interdependence of specimens which may be from one or many individuals. In an attempt to correct for the potential overrepresentation of certain taxa, a variation of the chi-square test of independence, Cochran’s test of linear trend, was applied to samples. Chi-square is calculated as follows: 𝛘2 = ∑ (observed – expected)2 expected A percentage of significance or reliability is then applied to the result based on the number of data groups (degrees of freedom) used in the calculation. NISP totals were entered into a chi-square calculator macro for Excel created by M. Cannon (Cannon 2012) and adjusted residuals were rounded to the hundredth. The results of the chi-square analyses 72 were compared by habitat type, because the restricted mobility assumed for both Middle and Late Period populations would have limited foraging to site specific catchment areas. Freshwater Marsh The sites representing the freshwater marsh habitat are SAC-67 (Middle Period) and SAC-329 (Late Period). First, catchments areas for each site are described. Next, subsistence patterns and chi-square analysis are compared. Last, results are interpreted in terms of site catchments and foraging efficiency. Site catchments were reconstructed for SAC-67 and SAC-329 based on the paleoenvironmental data used to create Figure 2 (see Chapter 2). The site catchment of SAC-67 indicates 59 percent of the habitat within five kilometers (three miles) is grassland, 28 percent is freshwater marsh, and 13 percent is riparian woodland. The catchment area of SAC-329 is 86 percent freshwater marsh and 14 percent riparian woodland habitat. No grassland habitat was located within five kilometers (three miles) of SAC-329. The SAC-67 faunal assemblage has an NISP of 1,087. Faunal diversity (richness) includes eight mammals, two birds, ten fish, and three herpetofauna taxa. The faunal assemblage from SAC-329 has an NISP of 2,690 and sample diversity includes 15 mammals, four birds, 11 fish, and four herpetofauna. Over time, diversity in the freshwater marsh habitat increased by 48 percent. Subsistence patterns were developed for each site using an arbitrary value system to measure taxon attributes important to prey choice, e.g., taxa weight, habitat preference, and procurement method. For fish, weight was replaced by standard length (length x girth). Taxon selection used the same criteria as that used to calculate richness, i.e., only specimens from the lowest level of taxonomic rank were included for analysis. A key showing values assigned to each taxon attribute and site specific calculation charts are presented in Appendix C. Freshwater marsh populations appear to have relied on medium sized animals primarily procured by mass harvesting techniques from the freshwater marsh habitat (𝜒2=156.38, df=28, p <0.001). Large slow-water fish, such as Sacramento perch, accounted for over 50 percent of the taxa targeted at both sites, suggesting such fishes remained a dietary staple for freshwater marsh populations over time. In spite of this consistency, changes in targeted taxa are apparent. First, the number of larger taxa incorporated in the diet 73 Figure 15. Freshwater Marsh Individual Site Catchments. 74 Figure 16. SAC-67 Middle Period Freshwater Marsh Subsistence Pattern. Figure 17. SAC-329 Late Period Freshwater Marsh Subsistence Pattern. 75 increases in the Late Period. There is also a slight increase in the number of riparian woodland taxa included in the diet. Finally, there is an increase in the number of taxa procured by single takes. The first and third differences will be considered in the chi-square analysis presented below. The second difference will be discussed in terms of site catchment area at the end of the freshwater marsh analysis. In spite of this consistency, changes in targeted taxa are apparent. First, the number of larger taxa incorporated in the diet increases in the Late Period. There is also a slight increase in the number of riparian woodland taxa included in the diet. Finally, there is an increase in the number of taxa procured by single takes. The first and third differences will be considered in the chi-square analysis presented below. The second difference will be discussed in terms of site catchment area at the end of the freshwater marsh analysis. Chi-square analyses of the terrestrial taxa identified in freshwater marsh assemblages indicate there is a significant increase in the amount of artiodactyl, carnivore (e.g., canids, raccoon, river otter, mink, and skunks), and beaver remains in the Late Period (𝜒2=337.40, df =6, p <0.001). Significant decreases are seen in lagomorphs, waterfowl, medium rodents (squirrels and gophers), and turtle. Figure 18. Diachronic Freshwater Marsh Terrestrial Taxa. 76 Thus, the chi-square results for terrestrial taxa indicate that not only were more taxa incorporated in the Late Period diet, but that more larger bodied taxa (e.g., artiodactyls, carnivores, and beavers) were being procured at the expense of previously exploited smaller bodied animals (e.g., waterfowl, medium rodents, and turtle). Further, as indicated in the subsistence pattern comparison, most of the animals which increase in frequency are best procured via single takes (see Ugan 2005:77). Figure 20 shows that fish NISP also increases over time, but it is not equally distributed across all taxa (𝜒2=106.46, df=10, p <0.001). Due to the significant increases in other species, Sacramento perch shows a decrease in abundance over time despite a nearly 200 percent increase in NISP. Similarly, Sacramento sucker shows a decrease in abundance, despite a 157 percent increase in NISP. All other resident fish show statistically significant increases, with the exception of Sacramento splittail, salmon, and sturgeon. The largest increase is in thicktail chub, which is second in magnitude Figure 19. Diachronic Freshwater Marsh Fish. only to the decrease in the frequency of Sacramento perch. Thus, although Sacramento perch remains an important constituent of the diet, a wider variety of fish was utilized by freshwater marsh occupants during the Late Period. 77 One explanation for the increased variety of Late Period fish is that the SAC-329 catchment contains approximately three times the amount of freshwater marsh habitat as SAC-67. This increased the habitat for slow-water, i.e., freshwater marsh, species essentially, three-fold. On the one hand, mass harvesting techniques likely procured a greater variety of fish simply because they were present. Support for this conclusion can be found in the limited difference of NISP (no more than 50 specimens) between the species intensified in presence during the Late Period. Such minor differences suggest that these species were not specifically, but generically targeted. On the other hand, it may be that the decrease in large bodied, slow-water fish indicates that foraging efficiency was achieved in other ways. The significant increase in larger bodied terrestrial taxa in the Late Period supports this conclusion as does the increase in single-take procurement. Many of the larger bodied terrestrial species added to the diet were best procured individually and found primarily in the riparian woodlands. These include carnivores and beaver, with deficiencies in the analysis of artiodactyl remains from SAC-67 and SAC-329 making it difficult to assess which, if any, species significantly increased over time. Scarcely any difference (one percent) in riparian woodland exists between the catchments of the Middle and Late Period sites, so increased use of this habitat in the Late Period supports a shift in prey choice. Subsistence patterns for both sites also show that the grassland habitat was largely ignored during both the Middle and Late periods. Grassland taxa account for less than one percent of each assemblage. This is not surprising for SAC-329, because no grassland habitat was located within five kilometers (three miles) of the site. However, 59 percent of SAC-67 catchment was grassland. One explanation for this incongruity relates to the seasonal characteristics of study area grasslands. During late winter/spring flooding, grasslands adjacent to the Cosumnes River (and other streams) became seasonal wetlands, leaving SAC-67 an island. In fact, archaeologists working at the site needed boats to reach the site during these seasons (J. Johnson 2010, personal communication). Thus, the formerly seasonal conversion of certain grasslands to wetland habitats would have reduced the availability of grassland resources for nearly a quarter of the year. 78 Grassland The two grassland habitat sites are SAC-133 (Middle Period) and SAC-267 (Late Period). Both have nearly identical catchments with 79 to 80 percent of the foraging radius in riparian woodland, 20-21 percent in grassland, and less than one percent freshwater marsh. Subsistence patterns reveal that exploitation of small to medium bodied freshwater marsh & riparian woodland animals procured by mass capture is the dominant subsistence strategy at both sites (𝜒2=92.34, df=29, p <0.001). This is largely reflected by the large abundance of Sacramento sucker in both assemblages, where it appears to have been a dietary staple over time. Apart from that, little else is similar between the two subsistence patterns. The most significant differences between the two subsistence patterns are reflected in the decrease of very small taxa (e.g., gopher), decrease in use of grassland & riparian woodland, and increase in the use of freshwater marsh habitat in the Late Period. Decreases in the first two patterns are reflected by an 81 percent decrease in Late Period gopher remains, and ambiguity surrounding the squirrel remains from Figure 20. SAC-133 Middle Period Grassland Subsistence Pattern. 79 Figure 21. Grassland Individual Site Catchments. 80 Figure 22. SAC-267 Late Period Grassland Subsistence Pattern. SAC-267. Botta’s pocket gopher is attracted to mesic environments with a continuously growing root system (Jameson and Peeters 2004:291). If root access is restricted due to prolonged flooding or habitat loss, Botta’s pocket gophers will decrease in numbers. These preferences suggest pocket gophers would have been most prevalent in riparian woodlands, but could also be found in moist areas of grassland. Thirty-nine squirrel remains were documented in the SAC-267 assemblage, but none were identified to species. This may be due to use of analysts from Arizona, who lacked familiarity with California ground squirrel remains. If the squirrel remains had been assigned to species, use of either grassland (California ground squirrel), or riparian woodland (grey squirrel), or both habitats may have increased along with that of small animals during the Late Period. The increase in freshwater marsh taxa relates primarily to waterfowl and slow-water fishes during the Late Period. The significance of other changes is evident in the results of the chi-square analysis. Chi-square analysis of terrestrial taxa (𝜒2=33.86, df=6, p <0.001) show significant increases in artiodactyls, waterfowl, and beaver in grassland subsistence economies over time. Of note is the addition of 81 geese, primarily Canada goose, to Late Period diets. Geese can be found in both grassland and freshwater marsh habitats, so travel beyond the catchment may have been unnecessary to procure geese. Significant decreases are likewise apparent in carnivores and medium-size rodents, but the numerical decrease in lagomorphs is not statistically significant. Figure 23. Diachronic Grassland Terrestrial Taxa. Figure 24. Diachronic Grassland Fish. 82 Figure 24 shows significant differences in grassland fish over time (𝜒2=100.62, df=10, p <0.001). These differences included significant changes in salmon and all slow-water species. Salmon increases by 325 percent in the Late Period and slow-water fish by 638 percent. The extremely large increase in slowwater fish is amplified by the inclusion of three previously unidentified species: hitch, thicktail chub, and tule perch. In terms of site catchment, both the subsistence pattern and chi-square data indicate that habitats within three kilometers (five miles) of the sites provided the bulk of faunal resources to occupants of grassland environments over time. Although riparian woodland and grassland taxa provided the bulk of the diet (e.g., lagomorphs, medium rodents, and Sacramento sucker), people living in grassland habitats were exploiting habitats beyond their catchment to obtain freshwater marsh resources (e.g., waterfowl and Sacramento perch). Intensification of this behavior during the Late Period is supported by a widening of diet breadth to include three additional species of slow-water fish and an 83 percent increase in waterfowl remains. At the same time, however, foraging efficiency also appears to increase, given a rise in large bodied taxa. Most of these taxa (e.g., artiodactyls, beaver, Canada goose, and salmon) were previously available in the site catchment, such that their increase implies improved success in encounter rates. Riparian Woodland The riparian woodland habitat is represented by SAC-42 and SAC-29. Both sites are located on the east side of the Sacramento River. The SAC-42 catchment was composed of 75 percent freshwater marsh and 25 percent riparian woodland, and the SAC-29 catchment was 75 percent freshwater marsh, 18 percent riparian woodland, and seven percent grassland (Figure 25). The subsistence pattern derived from each of the riparian woodland faunal assemblages is presented below. Despite similarities in catchment areas, significant differences exist between riparian woodland subsistence patterns over time (𝜒2=213.41, df=26, p <0.001). The Middle Period subsistence pattern indicates that inhabitants of SAC-42 practiced mass harvesting of medium sized animals from freshwater marsh habitats. Most of these are Sacramento perch, a large, slowwater fish, that comprises nearly a third of the assemblage. The next most numerous taxa are Sacramento 83 Figure 25. Riparian Woodland Individual Site Catchments. 84 Figure 26. SAC-42 Middle Period Riparian Woodland Subsistence Pattern. Figure 27. SAC-29 Late Period Riparian Woodland Subsistence Pattern. 85 sucker, artiodactyls, and waterfowl. Although no grassland habitat is present in the SAC-42 catchment, limited numbers of grassland taxa (e.g., pronghorn, badger, and lagomorphs) were identified. These indicate that inhabitants of SAC-42 probably ventured outside their catchment to supplement a fish based diet, much like the people at SAC-133. Unlike the inhabitants of SAC-42, the Late Period people at SAC29 acquired a wide range of medium- and large-sized taxa from freshwater marsh and riparian woodland habitats. These include large fish (e.g., salmon, Sacramento perch, and hardhead), large waterfowl (e.g., geese), and medium to large mammals (e.g., artiodactyls, canids, raccoon, badger, and beaver). Most of these taxa were best procured by individual rather than mass capture techniques. Both subsistence patterns include some grassland taxa (pronghorn and lagomorphs) despite the lack of grassland habitat in the SAC-42 catchment and only seven percent in SAC-29 foraging radius. As with the grassland habitats, inhabitants of the riparian woodland probably ventured outside their catchment area to supplement a fish based diet. Other diachronic changes in subsistence are highlighted by the chisquare analysis. It shows that changes in the frequency of some terrestrial taxa are not significant (𝜒2=60.73, df =6, p <0.001). Four of the seven taxa show no statistically significant changes in relative frequency. A significant increase is seen in waterfowl, which increase by seven percent. Conversely, carnivores and turtles exhibit a significant decrease over the same time. Figure 28. Diachronic Riparian Woodland Terrestrial Taxa. 86 Despite significant increases in the frequency of identified fish, fish remains actually decrease in the Late Period assemblage overall by 25 percent. This decrease is associated with a significant decline in Sacramento perch, coupled with a significant increase in anadromous fast-water species over time. (𝜒2=1732.08, df=10, p <0.001). As Figure 29 shows, Sacramento perch, staple of the Middle Period diet, decrease by 92 percent and Sacramento sucker by 98 percent. These decreases are overshadowed, however, by significant increases in the abundance of anadromous species (i.e., salmon and sturgeon). Salmon increase in frequency by over 2,000 percent and sturgeon by 700 percent. Hardhead, another fast-water species, shows a smaller yet still significant increase in frequency. The significant increase in all fast-water species supports the increased use of riparian woodlands during the Late Period, whereas fish from other settings show no consistent pattern. When the subsistence patterns and chi-square analysis are compared to the site catchment for each time period, several trends are apparent. First, although freshwater marsh is equally represented in both site catchments, the use of this habitat decreased over time with increasing use of large bodied taxa from the Figure 29. Diachronic Riparian Woodland Fish. 87 riparian woodland (i.e., salmon and sturgeon). Increases in the NISP of other large bodied riparian woodland taxa such as black-tailed deer, raccoon, and beaver support this Late Period trend, although the latter increases are not statistically significant (see Figure 29). Second, the limited grassland taxa in either assemblage reflect the likely absence/near absence of grassland habitat in both site catchments. Nevertheless, people in both time periods either foraged beyond their immediate catchments or traded in order to procure grassland resources. Third, in contrast to previous habitats, there is a decrease in diet breadth (richness decreases) and an increase in foraging efficiency (amount and kind of large bodied game) within the riparian woodland during the Late Period. This, too, is supported by the results of the reciprocal of Simpson’s Diversity Index presented in Table 11. It indicates that the observed decrease in taxonomic diversity is not a function of sample size because NISP varies independently. Lastly, the significant shift in procurement methods from group to individual hunting techniques demonstrates that Late Period changes in hunting technologies increased success in encounter rates with larger bodied taxa and, as a result, increased foraging efficiency. Summary and Discussion Quantitative analysis of diachronic faunal assemblages resulted in several conclusions relating to changes in prehistoric subsistence patterns over time and space in the study area. 1. A broad spectrum diet with fish serving as a staple supplemented by mammals and seasonal waterfowl characterized regional subsistence patterns Diversity among site samples ranges from 23 to 38 taxa. This includes between eight and 11 fish taxa, which comprise 60 to 79 percent of the assemblages. The most numerous fish in all samples were either Sacramento perch or Sacramento sucker. Sacramento perch was the largest minnow in the study area and its remains account for a quarter of all fish bone examined. Sacramento sucker is a medium sized fish and its remains account for eight percent of the fish bone. A total of 21 different mammals are represented in study area assemblages. Artiodactyls and lagomorphs are found in every sample, with the quantity of 88 medium rodents and carnivores varying. Although more than 11 bird taxa were identified, waterfowl comprise the most numerous bird remains in every sample, with ducks accounting for 54 percent of all identified birds. 2. The freshwater marsh was the most targeted habitat during both the Middle and Late periods The freshwater marsh habitat accounted for approximately 36 percent of the study area and 54 percent of the taxa identified. Taxa which prefer a variety of habitats, including the freshwater marsh, account for an additional 22 percent. Figures 30 and 31 depict the subsistence patterns derived for the Middle and Late period assemblages based on specimens from the lowest level of taxonomic rank (𝜒2=215.14, df=30, p <0.001). Even when site catchments included less than one percent of this habitat (e.g., SAC-133 and SAC-267), more than 50 percent of the taxa identified were associated with the freshwater marsh. While most taxa procured from the freshwater marsh were fish, 29 percent represented terrestrial taxa, such as tule elk, Figure 30. Study Area Middle Period Subsistence Pattern. 89 Figure 31. Study Area Late Period Subsistence Pattern. raccoon, beaver, ducks, and turtle. Conversely, grassland habitats, which comprised 52 percent of the study area, never account for more than 10 percent of the taxa exploited, even when taxa preference includes a grassland component. 3. Artiodactyl remains increase over time Artiodactyls are the largest taxa found in the study area and account for 39 percent of the mammals identified. A significant increase in artiodactyl remains is exhibited in both the freshwater marsh and grassland habitats. An increase in artiodactyls is also seen in the riparian woodland, although it is less significant than changes in other terrestrial fauna. Table 9. Diachronic Change in Artiodactyl Species. Species NISP Adjusted Residual Middle Late Tule Elk 62 327 2.44 Black-tailed Deer 33 176 1.44 Pronghorn 29 25 -6.78 90 The limited number of pronghorn in all assemblages most likely reflects the general lack of grassland targeting, while the significant increases in tule elk and black-tailed deer are more likely related to the next conclusion. 4. Changes in hunting technology increased foraging efficiency in the Late Period Although increases in taxonomic diversity were not identified in all habitats, increases in large prey are seen in every study area habitat over time, regardless of site catchment. These large bodied taxa prefer multiple habitats, and some were only seasonally present. Thus, increases in these animals cannot be attributed to changes in environment alone. The aridity and droughts experienced in the Late Period during the MCA appear to have precluded high water flows in the Sacramento River, which may have impacted salmon runs and reduced primary production for herbivores. Nevertheless, this is exactly the time period when both salmon and artiodactyl remains increase in assemblages. Increased flooding during the LIA in the latter half of the Late Period, could have also impacted primary production and reduced habitat for certain grassland species, such as lagomorphs and pronghorn, by increasing the size and duration of seasonal wetlands. Despite this possibility, changes in lagomorph frequency were typically not significant and the frequency of pronghorn specimens is currently impossible to assess with the data available. Enlarged seasonal wetlands may have provided increased forage for juvenile fish, increasing survivorship, and/or providing hunters with easy targets. Support for such a scenario is reflected in fish increases in Late Period freshwater marsh and grassland assemblages, but additional study (age grading) is needed before this scenario can be considered a direct influence on local subsistence economies. More certain, perhaps, is that the ubiquitous increase in Late Period artiodactyls, particularly tule elk and black-tailed deer, implies that introduction of bow and arrow technology increased hunting success. Corresponding conclusions have been proposed for the Bay Area in order to explain similar increases in highly ranked mammalian prey species (Wake 2012:31). Increases in individually procured medium mammals (i.e., canids, carnivores, and beaver) and geese, support the adoption of a more efficient hunting 91 technology. Similarly, changes in fishing technology (e.g., the toggle harpoon) allowed large bodied anadromous species to be exploited with greater success. Salmon and sturgeon were identified at all six sites regardless of time period. It is not until when the toggle harpoon was introduced during the Late Period that salmon and sturgeon surpass the stable dietary contribution of slow-water minnows and suckers.