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A Prehistoric Freshwater Mussel Assemblage from the Big Sunflower River, Sunflower County, Mississippi
Joseph Mitchell and Evan Peacock

Southeastern Naturalist, Volume 13, Issue 3 (2014): 626–638

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Southeastern Naturalist J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 626 2014 SOUTHEASTERN NATURALIST 13(3):626–638 A Prehistoric Freshwater Mussel Assemblage from the Big Sunflower River, Sunflower County, Mississippi Joseph Mitchell1,* and Evan Peacock2 Abstract - Zooarchaeological deposits present a unique insight into the trajectory of faunal communities through time. Freshwater mussels (Bivalvia: Unionidae) are one of the world’s most imperiled faunal groups, and, at the same time, constitute a significant component of the archaeological record in North America. Current conservation efforts can be informed by studies of prehistoric mussel assemblages that catalog communities as they existed prior to any extensive modern impacts, ultimately representing an ecological baseline against which current populations can be evaluated. Over 47,000 freshwater mussel valves were recovered from the late prehistoric (ca. A.D. 700–1200) Kinlock site (22SU526), in Sunflower County, MS. Analyses revealed that the Sunflower River once supported a mussel community with greater taxonomic richness and more evenness among all the major species than today. Additionally, the presence in the archaeological record of numerous species which are currently considered rare, endangered, or extinct in the Sunflower River is indicative of a habitat disturbed by increased human environmental impact in historic times. Introduction Bone and shell assemblages recovered from archaeological deposits are receiving increased attention from scientists interested in the evolution and current state of faunal communities (Frazier 2010; Lyman 1996, 2006; Peacock et al. 2011, 2014). The applied value of zooarchaeology for a number of fields, including conservation biology, has been noted (e.g., Anderson et al. 2010; Bailey et al. 2000; Crumley 1994; Frazier 2007; Lyman 1996, 2006) and has been demonstrated in a growing number of case studies (e.g., Haag 2009, Lyman and Cannon 2004, Wolverton and Lyman 2012). Identifying original species’ ranges and how they have changed over time is an important component of conservation efforts. The perspective provided by zooarchaeology in this regard is unique, allowing the characterization of faunal communities as they existed prior to extensive modern impacts thus establishing an ecological baseline against which data from current communities can be compared. Freshwater mussel (Bivalvia: Unionidae) shells are common in archaeological contexts in Mississippi; they are found at sites of various ages and sizes located along lakes, major rivers, and small channels (Peacock and Jenkins 2010, Peacock et al. 2011). Zooarchaeological studies in Mississippi (e.g., Bogan 1987, Hartfield 1993, Jones et al. 2005, Peacock and James 2002, Peacock and Mistak 2008, Peacock et al. 2011) and elsewhere (e.g., Barber 1982, Gordon 1983, Peacock et al. 2013, Randklev and Lundeen 2012) have shown that sites frequently contain species not previously 1Department of Geosciences, Mississippi State University, Starkville, MS 39762. 2Cobb Institute of Archaeology, Mississippi State University, Starkville, MS. 39762. *Corresponding author - Manuscript Editor: Alex Huryn Southeastern Naturalist 627 J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 known to have existed in a given water body. Accordingly, data derived from archaeological shell assemblages can be very useful for current conservation efforts, particularly those data derived from sites located along waterways where modern biological surveys have not been carried out or where modern coverage has been spotty (Peacock et al. 2011:3, Randklev and Lundeen 2012). Despite being historically diverse and abundant throughout much of North America, many freshwater mussel taxa are presently in steep decline, and mussels in general are considered one of the most imperiled faunal groups globally (Bogan 2008, Grabarkiewicz and Davis 2008, Haag 2009, Lydeard et al. 2004, Machtinger 2007, Neves et al. 1997). In Mississippi alone, there are over 23 endangered mussel species (Jones et al. 2005, Mississippi National Heritage Program 2011). Data obtained from shell-bearing archaeological sites can be used to establish the pre-industrial ranges and expected natural proportions of mussel species (Peacock 2012) in aquatic systems now extensively altered through impoundment, channelization, dredging, water pollution, other forms of habitat destruction, or hydrologic changes (Bogan 1993; Haag and Warren 1998; Strayer et al. 2004; Williams et al. 1993, 2008). Here, we present data on a large assemblage of mussel shells obtained from a prehistoric site located beside the Big Sunflower River in Sunflower County, MS, and discuss differences between that assemblage and modern faunas recorded in the waterway. Methods Specimens examined for this study are from the late prehistoric (ca. AD 700– 1200) Kinlock site (22SU526), located in southern Sunflower County, MS (Fig. 1). The site is in an agricultural field along the Big Sunflower River, a major stream in the Yazoo Basin, and consists of extensive deposits of habitation debris, a plaza, and as many as six earthen mounds (Carlock and Rafferty 2009, Phillips 1970). In the summer of 2009, the Mississippi State University (Mississippi State, MS) archaeology field school collected shells from the surface of the shell-ring portion of the site and from three excavation units—two in areas of high shell density and one in the plaza (Carlock and Rafferty 2009). Workers screened excavated deposits through 6.35-mm (quarter-inch) wire mesh to recover shells. Other details of collection strategies, unit sizes, and recovery methods are given in Mitchell (2012). Shells obtained from plowzones (surface-collected specimens) undoubtedly are biased by differential preservation, with thicker, heavier shells disproportionately represented (e.g., Peacock 2000, Peacock and Chapman 2001, Wolverton et al. 2010). Shells excavated from sub-plowzone contexts, however, were in excellent condition (see Mitchell 2012 for details). Ultimately, we were able to determine the species for 75% of the valves recovered from the site. We identified shells to genus and species level using various guides (e.g., Burch 1975, Cummings and Mayer 1992, Howells et al. 1996, Parmalee and Bogan 1998) and the freshwater mussel comparative collection housed at the Cobb Institute of Archaeology, Mississippi State University, from which representative specimens have been verified at a number of natural science collection locales. Taxonomy was assigned using Turgeon et al. (1998) with minor updates. Southeastern Naturalist J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 628 Results A total of 47,431 valves retaining umbos were recovered from the site (surface and excavated contexts combined), from which we identified 33 species. These species (left and right valves combined) are listed in descending order of abundance in Table 1. Figure 1. Location of the Kinlock Site (22SU526), Sunflower Coun ty, MS. Southeastern Naturalist 629 J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 Table 1. Archaeological freshwater mussel valves from the Kinlock site (22SU526) and modern mussel surveys in the Big Sunflower River, Sunflower County, MS. Data from modern surveys derived from Miller and Payne (1995, 1997, 2004) and Miller et al. (1992). # = number of valves, and % = percent identified valves. Modern Species # % surveys Pleurobema rubrum (Rafinesque) Pyramid Pigtoe 6968 19.682 X Amblema plicata (Say) Threeridge 6774 19.134 X Plectomerus dombeyanus (Valenciennes) Bankclimber 5062 14.298 X Fusconaia ebena (Lea) Ebonyshell 3714 10.491 X Fusconaia flava (Rafinesque) Wabash Pigtoe 3703 10.460 X Obliquaria reflexa (Rafinesque) Threehorn Wartyback 2730 7.711 X Quadrula pustulosa (Lea) Pimpleback 1879 5.307 X Quadrula quadrula (Rafinesque) Mapleleaf 1445 4.082 X Quadrula nodulata (Rafinesque) Wartyback 647 1.828 X Elliptio dilatata (Rafinesque) Spike 604 1.706 Lampsilis hydiana (Lea) Louisiana Fat Mucket 427 1.206 X Lampsilis teres (Rafinesque) Yellow Sandshell 351 0.991 X Cyprogenia aberti (Conrad) Western Fanshell 173 0.489 Megalonaias nervosa (Rafinesque) Washboard 166 0.469 X Truncilla truncata (Rafinesque) Deer Toe 149 0.421 X Quadrula apiculata (Say) Southern Mapleleaf 116 0.328 Ligumia recta (Lamarch) Black Sandshell 103 0.291 Quadrula verrucosa (Rafinesque) Pistolgrip 86 0.243 X Lampsilis cardium (Rafinesque) Plain Pocketbook 72 0.203 Quadrula cylindrica (Say) Rabbitsfoot 56 0.158 Lampsilis siliquoidea (Barnes) Fatmucket 47 0.133 Arcidens confragosus (Say) Rock Pocketbook 46 0.130 X Quadrula metanevra (Rafinesque) Monkeyface 18 0.051 Toxolasma parvum (Barnes) Lilliput 17 0.048 Plethobasus cyphyus (Rafinesque) Sheepnose 12 0.034 X Glebula rotundata (Lamarck) Round Pearlshell 11 0.031 X cf. Lampsilis ovata (Say) Pocketbook 8 0.023 Potamilus purpuratus (Lamarck) Bleufer 7 0.020 X Ellipsaria lineolata (Rafinesque) Butterfly 5 0.014 X Villosa lienosa (Conrad) Little Spectaclecase 4 0.011 Toxolasma texasiensis (Lea) Texas Lilliput 1 0.003 X Strophitus undulatus (Say) Creeper 1 0.003 Obovaria subrotunda (Rafinesque) Round Hickorynut 1 0.003 Actinonaias ligamentina (Lamarck) Mucket 0 .000 X Elliptio crassidens (Lamarck) Elephantear 0 .000 X Leptodea fragilis (Rafinesque) Fragile Papershell 0 .000 X Potamilus ohiensis (Rafinesque) Pink Papershell 0 .000 X Pyganodon grandis (Say) Giant Floater 0 .000 X Truncilla donaciformis (Lea) Fawnsfoot 0 .000 X Uniomerus declivis (Say) Tapered Pondhorn 0 .000 X Uniomerus tetralasmus (Say) Pondhorn 0 .000 X Anodonta suborbiculata (Say) Fat Floater 0 .000 X Utterbackia imbecillis (Say) Paper Pondshell 0 .000 X Total identified: 35,403 100 Unidentifiable: 12,028 Total analyzed valves: 47,431 Southeastern Naturalist J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 630 Discussion Comparison of archaeological and modern mussel assemblages is complicated by a number of factors. For example, work at a given archaeological site might produce tens of thousands of valves, as at Kinlock, so that the assemblage presumably represents long-term collecting over a relatively small area (see Peacock 2002, where spatially limited collection areas were demonstrated in an archaeological test case). Modern biological surveys, on the other hand, represent very time-limited collecting over stations that are widely (and usually not evenly) distributed in space, so that results from modern surveys can vary significantly from year to year. Even simple comparisons of richness and evenness are complicated by preservation (and possibly prehistoric collection) biases in archaeological shells and by the fact that some species have expanded their ranges in modern times. Accordingly, comparisons between archaeological and modern faunas have tended to be qualitative in nature (e.g., Lyons et al. 2007, Tevesz et al. 2002). While more statistically formal, drainage-scale comparisons may be made when enough archaeological assemblages have been reported (Haag 2009, Hughes and Parmalee 1999), such is not yet the case with the Yazoo Basin, although both the quantity and quality of data on archaeological mussel-shell assemblages is very much on the increase there (Peacock et al. 2011). In this paper, we discuss differences between prehistoric and modern faunas in a qualitative fashion. We identified 13 species in the Kinlock assemblages not found in modern surveys (Miller and Payne 1995, 1997, 2004; Miller et al. 1992). Of these, Lampsilis ovata (Pocketbook) must be considered a tentative identification within the archaeological assemblage, given the difficulty of separating this form from related species. Those from Kinlock are more inflated and have a more pronounced break along the posterior ridge than do the specimens identified as L. cardium (Plain Pocketbook). We note however, that L. cardium also is not reported in the modern surveys discussed in this article. While we believe we have consistently differentiated between Toxolasma parvum (Lilliput) and T. texasiensis (Texas Lilliput) based on size and surface sculpture (see Peacock et al. 2013), we acknowledge that archaeological specimens of these forms can be difficult to separate and they are not further discussed here. In contrast, although different species of Obovaria also can be hard to discriminate, we are confident that those we have categorized as O. subrotunda (Round Hickorynut) are correctly identified, based on a distinctive shell shape and extensive comparison with modern specimens in museum collections. The species is known historically from the Yazoo Basin (Jones et al. 2005), where it also has been found at over half a dozen archaeological sites (Peacock et al. 2011:map A-36). Elliptio dilatata (Spike) is frequently characterized as a clear-water species (e.g., Parmallee 1967, Starret 1971), and its presence at Kinlock and other archaeological sites in the Yazoo Basin (Hartfield 1993:table 1, Peacock et al. 2011:map A-10), emphasizes changes that likely have occurred as a result of increased turbidity (e.g., Dineen 1971, Hoeh and Trdan 1984, Klippel et al. 1978, Starrett 1971, Taylor and Spurlock 1982, White 1977). The same applies to Cyprogenia aberti (Western Southeastern Naturalist 631 J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 Fanshell), an Ozarkian species known to have had a wide prehistoric distribution in the Yazoo Basin (Bogan 1987, Hartfield 1993, Jones et al. 2005, Peacock and James 2002, Peacock et al. 2011). Both of these species also have been found in lowland archaeological contexts along western tributaries in the Mississippi River drainage (Peacock and Chapman 2001, Peacock et al. 2013, Vidrine 1993). Quadrula cylindrica (Rabbitsfoot) and Quadrula metanevra (Monkeyface) are historically very rare in Mississippi (e.g., Hartfield 1987:140); these species were widely distributed but similarly rare in alluvial valley settings in Mississippi and adjacent states during prehistoric times (Hartfield 1993, Jones et al. 2005, Peacock and Chapman 2001, Peacock et al. 2011:map A-28, Peacock et al. 2013, Vidrine 2008). Monkeyface is considered endangered nationally, but is currently presumed extinct in Mississippi (Mississippi Natural Heritage Program 2011). Strophitus undulates (Creeper), a critically imperiled species in Mississippi (Jones et al. 2005), is described as present, but extremely rare in the Yazoo drainage (Jones et al. 2005). Apparently, it was equally rare in the past, with only one other archaeological occurrence on the Big Sunflower River in Coahoma County documented for Mississippi (Peacock et al. 2011:53; see Peacock et al. 2013 for occurrences from Bayou Bartho lomew, AR). Species present in modern collections in the Big Sunflower River, but absent at Kinlock, include Actinonaias ligamentina (Lamark) (Brass Mucket), Elliptio crassidens (Lamark) (Elephantear Mussel), Leptodea fragilis (Rafinesque) (Fragile Papershell), Potamilus ohiensis (Rafinesque) (Pink Papershell), Pyganodon grandis (Say) (Giant Floater), Truncilla donaciformis (Lea) (Fawnsfoot), Uniomerus declivis (Say) (Tapered Pondhorn), U. tetralasmus (Say) (Pondhorn), Anodonta suborbiculata Say (Flat Floater), and Utterbackia imbecillis (Say) (Paper Pondshell). Of these, Brass Mucket, Elephantear Mussel, Fawnsfoot, Tapered Pondhorn, and Pondhorn have been reported from other archaeological sites in the Yazoo Basin, although all were apparently quite rare there in prehistoric times (Peacock et al. 2011). Fragile Papershell has been reported from archaeological contexts in numerous cases (see Peacock et al. 2011:table 3), and its absence at Kinlock may be due to preservation bias against thin-shelled species, a factor that also might explain the absence of Giant Floater, Flat Floater, and Paper Pondshell.1 Alternatively, these species may have spread into the drainage coincident with habitat change and/or the introduction of fish hosts in historic times, a commonly noted phenomenon (e.g., Casey 1987, Grace and Buchanan 1981, Taylor and Spurlock 1982, Williams et al. 2008:750–751). While it is possible that some of the archaeological specimens we have identified as Potamilus purpuratus (Bleufer) may be P. ohiensis, we do not think this is likely because our specimens have the inflated shells characteristic of the former species. If we are correct, then P. ohiensis seems to be a recent introduction into the Yazoo Basin. Modern surveys in the Sunflower River found that mussel beds tend to be dominated by one or two species. In one survey, for example, researchers found that Amblema plicata (Threeridge) accounted for ~90% of the total population, and the next most abundant taxon represented only 3.5% (Miller and Payne 1995:17). The prehistoric assemblage is much more balanced, with eight species representing at least 5 percent of the population and none accounting for more than 20 percent. Also, large numbers of small valves were recovered from archaeological deposits, Southeastern Naturalist J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 632 indicating healthy populations with consistent recruitment in all the major species, whereas modern surveys of the Big Sunflower River show no recent recruitment visible for several species (Miller and Payne 1995, 2004; Miller et al. 1992). This historic decline in evenness is typical of disturbed habitats and likely is related to increased human environmental impact (e.g., Haag and Warren 1998, Miller and Payne 1995, Peacock et al. 2011). The Big Sunflower River has been impacted in modern times by dredging activities of the US Corps of Engineers, the negative effects of which have been discussed by Miller and Payne (1995) and others (Miller et al. 1992, 1993). Hartfield (1993) suggested that siltation stemming from erosion is also a factor in mussel scarcity (Hartfield 1993). The modern assemblages referenced herein are generally similar, with differences between them more likely due to sampling error than actual conditions. Not surprisingly, silt-tolerant taxa dominate the modern fauna. One unexpected finding in the Kinlock assemblage is a single Rangia cuneata (G.B. Sowerby I) (Atlantic Rangia) valve (Fig. 2). Prior research on the geographic Figure 2. Left valve of Rangia cuneata (Atlantic Rangia), recovered from the surface of the Kinlock site. Southeastern Naturalist 633 J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 extent of Atlantic Rangia, a brackish-water bivalve (e.g., LaSalle and de la Cruz 1985, Wakida-Kusunoki and MacKenzie 2004; see also Cobb 2009), shows that it is primarily associated with coastal archaeological sites. The most northerly archaeological occurrence along the Gulf of Mexico is from the Lower Tombigbee River, slightly north of the Mobile–Tensaw Delta of south Alabama, which seems to have been the northern natural limit of the species in prehistoric times (Peacock et al. 2014). Thus, the identification of Atlantic Rangia from a site on the upper Big Sunflower River is curious. It is reasonable to assume that this shell is an import, as has been suggested for a few other prehistoric sites. For example, Hill (1983) discussed thousands of valves of Atlantic Rangia at the Arrowhead Farm site on the Ohio River in Jefferson County, KY, and small numbers of valves have been recovered from the famous Cahokia site in Illinois, near the confluence of the Mississippi and Missouri rivers (Baker 1941, Parmalee 1958). The hypothesis that such biogeographical anomalies represent imports can be tested and source areas can potentially be determined using elemental and stable isotope analyses (e.g., Peacock et al. 2010). Conclusions State and federal agencies are working hard to slow the decline of freshwater mussel species in the US. Efforts to restore areas where mussel populations have been seriously impacted have met limited success (Machtinger 2007). Unfortunately, biologists and malacologists are still in the developmental stage in understanding how to improve the survival rate, population growth, and recruitment of relocated individuals in affected mussel communities (Machtinger 2007:7). The lack of a true baseline in how we view and understand mussel communities is a perennial problem, one that can be addressed via reference to the abundant zooarchaeological record. To better understand how humans are affecting mussel communities today, it is useful to first establish an evolutionary trajectory from past to present. Zooarchaeology provides a unique perspective that can contribute to solving modern-day problems by informing species-reintroduction efforts, providing long-term data on endangered species, promoting conservation, and providing better understanding of the long-term trajectory of human environmental impacts. Notes 1Another explanation is that Indians were avoiding slack-water species due to taste. While this is a legitimate hypothesis, such species are, in fact, frequently found with other species in prehistoric middens (e.g., Peacock et al. 2013:table 3). There is an obvious danger in using contemporary, common-sense assumptions to explain the behaviors of prehistoric peoples from completely different cultures; such assumptions must be framed as hypotheses and tested. Acknowledgments We thank Bradley Carlock, Janet Rafferty, and the Mississippi State University fieldschool students and lab workers whose hard work yielded the interesting molluscan Southeastern Naturalist J. Mitchell and E. Peacock 2014 Vol. 13, No. 3 634 assemblage from Kinlock. The comments by an anonymous reviewer were very helpful, especially in pointing us to sources on the archaeological occurrence of Rangia cuneata. We also thank John Rodgers for providing the map used in Fig. 1. Literature Cited Anderson, E.N., N.J. Turner, E.S. Hunn, and D.M. Pearsall. 2010. Ethnobiology. 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