FranzQuitmyer 179 A FOSSIL AND ZOOARCHAEOLOGICAL HISTORY OF THE GOPHER TORTOISE (GOPHERUS POLYPHEMUS) IN THE SOUTHEASTERN UNITED STATES Richard Franz1 and Irvy R. Quitmyer1 Specimens of fossil gopher tortoises (Gopherus) were collected from five late Pliocene, two early Pleistocene, five middle Pleistocene, and 52 late Pleistocene sites in 18 counties in Florida, one county in Georgia, three in South Carolina, and one in Mississippi. Occurrences of fossil Gopherus polyphemus in Lowndes County, Mississippi, and Charleston, Colleton, and Horry counties, South Carolina, represent extralimital records outside the current geographic range of the species. The extensive fossil record indicates G. polyphemus has been part of the xeric-adapted fauna of the southeastern coastal plain for at least two million years. The majority of the Florida records are from Alachua and Marion counties. This concentration corresponds to the high frequency of late Pleistocene fossil deposits in solution features associated with limestone quarries at Arredondo, Haile, and Reddick. A query of 609 sites in the zooarchaeological database of the Environmental Archaeology laboratory at the Florida Museum of Natural History was made to determine the presence of G. polyphemus in 67 archaeological sites from the southeast- ern United States dating from the latest Pleistocene to the late 19th century. The zooarchaeology collections are heavily weighted in favor of sites from Florida and Georgia. These data are not all inclusive of sites from the Southeast, but present a representative record of the association of this species with humans for nearly 12,000 years. Specimens of G. polyphemus from archeological sites are known from 20 Florida counties and one in Georgia. Key Words: tortoise; Gopherus polyphemus; Pleistocene; zooarchaeology; southeastern United States 1Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800; Bull. Fla. Mus. Nat. Hist. (2005) 45(4): 179-199 INTRODUCTION The fossil and archaeological record for the gopher tor- toise, Gopherus polyphemus, is extensive but has never been fully reviewed. Hay (1916) named Gopherus praecedens based on a left xiphiplastron from the late Pleistocene at Vero (bed 2), Indian River County, Florida. However, Auffenberg (1974) considered this species a synonym of G. polyphemus. Other references for fos- sil records of G. polyphemus include Auffenberg (1974), Hay (1917), Holman (1958, 1959, 1995), Holman and Clausen (1984), Hulbert and Pratt (1998), Meylan (1982, 1984, 1995), Weigel (1962), and Young and Laerm (1993). Gopherus fossils are commonly collected from cave, sinkhole, fluvial, and estuarine deposits in the south- eastern United States (Fig. 1). Gopher tortoises also routinely occur in midden deposits and are often men- tioned in archaeological site reports (Fig. 2). Native Americans migrated into the southeastern United States near the end of the Pleistocene and apparently lived continuously in this area for at least 12,000 years (Milanich 1994). PaleoIndians probably encountered and foraged on gopher tortoises, as well as extinct tortoises of the genus Hesperotestudo. An early archaeological site at Little Salt Spring, Sarasota County, Florida, con- tained fossils of both G. polyphemus and H. crassiscutata (Clausen et al. 1979; Holman & Clausen 1984). The primary purpose of this paper is to review the fossil record of Gopherus polyphemus from Florida and the Southeast. It is not our intent to evaluate mor- phological variability within fossil gopher tortoise popu- lations of the Southeast, or to make taxonomic revisions. The zooarchaeological record presented here is not ex- haustive, but provides a representative record of the association of G. polyphemus with humans in the re- gion. The late Pliocene samples from the Inglis and Waccasassa sites may represent an undescribed dwarf population of a Gopherus polyphemus-like species, but its description lies beyond our present focus. We treat it 180 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb here as G. cf. polyphemus because of its general ap- pearance and probable close relationship to the recent species. We consider all late Pliocene and Pleistocene populations from the Southeast to be ecological equiva- lents of the modern species based on morphological simi- larities that we consider adaptations for burrowing. The most important conclusion reached here is that G. polyphemus has had a long association with the south- eastern coastal plain and probably played important roles in southeastern landscape dynamics and human history. METHODS Fossil and zooarchaelogical materials examined are de- posited in following collections: Florida Museum of Natu- ral History’s vertebrate paleontology (UF), Florida Geo- logical Survey (UF/FGS), and zooarchaeology (UFea) collections, Charleston Museum (ChM), and United States National Museum (USNM). Specimens in the private collection of Steve and Suzan Hutchens of Old Town, Florida, are designated SSH. We also examined collections at Auburn University, Georgia State College and University at Milledgeville, and the South Carolina State Museum, but failed to find additional material in their collections. Specific locations of sites are main- tained in site files at each depository and can be ob- tained through those listed institutions. Recent material used for comparison was obtained from the Florida Museum of Natural History’s herpetological (UFh) and zooarchaeological comparative collections. TERMINOLOGY AND MEASUREMENTS Shell Bone Abbreviations: NUCH, Nuchal; PYG, Pygal; EPI, Epiplastron; ENT, Entoplastron; HYO, Hyoplastron; HYPO, Hypoplastron; and XIPH, Xiphiplastron. Other Abbreviations: CL, Carapace Length; Ma, millions of years ago; MNI, minimum number of indi- viduals. Measurements: Measurements are in millimeters (mm). All measurements from external surfaces: HYO length (along midline suture); HYPO length (along mid- line suture); XIPH length (along midline suture); NUCH greatest length; NUCH width (across front of carapace from suture to suture); CL, carapace length (maximum straight line measurement with calipers). We follow the geologic time scale and faunal ages recommended for Florida by Hulbert (2001). Figure 1. Map of fossil records for Florida. Grey areas repre- sent the county and the number represents the number of sites within a county. Figure 2. Map of zooarchaeological records for Florida. Grey areas represent the county and the number represents the number of sites within a county. 181 CARAPACE LENGTH REGRESSIONS Paleontologists and zooarchaeologists are usually presented with only a portion of an animal skeleton and are unable to measure the whole organism. Fortunately, many skeletal elements scale allometrically with body size (Peters 1983). Allometry reflects the sutural and functional consequences of a change in size or scale among similarly shaped animals (Peters 1983; Schmidt- Nielson 1984; Reitz et al. 1987). Growth is a nonlinear process through ontogeny, and this allometric relation- ship is described by a mathematical power function y = a(Xb ) (Schmidt-Nielson 1984). This is transformed us- ing the common log in order to produce a straight-line regression. The resulting formula is log y = a + b(log X) with b as the slope of the line, a the y intercept, x the independent variable (skeletal measurement), and y the dependent variable, the estimated body size (or cara- pace length). Table 1 presents the results of the allometric rela- tionship of CL to each of four skeletal elements (NUCH length, NUCH width, HYO length, and XIPH length) that are often well preserved in the fossil record. HYPO length proved unreliable because of the changes in con- figuration as tortoises mature and, therefore, was not used. The regression constants are based on measure- ments of 28 modern gopher tortoises from the Florida Museum of Natural History herpetology and environ- mental archaeology collections. We obtained 109 esti- mated CLs from 78 separate fossil tortoise elements collected at Inglis 1A and 1C, Coleman 2A, Leisey Shell Pit 1A, Haile 8A, Reddick 1A, and Surprise Cave (Table 2). Each measurement was treated as a separate sample. Measurements (mm), means, standard deviations, and ranges for elements and for estimated CL are presented in Table 2. We did not calculate estimates of CL from any zooarchaeological material. DIAGNOSTIC CHARACTERS Salient morphologic features of the skeletons of the four species of extant gopher tortoises were exten- sively described by Auffenberg (1975). Data from his study supported two major evolutionary lines within go- pher tortoises: the G. flavomarginatus-polyphemus and G. agassizii-berlandieri groups. The features that dis- tinguish them were reviewed in Bramble (1971), Crumly (1994), and McCord (1997). Bramble (1971, 1982) pre- sented a phenetic analysis for gopher tortoises and erected the genus Scaptochelys for the G. agassizii- berlandieri group. The name Scaptochelys was later suppressed, and an older name, Xerobates, suggested as a replacement (Bour & Dubois 1984). Crumly (1994) argued for keeping the two clades in the genus Gopherus. McCord (1997, 2002) performed a stratocladistic analysis to evaluate relationships of re- cent and fossil gopher tortoise species, and his resulting cladogram supported a 2-group arrangement in the ge- nus Gopherus. Recent molecular data support the avail- able phenetic and cladistic conclusions and suggest that the two clades have been separated for about 18 million years (Lamb & Lydeard 1994). For this study, we used a variety of shell features to identify fossil Gopherus polyphemus. This method was greatly enhanced by direct comparisons with the extensive recent collections at UF. We relied heavily on the following features to separate Gopherus from Hesperotestudo and other southeastern fossil turtles: broad EPI beak and shelf, low-domed carapace, thin carapacial and plastral bones, clearly defined scute sulci Table 1. Allometry formula and constants used in carapace length estimate calculations for fossil Gopherus polyphemus. N r2 Intercept (a) Slope (b) Nuchal Length 27 0.84 0.79 0.98 Nuchal Width 27 0.88 0.75 1.01 Xiphiplastron Length 26 0.77 0.90 0.89 Hyoplastron Length 26 0.90 1.00 0.75 FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 182 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb Late Blancan Inglis 1A NUCH Length Estimated CL NUCH Width Estimated CL UF 211285 23.0 133.2 26.6 154.57 UF 211286 20.4 118.42 20.6 119.4 UF 211287 — — 31.1 181 UF 211288 34.1 195.93 33.3 193.94 UF 211289 32.8 188.6 34.0 198.06 UF 211290 38.0 217.87 43.7 255.50 UF 211291 31.1 179.02 33.9 197.47 UF 211292 36.4 208.87 39.2 228.68 UF 211293 31.1 180.15 31.2 181.59 UF 211294 36.2 207.75 34.5 201 UF 211295 30.6 176.2 31.9 185.71 UF 211296 36.4 208.87 36.1 210.42 UF 211297 31.4 180.71 31.6 183.94 UF 211298 26.2 151.33 30.2 175.71 UF 211323 — — 32.4 188.65 N= 13 13 15 15 Mean= 31.38 180.53 32.69 190.36 SD= 5.36 30.3 5.2 30.56 Min= 20.4 118.42 20.6 116.4 Max= 38.0 217.87 43.7 225.2 Inglis 1C NUCH Length Estimated CL NUCH Width Estimated CL SSH 43.7 249.85 40.1 233.98 SSH 34.6 198.74 35.5 206.89 SSH 40 229.1 39.6 231.03 SSH 42.2 241.44 42.7 249.31 SSH 38.8 222.36 38.7 225.73 SSH 46.2 263.85 43.7 255.2 N= 6 6 6 6 Mean= 40.92 234.22 40.05 233.69 SD= 4.06 22.8 2.94 17.3 Min= 34.6 198.74 35.5 206.89 Max= 46.2 263.85 43.7 255.2 Early-Middle Pleistocene Leisey 1A XIPH Length Estimated CL UF 80458 54 276.59 UF 69394 52.3 268.92 UF 83601 41.3 217.87 N= 3 3 Mean= 49.2 254.43 SD= 6.89 31.9 Min= 41.3 217.87 Max= 54 276.59 Table 2. Measurements (in mm) and descriptive statistics from individual fossil elements of Gopherus polyphemus from six sites in Florida. Coleman 2A NUCH Length Estimated CL NUCH Width Estimated CL UF 13390b 52.2 276.38 50.4 294.75 UF 13390h 62.4 354.23 59.3 347.36 UF 13390c 49.6 282.86 44.9 262.28 UF 13390m 63.1 358.12 59.5 348.55 183 Table 2. (Cont.) Haile 8A XIPH Length Estimate CL UF 3834 49 253.67 UF uncat. 64.2 322.63 UF 3476 49.9 257.82 UF uncat. 46 239.8 UF 3074 49.6 256.44 Coleman 2A NUCH Length Estimated CL NUCH Width Estimated CL UF 13390d 53.8 306.31 51.8 303.02 UF 13390e 47.4 270.56 42.9 250.49 UF 13390i — — 50.1 292.98 UF 13390l 55.6 316.35 51.3 300.07 UF 13390k 61.4 348.66 — — UF 13390j 53 301.85 49.6 290.03 UF 13390f — — 44.6 260.51 UF 13390a — — 50.2 293.57 N= 9 9 11 11 Mean= 55.39 315.15 50.42 294.87 SD= 5.71 31.82 5.35 31.58 Min= 47.4 270.56 42.9 250.49 Max= 63.1 358.12 59.5 348.55 Coleman 2A XIPH Length Estimated CL UF13391b 55.5 283.41 UF13391a 56.2 286.59 UF13391c 63.5 319.5 UF13397 48.3 250.45 N= 4 4 Mean= 55.88 284.99 SD= 6.21 28.22 Min= 48.3 250.45 Max= 63.5 319.5 Haile 8A NUCH Length Estimated CL UF uncat. 52.3 305.98 UF 9435 52.3 305.98 UF 3254 57.6 337.31 N= 3 3 Mean= 54.07 316.42 SD= 3.06 18.09 Min= 52.3 305.98 Max= 57.76 337.31 UF 3071 57.6 292.94 UF 3477 51.9 266.99 UF 2457 62.4 314.57 UF 3786 55.9 285.23 UF 9655 55.4 282.96 UF 3824 59.7 302.42 UF 3823 57.4 292.03 UF 3813 53.5 274.31 UF 9658 55.5 283.41 UF 9591 53.1 272.48 N= 15 15 FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 184 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb Table 2. (Cont.) Haile 8A XIPH Length Estimate CL Mean= 54.74 279.85 SD= 5.08 23.13 Min= 46 239.8 Max= 64.2 322.63 Late Pleistocene Reddick 1A NUCH Length Estimated CL NUCH Width Estimated CL UF 2706 70.8 400.9 65.3 382.88 UF 2706 58.0 329.73 55.6 325.48 UF uncat. 29.7 171.12 24.1 139.91 N= 3 3 3 3 Mean= 52.83 300.58 48.33 282.76 SD= 21.03 117.63 21.54 127 Min= 29.7 171.12 24.1 139.91 Max= 70.8 400.9 65.3 382.88 Surprise Cave NUCH Length Estimated CL NUCH Width Estimated CL UF 161885 60.8 345.32 60.5 354.46 UF 161674 — — 50.3 294.16 UF160157 — — 17.7 102.44 UF161886 — — 52 304.2 N= 1 1 4 4 Mean= — — 45.13 263.82 SD= — — 18.82 110.77 Min= — — 17.7 102.44 Max= 60.8 345.32 60.5 354.46 Surprise Cave XIPH Length Estimated CL UF 138002 64.5 323.97 UF 161921 50 258.28 UF uncat. 59.7 302.42 UF 150333 66.9 334.68 UF uncat. 50.4 260.11 UF 138060 54.7 279.77 UF 160332 89 431.48 UF 161669 59.6 301.97 N= 8 8 Mean= 61.85 311.59 SD= 12.55 55.73 Min= 50 258.28 Max- 89 431.48 Surprise Cave HYO Length Estimated CL UF uncat. 85 279.94 UF 138001 82.3 273.24 UF 161664 74.1 252.56 UF 160334 87 284.87 UF uncat. 84.2 277.96 N= 5 5 Mean= 82.52 273.24 SD= 5 12.54 Min= 74.1 252.56 Max= 89 284.87 often on raised bone, acute-edged peripherals, bony sul- cal spurs developed along the free edge of the peripher- als, distinctive shape and size of the XIPH (anal) pro- jections, squarish or broadly rectangular NUCH scale, and obvious bone scar from a vertebral strut on the un- derside of NUCH. Bramble (1982) described a suite of unique head, neck, shell, and limb characters that separated Gopherus flavomarginatus and Gopherus polyphemus from related Gopherus agassizii and Gopherus berlandieri. Among these features, the ventral strut, which extends from the base of the first dorsal vertebra onto the back of the NUCH, shows as a prominent bone scar on the underside of the NUCH plate in all fossil 185 and recent G. polyphemus we examined (Figs. 3-5). Meylan (1982) reported these strut scars in fossil Gopherus specimens from the late Pliocene Inglis 1A local fauna (Fig. 3). Strut scars are often 10-15 mm long (greater than 35% of the total width of anterior margin of the NUCH) in a sample of modern and fossil G. polyphemus adults. The strut is thought to reinforce the vertebral connection between the neck and body during “head bracing” behavior while tortoises dig with their front legs and stabilize their bodies with the hind legs (Bramble 1978, 1982). This strut (or scar) was present in all G. flavomarginatus we examined, but absent or nearly so in large samples (n=40) of modern G. agassizii and G. berlandieri. Thus, we consider the presence of strut scars as a strong indication that fossil populations dug extensive burrows similar to those as- sociated with modern G. polyphemus. We credit Dale R. Jackson who in the early 1970s originally suggested to us the importance of the bone scar character to sepa- rate fossil G. polyphemus from other Florida fossil turtles. We believe that the extensive encroachment of the strut (and its scar) onto the NUCH is a useful synapomorphy to define G. flavomarginatus, G. polyphemus, and other closely related fossil species. THE ZOOARCHAEOLOGICAL RECORD A total of 609 sites in the zooarchaeological data- base of the Environmental Archaeology laboratory at the Florida Museum of Natural History (UFea) were queried to determine the presence of Gopherus polyphemus (Table 4). This database is the largest of its kind presenting a temporal and geographic record of gopher tortoise identified in archaeological sites in the southeastern United States. Due to the heavy focus on Florida archaeology, the UF data are heavily weighted in favor of sites from Florida. GOPHER TORTOISES AS FOSSILS LATE PLIOCENE, FLORIDA (2.5-1.8 MA) Inglis 1A Local Fauna, Citrus Co.: The Inglis 1A sample includes 17 NUCH (UF 211285-211298, 211323) (Fig. 4), 7 right and 9 left EPI (UF 211324- 211339) (Fig. 6), 6 right and 8 left XIPH (UF 211340- 211350, 211401-211403), 4 right and 6 left HYO (UF 211405-211414), 1 right and 4 left HYPO (UF 211415- 211419), and numerous other shell fragments. No skull material was associated with the sample. We assume a MNI of 17, based on the number of NUCH bones. The sample consists of mostly small individuals with an esti- mated mean CL of 180.5 (using NUCH length) or 190.4 mm (using NUCH width) (Table 2). Growth annuli are visible on the HYO and HYPO bones. One HYO (UF 211412) has 11 prominent annuli plus the natal plate, sug- gesting an age of 12 years for this individual, based on known correspondence of annuli to age in modern indi- viduals; a second specimen (UF 211408) shows at least 7 or 8 annuli. The estimated age of Inglis 1A is about 1.9 Ma (Hulbert 2001). Inglis 1C Local Fauna, Citrus Co.: The UF sample from Inglis 1C includes two partial plastra (both from the left side) and a pair of costals with portions of the lower bridge attached (UF 211421). The most com- plete specimen in the UF sample is a reconstructed plas- tron that includes the left HYO and HYPO (UF 211284). These elements together measure 155 mm in length (measured as a straight line from EPI-HYO suture to the HYPO-XIPH suture). We estimate that this speci- men originally had a CL of 230 mm based on direct com- parison with a similarly-sized contemporary G. polyphemus from Alachua County (UFh 39639). The estimated length for this specimen falls within the upper range limits of the Inglis 1A sample. The second plas- tral specimen is an isolated HYPO (UF 211420)(59.8 mm along the midline suture). The estimated CL of this tortoise may have been larger than UF 211284. Neither of the two HYPO show the curvature associated with male plastral concavities, suggesting that both specimens are females. The plastral elements show growth annuli; however, only the annuli on one HYPO (UF 211284) were distinct enough to be counted (17 plus the natal plate). This suggests an age of 18 years for this speci- men, which would make it an adult female by modern standards. The SSH sample from Inglis 1C includes two an- terior parts of the carapace with NUCH and first dorsal vertebrae intact (Figs. 3, 7), 4 isolated NUCH, one par- tial plastron with both HYO, right side of lower plastron with HYPO and XIPH attached, and numerous carapacial and plastral bones and fragments. The ele- ments are identical to those from Inglis 1A, except the majority of bones are from slightly larger individuals. The Inglis 1A and 1C sites are considered of similar age (Ruez 2001), although 1C may be slightly younger (R. Hulbert pers. comm.). Inglis 1D Local Fauna, Citrus Co.: The site contained one NUCH and numerous carapace and plas- tral elements and fragments (SSH). Specimens repre- sent small individuals. Inglis 1F Local Fauna, Citrus Co.: Gopherus FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 186 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb is known from one HYO fragment (SSH). Waccasassa River 9A Local Fauna, Levy Co.: The sample includes a partial HYPO and several bridge elements (SSH). A late Pliocene age for the Waccasassaa River 9A local fauna is supported by the joint occurrence of Arctodus pristinus, Capromeryx arizonensis, and Hemiauchenia gracilis (UF collec- tion). EARLY PLEISTOCENE, FLORIDA (1.8-0.6 MA) Leisey Shell Pit 1A Local Fauna, Hillsborough Co.: The Leisey 1A sample of Gopherus consists of one partial NUCH (very small) (UF 144309), 12 carapacial elements, one associated pair of EPI (UF 80796), two left and one right HYO, five left and one right XIPH (UF 69394, 80458, 80796, 81063, 83601, 83602), one partial dentary (UF 144598), and one pha- lanx (UF 144599). Based on XIPH, the MNI of Gopherus represented at Leisey is five. Measurements from three left XIPH indicate CLs of 218-277 mm (Table 2). These estimated CLs are greater than those from the late Pliocene samples, but smaller than those from the late Pleistocene or modern samples. The fossils from Leisey 1A are considered to be about 800,000 years younger than those from Inglis. Haile 21A Local Fauna, Alachua Co.: Gopherus is represented by ten partial costals (e.g., UF 63616, 221261, 221851-221854), four neurals (UF 221095, 221303, 221354, 221355), a partial peripheral (UF 221054), and a PYG (UF 221355). Identification as Gopherus is based on bone shape and thickness of the specimens. MIDDLE PLEISTOCENE, FLORIDA (0.6-0.12 MA) Tri-Britton Local Fauna, Hendry Co.: The small sample includes one NUCH (UF 209990), several carapacial elements (costal, peripherals, and bridge ele- ments), one right EPI (UF 210862), one ENT (UF Figure 3. Internal view of the anterior carapace of Gopherus polyphemus from Inglis 1C, Citrus Co., Florida. Arrow points to the vertebral strut attached to the posterior edge of the nuchal bone (from the private collection of SSH). Figure 4. Nuchal bones of Gopherus cf. polyphemus from Inglis 1A, Citrus Co., Florida. A. UF 211288, internal view (smaller of two). Note the prominent bone scar on the under- side of the bone (A) from the attachment of the strut of the first dorsal vertebra. B. UF 211290 showing the shape of the nuchal scute in external view. 187 208986), two HYPO (UF 209990, 209098), and three XIPH (UF 209098, 209991). No skull material is repre- sented in the sample. The associated mammalian fauna indicates a late Irvingtonian age, ca. 0.3 to 0.5 Ma. Coleman 2A Local Fauna, Sumter Co.: This large sample includes 13 NUCH (UF 13390) (Fig. 5); five complete EPI (UF 13395a-c, e-f) (Fig. 6); 20 right EPI (UF13392a-t); and 14 left EPI (UF 13394a-m, UF 13395d); 18 PYG (including six with attached suprapygals) (UF 13402); two isolated first suprapygals; two paired XIPH (UF 13391a-b); 18 right XIPH (UF 13391c-e,13397, 13398); and 10 left XIPH (UF 13397, 13398). None of the isolated right and left EPI pre- cisely fitted together, suggesting a MNI of at least 39. No skull material was found in the sample. The esti- mated CL of tortoises in the Coleman sample ranged from 250-358 mm (Table 2). The associated mamma- lian fauna indicates a late Irvingtonian age, ca. 0.3 to 0.5 Ma. Sebastian Canal Local Fauna, Brevard Co.: Gopherus is represented by one peripheral (UF 12989). The associated mammalian fauna indicates a late Irvingtonian age, ca. 0.3 to 0.5 Ma. Haile 8A Local Fauna, Alachua Co.: The sample from Haile 8A contains one skull (UF 3147), a large number of nearly complete shells (UF 2988, 3127-3131, 3254, 3476-3478, 3786-3790, 3793-3794, 9435, 9575- 9582, 51610), 10 isolated NUCH (UF 3024, 3249, 3818, 3835-3839, 9436, 9583), two EPI (UF 9604), a first dor- sal vertebra (UF 9604), isolated PYG and suprapygal (UF 19021), XIPH (UF 3791), and assorted fragments (UF 3242, 3257, 9434, 9562, 9565, 9574, 9604, 9673- 9675, 9692). There were several neural bones with the first dorsal vertebrae in place. It was obvious that the anterior portion of the vertebrae had been attached to the NUCH in front of the neural. A sample of 28 indi- viduals ranged from 240-337 mm in CL, based on NUCH and XIPH measurements (Table 2). The associated mammalian fauna indicates an early Rancholabrean age, ca. 0.25 Ma. LATE PLEISTOCENE, FLORIDA (120,000-10,000 YR BP) Fossils of Gopherus polyphemus occur in 42 Florida sites of this age (Table 3). We have selected six localities as examples of late Pleistocene sites. Surprise Cave Local Fauna, Alachua Co.: The large sample consists of seven NUCH (UF 160156- 160157, 160208, 160209, 161885-161886, 161674), seven right and three left EPI (UF 160354, 161661-161663, 161914-161917, 161971, uncat.), ten right and six left HYO (UF 138001, 138002, 160334-160335, 161923, 161925, 161664, 161970, uncat.), 12 right and 14 left XIPH (UF 138002, 138060, 138063, 160207, 160332- 160333, 160336-160337, 160849, 161669-161670, 161735, 161788-161789, 161919-161922, uncat.), and several hundred shell and internal bone fragments (cat. and uncat. specimens). Two neural bones (UF 160855) have the first cervical vertebrae attached. We estimate a MNI of 14 based on left XIPH, with all being adults except for one very small individual. Estimated CL for a select sample of specimens ranged from 102-431 mm Figure 5. Nuchal bones of Gopherus polyphemus from Coleman 2A, Sumter Co., Florida. A. UF 13390B, internal view. Note the bone scar from the ventral strut. B. UF 13390A, showing the shape of the nuchal scute in external view. FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 188 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb Table 3. Fauna associates in sites with gopher tortoises. Other fossil tortoise species: Hesperotestudo (Caudochelys) crassiscutata, Hesperotestudo (Hesperotestudo) incisa, and Hesperotestudo (Hesperotestudo) mlynarski. The “Large species” from late Pliocene sites may represent an undescribed species of Hesperotestudo with a caudal buckler. An asterisk (*) denotes the presence of two Hesperotestudo species associated with gopher tortoises. Habitats: Xeric=Xe, Mesic hardwoods=Me, Pine flatwoods=Pf, Freshwater=Fw, Coastal marine=Cm. Use of parenthesis (Xe) indicates presumed xeric habitat based on the presence of Gopherus, but without other supporting associated xeric-adapted taxa. The genus name of Cnemdipohorus was recently changed to Aspidoscelis. Sites County Other Tortoises Upland Associates Habitats Florida Latest Pliocene *Inglis 1A Citrus Large sp. A H. cf. mlynarskii Rana capito, Geomys, Spilogale, Pituophis, Stilosoma, Heterodon nasicus Xe, Me, Pf, Fw Inglis 1C Citrus H. cf. mlynarskii Spilogale Xe, Fw Inglis 1D Citrus Spilogale Xe, Fw Inglis 1F Citrus Large sp. A Spilogale Xe, Fw *Waccasassa River 9A Levy Large sp. A H. cf. mlynarskii (Xe), Fw Early Pleistocene *Leisey 1A Hillsborough H. cf. crassiscutata Geomys, Podomys Xe, Me, H. cf. mlynarskii Fw, Cm *Haile 21A Alachua H. cf. crassiscutata Podomys Xe, Pf, Fw H. cf. mlynarskii Middle Pleistocene *Haile 8A Alachua Large sp. B Geomys Xe, Pf, Fw H. mlynarskii Sebastian Canal Brevard H. mlynarskii (Xe), Fw *Tri-Britton Hendry H. cf. crassiscutata Xe H. mlynarskii *LaBelle Highway Pit Hendry H. cf. crassiscutata (Xe), Fw H. mlynarskii *Coleman 2A Sumter H. cf. crassiscutata Geomys, Spilogale, Xe, Me, H. mlynarskii Pituophis, Heterodon Pf, Fw simus, Aspidoscelis, Scaphiopus Late Pleistocene Arredondo 1A Alachua H. crassiscutata Scaphiopus Xe, Me, Fw Arredondo 1B Alachua Geomys Xe, Fw Arredondo 1C Alachua H. crassiscutata (Xe), Fw *Arredondo 2A Alachua H. crassiscutata Geomys, Podomys, Xe, Me, H. incisa Scaphiopus, Stilosoma, Pf, Fw Heterodon simus Haile 1A Alachua H. crassiscutata Xe, Fw Haile 2A Alachua (Xe), Fw Haile 2D Alachua (Xe) Haile 11A Alachua H. incisa Geomys, Podomys Xe, Pf 189 Table 3. (Cont.) Sites County Other Tortoises Upland Associates Habitats *Haile 12A Alachua H. crassiscutata (Xe), Pf H. incisa Haile 13C Alachua H. crassiscutata Geomys Xe *Haile 14A Alachua H. crassiscutata Spilogale, Stilosoma, Xe, Pf, Fw H. incisa Aspidoscelis High Springs 1A Alachua (Xe) Hornsby Springs Alachua H. crassiscutata (Xe). Fw Kanapaha 1C Alachua H. crassiscutata (Xe), Fw Surprise Cave Alachua H. crassiscutata (Xe), Pf, Fw Wades Cave Alachua Geomys Xe, Pf Melbourne Brevard H. crassiscutata (Xe) Bone Cave Citrus H. crassiscutata (Xe), Fw Lecanto 2A Citrus H. crassiscutata Geomys, Thomomys, Xe, Me, Podomys, Scaphiopus Pf, Fw Sabertooth Cave Citrus Ichetucknee River Columbia H. crassiscutata Spilogale Xe Santa Fe 1 Columbia Santa Fe 2 Columbia H. crassiscutata Geomys Xe, Fw Santa Fe 8 Columbia H. crassiscutata (Xe), Fw Monkey Jungle Dade Podomys Xe, Me Nocatee Desota (Xe), Fw *Jacksonville Beach Duval H. crassiscutata (Xe), Fw, H. incisa Cm Vero Indian River H. crassiscutata Geomys, Peromyscus Xe, Fw, polionotus, Pituophis Cm *Devils Den Levy H. crassiscutata Geomys, Podomys, Xe H. incisa Spilogale, Pituophis, Heterodon simus, Scaphiopus Waccasassa 5A Levy H. crassiscutata (Xe) Williston 3A Levy Withlacoochee 7A Levy Eickelberger Cave Marion Geomys, Pituophis Xe, Me, Fw Kendrick 1A Marion Geomys, Spilogale Xe, Fw Medford Cave Marion H. crassiscutata Geomys, Heterodon simusXe, Me, Fw *Reddick 1A Marion H. crassiscutata H. incisa Geomys, Xe. Me, Podomys,Pituophis, Fw Aspidoscelis Reddick 1B Marion H. crassiscutata Pituophis, Heterodon Xe, Me, simus, Scaphiopus Fw Reddick 1C Marion (Xe), Me, Fw Reddick 1D Marion H. crassiscutata (Xe), Fw Reddick 1X Marion (Xe) Pratt Whitney Canal Palm Beach H. crassiscutata (Xe), Fw Seminole Field Pinellas (Xe), Fw FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 190 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb (based on XIPH, HYO, NUCH lengths and widths). Bone Cave Local Fauna, Citrus Co.: The sub- stantial sample includes 2 large NUCH (UF uncat.), 4 left and 3 right EPI (UF 6534, uncat.), 2 left XIPH (UF 6534, uncat.), one femur (UF 2256), a neural with at- tached first dorsal vertebra (UF uncat.), one anterior plastron with EPI (eroded) and ENT (UF2100), and many shell fragments (UF 6519, 6534, uncat.). Lecanto 2A Local Fauna., Citrus Co.: The sample includes one NUCH (UF 128231), two HYO and three peripherals (UF 128241-128245), a maxilla (UF 128235), and a series of cervical vertebrae, leg bones, and girdle pieces (UF 128294, 128230, 128238- 128244). Monkey Jungle Hammock Local Fauna, Dade Co.: One shell fragment (UF18708) is available. This Dade County site lies near Cutler Ridge, which is the southernmost locality for recent colonies on the Atlantic Coast. Reddick 1A Local Fauna, Marion Co.: The sample consists of a skull (UF 2401), three NUCH (UF 2461, 2706), EPI (UF 19066), XIPH (UF 19052), vari- ous other shell fragments (UF 2457, 2527, 2529, 2637, 2706) and a series of phalanges (UF 112190-112196). Carapacial fragments (UF 2527, 2529, 2706) are from large individuals. Wilson Quarry Local Fauna, St. Johns Co.: One peripheral (UF 11598) was found in a cemented marine shell hash associated with the Anastasia Formation, in- dicating a inshore origin, with the tortoise possibly origi- nating from the dune strand along a former coast line. Modern G. polyphemus currently occupy these habi- tats along the Atlantic coast, where they often live in large colonies. LATE PLEISTOCENE, GEORGIA Fragments of several fossil gopher tortoises have been found in coastal sites in Chatham County. A Charleston Museum specimen (ChMPV-1538) was probably picked from spoil piles along the Intracoastal Waterway (Al Sanders, Charleston Museum, pers. comm.). The Isle of Hope site lies “seaward (east) of Table 3. (Cont.) Sites County Other Tortoises Upland Associates Habitats Little Salt Springs Sarasota H. crassiscutata *Wilson Quarry St. Johns H. crassiscutata (Xe), Fw, H. incisa Cm Aucilla 3J Taylor H. crassiscutata (Xe), Cm St. Marks River Wakulla H. incisa Geomys Xe, Fw, Cm Georgia Late Pleistocene Savannah Chatham (Xe) Isle of Hope Chatham H. crassiscutata Pituophis Xe, Pf, H. incisa Fw, Cm Mississippi Late Pleistocene Catalpa Creek Lowndes H. crassiscutata (Xe), Fw South Carolina Late Pleistocene Charleston Charleston Edisto Beach Edisto H. crassiscutata (Xe) Myrtle Beach Horry (Xe) 191 the Pamlico barrier island complex.” The vertebrate fauna is associated with estuarine and neritic species of mollusks (Hulbert & Pratt 1998). The Isle of Hope fauna includes recent, as well as extinct, vertebrate taxa (e.g., Hesperotestudo crassiscutata, H. incisa, Equus, Mammut, and Mammuthus). Several scenarios on the possible depositional origin of this local fauna were dis- cussed by Hulbert and Pratt (1998). LATE PLEISTOCENE, MISSISSIPPI There is only one record of fossil Gopherus from Mississippi. This specimen is a water-worn XIPH from Catalpa Creek, near Columbus, Lowndes County (UF 23834). The site lies 190 km north of the nearest con- temporary locality for Gopherus in Mississippi (Wayne County), and 160 km northwest of the closest site in Alabama (Choctaw County). The specimen was col- lected in a stream deposit along with a number of other turtle species, including Hesperotestudo crassiscutata and the freshwater Macrochelys temminckii. LATE PLEISTOCENE, SOUTH CAROLINA Three fossil deposits, all late Pleistocene, contain remains of Gopherus polyphemus: Edisto Island (Charleston and Colleton counties.)(ChMVP-1537, 6414), and the Intracoastal Waterway site near Myrtle Beach (Horry County) (ChMPV-1539). The Horry County record was previously reported by Auffenberg and Franz (1978e, shown as star on range map). A fourth record for G. polyphemus was taken from an archaeological site near Myrtle Beach, where human transport may have been involved in its occurrence there (Jim Knight, South Carolina State Museum, Columbia, pers. comm.). The Charleston area and the Horry County sites lay 50 and 245 km, respectively, northeast of the most northern locations of extant populations in Hamp- ton and Jasper counties (Auffenberg & Franz 1982). Surprisingly, no Gopherus specimens are reported from the productive Ardis or Camelot sites (Jim Knight, pers. comm.). Both of these sites sample time periods when Gopherus was common in Florida. FOSSIL DISTRIBUTION AND THE ENVIRONMENT The fossil record for Gopherus polyphemus consists of 58 occurrences in Florida and six in three other south- eastern states (Table 3). The fossil and zooarchaeo- logical record is discontinuous during Plio-Pleistocene and Holocene times. All of the Florida fossil sites are restricted to the peninsula (east of Apalachicola River). Sites older than late Pleistocene account for only 20% of the sample and are limited to Alachua, Brevard, Cit- rus, Hendry, Hillsborough, Levy, and Sumter counties, Florida (Table 3). The oldest fossils are from Inglis 1A, 1C, 1D, and 1F in Citrus County, and Waccasassa River 9A in Levy County, Florida. We have yet to locate speci- mens from older late Blancan sites (ca. 2-2.5 Ma) in Florida, such as Haile 7C, Macasphalt Shell Pit, and Haile 15A. All of the Florida and Georgia records occur within the modern range of G. polyphemus. Records from Mississippi and South Carolina are extralimital, indicat- ing a slightly more northerly distribution in the south- eastern coastal plain in the late Pleistocene. The scar- city of Gopherus fossils from the early and middle Pleis- tocene may reflect a paucity of suitable habitats during this period, or simply the smaller pool of sites that repre- sent these ages. Conversely, their abundance in late Pleistocene faunas might indicate a time of expansion of xeric upland habitats and/or larger populations of tor- toises. Modern populations of Gopherus polyphemus are associated with upland longleaf pine (Pinus palustris)- oak uplands, dry oak-pine hammocks, scrubby pine flatwoods, and sand pine (Pinus clausa)-scrub oak ridges in the Southeast (Auffenberg & Franz 1982). Gopher tortoises also occur in ruderal settings, such as gardens, pastures, lawns, old fields, and road sides. Fire is demonstrably the most important natural force in the maintenance of modern gopher tortoise colonies and their xerophytic habitats. Fire acts to reduce the densities of woody species, remove leaf litter, and open the tree canopy (Myers 1994), which fosters light penetration and the growth of herbaceous forage species as well as creating areas of open soil used by tortoises for nesting. Often these fire-dominated pine forests have an open, savanna-like appearance with a continuous grassy and herbaceous low understory. We contend that pre-Columbian populations of gopher tortoises are similarly predisposed to droughty habitats and that they with other xeric-adapted special- ists common in Pleistocene deposits, e.g., Geomys, Spilogale, Podomys, Peromyscus (polionotus), Pituophis, Heterodon (nasicus/simus), Stilosoma, Aspidoscellis (=Cnemidophorus), Scaphiopus, and Rana (capito), are strong signatures for the presence of xerophytic pine landscapes in the Southeast in the past. We further suggest that these prehistoric habitats are similar, if not identical, to modern upland habitats and that frequent fires that regulate them today influ- enced them in the past. This implies, then, that contem- FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 192 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb FLMNH Site Name Site Designation # Cultural period and Date StateCounty Accession # Years BP x Litle Salt Spring 8So18 Late Pleistocene — ca. 11500 BP FloridaSarasota x Cutler Fossil Site 8DA2001 Early Archaic — ca. 8500 Florida Dade 0019 Tick Is. 8VO24 Middle Archaic — 7000-5000 BP Florida Volusia 0572 Lake Monroe Outlet Midden 8VO53 Middle Archaic — 7000-5000 BP Florida Volusia 0224 Palmer-Taylor 8SE18 Middle Archaic — 7000-5000 BP Florida 0025 Summer Haven 8SJ46 Late Archaic — 5000-2500 BP FloridaSt. Johns 0172 Boca Weir 8PB56 Late Archaic — 5000-2500 BP Florida Palm Beach 0447 Useppa Is. 8LL51 Late Archaic — 5000-2500 BP Florida Lee 0448 Horrs Is. 8CR209 Late Archaic — 5000-2500 BP FloridaCollier 0462 Cresent Beach Midden 8SJ43 Late Archaic — 5000-2500 BP FloridaSt. Johns 0587 Enclave Site 8PA1139 Late Archaic — 5000-2500 BP FloridaPasco 0296 Bay West Immokalee Archaic 8CR200 Archaic Florida Collier 0021 Cotton 8VO83 Archaic Florida Volusia 0160 Ft. Center 8GL13 ca. 2800—450 BP Florida Glades 0465 Piney Point 1 8NA3 ca. 2500-1650 BP Florida Nassau 0224a Alderman 8Vo135 St. Johns I - II — ca. 2400-1100 BP Florida Volusia 0511 Deer Island Causeway Site 8LA512 St. Johns I - ca. 2400-1800 BP FloridaLake 0188 Wightman Site 8L54 Glades I — ca. 2250-1850 BP Florida 0108 Key Marco 8Cr107 ca. 2350 BP Florida Collier 0179 Cumberland Is. 9Cam12 + 9Cam13Deptford — ca. 2450 BP Georgia Camden 0599 Brickell Point 8DA12 Glades I — ca. 1950-1600 BP FloridaDade 0254 Granada Site 8DA11 Glades I — ca. 1950-1450 BP FloridaDade 0294 Hontoon Is. 8VO202 St. Johns — ca. 1950 BP Florida Volusia 0131 Alexander Spring Bath House St. Johns II a — ca. 1850-450 BP Florida 0254 Granada Site 8DA11 Glades II — ca. 1450-750 BP FloridaDade 0142 Black Creek 1 8DA85 Glades II — ca. 1200-750 BP FloridaDade 0018 Jungerman 8BR136 St. Johns I — ca. 1200-640 BP Florida Brevard 0020 Goodman 8DU66A St. Johns I — ca. 1200-640 BP Florida Duval 0107 Palm Coast 8Fl15 St. Johns I — ca. 1200-640 BP FloridaFlagler 0024 Boynton Inlet 8 PB54 Glades II - III — ca. 1200-430 BP Florida Palm Beach 0254 Granada Site 8DA11 Glades II - III — ca. 1200-431 BP FloridaDade 0556 Remnant Mound (Shaw’s Point)8MA-7 Manasota — ca. 1055-540 BP FloridaManatee 0312 Jacksonville Electric Authority 8DU634/8DU669 Savannah — ca. 950-550 BP Florida Duval 0163 Maximo Point 8PI31 ca. 950-250 BP Florida 0254 Granada Site 8DA11 Glades IIIb — ca. 600-450 BP FloridaDade 0113 Melton 8A169 ca. 750-550 BP Florida Alachua 0294 Hontoon Is. 8VO202 St. Johns — ca. 450 BP Florida Volusia 0326 (429) Fig Spring’s 8CO1 ca. 450 BP Florida Columbia 0221 Baptizing Spr. 8Su65 Historic — ca. 400-300 BP Florida Suwanee 0222 Arrivas House 8SJ46 Historic — ca. 400-300 BP Florida St. Johns 0242 Palm Row 8SA 36-4 Historic — ca. 375-150 BP Florida St. Johns 0283 Ximenez-Fatio 19th C 8SA 34-2 Historic — ca. 375-150 BP Florida St. Johns 0518 DeBurgo-Pellicer SA 7-7 Historic — ca. 375-150 BP FloridaSt. Johns 0583 Ximenez-Fatio SA34-2 Historic — ca. 375-150 BP Florida St. Johns 0451 St. Francis Barracks 8 SA42A Historic — ca. 350-300 BP FloridaSt. Johns 0220 DeLeon 8SA26-1 Historic — ca. 300-present BP Florida St. Johns 0267 Useppa 8LL51 Historic — ca. 300 BP Florida Lee Table 4. The presence of Gopherus polyphemus in Zooarchaeological Assemblages from Florida and Georgia. 193 Table 4. (Cont.) porary longleaf pine-dominated scrubby flatwoods and sandhill, dry pine-oak hammocks, and sand pine-scrub oak ridge habitats have existed in the Southeast for at least two million years, the clock for modern G. polyphemus beginning with the fossils at Inglis and Waccasassa River. Most Florida sites with Gopherus include other herpetological species. Many of these taxa show a pro- pensity for specific habitat specialization. For example, Meylan (1982, 1984) listed 47 species (one salamander, 6 frogs, 7 turtles, 5 lizards, one amphisbaenid, 26 snakes, and the alligator) from the two-million-year-old Inglis 1A site. All of these species are present in modern Florida habitats except for seven extinct taxa and three extralimital species (Gerrhonotus sp., Liochlorophis vernalis, and Heterodon nasicus). By grouping these faunal elements, Meylan concluded that six modern Florida habitat types were represented at Inglis in the late Pliocene: longleaf pine, xeric hammocks, mesophytic hammocks, pine flatwoods, wetlands (swamps, marshes, and ponds), and rivers. He described the landscape as follows: “...the herpetofauna, other vertebrates, and sedi- mentary context suggest a mixed habitat of mature longleaf pine with xeric hammock interspersed. Xeric hammock would be expected in the depressions char- acteristic of a karst topography. That one or more of these depressions contained water at least seasonally...” (Meylan 1982:67). Following Meylan’s lead, we evaluated each of the Florida fossil sites that contained gopher tortoises (Table 3). We assumed that all 58 Florida sites had a xeric component based on the presence of gopher tortoises; 27 of these sites also included other xeric-adapted am- phibians (Scaphiopus holbrookii, Rana capito), rep- tiles (Aspidoscelis sexlineatus, Heterodon nasicus/ simus, Pituophis melanoleucus, Stilosoma extenuatum), and mammals (Geomys pinetis, Podomys sp., Peromyscus polionotus, and Spilogale putorius). Freshwater taxa were represented in 38 sites. Most of the aquatic taxa (e.g., Amphiuma, Alligator, Trachemys, Pseudemys, Nerodia, etc.) are characteristic of marsh and weedy pond habitats, most of which can be found in small wetlands in upland areas. Pine flatwoods and mesic (woodlands) taxa were found at 10 and 12 sites, respectively; coastal (estuarine) species occurred in only 6 sites. The presence of Rana capito at Inglis 1C and Podomys sp. at Leisey Shell Pit 1A and Haile 21A indi- cate that the typical commensal fauna associated with the burrows of modern gopher tortoises was already established by the late Pliocene and early Pleistocene, FLMNH Site Name Site Designation # Cultural period and Date StateCounty Accession # Years BP 0584 Segui/Kirby-Smith SA34-3 Historic — ca. 200-100 BP Florida St. Johns 0191 De Hita House 8SA7-4 Historic — ca. 230-187 BP Florida St. Johns 0227 De Mesa (Old Spanish Inn) 8SA 7-6 Historic — ca. 200-100 BP Florida St. Johns 0230 Acosta SA 13-5 Historic — ca. 200-100 BP Florida St. Johns 0234 De la Cruz 8SA 16-23 Historic — ca. 200-100 BP Florida St. Johns 0315 Ft. Matanzas 1980 exc. FOMA Historic — ca. 200 BP Florida St. Johns 0243b Kings Bay: Kings Bay 9CAM171 St. Simons - Historic PlantationGeorgia Camden 0243c Kings Bay: Plantation Site 9CAM172 Historic Plantation — ca. 200 BPGeorgia Camden 0243d Kings Bay 9CAM173 Historic Plantation Georgia Camden 0243e Kings Bay: Marianna Plantation 9CAM174 Historic Plantation Georgia Camden 0381 Sugar House 9Cam205 Historic - ca. 150 BP Georgia Camden 0422 Christ Church 8Es49 Historic — ca. 187-168 BP Florida Escambia 0430 Br. Commanding Officer’s 8Es1150 Historic — ca. 187-168 BP Florida Escambia 0243h Kings Bay: Araminta Sowerby Site9CAM178 Historic ca. 150 BP Georgia Camden 0313 Kingsley Plantation 8DU108 Historic — ca. 130-50 BP Florida Duval 0569 Segui/Kirby-Smith SA34-3 Historic — ca. 200-100 BP Florida St. Johns 0266 Castillo St. Augustine Historic Florida St. Johns FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 194 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb at least in Florida. All of the Plio-Pleistocene populations of Gopherus in Florida lived with elements of the extinct megafauna. Forty of the Gopherus sites include one or two extinct tortoise species in the genus Hesperotestudo (4 of 5 late Pliocene sites, 2 of 2 early Pleistocene sites, 5 of 5 middle Pleistocene sites, and 30 of 46 late Pleistocene sites) (Table 3). Gopherus was also found in associa- tion with Hesperotestudo at Isle of Hope (Georgia), Catalpa Creek (Mississippi), and Edisto Beach (South Carolina). One of the values of biotic reconstruction is that it provides an ecological framework in which to insert populations of extinct taxa. Examination of Table 3 shows that 20 Florida sites with Gopherus and Hesperotestudo had strong xeric species signatures (multiple upland species associates). Nine sites with Hesperotestudo had only terrestrial species represented in their faunas; 30 sites had a mixture of terrestrial plus freshwater species, which presumably meant that wa- ter was close by (habitat assessment not available for Little Salt Spring site). The presence of freshwater ap- pears to be the most important correlate that favors the presence of Hesperotestudo at any given sites. This is also true for many modern tortoise species, which are known to congregate at wet places. The relationship with water sources would assure proper water balance, water storage, and thermoregulation in more mesic- adapted tortoise species. The local distributions of mod- ern gopher tortoise populations are not known to track wetlands, probably because of their fossorial life style, which help them mitigate water and temperature stresses. Figure 7. Dorsal view of the anterior carapace of Gopherus cf. polyphemus from Inglis 1C, Citrus Co., Florida (from the pri- vate collection of SSH), showing the nuchal bone, first three neural bones, first three pairs of costals, and associated pe- ripheral bones. This is the most complete specimen from the Inglis series. Figure 6. Epiplastra of Gopherus polyphemus. A. UF 13395A, Coleman 2A, Sumter Co., Florida. Complete EPI showing prominent gular projection, deep excavation, and prominent lip. B. UF 211330, Inglis 1A, Citrus Co., Florida. Right EPI. C. UF 211329, Inglis 1A, Citrus Co., Florida. Right EPI (under- side). 195 WERE GOPHER TORTOISES LARGER IN THE PAST? Based on 109 measurements of 78 fossil shell elements, we estimate CLs for fossil gopher tortoises ranged from 102 mm at Surprise Cave to more than 400 mm at Reddick 1A and Surprise Cave (Table 2). Demographic studies of modern Gopherus polyphemus reported adult sizes (as CL) of 230-341, 238-368 mm (Auffenberg & Iverson 1979) and 232-291, 242-307, and 245-272 mm (Diemer 1992) for north Florida populations; and 335 mm (max. adult CL) (Landers et al 1982) for southwest Georgia. Auffenberg and Iverson (1979) and Diemer and Moore (1994) indicated 226 and 232 mm CL and 10-15 and 14- 18 years, respectively, for the minimum sizes and ages of females at first reproduction for north Florida popula- tions; Landers et al. (1982) listed 250-265 mm and 19- 21 years for southwest Georgia females. Size at hatch- ing is 49-52 mm CL (RF, pers. observ.). The largest CLs recorded for extant populations of this species are 368 mm (Conant & Collins 1998), 380 mm for a female from Citrus County, Florida (R. Ashton, pers. comm.), and 387 mm from Martin County, Florida (Timmerman & Roberts 1994). The largest fossils from Reddick 1A (401 mm based on NUCH length) and Surprise Cave (431 mm based on XIPH length) exceed the known maximum carapacial limits of modern G. polyphemus. Estimated CLs for 29 individual measurements from Reddick 1A, Coleman 2A, Haile 8A, and Surprise Cave are 300 mm or greater. A complete plastron, labeled as Gopherus praecedens (USNM 11999) from Melbourne, Brevard County, Florida, measured 335 mm along the midline suture (and 360 mm TL) indicating a CL of at least 360 mm. Mean CLs of samples from Coleman 2A (315 mm based on NUCH length), Haile 8A (316 mm based NUCH width), Reddick 1A (301 mm based NUCH length), and Sur- prise Cave (312 mm based on XIPH length) are 300 mm or greater. These data suggest that G. polyphemus frequently attained larger body sizes during the Pleis- tocene than in most modern populations today. Many of these Pleistocene individuals rivaled or exceeded the maximum sizes known for modern G. flavomarginatus (371 mm CL listed in Legler & Webb 1961), which is considered the largest living species of Gopherus. The entire sample from Inglis 1A and Inglis 1C contains only small individuals, most falling near or be- low the minimum size for sexual maturity expressed in modern female Gopherus polyphemus (Table 2). Samples from Inglis 1D, 1F, and Waccasassa River 9A (SSH) also consist of smaller individuals. Together, these samples suggest a small body size for all late Pliocene populations of Gopherus in Florida. Several explana- tions are possible to explain this size discrepancy: (1) the samples represent a yet unrecognized species or subspecies of small gopher tortoises, (2) some physical attribute of the site caused the selective preservation of only small individuals, or (3) certain ecological factors, such as selective predation, disease, several years of successful recruitment, or some catastrophic event, caused a temporary shift in the size (and age) structure of the population. Any of these scenarios are possible. The first scenario has the greatest merit given the smaller body sizes of the entire late Pliocene gopher tortoise sample. It remains possible that this population could eventually be recognized as distinct upon a more com- prehensive review (see above). The second scenario is more remote since faunal samples include extinct large tortoises (Hesperotestudo) and other megafaunal spe- cies. The ecological scenario remains plausible, although the causal agent is not readily apparent. This last expla- nation also implies that the samples were deposited dur- ing a very short time period. GOPHER TORTOISES IN THE ARCHAEOLOGICAL RECORD ZOOARCHAEOLOGICAL DATA The temporal and geographic distribution of Gopherus polyphemus in the zooarchaeological record of Florida and Georgia is presented in Table 4 and Fig- ure 2. We identified 67 dateable sites from 20 Florida counties and one county in Georgia containing Gopherus polyphemus remains. This accounts for approximately 12,000 years of human history and human culture in vary- ing stages of complexity; from fisher, hunter, and gath- erers to the 19th Century. Late Pleistocene: The first humans arrived in southeastern North America during the late Pleistocene, ca. 12,000 yrBP. The climate was cooler and drier and the late Pleistocene sea was well below its current level (Davis 1997; Randazo & Jones 1997). It is for this lat- ter reason that most of the zooarchaeological record of these earliest people is difficult to document. Most of their living sites were inundated by the rising Holocene sea, taking with it well preserved and intact midden de- posits. Some late Pleistocene sites are found along Florida’s rivers and springs. The zooarchaeological evi- dence of these locations tends to be rare relative to other archaeological periods, thus making their documenta- tion difficult. Little Salt Spring represents one such site that contains the remains of gopher tortoise and FRANZ and QUITMYER: History of the Eastern Gopher Tortoise 196 CENOZOIC VERTEBRATES: Paper to Honor S. David Webb Hesperotestudo crassiscutata (Clausen et al. 1979; Holman & Clausen 1984). It is unclear if these animals were the remains of subsistence resources or whether they were animals living around the site when humans occupied the region. Nonetheless, Little Salt Spring ap- pears to be the only locality in which the gopher tortoise is found in association with late Pleistocene period hu- mans. Early Archaic Period (10,000-8,000 BP): The early Archaic period residents of the southeastern United States faced a rapidly changing environment. The cli- mate was still cooler and dryer, and sea level lower than at present (Davis 1997). By the time the Cutler fossil (Table 4; Fig. 2) site was occupied by early Archaic people, the Pleistocene megafauna (e.g., mammoth, mastodon, giant sloth, and Hesperotestudo crassiscutata) was extinct. Humans were subsisting on smaller animals, and gopher tortoises were among the zooarchaeological remains identified from the Cut- ler site (Table 4; Fig. 2). Middle Archaic Period (8,000-7,000 BP): The archaeology of Florida indicates that humans had be- come more sedentary and their populations more nu- merous by the middle Archaic period (Russo 1991). Fish- ing, shellfishing, and gathering were the primary subsis- tence activities, while hunting of upland game was of secondary importance. Gopher tortoise remains are rep- resented in two middle Archaic period sites along the St. Johns River and one site in southwest Florida (Table 4; Fig. 2). Late Archaic Period (5,000-2,500 BP): Envi- ronmental conditions of the late Archaic Period had ap- proached modern conditions. Permanent settlements continued to be established and human populations con- tinued to grow. By ca. 4,000 BP, fiber-tempered ce- ramics were invented along the coast of the southeast- ern United States. Subsistence practices resembled those established in the middle Archaic period. Gopher tor- toise remains are known from six zooarchaeological as- semblages in six Florida counties (Table 4; Fig. 2) Woodland and Mississippian Period to Euro- pean Contact (2,800-450 BP): Table 4 and Figure 2 presents a list with remains known from 24 sites in 13 Florida and Georgia counties. These data show that gopher tortoises were clearly represented by a tempo- rally and geographically diverse group of archaeological cultures. Nonetheless, aquatic (marine and freshwater) resources were the focus of these people, whereas up- land game, such as gopher tortoise, was of secondary importance. Historic Period in Florida and Georgia (ca. A.D. 1513 – 19th Century): The date A.D. 1513 marks the discovery of Florida by Juan Ponce de Leon and repre- sents the start of European colonization of southeastern North America. The subsistence record from Florida and Georgia show a clear association of gopher tortoises with historic Native American people, early Spanish and British colonists, and plantations of the 18th and 19th cen- turies (Table 4; Fig. 2). THE ASSOCIATION OF GOPHER TORTOISES AND HUMANS As we indicated above, gopher tortoises are xeric habi- tat specialists. Fire is the most important natural vari- able in the maintenance of their populations. However, the appearance of humans near the end of the Pleis- tocene presented another important variable that affected and continues to affect gopher tortoise populations. It is known that pronounced anthropogenic alteration of the environment may occur with the incursion of pre-indus- trial peoples into a region (Rambo 1985; Redford 1990; Steadman 1995; Quitmyer & Jones 2000; Wing 2001). Such changes in the environment may not necessarily be a consequence of cultural complexity, but rather the interaction of human behavior with the environment. The rudiments of environmental degradation probably ap- peared earlier in human history than most scientists have realized (Jackson et al. 2000). Jackson et al. (2000) suggested that the deleterious effect on various aspects of the environment have been a prolonged phenomenon that started with pre-industrial people. Habitat destruction has had a clear influence on modern gopher tortoise populations, while in some cases gopher tortoise habitat can be created by human distur- bances of the environment, such as in gardens, pastures, lawns, old-fields, and roadsides (Auffenberg & Franz 1982). Linares (1976) reported a similar pattern for the American tropics where there is a positive correlation in the zooarchaelogical record between the appearance of pre-Columbian gardening and the increased frequency of animals that are attracted to garden plots where they were hunted and consumed. The mid-Holocene archaeological record of the southeastern United States indicates that humans were giving up their mobile hunter-gatherer strategies for a sedentary lifestyle (Russo 1991; Quitmyer et al. 1997). The zooarchaeological evidence shows that the adop- tion of a sedentary life style along the coastal plain was supported by the exploitation of marine and aquatic re- 197 sources (Reitz & Quitmyer 1988; Reitz & Wing 1999; Quitmyer & Massaro 1999; Quitmyer 2001, 2002). Sedentism would have contributed to greater disturbance of local environments. Gopher tortoise remains in the zooarchaeological record of most post mid-Holocene sites is typically a minor component of the overall midden assemblages (Larson 1980), a pattern that does not ap- pear to change for nearly 12,000 years of human his- tory. One explanation for their low numbers in faunal assemblages is that gopher tortoises were only opportu- nistically exploited during the course of everyday sub- sistence activity (Larson 1980). It is also possible that gopher tortoise habitat was created by disturbance around human living sites (e.g., reduction of fuel woods and land clearing). These scenarios remain a subject of fur- ther research, but it is clear that there has been an asso- ciation of humans and gopher tortoises since the end of the Pleistocene (Table 4). CONCLUSIONS The data for 64 fossil and 67 zooarchaeological sites show that Gopherus polyphemus has been part of the southeastern coastal plain fauna for about two million years. This tortoise first appeared in the Inglis and Waccasassa 9A local faunas in the late Pliocene. Its fossils are known from relatively few sites in the early and middle Pleistocene. The samples are more numer- ous in the late Pleistocene. We contend that gopher tortoises and other associated xeric-adapted species found in these sites are strong signatures for the pres- ence of longleaf pine sandhills, oak-pine hammocks, and sand pine-scrub oak ridges during the late Pliocene and Pleistocene. Presumably, as in the case today, frequent fires strongly influenced the aspect and species compo- sition of these pine landscapes. The fossil record also indicates that gopher tortoises lived with extinct portions of the Pleistocene megafauna, including extinct tortoises of the genus Hesperotestudo. Gopher tortoises and Hesperotestudo occurred together at 42 (68%) (Florida + other states) fossil sites. The record also suggests that earliest populations of gopher tortoises were rela- tively smaller than the middle-late Pleistocene or extant populations. The largest individuals in the Pleistocene samples rivaled or exceeded the CLs of their extant Mexican relative, Gopherus flavomarginatus, which traditionally has been thought to be larger. Near the end of the Pleistocene, humans entered the landscape of the southeastern United States, and this corresponds to the extinction of a variety of animals such as the horse, mammoth, mastodon, and tortoises of the genus Hesperotestudo. The zooarchaeological records from Georgia and Florida show that humans and gopher tortoises have remained in associated with each other for 12,000 years. The data indicate that during this time gopher tortoises represented a regular food item, albeit a minor one. The development of agriculture within native American cultures may have led to increased gopher tortoise populations accompanying disturbance from forest clearing, crop planting, and abandonment of food plots. ACKNOWLEDGEMENTS We are indebted to Richard Hulbert who weathered our most elementary questions about the Florida Museum of Natural History vertebrate paleontology collection (UF/FLMNH) and site information. We also wish to thank Walter Auffenberg and Dave Webb for many hours of tortoise chats; Bob Chandler (Georgia State College and University), Richard Hulbert (UF), Steve and Suzan Hutchens (SSH), F. Wayne King, Kenny Krysko, and Kitty F. Emery (Florida Museum), Jim Knight (South Carolina State Museum), and Al Sanders (Charleston Museum) for allowing us access to their collections; Joan Berish, Richard Hulbert, Dale R. Jack- son, Bob McCord, and Dave Steadman who reviewed various stages of the manuscript; Jeff Gaige for the pho- tographs; and Shelley Franz as traveling companion and keeper of our many miscellaneous notes about fossil tor- toises. We thank Bill Paine, Brian Worthington, Catherine Liu and Erin Thorton for their assistance in assembling the zooarchaeological data. 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