"Bit. .2 .1 .2113: {5:131 ‘IV I r.u«?L. — , :u‘éin 1211.3! . I '3. . ‘ . £31.31 .. amen?“ a Ih‘bg'firiqglgn‘ uh”... 2.14 .2... ,r 1333......" > v 5) iv .x. "K await... .11... u: i .1) II: 239...}? ,. .2 Luann LIBRARY Michigan State Ur IlVb'lSlly This is to certify that the thesis entitled BEHAVIORAL AND ECOLOGICAL STUDY OF THE SPOTTED TURTLE, CLEMMYS GUTTATA (SCHNEIDER) presented by DIANA JANE LUTZ has been accepted towards fulfillment of the requirements for the MS. degree in ZOOLOGY / ° / [11.144-I1'//11-“"/" ' Major ‘ ofe 3Vure 01/26/ / 02 Date MSU is an Affinnative Action/Equal Opportunity Employer —A—.—.-o-I-I-I-o-l-e-O- PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINES return on or before date due. MAY BE RECALLED with earlier due date if requested. DATE DUE DATE DUE DATE DUE 5/08 KIProj/Acc8Pres/ClRC/DateDue indd BEHAVIORAL AND ECOLOGICAL STUDY OF THE SPOTTED TURTLE, CLEMMYS GUTTATA (SCHNEIDER) By Diana Jane Lutz A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Zoology 2009 ABSTRACT BEHAVIORAL AND ECOLOGICAL STUDY OF THE SPOTTED TURTLE, CLEMMYS GUTTATA (SCHNEIDER) By Diana Jane Lutz The specific objectives of this study were to determine habitat requirements, movement patterns, population dynamics, demography and seasonal activities of the Spotted Turtle, Clemmys guffata in a southwestern Michigan fen. Habitat areas were identified with lower fen elevation areas which contained low-growing hydrophytes preferred. Population overlapping and homing were found to occur, with male home ranges larger than female home ranges. Population size appeared to be a 1:1 sex ratio. Morphometric measurements and male and female comparisons are presented, with females shown larger than males. Aestivation was found to occur. Communal hibemaculae and site fidelity also occurred with preferred hibemaculae habitats descfibed. ACKNOWLEDGEMENTS My passion and love for turtles started at a very early age. Every waking minute growing up at Rose Lake (in Branch County, MI) was spent catching, observing and then releasing turtles. This passion has never stopped-nor will it ever. Now that I am older, being able to spend my time working and studying these amazing creatures is a dream come true. I would like to express a big thank you to my committee members. With different areas of expertise and specializations, they were able to provide me with the tools to create and achieve a successful research project. I would like to thank Dr. Richard Snider, my advisor, for his help in providing me with the opportunity to “do my turtle research” at MSU. A special thanks to Jim Harding, who was instrumental in finding the location for my research and encouraging me throughout my education as a “non-traditional” student. The support and guidance given to me by Dr. Kelly Millenbah was greatly appreciated. Dr. J. Whitfield Gibbons-thanks for taking on a long distance student and taking the time to visit the project and being involved. I am most appreciative to Melinda Freeland. Melinda’s help over the last few years has been immeasurable. Her computer knowledge is like no others, as well as her science, math, and biology skills. She is a “natural” when it comes to teaching. The quality of my research and the final Thesis is due to her input. The endless hours (and days), we have spent together has blossomed into a true and cherished friendship. She has more patience than any person I have known. iii A special thanks to Dave Mifsud for his enormous help with the Arcview program. The maps created provided invaluable information about these animals. I owe you more brownies. A big thank you to Paul Dziepak, for his help in the research field. Thank you for allowing me the opportunity to study Spotties in your area. I appreciate all your work and knowledge in the field, with transmitters and telemetry. Thanks for all your input and for being such a good friend. Additionally, I would like to thank those who helped with field work. Thank you to Pete Wilson, Jerry Stampsky, and Jim and Harrison Barzyk. It is fun sharing days in the field with kindred spirits. Thanks also, to Dr. Thomas Burton for his help in the “bug lab” and for his insight with my project. A big thank you to my great friends, and fellow students, Edi Sonntag and Cole Provence. Edi, thanks for teaching me what is expected in the field of a real “techie”. Cole, I appreciate your field and lab help! Jeff Schier, thank you for all your late night assistance in the early days of returning to school. I would like to thank the Conservancy and the private landowners for allowing me to research the area. Thanks to Michigan State University, Michigan Society of Herpetologists and Chicago Herpetological Society for their financial suppon. Lastly, I would like to thank my family for their support throughout my college years. Thanks to Dece and Fred Lyons; Rindi, Mark, Brady and Taylor Baildon; Jodi, Bobby, Kylee and Jordan Dooley; Alice Conrad and Karin iv Kennedy. Special thanks and love to J.T., Lisonn, Addalynn and Dominic Delcamp. Sorry, I made you all so crazy. I love you! Aunt Jane Hurley, thank you for allowing me to grow up and spend so much time with you at Rose Lake, and for all our “turtle hunting” trips! Thanks for all your love and support. Most of all, I thank my mom, Helen Conrad. Thanks to her, I grew up with the appreciation for, and knowledge of, the wildlife all around me. She is the reason I returned to school-she was my biggest supporter and my biggest cheerleader! I cherish a small framed picture that was my mom’s. It reads: “The goal of life is to live in agreement with nature”. I’m sure trying..... TABLE OF CONTENTS LIST OF TABLES .............................................................................. vii LIST OF FIGURES ............................................................................ viii CHAPTER 1: HABITAT AND MOVEMENT OF CLEMMYS GUTTATA ............ 1 CHAPTER 2: POPULATION DYNAMICS AND DEMOGRAPHY ................... 49 CHAPTER 3: SEASONAL ACTIVITIES ................................................... 55 APPENDIX ....................................................................................... 69 LITERATURE CITED ......................................................................... 144 vi LIST OF TABLES Table 1. Specific habitat preferences documented throughout Spotted Turtle range. ......................................................... 70 Table 2. Non-woody plants located in southwestern Michigan fen. 71 Table 3. Woody plants located in southwestern Michigan fen. ...................... 73 Table 4. Wildlife documented in southwestern Michigan fen. 74 Table 5. Amphibians and reptiles documented in southwestern Michigan fen. ................................................................... 74 Table 6. Bird species present in southwestern Michigan fen. ........................ 75 Table 7. Example of Microsoft Excel individual turtle data worksheet. ............ 77 Table 8. Example of the Microsoft Excel radio telemetry data sheet. .............. 78 Table 9. Compilation of 2007 and 2008 data for the 187 turtles of southwestern Michigan fen. 79 vii Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6. Figure 7. Figure 8. Figure 9. Figure 10 Figure 11 Figure 12 Figure 13 Figure 14 Figure 15 Figure 16 Figure 17 Figure 18 Figure 19 Figure 20 Figure 21 LIST OF FIGURES Presettlement land cover of southwestern Michigan fen. ................ 84 Outlying regions of southwestern Michigan fen. ........................... 85 Open wetland in middle of fen. ................................................ 86 Thick growth of Chara in study site Chara pond. .......................... 87 Large transmitter attached to turtle carapace. ............................. 88 Alphabetical turtle marking system diagram. .............................. 89 Diagram of sample markings for turtle ABX. ............................... 90 Turtle with notched markings. ................................................. 91 Small transmitter attached to carapace. .................................... 92 . Labeled diagram of study site. 93 . Compilation of habitat utilization for transmitter turtles 2007. ......... 94 . Compilation of habitat utilizations for turtles without transmitters in 2007. .......................................................................... 95 . Compilation of habitat utilizations for transmitter turtles in 2008. ......96 . Compilation of habitat utilizations for turtles without transmitters in 2008. .......................................................................... 97 . Predated Nesting Site. .......................................................... 98 . Hatchling found in successful nesting location. ........................... 98 . Channel 0 male IKL habitat utilization in 2007. ............................ 99 . Channel 1 male KVW habitat utilization in 2007. ....................... 100 . Channel 2 male ABCK habitat utilization in 2007. ....................... 101 . Channel 3 female KVX habitat utilization in 2007. ....................... 102 . Channel 4 female HKU habitat utilization in 2007. ...................... 103 viii Figure 22 Figure 23 Figure 24 Figure 25 Figure 26 Figure 27 Figure 28 Figure 29 Figure 30 Figure 31. Figure 32. Figure 33. Figure 34. Figure 35. Figure 36. Figure 37. Figure 38. Figure 39. Figure 40 Figure 41 Figure 42 Figure 43 Figure 44 . Channel 6 female DKVWX habitat utilization in 2007. ................. 104 . Channel 7 female KLU habitat utilization in 2007. ....................... 105 . Channel 8 female HKV habitat utilization in 2007. ....................... 106 . Channel 10 male IKW habitat utilization in 2007. ....................... 107 . Channel 11 female KV habitat utilization in 2007. ...................... 108 . Channel 1 female HJKV habitat utilization in 2008. ..................... 109 . Channel 2 male KUV habitat utilization in 2008. ........................ 110 . Channel 3 female HJK habitat utilization in 2008. ....................... 111 . Channel 4 female ABKW habitat utilization in 2008. .................... 112 Channel 5 female CHJKLVW habitat utilization in 2008. ............. 113 Channel 6 female BDKW habitat utilization in 2008. ................... 114 Channel 7 female AKX habitat utilization in 2008. ...................... 115 Channel 8 female FKS habitat utilization in 2008. ...................... 116 Channel 9 female BKW habitat utilization in 2008. ..................... 117 Channel 11 female CKU habitat utilization in 2008. .................... 118 Channel 12 female ABCKW habitat utilization in 2008. ............... 119 Channel 13 female DKP habitat utilization in 2008. .................... 120 Channel 14 female BHK habitat utilization in 2008. .................... 121 . Channel 15 female ADKO habitat utilization in 2008. .................. 122 . Channel 16 male ABCK habitat utilization in 2008. ..................... 123 . Channel 17 male IKW habitat utilization in 2008. ....................... 124 . Habitat utilization for male ABCK in 2007 and 2008. .................. 125 . Habitat utilization for male IKW in 2007 and 2008. ..................... 126 ix Figure 45. Overlapping areas of all females tracked in 2007. ..................... 127 Figure 46. Overlapping areas of all males tracked in 2007. ........................ 128 Figure 47. Channel 1 overlapping with all transmittered females during 2007. ................................................................... 129 Figure 48. Male overlapping home ranges in 2008. .................................. 130 Figure 49. Female overlapping home ranges in 2008. .............................. 131 Figure 50. Picture of Sphagnum spp. moss mounds. ................................ 132 Figure 51. C. gulfata basking on top of sedge on Sphagnum spp. moss mounds. ........................................................................ 132 Figure 52. Channel 4 aestivation and hibernation locale. ........................... 133 Figure 53. Fallen Shrubby Cinquefoil petals with comparison Spotted Turtle. ................................................................ 134 Figure 54. Fallen Shrubby Cinquefoil petals. ........................................... 134 Figure 55. Deer trails in the study site. ................................................... 135 Figure 56. Total utilization area for all turtles in 2007 and 2008. .................. 136 Figure 57. 2007 and 2008 Spotted Turtle sex distribution. ......................... 137 Figure 58. Main aggregations of Spotted Turtles during 2007. .................... 138 Figure 59. Main aggregations of Spotted Turtles during 2008. .................... 139 Figure 60. 25 September 2007 possible hibemaculum. ............................. 140 Figure 61. 2 October 2007 possible hibemaculum. .................................... 141 Figure 62. 12 October 2007 possible hibemaculum. .................................. 142 Figure 63. 16 October 2007 possible hibemaculum. ................................. 143 *lmages in this thesis are presented in color. Chapter 1: Habitat and Movement of Clemmys guttata (Schneider) Introduction The Spotted Turtle, Clemmys guttata (Schneider), is a semi-aquatic North American turtle in the family Emydidae. Spotted Turtles are endemic to eastern North America and range from southern Great Lakes region (Illinois, lower Michigan, Indiana, Ohio, southern Ontario) east through Pennsylvania and New York to southern Quebec, and New England, and southward along the eastern seaboard (piedmont and coastal plain east of the Appalachian Mountains) to northern Florida (Ernst etal, 1994; Conant and Collins, 1991). C. guttata is a small turtle with a smooth, keeless, low, oval black/brown carapace overlaid with small rounded yellow spots. The carapace is widest at the posterior. Occasionally appearing rust colored from staining, which may result from dissolved tannins or iron deposits. Spot number varies with some specimens lacking spots, while others have up to 125 scattered across the carapace and face (Roach, 2006). The hingeless, orangish-yellow plastron may display variable black/brown blotching. Secondary sexual characteristics are usually marked: males typically with tan chins, brown eyes, slightly concave plastra, and long, thick tails with vent beyond posterior carapacial rim. Females with yellow chins, orange eyes, flat or convex plastra, and shorter, thinner tails with vent beneath posterior marginals (Blake, 1922; Roach, 2006). In both sexes, heads mostly black with usually a few yellow spots and laterally one or more irregular orange or yellow blotch. The outer surfaces of legs are black, usually with a few yellow spots, while the lower leg surfaces, neck and other soft parts often orange or pinkish mottled with black (Harding, 1997). At hatching, the blue-black young Spotted Turtle is about 2.8 cm long. Carr (1952) reports that its width may be up to 95% of its length, making it appear almost round. Coloration that of the adult, but with usually one yellow spot per carapace scute; although, initially, some hatchlings may lack carapace spots. The head always spotted, with possible neck spotting (Ernst et al, 1994). Plastron yellowish-orange with central dark blotch (Harding, 1997). Hatchling’s tail proportionally longer than adult. The egg tooth (caruncle) drops off by end of the first week (Ernst et al, 1994). Unfortunately, Spotted Turtles are small and colorful, as well as shy, timid creatures with an easy-going disposition, exhibiting no trace of aggressiveness, thus creating a high demand as an aquarium or terrarium pet (Cahn, 1937). Roach (2006) describes the Spotted Turtle as, one of God’s greatest creations; one of the prettiest turtles on the face of the earth.” This kind of testament contributes to its over-collection and pet trade popularity. In addition, overgrazing (Minton, 1972), agricultural equipment such as plows, excavators (Fowle, 2001) and mowers (Ernst, 1976), vandalistic shooting (Harding, 1997), predation of individuals and nestlings by raccoons (Procyon Iota!) and skunks (Mephitis mephitis) (Ernst, 1976; Cahn, 1937), along with road-crossing mortalities (Ernst, 1976) are additional major factors impacting population decline. Muskrat (Ondatra zibethica) predation has also been recorded in several populations of Spotted Turtles (Grant, 1936; Nemuras, 1966). Based on the slow growth rate, delayed maturity, small clutch size, and low egg and juvenile survivorship, the species is considered especially vulnerable (Wilson etal, 1999; Harding, 1997). These factors further exacerbate local population susceptibility extinctions (Oldham, 1991). C. guttata research has been conducted in other areas of its range, but remains poorly studied in Michigan. Prior to this study little has been documented on the Spotted Turtle in Michigan (Ruthven etal, 1928; Harding, 1997; Lutz, 2008). C. guttata studies and research have been conducted in Ontario (Litzgus etal, 1999; Litzgus, 1996; Chippindale, 1989; Haxton and Berrill, 2001), Maine (Joyal etal, 2001), Maryland (Nemuras, 1966; Ward etal, 1976), Massachusetts (Fowle, 2001; Graham, 1995), South Carolina (Lovich, 1990; Litzgus and Mousseau, 2004), Georgia (Folkerts and Skorepa, 1967), Connecticut (Perillo, 1997), Florida (Bamwell etal, 1997; Berry, 1978), Pennsylvania (Ernst and Zug, 1994; Ernst, 1970; Ernst, 1975), Ohio (Lewis and Ritzenthaler, 1997), Indiana (Minton, 1972) and Illinois (Cahn, 1937). In the United States, the Spotted Turtle is not currently listed under the US. Federal Endangered Species Act; however, it is listed as endangered and threatened where it occurs in several states and Canada. Nature Serve’s Conservation Status Ranks classify the species status anywhere from 85 (most secure) to SI (critically imperiled). In Michigan, not only is the Spotted Turtle threatened, but it is listed as 32, meaning imperiled (six to twenty occurrences or restricted range) (www.natureserve.org) helm Previous Studies Habitat requirements vary depending on range and topographic conditions. General habitat preferences include shallow ponds, wet meadows, tamarack swamps, bogs, fens, marsh channels, Sphagnum spp. seepages, slow streams and clear shallow water with mud or muck bottom and ample aquatic and emergent vegetation (Harding, 1997). Habitat preferences documented by other researchers are compiled in Table 1. Little data has been documented on preferred Michigan habitat requirements. Females typically produce one, occasionally two clutches, of one to eight eggs per year (Ernst, 1970; Wilson, 1989, Ernst and Zug, 1994; Litzgus and Brooks, 1998). Besides some nests being dug into Sedge spp. tussocks (Ernst, 2001 ), nest sites include hummocks of moist Sphagnum moss and loamy soil of marshy pastures (Ernst, 1970; Belmore, 1980; Chippindale, 1989). C. guttata is comparatively cold tolerant, actually preferring cooler environments (Ernst, 1982; Nemuras, 1966). It generally emerges earlier in spring, than other turtle species, often as soon as the snow cover melts (Ernst, 1982). It is considered most active in cool, early spring (Ward et al, 1976; Lovich, 1988; Litzgus and Brooks, 2000). Unlike other turtle species, the Spotted Turtle does not tolerate heat, with inactivity observed when an average mean temperature of 203°C is reached (Ernst et al, 1994). Basking for long lengths of time is also minimized with increased temperatures. Being heliotherrnic, it can often be observed first thing in the morning basking in the sun, but will disappear into the muck, mud or vegetation as the temperature begins to increase. Later, as temperatures fall, the turtle may once again emerge to bask. Spotted Turtles are specialized in habitat preferences. Habitat choice may vary in different portions of its range. Unpolluted, shallow, mud and muck bottomed water bodies, such as marshes, bogs, swamps, small streams, drainage ditches, and vernal ponds (Ernst and Zug, 1994; Graham, 1995; Ditmars, 1933; Harding, 1997) are preferred. Other habitats may include fens (Lewis and Ritzenthaler, 1997; Lovich, 1987), grassy areas (Ward etal, 1976; Ditmars, 1933) and terrestrial environments (Berry, 1978; Fowle, 2001). Many factors have led to decline of C. guttata populations correlating to their S2 ranking. Habitat destruction caused by development, marshland drainage, pollution, and fragmentation (roads, fences, curbs, railroad tracks, and retaining walls) are a few documented reasons for reported declines (Lovich and Jaworski, 1988; Cook etal, 1980; Harding, 1997; Conant and Collins, 1991; Fowle, 2001). Introduction of invasive plant species (Conant and Collins, 1991; COSEWIC, 2004) also plays a major role in habitat destruction and is becoming a major threat to turtle populations. Once a habitat becomes overgrown with later successional plant species, it may be unsuitable for Spotted Turtles (Burke etal, 2000; Graham, 1995). These invasives include, but are not limited to, Glossy Buckthom (Fihamnus frangula), Phragmites (Phragmites australis), and Purple Loosestrife (Lythrum salicaria). Movement Patterns Previous Studies Cagle (1944) defines three major turtle movement categories: first, local activity resulting from food foraging, seeking basking sites, or mating impulse; second, seasonal migrations; third, irregular periods of migration. Important studies in Pennsylvania of C. guttata, were conducted by Ernst (1976), who thought the annual activity cycle of C. guttata was apparently controlled by two major factors: water temperature and reproductive drive. Water temperature influenced all of the normal activities such as feeding, basking, and dormancy, and also possibly limited reproductive activity when too low. At normal seasonal temperatures in his study site, reproductive drive controlled many spring activities of C. guttata. Ernst (1967) also reported that feeding does not occur until water temperatures reach 14°C. Active feeding continued as long as water temperature remained above this temperature. Initial activity after cold weather dormancy was spent mostly basking to gain heat and maintain relatively high body temperatures. Ernst (1976) found daily periods of basking and foraging varied in length depending upon environmental conditions. In periods of cool weather, Spotted Turtles either bask for most of the day, and feed only sparingly or become inactive by burrowing into the mud or entering muskrat burrows. During rainy weather, few C. guttata were active. Activity was not only triggered by temperature, but also daylight hours. As darkness approached, the turtles burrowed into the mud bottom of some watenivay or crawled into muskrat burrows and became inactive after dusk (Ernst, 1976). Home Rage Previous Studies Home range is defined as the area in which turtles are observed during a given year (Ernst 1970). There are several methods to measure home range, constantly being revised and upgraded. Two common methods are Minimum Convex Polygon (MCP) and Kernel Home Range Analysis (KHRA), both were used for this study. Minimum Convex Polygon is the home range that includes outer boundaries (peripheral) of observation, including most outliers. It takes into account most location points, including 95% of those observed. MCP is well defined and straight forward. Worton (1989) described Kernel Home Range Analysis as the utilization distribution (how much area an individual animal is using in a given time period) based on points observed. Kernel Home Range Analysis tends to give the most accurate home range data compared to other analyse (i.e. Minimum Convex Polygon). The 95%, 85%, 75%, and 50% Kernel estimation based on utilization distribution was used to eliminate any “outliers” that would artificially inflate home range. Ernst (1970) used minimum and modified minimum home ranges which are similar to Minimum Convex Polygon and Kernel Home Range Analysis. The ind! her and SUE! and dele du'ir sulfa P0!!! hecte fang. Cher two methods were used to measure home ranges of both male and female individuals. His study concluded that females had a mean minimum range of 1.31 acres (0.53 hectares) and a modified minimum range of 1.25 acres (0.509 hectares), while males had a mean minimum range of 1.3 acres (0.53 hectares) and a modified minimum range of 1.23 acres (0.498 hectares). His data suggested that there was no significant difference between male and female home ranges. Home ranges studied by Ernst (1970) in Pennsylvania overlapped in time and space, but no territoriality was displayed. Territory defined as an area of defended space. Interest in other individual Spotted Turtles was shown only during the mating period. Home ranges of females usually did not include a suitable nesting area and individuals had to migrate out of their range to nest. Litzgus (1996) determined home range estimates using Minimum Convex Polygon and found that the mean home range size for females was 3.22 hectares and 3.58 hectares for males. No significant difference between home range sizes for males and females was reported. Breisch (2006), using combined data, stated that the mean home range for West Virginia male and female Spotted Turtles was 0.39 hectares using Minimum Convex Polygon. In Indiana, Barlow (1999) documented a home range of 2.03 hectares for males and 2.82 hectares for females also using Minimum Convex Polygon (MCP). No differences in home range sizes were found in her Indiana study, although overlapping of home ranges was extensive. On the other hand, Haxton and Berrill (1999) in Ontario, Canada found an average home range using MCP of 3.7 hectares, with females having a significantly larger home range than males. Differences in numbers, methodology, locale, and habitat could explain the varied findings by Spotted Turtle researchers. Homin Previous Studies In Michigan, Cagle (1944) reported a tendency for turtles to return to a given area, displaying homing behavior. The study showed seasonal movements away from the home range occurred during early spring and late fall, when either the need for hibernating quarters or “...the period of spring wandering led the turtle to new areas.” Turtles, forced from their home area by aberrant habitat changes, either followed the last remnants of water or moved at random in search of a suitable environment. Cagle (1944) found individuals forced from their home ranges may return to them when conditions are again suitable. Previous studies suggested some turtle species, including C. guttata, have homing ability even after being removed from their home range (Breder, 1927; Nichols, 1939; Medsger, 1919; Schneck, 1886; Grant, 1936). Ernst (1968) found small numbers of C. guttata in a Pennsylvania population returned to the original capture point 4 to 64 days after being moved 805 m upstream from his study site. He suggested that the turtles could have recognized the current, temperature gradient, and aromatic characteristics of a certain creek and followed it home, but provided no evidence as to which of these are most important. Objectives The specific goal of this study was to observe Spotted Turtles in their natural habitat with the purpose of determining: . Habitat preferences: 0 hydrophyte identification 0 Nesting 0 location 0 vegetation used 0 nest building activity 0 egg/hatchling observation 0 Movement and movement patterns: 0 home range 0 homing ability (does it occur?) 0 Overlapping of home range (does it occur?) 0 comparisons between male and female movement 0 Population dynamics and demography 0 Comparisons of male/femaIe/juvenile 0 Seasonal Activities: 0 Aestivation (does it occur?) 0 location 0 Hibernation . Locaflon o Communal hibemaculum (does it occur?) 10 iii! an AU Tan 31’ .,, 3‘! 0 Site fidelity (does it occur?) This was accomplished through radio telemetry, hand capturing, mark and recapture methods and thousands of hours of observation and recording data. Study Site Description This study was conducted in a southwestern Michigan fen. Defined by Chadde (2002), a fen is a peat covered wetland with a constant flow of mineral rich groundwater which may appear dry at the surface in later summer months. During drier periods, the water table is present just below the ground surface. Fen soil is characterized as alkaline (pH ranging from 7.9 to 8.3) and hydric which means it is flooded or saturated long enough during the growing season so that anaerobic conditions develop in the upper strata. Groundwater contributes both calcium and magnesium minerals to the wetland. Chadde (2002) also stated wetlands, which include fens, have at least one of the following three attributes regarding vegetation, soils and hydrology: (1) predominant plants are hydrOphytes, (2) soils are largely undrained, hydric soils, and (3) water will either permanently or periodically cover the area during some or all of the growing season each year. The fen in this study was approximately 30 hectares in size. In the early 1800’s (presettlement era) land descriptions were recorded by people walking and surveying section lines. Areas where no survey section lines were available, a “best guess” was used. Extrapolation was determined from what little information was provided by surveyors. The habitat and area where today this study took place, was referred to as a “Shrub Swamp/Emergent Marsh” by Arc 11 View 9.2 (Figure 1); said to contain taller, woody hydrophytes, including Dogwood and Willow plant species. This suggests that the survey may have commenced from a section line that was near where the main road is today. If that is the case, the surveyor may not have been able to see beyond the taller shrub species to record an existing area of short vegetation. The plant majority within the study site were hydrophytes, defined by Chadde (2002) as plants that grow in water or soil that is at least periodically deficient in oxygen resulting from high water saturation. Today, the study site center would be considered an open Sedge meadow; shorter hydrophytes (61cm) included Sphagnum spp., Sedge tussocks including Common Threesquare (Scirpus pungens), and Sh rubby Cinquefoil (Potenti/la fruticosa), which created safe havens for C. guttata. Beyond the Sedge meadow, a taller (2 to 5 m) growth of vegetation containing both woody and non-woody plants included, but not limited to, cattails (Typha Iatifolia and Typha angustifolia) both of which will be referred to as Typha throughout the paper, Red-osier Dogwood (Cornus sericea), Poison Sumac (Toxicodendron vernix), Northern Swamp Dogwood (Cornus racemosa), Swamp Rose (Rosa palustris), and many species of Sedges and Rushes (Table 2 and Table 3). At the lake’s edge, thick canopies of Glossy Buckthom (Rhamnus frangula) dominated making some areas practically impenetrable (Figure 2). Prescribed burns by The Southwest Michigan Land Conservancy were initiated in the past near the lake’s edge in attempt to eradicate the thick stand of Glossy Buckthom. 12 Surrounding this area in presettlement times were Mixed Oak Savanna and Oak-History Forests. These areas are still visible but housing developments and infrastructure have altered the original habitat. During this study, all nearby surrounding habitats, including ponds, lakes, marshes, and wetlands were surveyed, but turtles were only observed in the study site. The fen was surrounded by housing developments on two sides, a lake on the third side, and a main road on the fourth side. This was described as a “shelf” that attaches to a nearby lake, protected by The Southwest Michigan Land Conservancy and private land owners. In addition, the site included various protected species plants which included, but were not limited to, Northern Grass-Of-parnassus (Parnassia palustris) and Lesser Fringed Gentian (Gentiana procera). Wildlife sighted in the fen included Coyote (Canis Iatrans), Red Fox (Vulpes fulva), Raccoon (Procyon Iotor), Meadow Vole (Microtus pennsylvanicus) and Muskrat (Ondatra zibethica) (Table 4). Reptiles and amphibians documented included, but were not limited to, Eastern Box Turtle (Terrapene carolina carolina), Eastern Massasauga Rattlesnake (Sistrurus catenatus catenatus), Western Chorus Frog (Pseudacris triseriata) and Spring Peeper (Pseudacris crucifer) (Table 5). Lastly, the site included a variety of nesting wetland bird species including Willow Flycatcher (Empidonax trail/r), Swamp Sparrow (Melospiza Georgiana), Common Snipe (Gallinago gallinago), Common Yellowthroat Warbler (Geothlypis trichas), and Yellow Warbler (Dendroica petechia) (Table 6). 13 man dlStr i038: On the fen’s east side was a pond dominated by thick growths of algae (Chara spp.), characterized as a Chara pond (Michigan Dept. of Env. Quality Water Bureau, 2005). Chara spp. (common names: Stonewart, Muskgrass) is an advanced form of algae. It has a musky odor and gritty texture caused by mineral deposits on its surface. It grows in low, dense mats. In a Chara pond, water is clear and rich in calcium and some magnesium. The water remains relatively unaffected by intensive land use or other surface nutrients and most often found in areas supporting mosaics of semi-aquatic vegetation (Joint Nature Conservation Committee, 2005). Chara spp. grows densely because, like other algae filters dissolved nutrients out of the water instead of sediments. In this respect, Chara spp. are highly beneficial vegetation (Michigan Dept. of Env. Quality Water Bureau, 2005). However, Chara spp. develops undenivater, grows rapidly, and as a dominant species, is usually considered undesirable, causing oxygen depletion (Brinlee 2009). Anchored to the bottom, it develops into large green mats with thin leaf-like structures, and has proven difficult for fish maneuverability (Peterson and Lee, 2005). It is very common and widely distributed throughout Michigan. Muskrats and their burrows were present in the pond. This pond had loose, muddy, marl deposits which, when stepped on, seemed “bottomless” and made it very dangerous for humans and perhaps turtles. Chara spp. appeared to play an important role in the life history of this Spotted Turtle population. It was a factor in mortality of Spotted Turtles. Discussion in results section of Chapter 2. 14 Methods Ernst (1970) defined sexual maturity in both sexes as when the individual reached a carapace length of 8.0 cm. For purposes of sexual determination at the start of this project, Ernst’s 8.0 cm limit was used for obvious male and females. However, female characteristics are juvenile characteristics, and male characteristics (when it comes to secondary characteristics) are yet to be defined. Thus in this study, Ernst’s definition may not hold true. From March 2007 to March 2009, telemetry equipment was used for tracking movement of selected male and female Spotted Turtles. In early spring (late March, early April), adult turtles were equipped with transmitters. During the two year study period, 29 turtles with an average age of 12.3 years and carapace length of 9.1 cm were telemetered. In 2007, twelve transmitters (model # 51-2FT Holohil Systems Ltd., Ontario, Canada) were attached to the study turtles. Transmitter battery life was at least ten months, lasting an entire year. Two small holes on the carapace’s posterior left side were drilled into the marginal scutes. The transmitter with a 22.9 cm whip antenna was attached using fine gauge copper wire. The transmitter was additionally secured with a waterproof epoxy to keep out debris, or prevent catching on vegetation. Black permanent marker was used to camouflage the epoxy for better blending into carapace color. When complete, the transmitter weight was 12 g or approximately 10% to 11% of the turtle’s original body weight (Figure 5). This was within the range used by other researchers. For example, Breisch (2006) described transmitter weight to be 10.2% body weight. The receiver used in this study was a Telonics TR-4 15 (Telonics Inc. Products, Mesa, AZ). A hand-held Yagi 3 element folding antenna (Wildlife Materials lntemational, Murphysboro, IL) was also used in conjunction for tracking. Turtles were tracked two to three times a week until November, then once a month. When a turtle was located, morphometric data was recorded. Using dial calipers, carapace length, plastron width at the bridge, plastron length and height were recorded. Sex determinations, as well as weight calculated with Salter Housewares Ltd. Scale, and age (by counting annuli) were also recorded. Also documented were injuries, behavior at capture time, GPS coordinates at observed location (Garmin etrex) and vegetation description. Carapace temperature (Pro Exotics Tempgun), as well as air temperature (Kestrel 2000), and water temperature (if not available, substrate temperature was recorded); also recorded were general weather conditions. All data were recorded in a field notebook, then transferred to Microsoft Excel data sheet for each individual turtle (Table 7). Radio telemetry data for each of the transmittered turtles were also transferred to a Microsoft Excel data sheet (Table 8). All turtles were hand captured and data were obtained and recorded during daylight hours. Most turtles were located by traversing through the fen. During fall, turtles were found by “mucking”, which is the process of probing through the mud, muck and water by hand to locate turtles. Turtles were marked by filing notches in the marginal scutes. A marking system using the alphabet letters was used (Figure 6, Figure 7, and Figure 8). 16 For hatchlings, a pair of small scissors was used to clip a single triangular shaped notch into the marginal scute. In 2008, six additional transmitters (model SB-2 modified, Holohil Systems Ltd., Ontario, Canada) were employed. Transmitter attachment, with a 15.2 cm whip antenna, was done by drilling one hole into a marginal scute. A fine gauge copper wire was used to attach the transmitter to the carapace, with waterproof epoxy for final transmitter securement. The transmitter battery life was six months, thus, transmitters were removed before turtles proceeded with hibernation. Total transmitter weight was 6 g which was approximately 4.5% to 6% of the total body weight (Figure 9). Turtles were tracked one to three times per week until November, then one to two times per month. Morphometric protocol and data collection methods were the same as 2007. During 2007 and 2008, the transmitters (model # S1-2FT) provided pulses, which when counted, applied to a calculation, then plotted to a temperature graph (supplied by Holohil Systems Ltd.) identified the turtle’s hibemaculum temperature. Litzgus and Brooks (2000) suggested a correlation between turtle’s hibemaculum temperature and actual body temperature. Maps of Spotted Turtle distribution and analysis were created using Arc Map 9.2 and Arc View 3.3 (ESRI, Redlands, CA). The Arc View programs were used to determine MCP and Kernel Home Range Analysis for turtles with transmitters. Only MCP was generated for turtles without transmitters. MCP included outer boundaries of observation including most outliers, taking into 17 an ice! Eve account most location points. On maps, yellow dots represent turtle observations and may include more than one observation. Habitat Results One of the study objectives was to discover Spotted Turtle habitat preference. Aggregations and possible preferred habitat locations were indeed identified. A diagram of the entire fen, pointing out areas of heavy aggregations was presented (Figure 10). Turtles with and without transmitters, located during 2007, were plotted in the study area (Figure 11 and 12). Two 50% Kernel estimation areas were identified. The first area included was centrally located at the lowest elevation. Even with 2007’s drought-like conditions, the substrate was similar to saturated mud. This area, during drier times, tended to be more saturated than most other areas and aggregations of Spotted Turtles occurred here. The main hydrophytes varied in height from short (less than one meter) which included Sphagnum spp., Sedge, and Shrubby Cinquefoil to taller heights (2-3 meters) including Typha, Willow, Northern Swamp Dogwood and Red-osier Dogwood. Within the second 50% Kernel estimator area was an overflow channel, referred to as “the outlet.” This originated at the lake’s edge and allowed lake water to flow approximately 15 meters inland. Even during the driest part of 2007, the outlet generally had water in it or was thoroughly saturated, leaving a thick, organic, mucky substrate. Channel 4 (HKU- female) was tracked and located deep beneath this muck on several occasions. 18 The area around the outlet was populated by taller (2-5 m) woody and non-woody hydrophytes including Glossy Buckthom, Swamp Rose, Poison Sumac, Northern Swamp Dogwood, Red-osier Dogwood, Sedge, Fern and Sphagnum spp. Included within one of the four 75% Kernel estimator areas was a locale designated “the ditch.” The ditch was located at the fen’s southern end, running east and west perpendicular to a nearby road. Besides rainfall and overflow from the fen, it collected run-off water. In 2007, during the driest of times, 10 cm of water was recorded in the ditch. At no time during the two year study was this location completely dry. Adjacent to the edge, tall emergent vegetation included Typha, Willow, Poison Sumac, Red-osier Dogwood and Northern Swamp Dogwood, Sedge, Bulrush and Shrubby Cinquefoil. Turtles with and without transmitters, frequented this area. The other three 75% Kernel estimator areas were lower elevation areas, where turtle aggregations were documented. This included areas where substrate moisture was present, even during drier months. One of these areas contained approximately 3 m high, thick canopies of Typha. The second area was located in a more open area of the fen, just north of the ditch and contained Bulrush, Sedge and Typha. The third area was referred to as the Chara pond, dominated by dense Chara species. Maps of all turtles with and without transmitters found during 2008 study are found in Figures 13 and 14. This was a much wetter year, with rainfall far exceeding 2007. Activity and frequency of individuals with and without 19 transmitters were more scattered. This suggests larger area utilization may have been the result of additional precipitation. Using Kernel Home Range Analysis, the 50% Kernel estimator area, was located in two areas of the fen. The first included the study site center a low elevation area described earlier. The second area was a 2 m wide trail to the lake, located on the southeastern edge. This path appears to have been made years ago by the property owners as an access to the lake. On either side of this path, tall woody and non-woody plant vegetation was present. The hydrophytes included, but were not limited to, Swamp Rose, Poison Sumac, Typha, Red-osier Dogwood, Glossy Buckthom, Sedge and Bulrush. The mud and muck path was trampled down, forming depressions that allowed for water collection. Tall emergent vegetation, next to the trail, consisted of many deer trails. These trails also supported trapped water. These trails opened to the lake’s edge, where there was low growing vegetation including Sedge, Shrubby Cinquefoil and Bulrush. With the large amounts of rainfall in 2008, the lake was at capacity causing shore overflow. In 2008, the turtle activity level was heavy with and without transmitters in this area. The 75% Kernel estimation was determined in three fen areas in 2008. The first area was known as the ditch, which was described above in 2007. The second area contained short hydrophytes that were less than 1 m in height in a low elevation area. The main hydrophytes included Sedge, Sphagnum spp., and Shrubby Cinquefoil. The third area was located near the southwestern lake 20 edge, a low vegetation area which included Shrubby Cinquefoil, Sedge and Bulrush. Water was present throughout the year. Sphagnum mounds, with Sedge and Shrubby Cinquefoil growth on top of the mounds, appeared to provide nesting locations for Spotted Turtles. Although no turtles were observed nesting in this study, six presumable successful nest sites were located. Eggshells found deep in the soil indicated successful hatching seemed likely. All of these nests were found in the previously described vegetation. Nests were placed in well drained areas exposed to full sunlight. One predated nest was found in 2008, soon after the eggs had been deposited. This nest had been dug up, with eggshells on top of the substrate (Figure 15). On 16 June 2008, a single egg was observed lying on top of a Sphagnum spp. mound, with Shrubby Cinquefoil growing from the top of the mound. With further investigation, a nest with additional eggs was located buried in the Sphagnum spp. mound. The egg observed had a weight of 4 grams, length of 3 cm, and width of 1.7 cm. Major rain storms deluged the site on 2 July 2008 and most deer trails contained at least 20 cm of water. On 4 July 2008, the nest was observed underwater, submerged for at least three days. On 4 September 2008, a hatchling was located with the empty eggshell from one additional hatchling (suggesting successful emergence). Also included in this nest, was a dead embryo still in its shell (Figure 16). Ernst et al (1994) reported Spotted Turtle incubation periods between 70-83 days in length. The incubation period for this nest was 81 days which fell within this range. 21 Movement Results 2007 Radio Telemetry and Climatic Conditions Transmitters were attached to 12 turtles in 2007. This included four males and eight females. In 2007, the total annual rainfall was documented at 33.82 inches (approximately 86 cm). During summer months, the precipitation was well below normal, causing drought-like conditions throughout the fen. Only areas of lower elevation contained standing water during this drought period. Included were six areas previously described in Chapter 1: 1) outlet, 2) area containing approximately three meter high canopies of Typha, 3) open area in middle of fen, 4) Chara pond, 5) ditch, and 6) area just north of ditch which contained hydrophytes Bulrush, Sedge and Typha. A short summary of each radio telemetered turtles’ movements and habitat utilization is described below. Channel 0 - IKL (Male) Male IKL was first captured and telemetry attached 24 March 2007. He was observed a total of 72 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of habitat utilization was 1.60 hectares (Figure 17). Main activity occurred in the ditch. Hydrophytes occupying this area included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. Throughout this season, the ditch usually contained more than 10 cm of water. Male IKL was originally captured coming out of hibemaculum in the ditch. At the end of the 2007 season, he returned to the ditch to hibernate suggesting hibemaculum site fidelity. 22 Channel 1 — KVW (Male) Male KVW was first captured and telemetry attached 25 March 2007. He was observed a total of 76 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 2.37 hectares (Figure 18). Main activity occurred in the open area in middle of fen. This area of lower elevation held water more frequently than other areas. Hydrophytes occupying this area included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. Male KVW was originally captured coming out of hibemaculum in the fen’s open middle area. At the end of the 2007 season, he hibernated in same area from which he emerged earlier in spring, suggesting hibemaculum site fidelity. Channel 2 — ABCK (Male) Male ABCK was first captured and telemetry attached 24 March 2007. He was observed a total of 77 times throughout 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 3.30 hectares (Figure 19). Channel 2 movement was extensive throughout fen in 2007, covering a wide variety of habitats. Two main areas of activity included big trail to lake and lake’s edge, as well as fen’s open middle area. The big trail to lake was a wide trail that led from an open area to lake. Both sides of this trail had tall hydrophytes including Red-osier Dogwood, Poison Sumac, Typha and Glossy Buckthom. Hydrophytes occupying the middle open area included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. This area of lower elevation held water more frequently than other areas. 23 The exact initial spring emergence location of Channel 2 was not observed but he was found near the middle area of fen with a lower elevation. The hibernation locale was documented in this same area. This suggests possible hibemaculum site fidelity. Channel 3 — KVX (Female) Female KVX was first captured and telemetry attached on 20 April 2007. She was Observed a total of 65 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.48 hectares (Figure 20). Main activity occurred near the road. Hydrophytes included Typha, Red-osier Dogwood, Swamp Rose and Willow. This turtle was originally located in close proximity to the main road. After the transmitter was attached, she was released into a more open area, located some distance from the main road, to ensure safety. In less than 24 hours, the turtle returned to its original home range. This finding suggests that homing ability may be present in Spotted Turtles in this southwestern Michigan fen. At the end of the 2007 season, she was sighted going into a hibemaculum near the same location where originally captured in the spring. Further studies of this turtle need to be completed to assess hibemaculum site fidelity. Channel 4 - HKU (Female) Female HKU was first captured and telemetry attached 11 May 2007. She was observed a total of 58 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.35 hectares (Figure 21). Main activity occurred near the lake. Hydrophytes occupying this area included 24 Glossy Buckthom, Sedge, Fem, Purple Loosestrife, Sphagnum spp. and Willow. Even during drought-like conditions, water was evident below the substrate. Also near this location was the outlet, which generally contained standing water. Documented sightings of female HKU, occurred throughout most of the season in this area. Female HKU was not captured until late spring, making her initial spring emergence location unknown. At the end of the 2007 season, she was found hibernating in fen’s open area. Hydrophytes were predominantly Sedge, Typha and Shrubby Cinquefoil. Caused by low elevation, this area held water longer than other areas. Future studies need to be completed to determine if this turtle had hibemaculum site fidelity. Channel 5 - BKW (Female) Female BKW was first captured and telemetry attached 21 April 2007. She was observed a total of 22 times throughout the 2007 season. On 4 June 2007, the transmitter malfunctioned and was removed. Minimum Convex Polygon (MCP) was not conducted due to incomplete data. Channel 6 - DKVWX (Female) Female DKVWX was first captured and telemetry attached 17 May 2007. She was observed a total of 54 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.29 hectares (Figure 22). Main activity occurred in the outlet and open area in middle of fen. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. Due to lower elevation, the substrate was more saturated than other areas. This turtle was originally captured late spring, making her initial 25 spring emergence location unknown. At the end of 2007, documented hibernation was in fen’s open area. Future studies need to be completed for determination of hibemaculum site fidelity. Channel 7 - KLU (Female) Female KLU was first captured and telemetry attached 18 May 2007. She was observed a total of 55 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.40 hectares (Figure 23). Main activity occurred in the ditch. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, Bulrush, and Shrubby Cinquefoil. This turtle was originally captured late spring, making her initial spring emergence location unknown. At the end of 2007, documented hibernation was in the ditch. Future studies need to be completed for determination of hibemaculum site fidelity. Channel 8 — HKV (Female) Female HKV first captured and telemetry attached 18 May 2007. She was observed a total of 34 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 1.12 hectares (Figure 24). On 25 June 2007, she was found in the Chara pond, caught in Chara spp. The turtle was untangled, and it was noted the epoxy was loose and may have caused the transmitter to “catch” on the Chara spp. With repairs made, the turtle was released at a safer location, a distance from the Chara pond, in the outlet. Within 10 days of original move, female turtle HKV had returned to the Chara pond, suggesting homing ability. On 19 July 2007, the turtle was found dead with 26 its antenna tangled in Chara spp. Main activity occurred in the Chara pond. Main hydrophytes included Typha and Chara spp. Channel 9 — KLW (Female) Female KLW was first captured and telemetry attached 17 May 2007. This turtle was originally captured near the Chara pond, and observed nine times. After these nine sightings, female KLW was never observed again. The turtles’ telemetry location did not change positions after 30 May 2007. With the inter-connected networks of underground muskrat burrows, the turtle was believed to be in the burrows and inaccessible for observation. On 17 June 2007, lacking turtle movement for several weeks, the telemetry location was thoroughly investigated for whereabouts of the missing turtle. Remains were found in deep water of the Chara pond, with head missing and transmitter still attached to carapace. Two, possibly three eggs were observed within turtle. Considering water depth where turtle was found, it is believed that it was predated by a muskrat. Because data was incomplete, Minimum Convex Polygon was not conducted. Channel 10 - IKW (Male) Male IKW was first captured and telemetry attached 21 May 2007. He was observed a total of 42 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.45 hectares (Figure 25). Main activity occurred in the outlet and ditch. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. Both locations had documentation of water throughout the dry season. This turtle was originally 27 captured in late spring, making his initial spring emergence location unknown. The turtle was documented hibernating in the ditch at the end of 2007. Future studies need to be completed to determine hibemaculum site fidelity. Channel 11 — KV (Female) Female KV was first captured and telemetry attached 30 May 2007. She was observed a total of 46 times throughout the 2007 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.07 hectares (Figure 26). Main activity occurred in open area in middle of fen. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. This area was lower in elevation and held water longer than other areas. This turtle was originally captured in late spring, making her initial spring emergence location unknown. At the end of 2007, documented hibernation was in open area of fen. Female KV returned to the same hibemaculum at the end of the 2008 season, suggesting hibemaculum site fidelity. 2008 Radio Telemetry and Climatic Conditions In 2008, 17 turtles had telemetry attached. This included 14 females and three males. Unlike 2007, the 2008 season was much wetter with higher levels of rainfall documented throughout summer months. Total rainfall for entire year equaled 54.1 inches (approximately 137 cm). This was substantially higher than 33.82 inches (approximately 86 cm) documented for 2007. Most areas throughout the site contained standing water during 2008. The lower elevation areas had increased water levels, and higher concentrations of turtles than the higher elevation areas of the fen. These lower elevation areas included: 1) open 28 area in middle of fen, 2) wide trail to lake including lake’s edge, 3) ditch, 4) an area of short hydrophytes, and 5) far southwestern edge of lake. Channel 0- KMW (Female) Female KMW was first captured and telemetry attached 9 June 2008 near lake’s edge. She was observed a total of 15 times throughout the 2008 season. During the 15 times this turtle was observed, it was found moving towards the Chara pond. On 4 September 2008, she was rescued from the middle of the Chara pond where she was caught in Chara spp. Telemetry equipment was removed, in order to avoid a repeat situation, similar to the one which occurred in 2007, with Channel 8. Due to minimal number of sightings, Minimum Convex Polygon (MCP) was not generated for this turtle. Channel 1 - HJKV (Female) Female HJKV was first captured and telemetry attached 13 May 2008. She was observed a total of 39 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 1.20 hectares (Figure 27). Main activity occurred in area of big trail to lake, a wide trail that led from open area to lake. Both sides of this trail had tall hydrophytes including Red-osier Dogwood, Poison Sumac, Typha and Glossy Buckthom. Main hydrophytes included Sedge, Bulrush and Shrubby Cinquefoil. This turtle was originally captured in late spring, making her initial spring emergence location unknown. Documented hibernation was near lake’s edge. Future studies need to be completed to determine hibemaculum site fidelity. 29 Channel 2 - KUV (Male) Male KUV was first captured and telemetry attached 10 June 2008. He was observed a total of 13 times. The turtle was originally located in the ditch and migrated towards the Chara pond. On 23 July 2008, he was found dead in the Chara pond. Main hydrophytes in the ditch area included Typha, Sedge, Bulrush, Shrubby Cinquefoil and Gray Twig Dogwood. Hydrophytes in the Chara pond included Typha and Chara spp. Even though Channel 2 was only observed 13 times, total area of utilization (using Minimum Convex Polygon) was generated. Home range for this turtle was 0.65 hectares (Figure 28). Channel 3 - HJK (Female) Female HJK was first captured and telemetry attached 23 May 2008. She was observed a total of 35 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.71 hectares (Figure 29). Main activity occurred in an area which contained a tall canopy of Typha near main road. Hydrophytes included Typha, Swamp Rose, Willow, Fern, Sedge, and Shrubby Cinquefoil. This turtle was originally captured late in spring making her initial spring emergence location unknown. Documented hibernation included a tall canopy of Typha, Swamp Rose, Willow, Sphagnum spp. and Sedge. Future studies need to be completed to determine hibemaculum site fidelity. Channel 4 - ABKW (Female) Female ABKW was first captured and telemetry attached 31 May 2008. She was observed a total of 34 times throughout the 2008 season. Using 30 Minimum Convex Polygon (MCP), total area of utilization was 0.23 hectares (Figure 30). Main activity occurred in middle open area of fen. Main hydrophytes included Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. This turtle was originally captured in late spring, making her initial spring emergence location unknown. Documented hibernation was in the open area of the fen. Female ABKW was buried in a hibemaculum below a Sphagnum spp. moss mound covered with Shrubby Cinquefoil and Sedge. Investigations from last year showed this turtle hibernating in an area in close proximity to its 2008 hibemaculum, indicating hibemaculum site fidelity. Channel 5 - CHJKLVW (Female) Female CHJKLVW was first captured and telemetry attached 10 June 2008. She was observed a total of 26 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.22 hectares (Figure 31). Main activity occurred in an area consisting of a tall canopy of Typha, Swamp Rose, Willow, Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. Channel 5 was originally captured in late spring, making her initial spring emergence location unknown. Documented hibernation was in the same area with the tall canopy of Typha. Future studies need to be completed to determine hibemaculum site fidelity. Channel 6 - BDKW (Female) Female BDKW was first captured and telemetry attached 31 May 2008. She was observed a total of 34 times throughout the 2008 season. Using 31 Minimum Convex Polygon (MCP), total area of utilization was 0.60 hectares (Figure 32). Main activity occurred in open area in middle of fen, with several sightings occurring in thick areas of Glossy Buckthom near lake. Main hydrophytes included Typha, Willow, Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. Channel 6 was originally captured in late spring, making her initial spring emergence location unknown. Documented hibernation was in middle open area of fen. Future studies need to be completed to determine hibemaculum site fidelity. Channel 7 - AKX (Female) Female AKX was captured and telemetry attached on 12 June 2008. She was observed a total of 24 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 1.09 hectares (Figure 33). This turtle was initially found in 2008 near the edge of lake and migrated, as the season progressed, toward middle open area of ten. Main hydrophytes in this area included Typha, Bulrush, Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. In March 2007, Female AKX was observed leaving a hibemaculum in open area of fen. She was documented entering a hibemaculum in this same area at the end of 2008, suggesting hibemaculum site fidelity. Channel 8 - FKS (Female) Female FKS was first captured and telemetry attached 10 June 2008. She was observed a total of 26 times throughout the 2008 season. Using 32 Minimum Convex Polygon (MCP), total area of utilization was 1.33 hectares (Figure 34). Main activity occurred in lower elevation area which included Sphagnum spp., Sedge, and Shrubby Cinquefoil. Later in the season, female FKS migrated to the ditch, where hibernation was documented. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, Bulrush and Shrubby Cinquefoil. Channel 8 was captured in late spring; her initial spring emergence location was unknown. Future studies need to be completed to determine hibemaculum site fidelity. Channel 9 - BKW (Female) Female BKW was first captured and telemetry attached 12 June 2008. She was observed a total of 26 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.16 hectares (Figure 35). Main activity occurred on big trail that led from open area to lake. Both sides of this trail had tall hydrophytes which included Red-osier Dogwood, Poison Sumac, Typha and Glossy Buckthom. This turtle was originally captured in late spring, making her initial spring emergence location unknown. Female BKW was documented hibernating near entrance of big trail to lake. Future studies need to be completed to determine hibemaculum site fidelity. Channel 11 - CKU (Female) Female CKU was first captured and telemetry attached 10 June 2008. She was observed a total of 26 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 1.08 hectares 33 (Figure 36). Main activity occurred in a tall canopy of Typha, Swamp Rose, Willow, Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. This turtle was originally captured in late spring, making her initial spring emergence location unknown. Documented hibernation was in same area of a tall canopy which included Typha, Swamp Rose, Willow, Sedge, Shrubby Cinquefoil, and Sphagnum spp. moss mounds. Future studies need to be completed to determine hibemaculum site fidelity. Channel 12 — ABCKW (Female) ABCKW was captured and telemetry attached 11 May 2008. She was observed a total of 37 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.49 hectares (Figure 37). Main activity occurred in open area in middle of fen. Several sightings occurred near lake, in thick areas of Glossy Buckthom. Main hydrophytes included Typha, Bulrush, Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. Female ABCKW was first observed 29 March 2007 coming out of a hibemaculum in middle open area of fen. On 5 October 2007, ABCK was observed entering a hibemaculum, identified as same spring emergence location. In 2008, ABCKW was documented hibernating in middle open area of ten, suggesting hibemaculum site fidelity. Channel 13 - DKP (Female) Female DKP was captured and telemetry attached 11 May 2008. She was observed a total of 39 times throughout the 2008 season. Using 34 Minimum Convex Polygon (MCP), total area of utilization was 1.08 hectares (Figure 38). Main activity occurred in an area which included a tall canopy of Typha, Willow, Swamp Rose, and Glossy Buckthom. On 3 October 2008, the turtle was found dead in deep water surrounded by a tall canopy of Typha and Willow. No visible signs or cause of death were noted. Channel 14 — BHK (Female) Female BHK was first observed 24 March 2007 emerging from a hibemaculum in middle open area of fen. She was captured and telemetry attached 11 May 2008. She was observed a total of 35 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.79 hectares (Figure 39). Main activity occurred in open area in middle of fen, with several sightings occurring near lake in thick areas of Glossy Buckthom. Main hydrophytes included Typha, Bulrush, Sedge, Shrubby Cinquefoil and Sphagnum spp. moss mounds. In September of 2008, the transmitter was removed; preferred hibemaculum was undetermined. Future studies are needed to determine hibemaculum site fidelity. Channel 15 — ADKO (Female) Female ADKO was captured and telemetry attached 15 May 2008. She was observed a total of 36 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 0.22 hectares (Figure 40). Main activity occurred in an area of Shmbby Cinquefoil, Sedge, Glossy Buckthom and Sphagnum spp. moss mounds. This turtle was 35 documented in a possible hibemaculum buried deep in mud and water, beneath thick brush and Sedge pile. Future studies need to be generated to determine hibemaculum site fidelity. Channel 16- ABCK (Male) Male ABCK was Channel 2 in 2007. He was captured again and telemetry reattached on 16 May 2008. He was observed a total of 30 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of utilization was 1.81 hectares (Figure 41). Main activity occurred in big trail to lake and its edge, and open area in middle of fen. Both sides of this trail had tall hydrophytes which included Red-osier Dogwood, Poison Sumac, Typha and Glossy Buckthom. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, and Shrubby Cinquefoil. This area of the fen was lower in elevation, which held water longer than other areas. In 2007, ABCK was documented emerging and hibernating in the middle open area with lower elevation. In 2008, he was documented emerging from the same locale as 2007, but hibernated in big trail near lake. Site fidelity was suggested from 2007 data but was not found in 2008. Channel 17 - IKW (Male) Male IKW was Channel 10 in 2007. He was recaptured and telemetry reattached 1 June 2008. He was observed a total of 31 times throughout the 2008 season. Using Minimum Convex Polygon (MCP), total area of habitat utilization was 1.33 hectares (Figure 42). Main activity occurred along the lake’s 36 edge and ditch. Main hydrophytes included Typha, Red-osier Dogwood, Sedge, Willow, Sphagnum spp. moss mounds and Shrubby Cinquefoil. In 2007, IKW was documented emerging from a hibemaculum in the ditch. He was recorded later in the season returning to ditch to possibly hibernate. In 2008, he was recorded returning again to ditch for possible hibernation. This information suggested hibemaculum site fidelity. The area of utilization of all turtles with and without transmitters in 2007 and 2008 was 13.30 hectares. Area of preferred habitat was described in Chapter 1. Each individual turtle’s home range included nesting locations, areas of aestivation (when applicable), and hibernation. Radio telemetry was used to track turtles in the 2007 and 2008 season. Radio telemetry turtles were observed more than 1500 times. More females than males had transmitters attached to them with hopes of acquiring a better understanding of nesting activity. This proved to be difficult, with no females being observed nesting. The home ranges of males were larger than females. The mean home range in 2007 using Minimum Convex Polygon (MCP), for males, was 1.9 hectares and for females 0.5 hectares. The mean home range in 2008 using Minimum Convex Polygon (MCP) for males, was 1.3 hectares and for females 0.7 hectares. Barlow (1999) and Litzgus (1996) found no significant difference in home range sizes between males and females. Haxton and Berrill (1999) found females to have a significantly larger mean home range. 37 In 2007, the home range of ABCK Channel 2 male was 3.30 hectares. Male ABCK, again, was equipped with telemetry in 2008, this time as Channel 16. His home range was observed to be 1.81 hectares (Figure 43). The 2007 home range was distinctly larger than 2008. It should be noted that in 2007, ABCK’s transmitter was a larger model. In 2008, a smaller transmitter was attached to the turtle. In 2007, Male IKW was equipped with a larger model transmitter (Channel 10). In 2008, the smaller transmitter (Channel 17) was attached. The home range in 2007, with the larger transmitter, was 0.45 hectares. In 2008, the distance covered with the smaller transmitter was 1.33 hectares (Figure 44). The 2008 home range was distinctly larger than the 2007 home range. Homingand Overlapping Results Homing behavior is the ability of an animal to return to its home range when removed (Cagle 1944). This was documented on two occasions in this study. In April 2007, female KVX was captured in close proximity to a main road. After attachment of transmitter, the turtle was released in another area of the habitat, out of her home range, to ensure the turtle’s safety. In less than 24 hours, the turtle had returned to its home range where originally captured. The second instance of homing behavior occurred with Channel 8. Female HKV was found tangled in Chara pond, 25 June 2007. She was untangled; repairs were made to transmitter (new epoxy) and released out of her home range from Chara pond. On 15 July 2007, Channel 8 was tracked and located back at Chara pond. This suggested homing behavior. 38 In 2007, overlapping of female and male home ranges was documented (Figure 45 and Figure 46). The largest male home range (Channel 2) was 3.30 hectares. Overlapping occurred with four transmittered females. The home range of Channel 0 (male) was 1.6 hectares which overlapped with four transmittered females. Channel 10 (male) had the smallest home range with 0.45 hectares. Although home range was small, it overlapped with five transmittered females. Home range of KVW Male (Channel 1) was 2.37 hectares. Although home range for Channel 1 was not as large as Channel 2 (3.30 hectares), what is noteworthy is home range overlapped with all eight transmittered females (Figure 47). This suggested that searching and encountering of females would have been more productive. Although Channel 5 (BKW) and 9 (KLW) were not tracked for the entire 2007 season, their home ranges for the beginning of the season were documented to overlap with Channel 1 and Channel 0. Channel 10 (IKW) overlapped with Channel 5. The home range of Male ABCK (Channel 16) was 1.81 hectares, and overlapped with seven transmittered females in 2008. Male IKW (Channel 17) home range was 1.3 hectares, and overlapped with seven transmittered female turtles in 2008. Home range for Male KUV (Channel 2) was documented as 0.65 hectares, and overlapped with two transmittered female turtles in 2008 (Figure 48 and Figure 49). Habitat Discussion In this study, the area of distribution of all Spotted Turtles with and without transmitters was observed for two years (2007 and 2008). Areas of preferred 39 habitat were generally those with lower elevation, and standing water or mucky, wet substrate. Shorter, more accessible vegetation was generally preferred. Areas of tall, thick vegetation were avoided. Fen areas that included massive, thick stands of Glossy Buckthom were generally uninhabited by C. guttata. Only Channel 2, appeared to use the deer trails for providing quick and easy movement through the thick Glossy Buckthom canopy. Drier areas were generally avoided throughout the fen. It is possible that turtles avoided these areas lacking water for feeding, mating, and risk of desiccation. The prescribed burn area was also avoided, with no turtles found. The conservancy owns only a small portion of the study site, with the remaining portion owned by private land owners. Thus, these controlled burns have proven to only slow the growth of the Glossy Buckthom, due to dominance and ability of Glossy Buckthom to encroach from the private land owner's property. This area contained thick stumps of Glossy Buckthom that were impenetrable for turtles. The burn area was extremely dry, also explaining the absence of turtle sightings. Throughout the area of preferred habitat, where standing water or muck was available, shorter vegetation was present. This vegetation included Sphagnum spp., Sedge tussocks (Scirpus pungens), Typha, and Shrubby Cinquefoil (Potentilla fruticosa). Small mounds of Sphagnum spp. were observed with Sedge and Shrubby Cinquefoil growing on top of these mounds, with a total height up to 61 cm (Figure 50). There appeared to be a positive relationship between this type of vegetation and the survival of the Spotted Turtle. Spotted Turtles hide within this 40 vegetation, burrowing beneath the Spaghnum spp. mounds. With the growth of Sedge and Shrubby Cinquefoil, root systems in the Sphagnum spp. mounds and in the mucky substrate were prevalent. These root systems, along with groundwater flow, provided small, below surface tunnels and air pockets. These areas provided a means of movement and protection from predators in addition to locations for aestivation and hibernation, which are further discussed in Chapter 3. Turtles were found basking on mounds (Figure 51). In 2008, after heavy rainfall, most vegetation in the low areas was undenivater, providing fewer basking locations. Shrubby Cinquefoil was observed in some areas above the water line. Channel 4 was located basking near the top branches of this hydrophyte, sharing prime basking space with a Northern Water Snake. In late summer, with elevated air temperatures, Spotted Turtles needed protection from heat. Crawling under the “umbrella-like” growth of Shrubby Cinquefoil provided protection from the sun for C. guttata. Not only was Channel 4 observed using Shrubby Cinquefoil for sun protection, but also for both aestivation and hibernation, by burrowing deep into the mud beneath these Sphagnum spp. mounds (Figure 51). Nesting in this type of vegetation was also documented in these mounds covered with the growth of Sedge and Shrubby Cinquefoil. Sphagnum spp. mounds, Sedge and Shrubby Cinquefoil are important vegetative components needed to provide a successful habitat for C. guttata. A positive correlation between this vegetation and turtle movement, predation 41 avoidance, camouflage, basking, nesting, temperature control, aestivation and hibernation have been found during this study. Duckweed (family Lemnaceae), is a floating, leaved plant found in most C. guttata habitats (Ross and Lovich, 1992). Ross and Lovich (1992) proposed that the carapacial cryptic patterns of C. guttata mimicked the color and form of this plant species. Natural selection, with the use of cryptic coloration, helped to protect this species from predation. Although Lemnaceae occurs throughout most of Michigan (Voss, 1972), it was absent from this study site. Since duckweed was not present in this C. guttata site, camouflage may have been accomplished with Shrubby Cinquefoil (Potentilla fruticosa), a predominant fen hydrophyte. Shrubby Cinquefoil is a much-branched shrub that is 0.5 -1.0 m tall which when it blooms produces yellow summer flowers (Chadde, 2002). As the flower petals drop, the mud and substrate was covered with yellow polka dots. Distinguishing carapacial patterns of C. guttata from that of fallen petals might be difficult for would be predators (Figure 53 and Figure 54). Shrubby Cinquefoil may have provided camouflage for Spotted Turtles and help protect from predation. No previous studies support this finding. Bloom length of Shrubby Cinquefoil may not be as long as the duckweed season, but may have a positive effect on C. guttata. Ross and Lovich (1992) stated that since juveniles and sub-adults of both species are patterned like adults, it is possible that the duckweed-like pattern effectively concealed younger turtles from would-be predators. Shrubby Cinquefoil appeared to provide this same concealment for C. guttata. 42 Whitetail deer trails were prevalent throughout the research site (Figure 55). These trails appeared to facilitate C. guttata movement. Constant use by deer created depressions in the hydric soil. Saturation and standing water in these depressions provided “mini highway” systems for Spotted Turtles. These systems allowed for movement with ease through the fen. Movement may have increased food availability by exposing more foraging area and opportunities for mating and better population dispersal. For Spotted Turtles, predation is a major cause of mortality. The opportunity of escape using deer trails may have “hedged the bet” for survival. Also, the submergence of C. guttata in water may have provided a scent cover-up, keeping it safe from predator detection. In times of low precipitation or drought, deer trails are one of the last resorts for the provision of water. Spotted Turtles feed almost entirely underwater (Ernst etal, 1994; Harding, 1997). This allows for extended availability of open water allowing continuation of feeding. Also, mating generally occurs underwater (Ernst, 1967). When no water was present in the deer trails, the substrate remained the consistency of mud or muck. This appeared to allow for easier burrowing or movement for C. guttata. Disturbed areas of a habitat are more vulnerable to invasive plant species. In this study site, one invasive plant species of most concern is the Glossy Buckthom, found in human-impacted areas. Movement through Glossy Buckthom is difficult for Spotted Turtles. Monitoring and controlling invasive species must be a priority. If habitats are entirely encroached by invasive species, turtle populations will suffer. 43 One of the most effective techniques for Glossy Buckthom removal was suggested by Nate Simons, Executive Director and plant ecologist of Blue Heron Ministries in Indiana. Simons (pers. comm.) suggested using a hatchet to create a wound at the base of the plant. Wounds then should be treated using a squirt bottle filled with a strong herbicide. A onetime treatment of the herbicide performed during the winter season was found effective at eradicating Glossy Buckthom. Movement Discussion In 2007, initially, time was spent traversing the entire habitat in search of C. guttata. It became apparent there were areas where no turtles were observed. Efforts were reduced in these areas, and focused more on habitat where turtles were observed. Telemetry data supported this decision of not equally covering the habitat. Figure 56 represents a reasonable assessment of turtle utilization of the area. In this study, Minimum Convex Polygon (MCP) and Kernel Home Range Analysis were used in determining home range and its specific preferred habitat, homing, and overlapping. Cagle (1944) referred to the “homing behavior” of Spotted Turtles after being “forced” from their home areas during spring and late fall, later returning to their home range. He states the reason for these peripheral movements may have been the attempt of mate locating and/or hibernation. With the use of MCP or Kernel Home Range Analysis, these peripheral movements or “outliers” have been included in this study’s home range, and not considered homing behavior. Using spatial analysis, the animal’s entire 44 utilization distribution of an area can be documented, all providing more knowledge needed for C. guttata preservation. Comparing 2007 to 2008 data from the Spotted Turtle Distribution and Analysis of All Transmittered Turtles, the utilization of habitat in 2008 increased by approximately 1.3 hectares, which was not significant (Figure 11 and Figure 13). It is important to note that an additional six turtles were outfitted with telemetry equipment in 2008. This may be one explanation for small, but slightly increased area covered by Spotted Turtles in 2008. Standing water is necessary for Spotted Turtles to forage and mate, both occurring underwater. High amounts of rainfall and standing water were documented in 2008. With the addition of six telemetered turtles in 2008 and additional rainfall, one would surmise that the area of distribution would be substantially larger than 2007. This was not the case. Even with additional water in 2008, turtles did not venture far from 2007 data coordinates. This would suggest that basic needs were met without traveling any additional distance. Comparing 2007 to 2008 data from the Spotted Turtle Distribution and Analysis of All Nontransmittered Turtles in 2007, showed a slight increase in habitat utilization in 2008 by approximately 0.9 hectares, which was also not significant (Figure 12 and Figure 14). In 2007, male ABCK equipped with large transmitter (Channel 2), displayed a home range of 3.30 hectares. In 2008, male ABCK was again equipped with a transmitter (Channel 16), this time the small model. Home range recorded was 1.81 hectares. 45 In 2007, male IKW equipped with large transmitter (Channel 10), displayed a home range of 0.45 hectares. In 2008, Male IKW was again equipped with a transmitter, this time the small model (Channel 17). Home range recorded was 1.33 hectares. ABCK’s home range varied significantly in 2007 compared to 2008. This was also true for male IKW. Both individuals may be demonstrating home range site fidelity. Although ABCK’s 2007 home range was larger than 2008, the 2008 home range was embedded within the 2007 home range. Although lKW’s 2008 home range was larger than 2007, areas of these two home ranges overlapped. Having results from only two turtles using both small and large transmitters made it difficult to conclude whether or not there was a negative impact for turtles using large versus small transmitters. It is not necessarily true that a large transmitter impeded the turtle. With the large water volume differences throughout the habitat in 2007 versus 2008, it could be suggested that movement was less confined in 2008. This suggests that with more water present, areas for foraging and mating were more accessible. It is also conceivable with available standing water throughout the site; traveling great distances for activities (such as foraging or mate-seeking) were not needed. In 2007, home range sizes varied significantly, not only between males and females, but also among same sex individuals. Home ranges of male transmittered turtles varied from 0.45 hectares to 3.30 hectares. During the same season, female home ranges varied from 0.07 hectares to 1.12 hectares. 46 Each Spotted Turtle appeared to be individualistic, with no set movement pattern, covering the amount of land needed to fulfill requirements for survival. These requirements may have included foraging, mating, nesting, aestivation and hibernation. Ernst’s (1970) findings were that home ranges of Spotted Turtles usually did not include a suitable nesting area; therefore, females had to migrate out of their home range. Not true in this study. Home ranges included not only the females’ nesting area, but also areas for both male and female foraging, mating, aestivation (if applicable) and hibernation. When comparing male 2008 home ranges, ABCK (Channel 16) and IKW (Channel 17) were similar in size. Male KUV’s (Channel 2) recorded home range, in comparison, was smaller as he was found dead in July 2008. This small sample size makes it difficult to speculate commonalities of 2008 male home ranges. However, home ranges for females in both 2007 (0.05-1.12 hectares) and 2008 (0.12-1.33 hectares) varied significantly. Because of large female sample size versus small male sample size, it is difficult to make any conclusions about home range without further study and data collection. One noteworthy observation takes into consideration that habitat utilization by both transmittered males and females was within preferred habitat area previously discussed in detail. Homing_a_nd Overlapping Discussion During this study, two turtles (female KVX and female HKV) returned to their original home ranges, suggesting homing ability. This supports the findings 47 of Breder (1927), Nichols (1939), Medsger (1919), Schneck (1886), Grant (1936) and Ernst (1968). Continued research in this area would be useful in determining if more turtles utilize homing ability. Overlapping of home ranges by both males and females were documented in 2007 and 2008. This suggests there was no territoriality in this study site, which agrees with Ernst’s (1970) findings. This may prove to be advantageous to both males and females, keeping the sexes in close proximity to one another for mating purposes. Also, encountering of food may prove to be easier when foraging within close proximity to other individuals. Locating of communal hibemaculae may prove to be a positive result with overlapping of home ranges. A disadvantage of overlapping Spotted Turtle home ranges makes them vulnerable to predators and collectors. 48 Chapter 2: Population Dynamics and Demography Introduction Population breakdown of males, females and juveniles vary in C. guttata studies. Breisch (2006) found in her West Virginia study, a population of 21 Spotted Turtles. This population included 38% males, 19% females, and 43% juveniles. In Ontario, Litzgus (1996) found males and females to be consistent at 47% each, and juveniles were 6% of the population of 171 turtles. In 2004, Litzgus and Brooks (1997) reported 118 Spotted Turtles in their Georgian Bay, Ontario site. This population included 42% males, 49% females, and 9% juveniles. In South Carolina, Litzgus and Mousseau (2004) studied a population of 44 Spotted Turtles. This study included a population breakdown of 39% males, 48% females, and 14% juveniles. In previous studies, the sex ratio for male and female C. guttata was found to be of approximately equal distribution, with a 1:1 ratio. These studies included: Litzgus (1996), Litzgus and Brooks (1997), Graham (1995) and Litzgus and Mousseau (2004). Breisch (2006) found a sex ratio of males to females of 2:1. The occurrence of sexual dimorphism varies with populations. Documented average carapace length for males and females varies depending on the population. Average carapace lengths for males have been recorded as 10.26 cm (Breisch, 2006), 11.69 cm (Litzgus, 1996) and 11.33 cm (Graham, 1995). Average carapace lengths documented for females were 9.92 cm (Breisch, 2006), 11.517 cm (Litzgus, 1996) and 11.02 cm (Graham, 1995). 49 Gibbons and Lovich (1990) found that on average, female C. guttata were larger than males, but Breisch (2006), Litzgus (1996) and Graham (1995) found male dominated sexual size dimorphism. Ernst (1970) stated that sexual maturity in both sexes of the Spotted Turtle is obtained when the carapace length of the turtle reaches 8.0 cm. Ditmars (1933) stated that C. guttata were fully mature when the carapace length reached four inches long (10.16 cm). Sexual maturity was obtained in both sexes between 7 and 14 years of age (Harding, 1997; Ernst, 1975; Graham, 1970). Population density has been studied in areas of the Spotted Turtles’ range. In West Virginia, Breisch (2006) reported a density of 6.8 turtles per hectare. Litzgus and Mousseau (2004) estimated the population density in South Carolina as 0.36 turtles per hectare. Litzgus (1996) reported a population density of 0.62 turtles per hectare in Ontario. Because Spotted Turtle research is ongoing, mortality rates of populations have been poorly documented. Litzgus (1996) found a relatively “high adult mortality rate” in her Ontario population. Dialecti—vee: The specific objectives contained in this chapter were to determine Spotted Turtle population dynamics and demography. Population size, population breakdown of males, females and juveniles and their morphometric data were specifically researched. Obtaining sex ratios, understanding sexual dimorphism and stability were also investigated. 50 Methods Methods were previously described in Chapter 1. Bfiulfi From March 2007 until March 2009, 187 Spotted Turtles were documented in this study site (Table 9). This number represented 72 juveniles (38%), 61 males (33%) and 54 females (29%) (Figure 57). The sex ratio for males and females appeared to be 1:1, further supporting the approximately equal sex distribution ratio stated by Litzgus (1996), Litzgus and Brooks (1998), Graham (1995) and Litzgus and Mousseau (2004). At the onset of this study 79 juveniles were documented with seven being identified as males. For purposes of sexual identification, Ernst’s (1970) defined sexual maturity as carapace length of 8.0 cm and above. However, female characteristics (for example, flat plastron and shorter tail) are juvenile characteristics and male characteristics (concave plastron and longer tall) are obtained during the maturation process. Current data showed sexual maturity of these seven males were obtained before reaching this 8.0 cm carapacial length. Thus, these seven juvenile males were documented as adult males, lowering the number of juveniles to 72. Annuli were counted to determine the minimum age of the Spotted Turtle (Litzgus and Brooks, 1998). Older adult turtles would show worn or smooth annuli and cessation of annuli growth; making it difficult to calculate their exact age. Therefore, the age of older adults may be the minimum age, not the maximum. The minimum possible age of the oldest turtle documented at the site 51 was a 22 year old female. The lowest mean minimum age of an adult Spotted Turtle was 9.2 years in this site. Mean morphometric measurements were documented for most Spotted Turtles observed in the study site. If strictly conforming to Ernst’s (1970) definition of sexual maturity (adult having the 8 cm or above carapacial length), the seven turtles documented as males would have been juveniles. In this study, turtles with male characteristics (concave plastron and longer tails) were recorded as males, making the average female larger than the average male. This supports the findings of Gibbons and Lovich (1990) which show female Spotted Turtles to be larger than male Spotted Turtles. The mean carapace length for male Spotted Turtles was 8.9 cm, with the largest male having a carapace length of 10.9 cm (range 65-109 cm). The mean carapace length for females was 9.0 cm, with the largest female having a carapace length of 10.3 cm (range 8.0 — 10.3 cm). Breisch (2006), Litzgus (1996) and Graham (1995) found males to have a larger average carapace length than females. The mean carapace width for males was 6.4 cm (range 5.1 — 7.7 cm), with females having a mean of 6.6 cm (range 5.9 - 7.8 cm). Males recorded a mean plastron width of 7.5 cm (range 5.7 — 8.6 cm) while a mean of 8.0 cm was recorded for females (range 7.2 — 8.5 cm). Mean height in males and females was 3.3 cm (range 2.6 — 3.7 cm) and 3.7 cm (range 3.0 — 4.1 cm), respectively. Males recorded an average weight of 0.10 kg (range 0.05 - 0.16 kg). Females recorded an average weight of 0.11 kg (range 0.08 — 0.14 kg). 52 This study took place in a fen with an approximate size of 30 hectares. Representative sampling of the entire fen was accomplished at the onset of the study. It was noted earlier in Chapter 1, that the outlying areas of the fen were void of turtles, and as stated earlier, continuing to traverse through these areas was not time well spent. Population density was based on 30 hectares, even though the area of actual turtle concentration was less. The turtle population density was calculated at approximately 6.2 turtles per hectare. During the 2007 and 2008 research, a total of 10 turtles (with and without transmitters) were found dead due to various reasons. Two Spotted Turtles died by becoming entangled in the Chara Pond. One turtle found dead in the Chara Pond had been predated upon by a probable Muskrat. Two turtles were found dead due to unknown causes. Two empty, fully intact shells were found during the two year study, along with shell fragments (which may have been very old) of three other turtles. The mortality rate of this study site was calculated to be 0.05%. Discussion Females were larger than males in this study site. The results of this study support findings of Gibbons and Lovich (1990). It should be noted that turtles in this Michigan study site appear to be smaller overall than findings of other Spotted Turtle researchers. One explanation may be turtles are maturing at an earlier age than in other sites, therefore, not reaching their maximum size capability. 53 It was recorded that males and females have a similar average weight. Although, the mean average was approximately equal, one must take into consideration gravid female’s weight fluctuation due to egg storage. If females were recorded prior to ovipositioning, their weights would be higher than after they have nested. The population density calculated from this study was similar to findings found by Breisch (2006) but was larger than those recorded by Litzgus and Mousseau (2004). With a high hatchling and juvenile rate of 38%, and overall adult mortality rate of 0.05%, the mortality rate of C. guttata appeared to be low. This study was conducted for two years. More long term data is needed to determine if this mortality rate remains consistent. Considering only ten dead animals were found and the large percentage of juveniles and hatchlings sighted, C. guttata appeared to be a stable or growing population in this study site. 54 Chapter 3: Seasonal Activities Introduction Aestivation Ernst et al (1994) noted activity levels reached a peak when mean monthly air temperatures were between 13.1 °C and 180°C (mean 155°C); at least two months before the highest mean air temperature month. Activity then declined when mean monthly air temperature was between 178°C and 223°C (mean 203°C), and then approached or reached a minimum level during the month with highest mean air temperature. This decline in activity was referred to as aestivation. Perillo (1997) defined aestivation as a period of dormancy punctuated by periods of brief activity. Spotted turtles were documented to be able to survive months in this dormant state, which is considered a summer equivalent of hibernation (Carroll, 1991). It had been suggested that “summer dormancy” may be a more appropriate term than aestivation to describe the behavior of northern populations of Spotted Turtles when they become inactive in the late summer (COSEWIC, 2004). Litzgus (1996) stated that aestivation is like a summer version of hibernation, in theory, to avoid increased temperatures rather than decreased temperatures. This behavior typically involved burying under leaf litter and pine needles in the shade of junipers or rock outcroppings or in forests. Populations of Spotted Turtles that live at more southern latitudes aestivated to avoid prolonged high summer temperatures. However, in northern populations, the 55 advantage of this behavior was less obvious. Litzgus (1996) found that turtles were not decreasing their body temperature by aestivating; therefore, they were not using aestivation to avoid increased temperatures. This behavior by Spotted Turtles in Georgian Bay, Ontario, suggested it may be carried over from a time when it served a completely different function. In Pennsylvania, Ernst (1976) found that Spotted Turtles become inactive when water temperatures reached 32°C, seeking out cool refuges such as muskrat burrows or mud at the bottom of pools of running water. Most turtles remained inactive until the following March but some emerged during the cooler days of July, August and the fall, and the warmer days of winter. Ernst (1982) stated C. guttata thermoregulate in their northern range by basking during cooler weather and by burrowing into the soft bottom of a watenivay or by entering muskrat burrows and lodges to avoid extreme hot (aestivation) and cold conditions (hibernation). Water and substrate covering C. guttata formed an effective thermal shell which provided adequate insulation for avoidance of extreme temperatures both hot and cold. Muskrat burrows and lodges contained underwater entrances. In the summer, the evaporation of the water in these burrows and lodges provided a cooler microhabitat and high humidity that helped prevent moisture loss. In Maryland, Ward et al (1976) stated aestivation sites varied depending on the location of the range of the Spotted Turtle. Aestivation sites included . moist areas under dead and loosely matted reeds, grasses and ferns. He also documented turtles in early successional paludal woods. This form of 56 microhabitat was characterized by heavy organic soils, shallow water, and an overstory of dense leaves, grasses and ferns. lndentations in the substrate, referred to as “forms” used by C. guttata are filled with water so that only the top of the carapace are visible. Haxton and Berrill (2001) noted in central Ontario, turtles were found in forms similar to those discussed by Ward et al (1976). They found that the temperature of the turtles was no different than the water temperature, indicating avoidance of warm air temperatures may not be the primary objective of aestivation as suggested by Ernst (1976, 1982). Other aestivation sites documented are muskrat burrows in banks of streams and lodges (Ernst, 1982), predominantly bogs (Haxton and Berrill, 1999), upland habitats and the dried edge of a vernal pool (Perillo, 1997), and terrestrial sites (Litzgus and Brooks, 2000; Ultsch, 2006; Barlow, 1999). Perillo (1997), whose Spotted Turtle research was conducted in Connecticut, noted dry summers with little rainfall, may stimulate turtles to aestivate upland for longer amounts of time than during a summer with more normal precipitation. Time spent in aestivation varied depending on site location. In Massachusetts, Graham (1995) found aestivation to last from one to three weeks. In Indiana, Barlow (1999) found turtles aestivating from two days to two weeks, with some turtles not aestivating at all. Hibernation Ultsch (2006) found that most species of turtles faced harsh winters causing them to hibernate for extended lengths of time that may exceed over half of their lives (more than six months per year). The ability or lack of it, to 57 ovenNinter in hypoxic/anoxic conditions places limitations on where aquatic turtles can overwinter, and perhaps on where they can occur at all. Litzgus et al (1999) and Lewis and Ritzenthaler (1997) documented turtles entering a hibemaculum as early as mid-September and as late as April if ice had not sealed the entrance to the hibemaculum. Lewis and Ritzenthaler (1997) found that turtle departure positively correlated to the warming of water in hibemaculum and melting of ice in the hole. Spotted Turtle hibemaculae were typically aquatic beneath a layer of ice and mud (Ward et al, 1976; Ultsch, 2006). These sites never froze completely, but were shallow enough to thaw quickly in spring (Ernst, 1982). The majority of hibernation took place in wetlands and small pools in various microhabitats; one common place being beneath hummocks in swamps (Ward et al, 1976; Graham, 1995; Perillo, 1997; Joyal etal, 2001; Litzgus etal, 1999; Milan and Melvin, 2001). Hibemaculae have been known to contain a saturated, organic muck layer (Lewis and Ritzenthaler, 1997). Communal aggregations have been documented in central Ontario (Haxton and Berrill, 1999), Ohio (Lewis and Ritzenthaler, 1997), Pennsylvania (Ernst, 1967), Connecticut (Perillo, 1997) and northern Ontario (Litzgus et al, 1999). Small aggregations of Spotted Turtles were common, but groups have been documented to contain as many as 34 turtles (Lewis and Ritzenthaler, 1997). Hibemaculae near its northern limits in Ontario contained single turtles up to nine turtles. Litzgus et al (1999) noted that hibemaculae were of two types both in swamps: elevated Sphagnum spp. moss hummocks reinforced by roots 58 and stems of vegetation, including trees and shrubs; rock caverns near shore with a measured water depth of 30-40 cm. Lewis and Ritzenthaler (1997) determined hibemaculae that contained three or less turtles were simple vertical holes that were approximately seven cm in diameter and no deeper than 70 cm. Larger hibemaculae were 15 by 20 cm at the surface entrance and contained some horizontal passages in addition to the primary vertical hole. Further, Lewis and Ritzenthaler (1997) found no sex biased hibemaculum use. Although large aggregations present obvious vulnerabilities to predators and collectors, benefits were also found. Communal hibernation provided reproductive advantages by putting otherwise solitary turtles in close proximity during early spring for mating (Ernst, 1970; Lewis and Ritzenthaler, 1997; Perillo, 1997). Barlow (1999) found aggregations may have made it easier for males to find females and that these oven~intering locations may have had some historical value of C. guttata when located near favorite spring breeding locations. Communal hibernation may have improved fitness of individuals by synchronizing emergence (Gregory, 1982; Ultsch, 1989). Hibemaculum site fidelity was observed in Spotted Turtles. Research conducted in Central Ontario (Haxton and Berrill, 1999), Ohio (Lewis and Ritzenthaler, 1997), Northern Ontario (Litzgus et al, 1999), and Connecticut (Perillo, 1997), showed that this turtle species displayed high fidelity to hibemaculae. 59 Objectives The objectives of this study were to determine seasonal activities of the Spotted Turtle which included aestivation (determining if it does occur, and if so, when, how long, habitat temperature, and type of habitat preferred). Hibernation of Spotted Turtles was also studied to determine the preferred hibemaculae. Also researched was whether or not the hibemaculae were communal, substrate properties, location, hibemaculae temperature, and possible site fidelity. Methods Methods were previously described in Chapter 1. Aestivation Results and Discussion Aestivation is a period of brief inactivity. In this fen, aestivation coincided with high air temperatures. In 2007, turtles were tracked throughout late summer, with rising air temperatures causing the land to warm and turtles to become inactive. The majority of telemetered turtles did aestivate. Length and period during which aestivation occurred varied between 4 and 26 days, with some turtles coming out of aestivation for a short period of time and then returning again to inactivity. Aestivation in this southwestern fen occurred within the months of June, July and August. Nine turtles aestivated, with six aestivating twice. Aestivation coincided with the drier parts of summer. Field data entries in mid June and July showed at these aestivation times, the substrate consisted of “dried mud” with only traces of precipitation recorded. In 2007, turtles aestivated in areas which corresponded with areas shown in Figure 58. Areas included the open middle fen area, ditch, outlet, area of short 60 height hydrophytes, and a more open area just north of ditch. These areas were lower in elevation, with water levels just beneath the substrate, even during drier penods. With water absent, mud temperatures were recorded. In June and July Of 2007, seven turtles aestivated (some more than once) with mud temperatures ranging from 155°C to 290°C. Air temperatures ranged from 179°C to 324°C. Two turtles aestivated in August when the mud temperatures were recorded between 19°C and 26°C and the air temperatures recorded were between 21 .4°C and 32°C. There was more precipitation in 2008 than 2007. Seven turtles aestivated, most occurring late July thru early September. All turtles aestivated once during this season, except for one turtle (aestivation occurred twice). The majority of turtles aestivated for nine days when the mud temperature was 205°C to 29.1 °C and the air temperature was 245°C to 37°C. Two turtles aestivated for a period of 20 days during which the mud temperature was 21 .6°C to 27°C, air temperature was 235°C to 345°C, and water temperature (in one location) was 31°C. Turtle aestivation locations in 2008 were found in areas of habitat where large turtle aggregations were located, as shown in Figure 59. These areas included the open middle fen area, ditch, and big trail to lake including the lake’s edge and the area of short height hydrophytes. These areas were lower in elevation and contained water even throughout the drier months. 61 In both 2007 and 2008, turtles aestivated by burying themselves in mud covered with a variety of hydrophytes. These hydrophytes included Sedge, Shrubby Cinquefoil, Typha, Bulrush, Red-osier Dogwood, Willow and Fern. Hibeflation Results and Discussion Radio telemetry was used to track fall season movement and location of Spotted Turtles. Locations were identified and documented as possible hibemaculae. Turtles were tracked late in summer and early autumn (September and October). Two areas were found to have a heavier aggregation which appeared to be hibemaculae locations. In spring (March), larger numbers of C. guttata were observed and captured in these same areas, suggesting emergence from close-by hibemaculae. With less recorded precipitation during 2007, there was no standing water throughout the site; water table present just below substrate surface. At this time, turtles were no longer visible at surface and radio telemetry was used to locate turtles. On 25 September 2007, using radio telemetry to locate transmittered female Channel 11, a possible hibemaculum was identified in a lower elevation fen area. The hydrophytes in this area included Typha, Sedge (Scirpus pungens) and Bulrush (Scirpus acutus). While attempting to locate Channel 11 (female KV) by its frequency, a hole, approximately 15 cm deep, was located in the substrate containing no water, only damp mud. Reaching into the hole, four turtles were stacked, one below the other, all in horizontal positions. Channel 11 was found with two females (BDKW and HKP) and a male (AJKX) (Figure 60). 62 Another hole, similar to the one which contained the four turtles, was observed nearby. Reaching approximately 13 cm into this damp, muddy hole, three turtles were collected, each on top of the other. All three Spotted Turtles were male (CDKV, FJKS, CDKVW). With the use of telemetry, Channel 2 (male ABCK) was found only one foot from the hole where the three males were found. With further investigation, female CKX was located nearby buried in mud, Spaghnum spp., and Sedge (Figure 60). A week later, 2 October 2007, while attempting to locate Channel 11 (female KV), another possible hibemaculum was sighted. By locating Channel 11 using its radio telemetry frequency, a hole in the substrate was observed. Reaching into the hole, four turtles were collected; three females (HKP, CKX, KV Channel 11) and 1 male (AJKX). Turtles were found stacked on top of each other ranging from 13-33 cm downward. The hole contained both mud and water (Figure 61). AJKX, KV, and HKP had previously been located (25 September 2007) in a different hibemaculum locale. On 12 October 2007, a possible hibemaculum was reinvestigated known to contain turtles a week prior. Standing over the entrance to the possible hibemaculum, no turtles were visible from the surface, but the small 7.62 cm hole was observed. Reaching down into the vertical hole, filled with water, mud and muck, a non-transmittered Spotted Turtle was located. Continuing to reach down into this hole another conspecific turtle, a little deeper horizontally to the first turtle was retrieved. Small tunnels, approximately three to six cm in size, were filled with water and were intertwined with roots of Sedge spp., Typha and other 63 hydrophytes. It is believed that these tunnels allowed easy movement for Spotted Turtles from surface predators. Within this hole three non-transmittered turtles were identified. The first turtle removed, a male (BCKV), was located approximately nine cm from the ground level and had a recorded carapace shell temperature of 118°C. Reaching back into the hole, a second male turtle (IKP) was located just below the first turtle. The shell temperature was IKP was recorded at 123°C. Reaching again into the hole, a third turtle, a female (KUW) was located slightly lower in the hole and horizontally to the right approximately 15 cm from the original vertical hole. The third turtle had a shell temperature of 13.3 ’0 (Figure 62). On 16 October 2007, another possible hibemaculum was found in an area where Sandbar Willow was the main hydrophyte. While traversing through the site, a male Spotted Turtle (CDKV) was observed with only its carapace showing. Upon further observation, it was determined that the male turtle was located over a hole of possible hibemaculum. With further investigation, it was determined that three additional turtles were located stacked below the substrate in water and mud in a hole approximately 25 cm deep. The carapace shell temperature for the turtle located on top of the substrate was 14.5°C. Downward in the hole, a juvenile (KOQ) was collected with shell temperature of 145°C. Beneath the juvenile, a juvenile (IKNP) was documented with a shell temperature of 155°C. Lastly, another juvenile (AKP) was documented with a shell temperature of 167°C (Figure 63). 64 As turtles were collected deeper in the possible hibemaculum, carapace shell temperatures increased. Although, Lewis and Ritzenthaler (1997) found no evidence that communal hibemaculae serve any thermal function, this study suggests the opposite. Evidence from this study implies that the deeper into the hibemaculum a turtle is found, the warmer the body temperature. These possible hibemaculae, which were in lower elevation areas of the fen, were found with both single turtles and aggregations. Most hibemaculae ranged from eight to 35 cm in depth with some containing small tunnels that formed interconnected networks under the substrate. These tunnels were too small in size to be considered muskrat burrows. A possible explanation for the cause of these undenivater passages may be the steady flow of groundwater throughout the root systems. Another area of heavy Spotted Turtle concentration known as the “ditch” was to the south of the fen, running east and west. It handled overflow from the wetland and a nearby road. The ditch was generally filled with approximately 40- 60 cm of water and hydrophytes which included Typha, Red-osier Dogwood, Sedge and Bulrush. In spring of 2008, a heavy concentration of turtle emergence was observed from the ditch. On 3 April 2008, four turtles (CKW-juvenile, KLN-male, AKM-female, DKO- male) emerged and captured from the ditch and GPS coordinates were recorded. Three days later, 6 April 2008, four different individual turtles (KMW-female, KUV-male, KLM-male, and AK-male) were observed and documented close to this same location. Turtles were found 65 emerging periodically during the next several weeks in this same locale. These observations suggest the ditch was a heavily populated hibemaculum. Spotted Turtles selected areas of the wetland for hibernation that are generally underwater or with the water table at ground level. Turtles preferred areas that had the deepest concentrations of water in the fen. Both possible hibemaculae were located within these areas. In spring, when C. guttata emerged from hibernation in these two known hibemaculae, both sites were inundated with water from 40 to 60 cm deep. Shortly after emergence, turtles were observed moving from deep to shallow water. The deep water temperature ranged from 60°C to 85°C while the temperature in the shallow water ranged from 140°C to 185°C. The observation of Spotted Turtles migrating to shallower water confirms that C. guttata, being ectotherrnic, may be moving to a warmer habitat to raise metabolic rates. In late summer and early fall of 2008 (September and October), with heavy amounts of rainfall, 2-40 cm of standing water covered the fen. Water levels in the first known hibemaculae (the area in the middle of the fen) were approximately 40 cm. Although tracking the transmittered turtles to these areas was no problem, locating holes and reaching into the substrate was a difficult undertaking. With deep water, locating turtle aggregations proved difficult. When radio telemetry was conducted, and turtles located, these potential hibemaculae contained some of the same and new conspecifics than previously documented, including different combinations of males, females and juveniles. 66 This infers that turtles may be “staging” or trying to locate the most viable locations for future courtship which may yield genetic diversity. On 12 January 2007, four Spotted Turtles were observed during a “January thaw” in the ditch. Two individual male turtles were observed, as well as a copulating pair (Lutz, 2008). This observation coincides with Ernst’s (1970) suggestion that communal hibemaculae serve a purpose for turtles in their relationship to mating preference. Males, females and juveniles were all found within the same hibemaculum, suggesting that hibemaculae are not sex biased. This evidence aligns with a previous study by Lewis and Ritzenthaler (1997). Previous suggestions that the two main areas discussed (ditch and open middle area of fen) were hibemaculae areas for the Spotted Turtle were further supported by evidence shown using Arc View© and Arc Map©. Figure 56 (Kernel Home Range Analysis) also substantiates these findings. Not only were transmittered turtles found to hibernate in these two main regions, but also non- transmittered turtles were recorded to hibernate there, as well. This evidence further supports the findings that Spotted Turtles do communally hibernate. Site fidelity of a hibemaculum was documented in 2007 and 2008 in the ditch and open middle area of fen. Eight of 23 (35%) transmittered turtles were documented having site fidelity. It should be noted that 29 turtles were originally equipped with transmitters, with two transmitters malfunctioning, and four turtles found dead. Further research is needed to determine whether the other 15 turtles show site fidelity. 67 With the telemetry equipment used in this study, hibemaculae temperatures were obtained (refer to Methods). A low winter hibemaculum temperature of 13°C was reported in the beginning of March 2008. In Ohio, Lewis and Ritzenthaler (1997) recorded low winter hibemaculae temperatures of 2.21 and 273°C. The findings of this study are in close proximity to the Ohio findings. 68 APPENDIX 69 Table 1. Specific habitat preferences documented throughout Spotted Turtle range. Habitat Type Location Source Forested Swamps Maine Joyal et al 2001 Boreal Fen Ohio Lovich 1989 Terrestrial Sites Ontario Haxton and Berrill 2001 Florida Berry 1978 Rock Outcrops Ontario Haxton and Berrill 2001 Ontario Litzgus et al 1999 Low Acidic Swamps Ontario Litzgus et al 1999 Small Bodies of Water Salt Marshes/Brackish Meandering Brooks Little Bog Holes Ditches Vernal Pools Early Successional Fields Upland Fields Older, more established woodlands Cypress swamps Freshwater marshes Mixed Hardwood Wetland Cypress Tupelo Ponds Ephemeral Marshes Upland Pine-Hardwood Forest throughout range Illinois Maryland throughout range throughout range throughout range Indiana Connecticut Massachusetts Maine Connecticut Connecticut Massachusetts Connecticut Florida Florida Pennsylvania Florida South Carolina South Carolina South Carolina South Carolina South Carolina Carr 1952 Cahn 1937 Nemuras 1966 Carr 1952 Carr 1952 Conant and Collins 1998 Minton 1972 Perillo 1997 Fowle 2001 Joyal et al 2001 Perillo 1997 Perillo 1997 Fowle 2001 Perillo 1997 Barnwell et al 1997 Barnwell et al 1997 Ernst 1976 Barnwell et al 1997 Lovich 1990 Litzgus and Mousseau 2004 Lovich 1990 Lovich 1990 Litzgus and Mousseau 2004 Flooded Mowed Fields Maryland Ward et al 1976 Maine Joyal et al 2001 Indiana Minton 1972 Woodland/Marsh Ecotones Maryland Ward et al 1976 Emergent Marshes Massachusetts Fowle 2001 Unpolluted Bodies of Water Massachusetts Graham 1995 throughout range Harding1997 7O Common Name Table 2. Non-woody plants located in study site. Scientific Name Small-flowered Gerardia Bugle Common Burdock Mugwort Swamp Milkweed Northern Bog Aster Bushy Aster New England Aster Frost Aster Willow Aster Tickseed Sunflower False Nettle Downy Chess Marsh Bellflower Chicory Canada Thistle Swamp Thistle Twig Rush Queen Anne's Lace Water Willow Common Wood Fern Daisy F leabane Spotted Joe Pye Weed Boneset Lesser Fringed Gentian White Avens Ground Ivy Spotted Touch-me-not Blue Flag Iris Canadian Rush Torrey's Rush Rice Cutgrass Marsh Blazing Star Kalm's Lobelia Northern water Horehound Tufted Loosestrife Purple Loosestrife Sweet-scented Waterlily Common Evening Primrose Cowbane Panic Grass Switch Grass Agalinis tenuifolia Ajuga reptans Arctium minus Artemisia vulgan's Asclepias incarnate Aster borealis Aster dumosus Aster novae-angliae Aster pilosus Aster praealtus Bidens coronatus Boehmeria cylindrical Bromus tectorum Campanula aparinoides Cichon‘um intybus Cirsium arvense Cirsium muticum Cladium man'scoides Daucus carota Decodon verticillatus Dryopten‘s spinulosa En'geron annuus Eupaton'um maculatum Eupaton'um pen‘oliatum Gentiana procera Geum canadense Glechoma hederaceae Impatiens capensis Iris versicolor Juncus Canadensis Juncus torreyi Leersia oryzoides Liatris spicata Lobelia kalmii L ycopus uniflorus L ysimachia thiysiflora L ythrum salicaria Nymphaea odorata Oenothera biennis Oxypolis rigidior Panicum flexile Panicum virgatum Swamp-Betony Pediculan’s lanceolata Timothy Phleum pretense Water Smartweed Polygonum amphibium 71 Table 2 (cont’d). Common Name Scientific Name Water Pepper Meadow Spikemoss Canada Goldenrod Ohio Goldenrod Swamp Goldenrod Indian Grass Peat Moss Marsh Fern Marsh St. Johnswort Cattail Cattail Small Purple Bladderwort Blue Vervain Rattlesnake Root Black-eyed Susan Bouncing Bet Hardstem Bulrush Common Threesquare Polygonum h ydropiper Selaginella apoda Solidago Canadensis Solidago ohiensis Solidago patula Sorghastrum nutans Sphagnum spp. Thelypten's palustn's Triadenum virginicum Typha angustifolia Typha Iatifolia Utn'cularia resupinata Verbena hastate Prenanthes racemosa Rudbeckia hirta Saponaria ofi'rcinalis Scirpus acutus Scirpus pugqens Table 3. Woody plants located in study site. Common Name Box Elder Red Maple Sugar Maple Shad spp. Bog Birch Oriental Bittersweet Northern Swamp Dogwood Red-Osier Dogwood White Ash Winterberry Honeysuckle spp. Virginia Creeper Cottonwood Bigtooth Aspen Trembling Aspen Shrubby Cinquefoil Wild Black Cherry Red Oak Black Oak Alder-leaf Buckthom Glossy Buckthom Black Locust Carolina Rose Multiflora Rose Scientific Name Acer negundo Acer rubrum Acer saccharum Amelanchier spp. Betula pumila Celastrus orbiculata Camus racemosa Cornus stolonifera Fraxinus Americana llex verticillata Lonicera spp. Parthenocissus quinquefolia Populus deltoids Populus grandidentata Populus tremuloides Potentilla fruticosa Prunus serotina Ouercus rubra Quercus velutina Rhamnus alnifolia Rhamnus frangula Robinia pseudoacacia Rosa Carolina Rosa multiflora Swamp Rose Rosa palustris Common Blackberry Rubus allegheniensis Black Raspberry Rubus occidentalis Dewberry Rubus spp. Blue-leaved Willow Salix myricoides Willow spp. Salix spp. Bittersweet Nightshade Solanum dulcamara Poison Ivy Toxicodendron radicans Poison Sumac Toxicodendron vernex Elm spp. U/mus spp. G rape Vitis riparia 73 Table 4. Wildlife documented in Common Name Meadow Vole White-Tailed Deer Muskrat Eastern Cottontail Rabbit Coyote Red Fox Raccoon study site. Scientific Name Microtus pennsylvanicus Odocoileus virginianus Ondatra zibethica Sylvilagus floridanus Canis Iatrans Vulpes fulva Procyon lotor Table 5. Amphibians and reptiles documented in study site. Common Name Western Chorus Frog Spring Peeper Wood Frog Brown Snake Eastern Garter Snake Northern Water Snake Northern Ribbon Snake Midland Painted Turtle Queen Snake American Toad Green Frog Bullfrog Common Snapping Turtle Spiny Softshell Eastern Hog-nosed Snake Eastern Box Turtle Eastern Grey Treefrog Common Musk Turtle Blanding's Turtle Spotted Turtle Eastern Massasatm Rattlesnake Scientific Name Pseudacris triseriata triseriata Pseudacris crucifer Rana sylvatica Storeria dekayi Thamnophis sirtalis sin‘alis Nerodia sipedon sipedon Thamnophis saun'tus Chrysemys picta Regina septemvittata Bufo americanus Rana clamitans Rana catesbeiana Chelydra serpentine Apalone spinifera Heterodon platiminos Terrapene carolina carolina Hyla versicolor Sternotherus odoratus Emydoidea blandingii Clemmys guttata Sistrurus ca tenatus ca tenatus 74 Table 6. Bird species present in study site. Common Name Scientific Name Red-winged Blackbird Swamp Sparrow Song Sparrow American Goldfinch America Robin Blue Jay Sandhill Crane Mute Swan Canada Goose Common Snipe Common Grackle American Crow Turkey Vulture Eastern Phoebe Wood Duck Mallard Duck Mourning Dove Hairy Woodpecker Downy Woodpecker Belted Kingfisher Tree Swallow American Woodcock Double-Crested Cormorant Carolina Wren Tufted Titmouse Black-capped Chickadee Northern Cardinal Ring-billed Gull Yellow Warbler Common Yellowthroat Warbler Yellow-rumped Warbler Palm Warbler Willow Flycatcher Red-tailed Hawk Virginia Rail Eastern Bluebird Rose-breasted Grosbeak Warbling Vireo Eastern Wood Pewee Baltimore Oriole Eastern Kingbird Cedar Waxwing Rough-winged Swallow Spotted Sandpiper Gray Catbird Black-billed Cuckoo Green Heron 75 Agelaius phoenlceus Melospiza Georgiana Melospiza melodia Carduelis tristis Turdus migratorius Cyanocitta cristata Grus Canadensis Cygnus olor Branta Canadensis Capella gallinago Quiscalus quiscula Corvis brachyrhynchos Cathartes aura Sayornis phoebe Aix sponsa Anas plalyrhynchos Zenaida macroura Picoides villosus Picoides pubescens Megaceryle alcyon Tachycineta bicolor Philohela minor Phalacrocorax auritus Thryothorus ludovicianus Parus bicolor Parus atricapillus Cardinalis cardinalis Larus dela warensis Dendroica petechia Geothlypis trichas Dendroica coronata Dendroica palmarum Empidonax trail/ii Buteo jamaicensis Rallus Iimicola Sialia sialis Pheucticus ludovicianus Vireo gilvus Contopus virens lcterus galbula Tyrannus tyrannus Bombyci/la cedrorum Stelgidopteryx ruficollis Actitis macularia Dumete/la carolinensis Coccyzus erythropthalmus Butorides striatus Table 6 (cont’d). Common Name Scientific Name Yellow-throated Vireo Great Crested Flycatcher White Throated Sparrow Scarlet Tanager Brown-headed Cowbird Ruby-throated Hummingbird Wood Thrush Cooper's Hawk Great Blue Heron Broad-winged Hawk Blue Gray Gnatcatcher Eastern Meadowlark Osprey Northern Waterthrush Veery Vireo fla vifrons Mylarchus crinitus Zonotrichia albicollis Piranga olivacea Molothrus ater Archilochus colubris Hylocichla mustelina Accipiter cooperii Ardea herodias Buteo playpterus Polioptila caerulea Sturnella magna Pandion haliaetus Seiurus noveboracensis Catharus fuscescens 76 Table 7. Example of Microsoft Excel individual turtle data worksheet. 835E 33!. $2 8.8 mm 9 9.0 in. NN— g Eo§3£mhmm2§=g3ousl - . - - ; r 39.9, >8: .EQSEE SSEESS. , . 9rd , In. new g 52.3 4 ounce E0223 0.9 3 mg in. 8N g EEoE 82 82.3.6.3 Eu m N I . ,,-..-l:l,i 388 E; E53852 E_ 28E 88E... SEES-22,2233 .. EN. E. EEG . 2.2me g .222 Eu mm a 93.. .225 .5 m— s .823 a E ESE 2 EEEEEE _ - 828932.... a... . 0. E.» o. 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E9; E2... .E 83m 22532 d 8.8 BNN 2< m.“ I 88 52:20.5 .3an 02.3 .5...) .EE .EEoE .22... 3.52 done 93 EE .2... 3.. Eu .8282 In. 8N 8:3 .2. £92.. 26.. 2.2.8.... 35.232. .8. 2E2 uaE .Eofixosn SEN 8.8 9:. En. SN 82% $83 a. 5.532 62...... £5. EucaoE ES .32.. E. 3...... mm? E22 .3: nausea §H_EauHu-Hdaflu.ldafl 82E Hum 8.233 «ad 5...; E< 5...... __.EE 29.2. E:: 3.3. $2.25.... ”2.22.5222. 2.... 2.. .255. .5... 3.23.. 2a: P2222231, 78 Table 9. Compilation of 2007 and 2008 data for the 187 turtles of southwestern Michigan fen. Age Male/ Adult! Carapace Width Plastron Height Weight Length Length Turtle (yrs) Female Juvenile (cm) (cm) (cm) (cm) (Ig)__ A (2006) 1 J 3.020 2.707 2.599 ABCK 13 M A 9.264 6.556 7.595 3.331 0.118 ABCKVW 12 M A 10.930 6.918 8.223 3.434 0.130 ABCKW 12 F A 9.580 6.765 8.383 3.954 0.134 ABDK 1 1 F A 9.368 6.350 8.432 3.852 0.1 14 ABKW 14 F A 8.848 6.245 7.699 3.748 0.1 10 ABKWX 15 M A 10.305 7.180 8.640 3.538 0.158 ACDK F A 9.388 6.391 8.352 AClK 1 1 F A 8.536 6.380 7.596 3.434 0.102 ACK 7 J 7.807 5.830 6.765 3.020 ACKVWX 7 M A 7.983 5.954 6.970 ACKW 8 F A 9.054 6.867 8.014 3.744 0.1 14 ADKO 12 F A 9.262 6.454 8.010 3.851 0.140 ADKV 1 1 F A 8.341 6.245 7.702 3.332 0.088 AHK 6 J 4.944 4.061 4.164 2.078 0.024 AJK 1 1 M A 9.073 6.525 7.742 AJKX 1 1 M A 9.678 6.868 8.119 3.437 0.120 AK 15 M A 9.056 6.462 7.598 3.320 0.1 10 AKLMP 8 J 7.710 6.620 6.665 2.813 0.072 AKM 12 F A 8.811 6.347 7.705 3.747 0.112 AKN 7 M A 8.488 6.047 7.285 3.333 0.092 AKO 12 F A 8.540 6.140 7.495 3.750 0.106 AKOQ 7 J 7.810 6.040 6.868 2.81 1 0.080 AKP 7 F J 7.708 5.620 6.766 3.018 0.066 AKU J 6.640 5.420 5.839 AKW 13 M A 9.570 6.574 8.223 3.740 0.1 16 AKWX 13 M A 8.354 5.944 7.282 3.021 0.084 AKX 15 F A 8.848 6.350 7.810 3.645 0.135 B (2006) 2 J 3.123 2.706 2.602 0.004 BCEKL 2 J 6.240 5.100 5.520 2.500 0.042 BCK F A 9.471 6.661 8.329 0.132 BCKV 12 M A 9.368 7.705 7.91 1 3.436 0.120 BCKW M A lBDK 10 F A 8.745 6.455 7.914 3.746 0.110 BDKV 10 M A 9.580 6.870 8.010 3.430 0.120 BDKW 10 F A 8.950 6.660 8.1 18 3.850 0.135 BDKX 14 M A 9.570 6.660 7.910 3.644 0.112 BHK 12 F A 8.535 6.557 7.705 3.850 0.126 BlK J 7.708 5.517 6.454 BIKX 8 M A 8.015 6.039 7.787 3.226 0.084 BJK 8 F A 8.120 6.038 7.390 3.123 0.084 BJKW 13 M A 9.246 6.507 7.809 3.435 0.114 BK (Juv.) J 5.622 4.889 4.993 79 Table 9 (cont’d). Age Male! Adult] Carapace Width Plastron Height Weight Length Length Turtle (yrs) Female Juvenile (cm) (cm) (cm) (cm) 33L BK F A 8.435 6.455 7.704 3.852 0.112 BKM 9 J 6.663 5.210 5.832 2.500 0.050 BKN 6 J 6.670 5.204 5.932 2.602 0.050 BKOV J 5.414 4.475 4.788 BKP J 6.208 5.103 5.512 BKU 13 F A 8.920 6.294 7.772 3.748 0.1 12 BKV F A 9.055 6.589 8.123 BKW 12 F A 9.471 6.768 8.329 3.642 0.144 BKX 7 M A 8.534 6.063 7.078 3.029 0.090 C 2 J 3.021 2.707 2.599 0.004 CDK F A 8.325 6.141 7.492 CDKV 12 M A 9.387 6.453 7.724 3.434 0.102 CDKVW 1 1 M A 9.992 7.280 8.235 3.540 0.130 CHJKLVW 1 1 F A 9.473 6.558 8.225 3.851 0.152 CHK 13 M A 8.848 6.039 7.596 CHKW J 5.623 4.788 5.099 CHKZ 10 J 6.038 4.889 5.312 2.495 0.036 CIJKP 6 M A 6.560 5.120 5.730 2.600 0.052 CJK 1 1 F A 8.328 6.139 7.699 CK 7 J 7.181 5.417 6.247 2.915 0.062 CKM J 6.973 5.413 6.038 CKN 7 F J 6.454 5.100 5.728 2.610 CKO J 7.153 5.893 6.223 CKP 5 J 4.995 4.372 4.265 1 .978 0.022 CKU F A 9.161 6.765 8.326 3.748 0.138 CKW 6 J 7.491 5.829 6.558 2.913 CKX 17 F A 9.163 6.520 8.118 3.643 0.116 CRV M A 10.635 DHK 1 1 M A 8.950 6.455 7.720 3.640 0.1 12 DIK F A 9.301 6.541 8.131 DJK M A 8.816 6.264 7.549 DKL F A 8.758 6.586 7.691 DKLM 8 M A 6.450 5.410 5.720 2.603 0.048 DKM 16 F A 9.368 6.555 7.912 3.539 0.106 DKN 9 M A 8.432 6.556 7.285 3.021 0.098 DKO 8 M A 7.705 6.036 6.869 3.123 0.070 DKP 13 F A 8.750 6.554 7.720 3.540 0.122 lDKU M A 8.593 6.245 7.288 3.333 0.102 DKVWX 12 F A 8.750 6.770 7.846 3.440 0.112 DKX 1 1 F A 9.050 6.770 8.174 3.640 0.1 14 FGK 14 F A 8.265 6.185 7.325 3.440 F lKS 6 J 5.839 4.892 5.000 2.495 0.038 FJKS 10 M A 9.060 6.350 7.500 3.230 0.1 12 8O Table 9 (cont’d). Age Male! Adult/ Carapace Width Plastron Height Weight Length Length Turtle (yrsl Female Juvenile (cm) 4cm) (cmL (cm) (kg)__ FKS 13 M A 9.160 6.680 7.601 3.539 0.122 GK 7 J 5.932 4.287 5.102 2.498 0.036 GKX 10 M A 8.535 6.036 7.077 3.125 0.096 H 2 J 4.265 3.750 3.643 1.665 0.012 HIK 12 M A 9.264 6.661 7.808 3.124 0.112 HJK 12 F A 9.158 6.560 8.222 3.954 0.136 HJKV 12 F A 8.640 6.870 7.705 3.539 0.136 HK 2 J HKL J 6.245 5.205 5.517 2.705 0.046 HKLQ 12 A 9.158 6.868 8.223 3.332 0.114 HKM 22 A 10.306 7.600 9.056 4.579 0.198 HKN J 6.558 5.412 5.727 HKNQV 1 1 M A 8.744 6.452 7.363 3.124 0.098 HKO 6 M A 7.388 5.620 6.452 2.709 0.056 HKP 13 F A 9.368 7.283 8.535 3.850 0.136 HKQ M A 8.580 5.987 7.328 HKQW 6 J 6.868 5.312 6.141 2.708 0.054 HKQX J 6.807 5.438 6.162 HKU 1 1 F A 9.265 7.077 8.324 3.850 0.134 HKV 18 F A 9.471 6.973 8.535 4.059 0.160 HKW (F) 5 J 5.933 4.890 5.207 2.394 HKW (M) M A 9.368 6.453 7.704 3.537 0.132 HR 1 J 3.850 3.230 3.540 3.230 0.010 I 5 J 6.247 4.892 5.623 2.707 0.044 lF J 5.525 4.717 4.770 U K M A 8.575 6.030 7.135 IK (2006) 2 J 3.995 3.230 3.228 1 .460 0.008 lK 5 yr. )( 5 J 4.893 4.164 4.268 1 .870 lKL 11 M A 9.413 6.599 7.705 3.332 0.118 IKM 10 M A 9.262 6.452 7.596 3.332 0.1 10 lKNP 8 J 7.821 5.932 7.389 3.226 0.084 lKNX 14 M A 9.252 6.140 7.390 3.225 0.096 lKO 5 J 5.519 4.682 4.994 2.287 0.030 IKOPQ 11 M A 8.326 6.140 7.290 3.124 0.092 IKP 10 M A 9.159 6.558 8.016 3.125 0.108 IKW 11 M A 9.327 6.765 7.869 3.644 0.124 lMX M A 8.966 6.203 7.356 J 1 J 3.227 2.813 2.707 1.249 0.006 J KL 12 M A 8.745 6.245 6.973 3.435 0.098 JKOP 7 M A 7.079 5.519 5.935 2.708 0.060 JKP 4 J 6.245 4.787 5.518 2.500 0.038 JKW 10 M A 9.991 6.454 8.225 3.540 0.138 81 Table 9 (cont’d). Age Male] Adult! Carapace Width Plastron Height Weight Length Length Turtle (yrs) Female Juvenile (cm) (cm) (cm) (cm) (kg)__ JKX 7 J 5.725 4.786 4.788 K 6 J 6.765 5.204 6.038 2.710 0.048 KL 4 J 4.371 3.645 3.539 KLM 12 M A 9.369 6.348 7.705 3.333 0.132 KLN 8 M A 9.053 6.244 7.390 3.227 0.102 KLO M A 9.522 6.789 7.887 KLP 6 J 6.035 5.205 5.205 2.598 0.034 KLQ 17 F A 9.367 6.455 8.265 3.745 0.126 KLU 9 F A 8.744 6.557 7.808 3.539 0.1 18 KLW 19 F A 8.898 6.650 7.717 3.740 KLX F A 9.091 6.746 8.120 3.540 0.122 KMN 12 F A 9.056 6.558 7.808 3.953 0.128 KMOQ 19 F A 9.159 6.558 8.118 3.850 0.126 KMP J 6.868 5.208 6.037 2.599 0.048 KMU 8 J 5.830 4.890 5.206 2.602 0.032 KMW (J UV) 1 1 J 7.600 5.934 6.675 2.915 0.070 KMW 13 F A 9.886 7.825 8.536 4.059 0.168 KMX 9 F A 8.750 5.921 7.500 3.021 0.096 KN 4 J 4.370 3.540 3.542 1.665 0.014 KNO 9 M A 9.262 6.143 7.597 3.226 0.100 KNP 6 J 6.660 5.210 5.930 2.810 0.052 KNU 14 M A 8.534 5.934 7.285 2.910 0.084 KNV 5 J 5.392 4.890 5.100 2.290 0.036 KNW 15 M A 9.161 6.452 8.385 3.331 0.104 KNX 12 M A 9.885 6.952 8.285 3.022 0.120 KO 3 J 4.268 3.747 3.332 1.660 0.014 KOP 5 J 6.558 5.206 6.038 2.604 0.048 KOPR 6 J 7.285 5.820 6.246 2.813 0.062 KOQ 6 M A 6.767 5.204 5.934 2.602 0.052 KOU 13 M A 9.158 6.555 7.910 3.339 0.110 KOV 7 J 7.390 5.516 6.351 2.917 0.064 KOW J 7.503 5.875 6.485 KOX J 7.226 5.517 6.383 KP 7 J 5.375 4.465 4.737 2.189 0.026 KPQ 8 F A 9.160 6.453 8.120 3.850 0.128 KPU 5 J 5.830 4.990 5.100 2.390 0.036 KPV 7 M A 7.492 5.729 6.659 2.708 0.068 KPW 6 J 6.145 5.205 5.518 2.705 0.044 KPX 9 F A 8.140 6.765 7.206 3.245 0.110 KG (2006) 2 J 3.123 2.707 2.602 1.146 0.006 KQU J 7.071 5.453 6.200 KU M A 82 Table 9 (cont’d). Age Male! Adult! Carapace Width Plastron Height Weight Length Length Turtle (yrs) Female Juvenile (cmL (cm) (cm) JemL (I59) KU 14 F A 9.158 6.763 8.224 KUV 12 M A 8.950 6.570 7.590 3.330 0.134 KUVX 9 J 6.878 5.271 6.140 2.708 KUW 8 F A 8.014 5.934 7.078 3.230 0.084 KV 21 F A 9.158 6.869 8.293 3.641 0.136 KVW 9 M A 9.055 6.455 7.390 3.125 0.1 10 KVWX J 6.713 5.177 6.033 KVX 14 F A 8.952 6.765 8.013 3.750 0.128 KW F A 9.083 6.871 8.202 KWX 4 J 5.208 4.578 4.580 2.291 KX (2006) 2 J 3.227 2.812 2.605 1 .145 0.006 LMN M A 10.066 0 2 J 4.058 3.437 3.332 1.560 0.010 P 1 J 3.228 2.915 2.603 1.147 0.006 R 1 J 3.330 3.020 2.610 1 .350 0.008 V 3 J 4.060 3.538 3.228 1 .562 0.012 W 1 J 3.123 2.810 2.501 1.145 0.004 83 Figure 1. Presettlement land cover of southwestern Michigan fen. .2922 Lancet 5’52““ I 8.2 wEgsetaw use «0.3 ream “5.5a. §i$§§sl 50325 Le 89¢. 3.592.1— 23.5308...— _ S35? I m7 i20< on: 39.88: Hectares:~30; Acresz~75 84 Figure 2. Outlying regions of study site. 85 Figure 3. Open wetland in middle of fen. 86 Figure 4. Thick growth of Chara spp. in study site Chara pond. 87 Figure 5. Large transmitter attached to turtle carapace. 88 Figure 6. Alphabetical turtle marking system diagram. Head /v 89 Figure 7. Diagram of sample markings for turtle ABX. Head Tail 90 Figure 8. Spotted Turtle with notched markings. 91 Figure 9. Small transmitter attached to turtle carapace. Figure 10. Labeled diagram of study site. I. l n.3’lt. ‘II3'5v 83... 1| 83.306»: East—03m tosm tame—O econ r820 zon— mo 23:2 «2.4 :30 8:3 0 59.3.25 :8... Em Eofixozm >320 Ammo— owem read 83 .55 contomoi 8.3 *Hand-drawn diagram by Diana Lutz. 93 Figure 11. Compilation of habitat utilization for transmittered turtles 2007. « 'Hectare.s€‘1 Acres:26.i - ' r. ‘3 ’ Spotted Turtle Distribution and Analysis For All Transmittered Turtles 2007 *Hectares:10.69; Acres: 26.42 94 Figure 12. Compilation of habitat utilizations for turtles without transmitters in 2007. Spotted Turtle Distribution and Analysis I for All Nontransmittered Turtles 2007 *Hectares: 9.81; Acres: 24.245 95 Figure 13. Compilation of habitat utilizations for transmittered turtles in 2008. Spotted Turtle Distribution and Analysis For All Transmittered Turtles 2008 *Hectares: 12.02; Acres: 29.71 96 Figure 14. Compilation of habitat utilizations for turtles without transmitters in 2008. Spotted Turtle Distribution and Analysis for All Nontrsnrnittered Turtles 2008 *Hectares: 10.71; Acres: 26.47 97 Figure 15. Predated Nesting Site. \ ‘\ “W‘TMQ . \\ 4" _ \ N 9 \V // I ' 98 ~,-' “‘1“ w_ n ‘\\I‘ ' ‘l“ 4 9.5“ \ ' Figure 17. Channel 0 male IKL habitat utilization in 2007. d: .3. a$§< ...: 823:5: 2:2. 8:25 '3? ... so; 0 s. .....w .1........._. ..~ .... . MUM... and.» s x h .Qccwfihk... *Hectares: 1.60; Acres: 3.95 99 Figure 18. Channel 1 male KVW habitat utilization in 2007. Hectares'1237o 1x _ Acres: 5.86 Spotted Turde Distribution and Analysis For KVW Legend MCP Spotted Turtle Observations O *Hectares: 2.37; Acres: 5.86 100 Figure 19. Channel 2 male ABCK habitat utilization in 2007. Spotted Turtle Distribution and Analysis For ABCK 2007 nd (MCP Spotted Turns Observations 0 *Hectares: 3.30; Acres: 8.16 101 Figure 20. Channel 3 female KVX habitat utilization in 2007. .23 388 o E s... ".223. ...: as. 8:353: 3:5 Beam F83 Viiv 3F... \ .5 .....ufi. . m ... a o... 4 ......w... x w... a: 8.9.3“ 3.6 M228: 2 's I ‘3 "0.1 ' ‘ . .5 0" «a. w, .’/ *Hectares: 0.48; Acres: 1.19 102 Figure 21. Channel 4 female HKU habitat utilization in 2007. ":: Hectares: Q35 Acres 06: Spotted Turtle Distribution and Analvsis For HKU WP Spotted Turtle Observations 0 *Hectares: 0.35; Acres: 0.86 103 Figure 22. Channel 6 female DKVWX habitat utilization in 2007. as '3! O‘- ..I~_ 30 Eli ‘52 00 I< E...” 33 5"...» hat-ht. to.“ .1 .... 913, AV . ,. ts . " A. .4..- .33-9. -( Spotted Turtle Distribution and Analysis For DKVWX MCP Spotted Turtle Observations O *Hectares: 0.29; Acres: 0.71 104 Figure 23. Channel 7 female KLU habitat utilization in 2007. 1. ‘t '1 -‘ i 3.11.! ' 6 iii 0 b a: .. U a: 3: Acres: 0.98 Spotted Turtle Distribution and Analysis For KLU WP o SpottsdTutle *Hectares: 0.40; Acres: 0.98 105 Figure 24. Channel 8 female HKV habitat utilization in 2007. 2 0"? ‘1: 'd) b ts .. 0 d: I Acres: 2.77 . Spotted Turtle Distribution and Analysis For HKV Lem MCP Spotted Turtle Observations O *Hectares: 1.12; Acres: 2.77 106 Figure 25. Channel 10 male lKW habitat utilization in 2007. [N 8 «3 u. in? =8 .. h .22 DO NO} I]: ‘— P. ‘— El; 0 s. U < i =5 gs ii Eh 5% 3'» e? 'D 5 Legend MCP o SpottedTutle *Hectares: 0.45; Acres: 1.11 107 Figure 26. Channel 11 female KV habitat utilization in 2007. [N O. can ..‘1 3° ._.. 2m 03 ea I< O. 0 es I = '3; = .9 '5: In a: on > Ea h: < ‘3: g. a. U) Legend MCP Spotted Turtle Observations O *Hectares: 0.07; Acres: 0.18 108 Figure 27. Channel 1 female HJKV habitat utilization in 2008. 9.? v n Sea->380 ......» 3.88 3:: 3m 2.2.5. a... .6: 525.53: 2:3. 8:25 E: a cites... e Pm... ....“ $63123 ... ...” ... ._ Wmuwwfifivz e ,e .. ... J . *Hectares: 1.20; Acres: 2.96 109 Figure 28. Channel 2 male KUV habitat utilization in 2008. In us oo ..CD. 3;- b“ sin 02 00 I< Spotted Turtle Distribution and Analysis For KUV WP Spotted Tube Observations O *Hectares: 0.66; Acres: 1.60 110 Figure 29. Channel 3 female HJK habitat utilization in 2008. ._ ". CID ..l\. 3.- ... EU) as 00 I< E .6 ’35 5 be. as .a E> :1 Pl: 4 Be 8: 8. m nd MCP Spotted Turtle Observations O *Hectares: 0.71; Acres: 1.75 111 Figure 30. Channel 4 female ABKW habitat utilization in 2008. n N. can 5"! m° ._.. 2m 02 mo I< e 55 '5 5 33% 0 2'3 ["3 34! '5 8.5 CD nd MCP Spotted Turtle Observations O *Hectares: 0.23; Acres: 0.58 112 Figure 31. Channel 5 female CHJKLVW habitat utilization in 2008. Spotted Turtle Distribution and Analysis For CHJKLVW Hectares: 0.22 Acres: 0.55 ts Leg—end 352.. a S; y, wavy"? u“. '1 ‘ 6.- .l , .. v “' I. :\‘- . ’9‘ J _; P ‘ . i" “ I ”5'! MCP Spotted Turth Observations O *Hectares: 0.22; Acres: 0.55 113 Figure 32. Channel 6 female BDKW habitat utilization in 2008. . O '1‘ O .553 2.. 53 UL QIU :< ea 2: “.9 3:. £5 LOW MCP Spotted Turth Observations O *Hectares: 0.60; Acres: 1.48 114 Figure 33. Channel 7 female AKX habitat utilization in 2008. case-Q . . ... . ... $1“f3’0 . é ‘ ' .. a ., Spotted Turde Distribution and Analysis For AKX MCP Spotted Tutle Observations O *Hectares: 1.09; Acres: 2.70 115 Figure 34. Channel 8 female FKS habitat utilization in 2008. (’3 ‘7 1- iii 0 L- 1V ... U Q! I Acres: 3.28 Spotted Turtle Distribution and Analysis For FKS Legend MCP o SpottedTutie *Hectares: 1.33; Acres: 3.28 116 Figure 35. Channel 9 female BKW habitat utilization in 2008. 412' ' O D o Hectaresziqés Acr‘es:‘0 .8 O Spotted Turtle Distribution and Analysis For BKW 2008 Le97nd MCP Spotted Turtle Observations O *Hectares: 0.16; Acres: 0.41 117 Figure 36. Channel 11 female CKU habitat utilization in 2008 co °. i-rs a“? ”N ... 3‘” 82 :E O Spotted Turtle Distribution and Analysis For CKU MCP Spotted Tutie Observations O *Hectares: 1.08; Acres: 2.67 118 Figure 37. Channel 12 female ABCKW habitat utilization in 2008. Spotted Turtle Distribution and Analysis For ABCKW ' C a. ‘2 ON “(‘5 3‘- Eli ‘62 0.1L) I< ) bx‘? . .4 a. Legend '. x MCP Spotted Turtle Observations O *Hectares: 0.49; Acres: 1.22 119 Figure 38. Channel 13 female DKP habitat utilization in 2008. Hectares: 1.08 Acres: 2.67 Spotted Turtle Distribution and Analysis For DKP nd MCP Spotted Turtle Observations O *Hectares: 1.08; Acres: 2.67 Figure 39. Channel 14 female BHK habitat utilization in 2008. - a N. O 111 2 (l .. o a; I Acres: 1.95 Spotted Turtle Distribu ' n and Analysis For BHK Legs—rid WP Spotted Tutle Observations O *Hectares: 0.79; Acres: 1.95 Figure 40. Channel 15 female ADKO habitat utilization in 2008. Hectares: 0.22 Acres: 0.53 ‘ ' .‘5 ‘5. .ofl“ 1‘ -./ w}. l a: 'cg I. ‘ I $34.1“ Spotted Turtle Distribution and Analysis For ADKO Legend MCP Spotted Turtle Obeervations O *Hectares: 0.22; Acres: 0.53 122 Figure 41. Channel 16 male ABCK habitat utilization in 2008. Habitat Use 2008 Hectares: 1.81 Acres: 4.48 Spotted Turtle Distribution and Analysis For ABCK 2008 Legend MCP Spotted Turtle Observations O ‘Hectares: 1.81; Acres: 4.48 123 Figure 42. Channel 17 male lKW habitat utilization in 2008. and Analysis For IKW Spotted Turtle Distribution *Hectares: 1.33; Acres: 3.29 124 Figure 43. Habitat utilization for male ABCK in 2007 and 2008. I .i o“. . ‘ " ‘3}.4‘3. .‘I . .5 if 1 I.‘ \ Habitat U ° Hectares Habitat Use 2008 Hectares: 1.81 Acres: 4.48 Spotted Turtle Distribution and Analysis For ABCK 2007 and 2008 Lem sopzoos M02007 SposedTunIe W O *2007 Hectares: 3.30; Acres: 8.16 *2008 Hectares: 1.81; Acres: 4.48 125 Figure 44. Habitat utilization for male IKW in 2007 and 2008. 130,6" . I Hectates Acres: 3.29' . ‘ er Habitarfu‘sf f. .s t q, 031. 03. e Spotted Turtle Distribution and Analysis For IKW 2007 and 2008 Habitat Use 2007 Hectares: 0.45 Acres: 1.11 Legend MCP Spotted Tutle Observations O *2007 Hectares: 0.45; Acres: 1.11 *2008 Hectares: 1.33; Acres: 3.29 126 Figure 45. Overlapping areas of all females tracked in 2007. co '5') c c (TS .: O Spotted Turtle Minimum Convex Polygon Showing All Females Tracked 2007 Roads Ch. 3: Hectares: 0.48; Acres: 1.19 Ch. 4: Hectares: 0.35; Acres: 0.86 Ch. 5: Hectares: 0.37; Acres: 0.92 Ch. 6: Hectares: 1.08; Acres: 2.67 Ch. 7: Hectares: 0.40; Acres: 0.98 Ch. 8: Hectares: 1.12; Acres: 2.77 Ch. 9: Hectares: 0.05; Acres: 0.11 Ch. 11: Hectares: 0.07; Acres: 0.18 127 Figure 46. Overlapping areas of all males tracked in 2007. Spotted Turtle Minimum Convex Polygon Showmg All Males Tracked During 200' T 1 chmo DCH2 8l 3 .— o I :r: 0 0 F Ch. 0: Hectares: 1.60; Acres: 3.95 Ch. 1: Hectares: 2.37; Acres: 5.86 Ch. 2: Hectares: 3.30; Acres: 8.16 Ch. 10: Hectares: 0.45; Acres: 1.11 128 Figure 47. Channel 1 overlapping with all transmittered females during 2007. «emanate 55.5 s36 _ 1555 Euro.— aosaeo 5.55: at; Baa—m m 85.5 Figure 48. Male overlapping home ranges in 2008. Showing All Males Tracked During zoos Spotted Turtle Minimum Convex Polygon N I 0 0 v- I O Ch. 2: Hectares: 0.65; Acres: 1.60 Ch. 16: Hectares: 1.81; Acres: 4.48 Ch. 17: Hectares: 1.33; Acres: 3.29 130 Figure 49. Female overlapping home ranges in 2008. a . 2% in 93 H (32 [- E ‘8 E .55 Eu. 25 to: 35 H'; '8: =85 w 9!! 58% [gm] m 3:3 : -1 I (D --.: E 6666 g l— o Ch. 1: Hectares: 1.20; Acres: 2.97 Ch. 8: Hectares: 1.33; Acres: 3.28 Ch. 3: Hectares: 0.71; Acres: 1.75 Ch. 9: Hectares: 0.17; Acres: 0.41 Ch. 4: Hectares: 0.23; Acres: 0.58 Ch. 11: Hectares: 0.17; Acres: 0.42 Ch. 5: Hectares: 0.22; Acres: 0.55 Ch. 12: Hectares: 0.50 Acres: 1.22 Ch. 6: Hectares: 0.60; Acres: 1.48 Ch. 13: Hectares: 0.12; Acres: 0.29 Ch. 7: Hectares: 1.09; Acres: 2.70 Ch. 14: Hectares: 0.79; Acres: 1.95 Ch. 15: Hectares: 0.22; Acres: 0.53 131 Figure 50. Sphagnum spp. moss mounds. 13599.n- -... .-. " "1 >11, . ' egg-v p -‘ "7;“ 7 “Q a X 1' . \ » , c - g- \\ I - 1‘ ")4; ' 40-: . . . . '- 132 Figure 52. Channel 4 aestivation and hibernation locale. C k A“ 3' (:11 l t1 . ... a r": 2” 1 r ‘ ' ‘: a . .1. 7.7/"3 “(z/1".“ ” ’ , Ale/Empty" 1L1; -‘ . .z‘ .- 1.4: 3w 1‘ a Mr" F015 hi". 1 l A (A l) L.- ’ 0' \\_ ,, . _ 117436 V/— ____ \~ , .. ' ‘ mo“ Noam; a-“ . “x he i Q -’ .\ , g g. .1 _. . z..;..---«.u,/ ~ - a»- - 4. 15.24 Cm Mafia mdemch/K CAI“ A? \ q Wafel—w’r’nf) [0°C 411’ 16m? l5-7 0L *Hand-drawn diagram by Diana Lutz. 133 ..y. . . . . a . . 1 . . . lisp. t nil i... 9‘ .v . .ll't. ' states has... bl. Waist. Figure 55. Deer trail in study site. 135 Figure 56. Total utilization area for all turtles in 2007 and 2008. Spotted Turtle Distribution and Analysis For All Turtles 2007 - 2008 *Hectares: 13.30; Acres: 32.85 136 Figure 57. 2007 and 2008 Spotted Turtle sex distribution. he. ”nous—HE.» 35852 .80... 2.52:. I ...: I 0.2:.“— I 5:39.85 xom 2:2. 8:26 83 can 33 137 Figure 58. Main aggregations of Spotted Turtles during 2007. Spotted Turtle Distribution and Analysis For All Transmittered Turtles 2007 *Hectares: 10.69; Acres: 26.42 138 Figure 59. Main aggregations of Spotted Turtles during 2008. ' Spotted Turtle Distribution and Analysis For All Transmittered Turtles 2008 *Hectares: 12.02; Acres: 29.71 139 Figure 60. 25 September 2007 possible hibemaculum. 77/656101] , 255€p+€mb€f 2007 ) 0% 9.16m 0 W1 ZQBCK—Wk) ‘bUl’iEA in mud /\ bunpd m mud "' C KY (female \ 3.48m Bulrush, 661136, .1 “r! 4",“ ’7‘ ' is ficATur‘HfS ’ciacbzat" horizonletli ‘ - f cDKV(mle). ij5(mle))coi ’ ' i i‘ 91"” J (06" awv’ ' “a" ’ é-WoZan-D “a? --. .2 ‘ . .- M M“ s ‘9 ‘2... ’25 C "1.....-m ‘- M~~ .- .——- 4m: w. . fl .. (£9 ' “'7' . a‘um““%~--“‘:§~:‘: , t Noie: CM 5 in swimsngs Bax/6m) 11.0‘6 shell Rmp, (firms .2: TKPka) 12.37. sAeIi imp, r L , seismic. Kuwgimk) i337. shell w In W! War andn‘wd) among Soundbar mum mo+ 35mm- L49!- WK (5 amber wan) 9C KVGMJC) was «Ln-Hu's hole weih 2. drfjferem- 5PM“ TM‘€S((LDKVm41e , KLX fanatic). *Hand-drawn diagram by Diana Lutz. I42 Figure 63. 16 October 2007 possible hibemaculum. ”Galaxy up adobe!“ 2007 WW “AU-0w —-; “ " «titrfi. walcr and mud (€727Mi- .. in M2. 4:_ 1::M --. ...? .....qu ”’1':- Nolc' change in com (rule) l~l$°c shell lame 6mm? 5+4: ’ KO? (330 Il‘f.$‘C shell Rm?- daggm. IKNPQM/m) ls.s°c shell lth- AKP gunk/3W) "...-1°C shell “MP- *Hand-drawn diagram by Diana Lutz. 143 LITERATURE CITED 144 I- sank-m LITERATURE CITED Barlow, C. 1999. Habitat Use and Spatial Ecology of Blanding’s Turtles (Emydoidea blandingil) and Spotted Turtles (Clemmys guttata) in Northeast Indiana. Master of Science Thesis. Purdue University, Indiana. Barnwell, M.E., Meylan, PA, and T. Walsh. 1997. The Spotted Turtle (Clemmys guttata) in Central Florida. Chelonian Conservation and Biology. 2(3):405-408. Blake, SF. 1922. Sexual differences in coloration in the spotted turtle, Clemmys guttata. Proc. US. Natl. Mus. 59:463-469. Berry, J.F. 1978. Spotted Turtle. IN: McDiarmid, R.W. (eds). Rare and Endangered Biota of Florida. 3:47-49. Breder, RB. 1927. Turtle Trailing: a New Technique for Studying the Life Habits of Certain Testudinata. Zoologica. 9(4):231-243. Breisch, AM. 2006. The Natural History and Thermal Ecology of a Population of Spotted Turtles (Clemmys guttata) and Wood Turtles (Glyptemys insculpta) in West Virginia. MS. Thesis. Marshall University. Brinlee, C.J. Herbicide Demonstration for Aquatic Vegetation Control [intemet] Available from: lnttpzfanninswcd.com/Newsletters/nletter-13-3-3-Pond. Accessed 2009. Burke, V.J., J.E. Lovich, and J.W. Gibbons. 2000. Conservation of freshwater turtles. IN: M.K. Klemmens (ed.). Turtle Conservation. Smithsonian Institution Press, Washington, DC. pp. 156-179. Cagle, PR. 1944. Home Range, Homing Behavior, and Migration in Turtles. Misc. Publ. Mus. Zool., Univ. Mich. 6121-33. . 1939. A System of Marking Turtles for Future Identification. Copeia. 1939(3):170-173. Cahn, AR. 1937. The turtles of Illinois. Contribution from the Zoological - Laboratory of the University of Illinois. 492:68-75. Carr, A. 1952. Handbook of Turtles: The Turtles of the United States, Canada and Baja California. Cornell University Press, Ithaca and London. pp.111-117. Chadde, SW. 2002. A Great Lakes Wetland Flora. Second Edition Pocketflora Press. Calumet, MI. 145 Chippindale, Paul. 1989. Courtship and Nesting Records for Spotted Turtles, Clemmys guttata, in the Mer Bleue Bog, southeastern Ontario. The Canadian Field Naturalist. 103:289-291. Conant, R. and J.T. Collins. 1991. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Houghton Miffin Co., Boston. pp.50- 51. Conant, R. and J.T. Collins. 1998. A Field Guide to Reptiles and Amphibians of Eastern and Central North America, Third Edition Expanded. Houghton Miffin 00., Boston. pp.158-159. Cook et al. 1980. Spotted Turtles (Clemmys guttata) in Eastern Ontario and Adjacent Quebec. The Canadian Field Naturalist. 94:411-415. COSEWIC. 2004. COSEWIC assessment and update status report on the Spotted Turtle, Clemmys guttata, in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. Vi + 27 pp. Available from: www.sararegistry.gc.ca/status/status e.cfm. Accessed 2009. Ditmars, R.L. 1933. Reptiles of the World. MacMillan Company, Norwood, Mass, USA. pp.285-286. Ernst, OH. 1967. A Mating Aggregation of the Turtle Clemmys guttata. Copeia. 1 967(2) :473-474. . 1968. Homing Ability in the Spotted Turtle, Clemmys guttata (Schneider). Herpetologica. 24(1):77-78. . 1968. Evaporative Water-Loss Relationships of Turtles. Journal of Herpetology. 2(3/4):159-161 . . 1970. Reproduction in Clemmys guttata. Herpetologica. 26:228- 232. . 1970. Home Range of the Spotted Turtle, Clemmys guttata (Schneider). Copeia. 970(2):391-393. . 1972. Clemmys guttata. Cat. Amer. Ambhib. Rept. 124.1-124.2. . 1975. Growth of the Spotted Turtle, Clemmys guttata. Journal of Herpetology. 9(3):313-318. . 1976. Ecology of the Spotted Turtle, Clemmys guttata (Reptilia, Testudines, Testudinidae), in Southeastern Pennsylvania. Journal of Herpetology. 10(1):25-33. 146 . 1982. Environmental Temperatures and Activities in Wild Spotted Turtles, Clemmys guttata. Journal of Herpetology. 16(2):112-120. . 2001. An Overview of the North American Turtle Genus Clemmys Ritgen, 1828. Chelonian Conservation and Biology. 4(1):211-216 , J.E. Lovich, and R.W. Barbour. 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washington and London. pp. 205-212. and G. Zug. 1944. Observations on the Reproductive Biology of the Spotted Turtle, Clemmys guttata, in Southeastern Pennsylvania. Journal of Herpetology. 28(1):99-102. Fitch, HS. 1958. Home range, territories, and seasonal movements of vertebrates of the University of Kansas Natural History Reservation. Univ. Kans. Pub, Mus. Nat. Hist. 11(3):63-326. Folkerts, G.W. and AC. Skorepa. 1967. A Spotted Turtle, Clemmys guttata (Schneider) from southeastern Georgia. Herpetologica. 23(1):63. Fowle, S. 2001. Guidelines for Protecting Spotted Turtles and Their Habitats in Massachusetts. Natural Heritage and Endangered Species Program, Massachusetts Division of Fisheries and Wildlife. pp. 1-11. Gibbons, J.W. and J.E. Lovich. 1990. Sexual dimorphism in turtles with emphasis on the Slider Turtle (Trachemys Scripta). Herpetological Monographs. 4:1-29. Graham, Terry E. 1970. Growth Rate of Spotted Turtle, Clemmys guttata, in Southern Rhode Island. Journal of Herpetology. 4(1/2):87-88. Graham, Terry E. 1995. Habitat use and population parameters of the Spotted Turtle, Clemmys guttata, a Species of Special Concern in Massachusetts. Chelonian Conservation and Biology. 1(3):207-214. Grant, C. 1936. The Southwestern Desert Tortoise, Gopherus agassizii. Zoologica. 21(4):225-229. Harding, J. 1997. Amphibians and Reptiles of the Great Lakes Region. University of Michigan Press. Haxton, T. and M. Berrill. 2001. Seasonal Activity of Spotted Turtles (Clemmys guttata) at the Northern Limit of Their Range. Journal of Herpetology. 35(4) :606-61 4. 147 Joint Nature Conversation Committee. 2005. 3140 Hard oligo-mesotrophic waters with benthic vegetation of Chara spp. Available from: www.jncc.gov.uk/protectedsites/sacselection/habitat.asp?FeaturelntCode= H3140-31K-. Accessed 2009. Joyal et al. 2001. Landscape Ecology Approaches to Wetland Species Conservation: A Case Study of Two Turtle Species in Southern Maine. Conservation Biology. 15(6):1755-1762. Kiester, A.R., C.W. Schwartz, and ER. Schwartz. 1982. Promotion of gene flow by transient individuals in an otherwise sedentary population of box turtles (Terrapene carolina). Evolution. 36:617-619. Lewis, TL. and J. Ritzenthaler. 1997. Characteristics of Hibemacula Use by Spotted Turtles, Clemmys guttata, in Ohio. Chelonian Conservation and Biology. 2(4):611-615. Litzgus, JD. 1996. Life-History and Demography of a Northern Population of Spotted Turtles, Clemmys guttata. Master’s Thesis, University of Guelph, Ontaria. . 1996. Cryptic Survivor: The Elusive Spotted Turtle. Bruce Trail News. pp.16-19. . 2006. Sex Differences in Longevity in the Spotted Turtle (Clemmys guttata). Copeia. 2006(2):281-288. and R.J. Brooks. 1998. Growth in a cold environment: body size and sexual maturity in a northern population of spotted turtles, Clemmys guttata. Can. J. Zool. 76:773-782. and R.J. Brooks. 1998. Reproduction in a Northern Population of Clemmys guttata. Journal of Herpetology. 32(2):252-259. and R.J. Brooks. 2000. Habitat and Temperature Selection of Clemmys guttata in a Northern Population. Journal of Herpetology. 34(2): 1 78-1 85. et al. 1999. Phenology and ecology of hibernation in spotted turtles (Clemmys guttata) near the northern limit of their range. Can. J. Zool. 77: 1 348-1 357. and T. Mousseau. Demography of a Southern Population of the Spotted Turtle (Clemmys guttata). Southeastern Naturalist. 2004. 3(3):391-400. 148 and T. Mousseau. Home Range and Seasonal Activity of Southeastern Spotted Turtles (Clemmys guttata): Implications for Management. Copeia. 2004. 2004(4):804-817. Lovich, J.E. 1987. The Savannah River Ecology Laboratory Museum. Herpetol. Rev. 20:37-38. . 1988. Geographic Variation in the Seasonal Activity Cycle of Spotted Turtles, Clemmys guttata. Journal of Herpetology. 22(4): 482-485. .1989. The spotted turtles of Cedar Bog: historical analysis of a declining population. Proceedings of Cedar Bog Symposium II, Glotzhober, R.C., A. Kochman, and WT Schultz (eds.). pp.23-28. . 1990. Spring Movement Patterns of Two Radio-tagged Male Spotted Turtles. Brimleyana. 16:67-71. and TR. Jaworski. 1988. Annotated check list of amphibians and reptiles from Cedar Bog, Ohio. Ohio J. Sci. 88:139-143. Lutz, D.J. 2008. Clemmys guttata Winter Copulation in Michigan, USA. Bull. Chicago Herp. Soc. 43(11):173-174. Medsger, OF. 1919. Notes on the First Turtle I Ever Saw. Copeia. 69:29. Milam, J. and S. Melvin. 2001. Density, Habitat Use, Movements, and Conservation of Spotted Turtles (Clemmys guttata) in Massachusetts. Journal of Herpetology. 35(3):418-427. Minton, S.A. Jr. 1972. Amphibians and reptiles of Indiana. Indiana Academy of Science. Indianapolis, Indiana. pp.155. Nature Serve. Available from: www.natureserve.org. Accessed 2008. Nemuras, K. 1966. Genus Clemmys. lntemational Turtle and Tortoise Society Journal. 1:26-27, 29, 44. . 1966. Spotted Turtles in Maryland. Bull. New York Herpetological Society. pp. 6-9. Netting, MG. 1936. Hibernation and Migration of the Spotted Turtle, Clemmys guttata (Schneider). Copeia. 2:112. Nichols, J.T. 1939. Range and Homing of Individual Box Turtles. Copeia. 3:125-127. 149 Oldham, M.J. 1991. Status of the Spotted Turtle, Clemmys guttata, in Canada. COSEWIC, Ottawa. Unpublished draft report. 90pp. Perillo. KM. 1997. Seasonal movements and habitat preferences of Spotted Turtles (Clemmys guttata) in North Central Connecticut. Chelonian Conservation and Biology. 2(3):445-447. Peterson, DE. and CD. Lee. 2005. Aquatic Plants and Their Control. Kansas State University Agricultural Experimental Station and Cooperative Extension Service. pp. 1-2. Roach, A. 2006. The Spotted Turtle: North America’s Best. Living Art Publishing, Canada. pp. 2-3. Ross and Lovich. 1992. Does the Color Pattern of Two Species of Turtles lmitate Duckweed? Journal of the Pennsylvania Academy of Science. 66(1):39-42. Ruthven et al. 1928. Herpetology of Michigan. University of Michigan, Michigan Handbook Series, No. 3. pp.143-145. Schneck, J. 1886. Longevity of Turtles. Amer. Nat. 20(9):897. Scribner, K.T., M.H. Smith, and J.W. Gibbons. 1984. Genetic differentiation among local populations of the yellow-bellied slider turtle (Pseudemys scripta). Herpetologica. 40:382-387. State of Michigan Department of Environmental Quality Water Bureau. 2005. Aquatic Plants of Michigan. Accessed 2009. Ultsch. GR. 2006. The Ecology of Overwintering Among Turtles: Where Turtles Overwinter and Its Consequences. Biological Reviews. 81:339-367. Voss, E.G. 1972. Michigan Flora, Part I. Gymnosperms and monocots. Cranbrook Institute of Science Bulletin 55 and University of Michigan Herbarium. Ward, F.P., C.J. Hohmann, J.F. Ulrich and SE. Hill. 1976. Seasonal Microhabitat selections of spotted turtles (Clemmys guttata) in Maryland Elucidated by Radioisotope Tracking. Herpetologica. 32:60-64. Wilson, RR. 1989. Clemmys guttata (Spotted Turtle) Reproduction. Herp. Review. 20(3):69-70. Wilson, T.P., J.C. Mitchell, and TS. Akre. 1999. Status and conservation of the genus Clemmys in Virginia: Prospects for the future. Abstract Symposium 150 on Conservation and Ecology of the Mid-Atlantic Region. Patuxent Research Refuge and National Wildlife Visitor Center, Laurel, Maryland. Worton, B.J. 1989. Kernel methods of estimating the utilization distribution in home range studies. Ecology. 70(1):164-168. 151 Illlllllll l 1293 03062 9916 l lllllilllllll