SPECIES FACT SHEET Common Name: Western Pond Turtle Photo by Chris Brown Scientific Name: Actinemys marmorata marmorata Phylum: Class: Order: Family: (Baird and Girard, 1852) Craniata Chelonia Cryptodeira Emydidae The taxonomic status of the western pond turtle is in a state of fluctuation. First classified as Emys marmorata by Baird and Girard when, in 1841, they collected the first specimen in the vicinity of Puget Sound (Baird and Girard 1852). Strauch (1862) changed the genus to Clemmys based on examination of 158 specimens. In 1945, the two subspecies; northwestern pond turtle (Clemmys marmorata marmorata) and southwestern pond turtle (Clemmys marmorata pallida) were formally recognized (Seeliger 1945). In 2001, a new arrangement for the genus Clemmys was published by Holman and Fritz (2001), placing it into the monotypic genus Actinemys. But in 2002, Feldman and Parham (2002) placed it back to its earliest genus Emys because they did not recognize Actinemys as a monotypic genus. In 2003, the Society for the Study of Amphibians and Reptiles used Actinemys and Pacific pond turtle (Crother et al. 2008). Recent genetic studies indicate the presence of four groups or clades within the species (Bury and Germano 2008). OR/WA BLM and FS Region 6 Units where Documented: Both the Forest Service and BLM have listed the pond turtle as a Sensitive Species in Oregon and Washington. This species has been documented occurring on six National Forests/Scenic Areas (CRG, FWI, RRS, SIU, UMP, WIL) and six BLM districts in Oregon (CB, EU, KF, LV, MD, RO). Natural Heritage Program Rank and Status: Oregon State Rank: S2; imperiled in Oregon. Washington State Rank: S1; critically imperiled. National Rank: N3; Global Rank: G3G4T3Q. 1 State Status: Oregon State Status: SC; sensitive critical. Washington State Status: E; endangered. In danger of becoming extinct or extirpated from Washington. Federal Status1: SOC2; species of concern; appears to be in jeopardy, but insufficient information to support listing. Figure 1. NatureServe (2007) map of the conservation status of the western pond turtle in the United States and Canada. A species is not shown in a jurisdiction if it is not known to breed in the jurisdiction or if it occurs only accidentally or casually in the jurisdiction. Thus, the species may occur in a jurisdiction as a seasonal non-breeding resident or as a migratory transient but this will not be indicated on this map. Technical Description: 1 It was petitioned in 1992 for federal listing, but in 1993 USFWS found that listing was not warranted. Species of Concern: Taxa whose conservation status is of concern to the U.S. Fish and Wildlife Service (many previously known as Category 2 candidates), but for which further information is still needed. Such species receive no legal protection and use of the term does not necessarily imply that a species will eventually be proposed for listing. (http://www.fws.gov/oregonfwo/Species/Lists/default.asp) 2 2 Although described as an aquatic turtle found in streams, ponds, lakes, and ephemeral wetlands, the pond turtle spends a large amount of time away from water. Even though pond turtles spend much of their lives in water they require terrestrial habitats for nesting. They also often overwinter on land, disperse via overland routes, and may spend part of the warmest months in aestivation on land (Hays et al 1999). Individuals in one study spent seven months of the year at overwintering sites in Northern California (Reese and Welsh 1997). The western pond turtle is a moderate-sized turtle with a relatively low carapace (Lovich). It reaches a maximum length of 18-24 centimeters and its weight may exceed 1200 grams (Holland 1994, NatureServe 2007; Lubcke and Wilson 2007). Turtles become sexually mature at a carapace length (CL) of about 120 mm (Nussbaum et al. 1983). Maximum size varies geographically, with the largest animals (210 mm) occurring in the northern part of the range (Hays et al. 1999). Their color and markings vary geographically and sexually (Holland 1994). They are usually dark brown to olive dorsally, or blackish with darker reticulations. There is usually a pattern of dark radiating spots or lines on each scute. The plastron is yellowish, often with dark blotching in the center of the plasteral shields (Storm et al. 1995). The top of the head has black spots or lines (NatureServe 2007). The western pond turtle displays sexual dimorphism at maturity. Holland (1994) found over 10 significantly different sexually dimorphic characteristics that allow the gender of adults to be distinguished. The degree of dimorphism is variable for each character and each individual. No single characteristic is 100% reliable, so it is best to look at several characters per individual for gender determination. Adult males tend to have: flatter carapace concave plastron posteriorly thicker tail base the cloacal opening at or beyond the margin of the carapace larger head with a longer nose and pointier snout larger neck with yellow or whitish chin and throat Adult females tend to have: higher, domed carapace flatter plastron thinner tail base with the cloacal opening at or within the margin of the carapace blunt snout mustache-like markings on the upper jaw dark markings on the chin and throat Juveniles Juveniles (< 120 m CL) of both sexes tend to resemble females. 3 Hatchlings measure 25 - 31 mm CL weigh 3 - 7 grams at the time of emergence from the nest tend to be a lighter brown, darkening with age shell is soft and pliable tail is relatively long In general, the female has a smaller head, less heavily angled snout, relatively higher and rounder carapace, and a thinner tail. Males have a slightly concave plastron. Life History: The western pond turtle is one of only two native west coast aquatic turtles (the other being the western painted turtle (Chrysemys picta belli) and is one of only a few North American aquatic turtle species that has populations that routinely overwinter on land (Ultsch 2006). They live at least four decades and are most active when water temperatures are above 15ºC (Bury and Germano 2008). Females may reproduce at about 4-6 years of age in the southern and 7-10 years in the northern portions of their range (Bury 1979, Holland 1993). The time required for males to achieve sexual maturity is not known, but is thought to be at least 10-12 years in Washington (Hays et al. 1999). Mating takes place underwater and may occur in early spring (Reese 1996) or April-May in central coastal California (Rathbun et al. 1992). In general, courtship and mating behavior have been observed from February to November (Holland 1988). Possible courtship behavior reported by Holland (1988) describes an interaction between a male and a female turtle in late August that involved scratching of the anterior edge the female’s carapace by the male’s fore limbs, followed by the female raising her posterior end up off the substrate. Copulation has been observed in the field in mid-June in Southern California and in captive specimens in late August and early September (Holland 1998). Adult females of several turtle species can store sperm for months or years (Galbraith 1993, Gist and Jones 1989). Most mature females oviposit every year, and some may “double-clutch” (Holland 1994, Goodman 1997, Reese 1996). Clutch sizes range from 3-13, with most clutches containing 4-7 eggs (Holland 1994). Eggs are deposited from April to August, depending on the location, and probably peaks in June-July throughout most of the range (NatureServe 2007). Oviposition occurs on land, usually above the flood plain, from 1-50 meters from water’s edge (Holland 1994) although some females have moved as far as 402 meters from water and up to 90 meters in elevation above it (Storer 1930; Rathbun et al. 1992). For nesting, gravid (with eggs) females tend to seek out open areas with sparse, low vegetation (annual grasses and herbs), low slope angle, and dry hard soil. After voiding her bladder to soften the soil, the female excavates a pear-shaped nest chamber (scrape) with her hind feet. Females may have many “false scrapes.” Eggs are deposited and the 4 nest chamber is plugged by kneading wet soil and vegetative fragments into the throat of the nest chamber (Holland 1994, Reese 1996). Nest site philopatry (the tendency of an animal to return to a specific location in order to breed) has been observed in several species of turtles and there is some evidence to suggest nest site philopatry is exhibited by this species (Holland 1994). Eggs are hard shelled and oval in shape, measuring 31-38 mm long by 20-24 mm wide and weighing 8-10 g (Holland 1994). Incubation takes about three months and hatching rates are about 70% (Holland 1994). Upon hatching, hatchlings have an umbilical yolk sac, which provides nourishment in the early stages of growth (Holland 1991, Voight 1963). Hatchlings either emerge in the fall or overwinter in the nest chamber and emerge in the spring (Holland 1994, Reese 1996). Hatchlings over-wintering in the nest chamber receive nourishment from their umbilical yolk sack (Holland 1994, Voight 1963). Variable growth rates are seen in this species. Growth rates are highest in hatchlings, which can almost double in size by the end of the first growing season (Holland 1991). Growth rates may be regulated by environmental factors, such as water temperature and prey base (Goodman 1997). Age structure studies often rely on counting annuli on the plastral scutes (Zug 1991, Germano and Bury 1998). Annuli are the result of alternating periods of maximum growth and minimum growth; counting annuli is based on the assumption that one growth ring is added each season (Ashton et al. 1997). Numerous studies show that growth rings on scutes match age in young turtles (Germano and Bury 1998). Western pond turtles locate food by sight or smell. Under normal conditions feeding behavior is solitary (Hays et al. 1997) and foraging apparently does not occur on land (Rathbun et al. 2002). The pond turtle is considered a dietary generalist (Bury 1986). They prefer live prey, which they capture by opportunistic foraging tactics, but will also scavenge carrion and browse on plant material. Prey items are ingested in the water and it appears this species is unable to swallow in air (Holland 1994). Preferred food items include aquatic insect larvae, crustaceans, and annelids. On occasion, small vertebrates (including Rana boylii tadpoles and egg masses) have been found in its diet (Bury 1986; Holland 1994). Western pond turtles spend several hours per day basking, and are more abundant in habitats that have basking sites. Basking has been noted in all months of the year, but generally increases in frequency through the spring to a peak in early to mid-June (Holland 1994). Turtles use a variety of sites for basking, such as rocks, mud, downed logs, branches of near-shore vegetation, and emergent or submerged aquatic vegetation (Hays et al. 1999). Turtles engage in aquatic basking (Holland 1985a) by concealing themselves in or under floating vegetation or cover in shallow water. Overwintering is a period of reduced or no activity, which may include periods of a hibernation-like state of reduced physiological activity, from mid-October or November to March or April the following year (Hays et al. 1999). According to Holland (1994), there is a tendency for turtles from ponds and lakes to hibernate underwater while turtles from 5 rivers and streams overwinter on land (possibly to avoid being swept away by winter and early spring floods). They can overwinter on land up to 500 m from the nearest watercourse (Reese and Welsh 1997), sometimes changing sites during the season (Holland 1994, Slavens 1992). When overwintering terrestrially, turtles will burrow in duff or soil (Ashton et al. 1997) where the duff and leaf or needle litter is 2–20 cm thick (Holland 1994). Radio-telemetry studies have shown individuals often return to the same terrestrial overwintering site each fall (Reese 1996, Goodman 1996). Movement to overwintering sites occurs from September to November, while emergence from terrestrial overwintering sites occurs from March to June. Occasional overland movements (usually less than 3 km) occur. Turtles can overwinter in the mud at the bottom of ponds, sometimes communally. If there is no suitable terrestrial habitat available for overwintering, then they will not leave the water during winter (Lovich & Meyer 2002). At one site in Oregon, there were 43 turtles found within one square meter. There is also some degree of winter activity in aquatic hibernacula; they have moved freely in one lake in Oregon at temperatures down to 1°C, and basked in air temperatures as low as 6°C (Holland, 1994). Most western pond turtles are somewhat sedentary, although they are capable of moving significant distances and occasionally travel several hundred meters in just a few days (Bury 1979). Most of their movements occur along river and stream courses (Rathbun et al. 1992). Reese (1996) found that over the summer months (May-September) juvenile turtles have an average maximum movement of approximately 84 meters. Their mean weekly aquatic travel is 19.9 meters. Juveniles sometimes travel back and forth between low-flow portions of the river and adjacent ponds. Their home range is smaller than adults and also includes terrestrial components (Reese 1996). Western pond turtles in a stream environment in northern California had average home ranges of about 1 ha for adult males, 0.25 ha for adult females, and 0.4 ha for juveniles (Bury 1979). Considerable overlap in home ranges of individuals of both sexes occurred in this area. Individuals may occasionally make sporadic long-distance aquatic movements outside their home range (Holland 1994). Range, Distribution, and Abundance: The western pond turtle has an appreciable geographic range, from Washington (historically British Columbia) to northern Baja California (NatureServe 2007). It appears to be declining in numbers in parts of its range, particularly in Washington, northern Oregon, southern California, and Baja. It remains fairly common in the center of its range from southern Oregon to central California (Bury and Germano 2008). It is now reduced from most of its range in Washington. Two populations remain in the Columbia River Gorge. The total number of western pond turtles in Washington was estimated at only 250-350 individuals in 1999 (Hays et al. 1999). Additional turtles were believed to still occur in wetlands that have not been surveyed in western Washington and the Columbia Gorge (Hays et al. 1999). Today, there are an estimated 1,250 wild turtles in Washington, according to the Woodland Park Zoo in Seattle, Washington, as a result of an active captive breeding and re-introduction program. 6 Figure 2. Approximate historic range of the western pond turtle in Washington (circa 1850) and in North America (figure from Hays et al. 1999). Distribution and abundance of the pond turtle have declined as a result of past commercial exploitation, habitat loss and degradation, introduced species, and disease. The pond turtle is discontinuously distributed and uncommon in parts of its range. Isolated populations occur in the Mojave River in California, the lower Columbia River (two populations in Washington, one in Oregon), the Puget Sound Trough in northern Washington (rarely observed), and in areas south of the Transverse Ranges in southern California and adjacent Baja California (NatureServe 2007). An isolated population east of the Sierra Nevada in extreme western Nevada may be an introduction (Cary 1887), although this has never been formally examined and a better review of the literature is needed. 7 Figure 3. A compilation of northwestern pond turtle observations in OR and WA from the NRIS, GeoBob and ORHIC databases (as of the date of this document). Additional observations outside of these databases are not represented here. 8 Pond turtle densities range from a few to an estimated 3,700 turtles/ha (this from one small pond; Holland 1991). Typically, they are found at much lower densities. In western Oregon, Nussbaum et al. (1983) stated that one oxbow lake of 1.5-2.0 ha contained an estimated 75 individuals. Another oxbow lake of about the same size contained an estimated 180 individuals. There are estimates that population declines may be occurring over 80% of its range (Holland 1994; Hays et al 1999), but field surveys are lacking. Northern California and southern Oregon support extensive populations (Ashton et al. 1997), representing over half the species’ range. Habitat Associations: The name "pond" turtle is something of a misnomer, as ponds are relatively scarce throughout most of the range of this species, and the turtles are more often associated with rivers and streams (Hays et al 1999). They occur in rivers, streams, lakes, ponds, reservoirs, stock ponds, and permanent and ephemeral wetland habitats (Bury et al 2001). They are usually rare or absent in reservoirs, impoundments, canals, or other bodies of water heavily altered by humans (Hays et al. 1999), but there are notable exceptions. Further research is needed on this topic. They can be found from sea level up to 1,370m or even at higher elevations in the southern part of their range (Bury et al 2001). Favorable pond turtle habitat is characterized by deep, slow-flowing pools with underwater cover and emergent basking sites, warm water, or both (Reese and Welsh 1998). Habitats used by western pond turtles may have a variety of substrates including solid rock, boulders, cobbles, gravel, sand, mud, decaying vegetation, and combinations of these. In many areas turtles are found in rocky streams with little or no emergent vegetation. In other areas they occur in slow-moving streams or backwaters with abundant emergent vegetation such as cattails or bulrush (Holland 1991). Terrestrial basking sites include mud banks, rocks, logs, and root wads on the bank and are within or immediately adjacent to water (Ashton et al. 1997). Pond turtles nest on sandy banks near water or in fields or sunny spots up to a few hundred meters from water (Nussbaum et al. 1983, Storer 1930). The nest site has good exposure to the sun and compact soil (Holland 1994, Reese 1996). Aspect is usually south or west facing and on a slope of 25 degrees or less (Bury et al 2001). Overwintering can be aquatic or terrestrial (Holland 1994). Terrestrial overwintering site characteristics are highly variable, but the microsite usually includes a thick duff layer (Holland 1994). Aquatic habitats with access to areas of deep slow water with underwater refugia and emergent basking sites are favored by adult and juvenile turtles. Hatchlings are relatively poor swimmers and tend to seek areas with slow, shallow, warmer water, often with emergent vegetation (Ashton et al. 1997). Bodies of water can be classified based on the 9 suitability of aquatic habitat. Those habitat quality classifications may be considered from a range from unsuitable to favorable. Three major habitat features to consider are (Bury et al. 2001): Size of water - Large bodies of standing water tend to have low mean temperatures, and thus generally provide less suitable habitat than smaller bodies of water. Depth of water - Bodies of water where most of the system has a depth less than 2 meters provide accessible underwater refugia for turtles. Basking structures – Basking is an important activity and lack of emergent basking structures can indicate poorer habitat. Threats: The greatest single threat to the pond turtle is habitat destruction, alteration and fragmentation (Ashton et al 1997, NatureServe 2007). Habitat impacts can be caused by conversion of wetlands to farmland, water diversions and dams, channelization, mining, logging, and urbanization. Associated with habitat fragmentation is the effect of genetic isolation, although genetic inbreeding is unproven. Basking, foraging, nesting, and other behaviors can be significantly limited by human presence, including foot, bicycle, boat, and motor vehicle traffic (Holland 1991). Incidental catches by fisherman may account for significant losses in some areas (Holland 1991). Turtles that don’t die from trauma associated with ingestion of fishing tackle probably starve to death. Encounters with automobiles may also contribute significantly to mortality. Turtles are hit by traffic when moving to and from the watercourse during terrestrial journeys, such as overwintering or nesting movements (Holland 1991). Hatchlings on their way from the nest to water are also at risk from vehicular traffic (Holland 1991). Commercial harvest of pond turtles was extensive in California from at least the 1870’s to ca. 1930’s. Many turtles were sold in San Francisco markets for human consumption, probably resulting in significant declines in many populations (Holland 1991; Bettelheim 2005). Commercial harvest of pond turtles is now prohibited in all states where the species lives. Illegal collection of pond turtles for the pet trade still occur, but at an undetermined level. Death by gunshot wound is also a significant threat in some areas (Holland 1991). Chemical spills pose a threat, especially near highways, railways and industrial areas. Holland (1994) reports there were several contaminant spills into aquatic habitats inhabited by this species in Oregon during 1993. One was a diesel oil spill that impacted 50 to 100 turtles, resulting in the death of some animals and physiological complications in many others. Chemical spills and pollution can kill turtles directly, or indirectly by removing prey base, degrading habitat quality, and/or increasing the risk of disease (Holland 1991). 10 Raccoon, spotted skunks, coyote, fox, feral and domestic dogs, black bear, river otter, mink, osprey, bald eagle, bullfrog, and largemouth bass are known to prey on pond turtles (Holland 1994). In some areas, most of the turtle nests are lost to predation (Holland 1994). Invasive species such as bullfrogs (Rana catesbeiana) and largemouth bass (Micropterus salmoides) may eat hatchlings (Holland 1994). As many large populations of turtles occur in the presence of these predators this is not a universal threat. Other suspected predators include herons, golden eagle, beaver, and nutria (Holland 1994). Adult turtles often show scarring on the shell and/or missing limbs, indicating attempted predation. Holland (1994) found a six-fold greater scarring rate on females and attributed it to greater exposure to predators during nesting movements. Hatchlings are especially vulnerable to predators because their shell is soft and they can be swallowed whole. In many cases only nests that are dug up are observed and so it is difficult to determine how many nests survive. Reports on disease epidemics are limited. In 1990, an upper respiratory disease (URD) outbreak in Washington resulted in losses of up to 40% (Holland 1994), leaving a total population of fewer than 100 individuals in the wild (Hays et al. 1999). In 1993, a single pond turtle carcass was found near Eugene, Oregon, exhibiting symptoms of URD. The method of disease introduction to these populations is unknown, but exotic turtles are known to have been introduced into Washington and Eugene waters with native western pond turtles and may have acted as vectors (Holland 1994). Severe drought can have a significant negative impact on pond turtle populations. The 1987-1992 drought in California reduced southern and central state populations by up to 85% (Holland 1994). Estivation occurs in response to drought. Pond turtles utilizing a vernal pool in Napa County, California moved upland and burrowed into leaf litter when the pond dried up (Reese 1996). Estivation may also occur in response to unseasonally cold water releases from upstream dams (Goodman 1997). Pond turtles are not especially strong swimmers and can be washed downstream in high flows. Over-wintering on land may be an adaptation to escape winter high flows. Turtle nests may be inundated during floods (Ashton et al 1997). Loss of lakeside emergent wetland vegetation to grazing and trampling may have made habitat less suitable for hatchlings and juveniles. Fire can kill turtles over-wintering on land and hatchlings still in the nest. Fire can also have adverse effects on the aquatic environment and macroinvertebrate prey base by increasing siltation (Holland 1991). Successional changes through fire suppression on native grasslands may have resulted in excessive shade on nesting grounds (Hays et al. 1999). 11 Conservation Considerations: NatureServe (2007) recommends protecting known large3 populations and also suggests focusing on protection of natural ecosystems, rather than on captive breeding or translocation. For adequate protection, these areas should include aquatic habitat and adjacent upland habitat for nesting (NatureServe 2007). The Washington Department of Fish and Wildlife is actively engaged in management and recovery efforts for the western pond turtle. These conservation activities are being conducted in cooperation with the Woodland Park Zoo in Seattle, Washington and the Oregon Zoo in Portland. This program is called the “Western Pond Turtle Recovery Project” and is focused on the Puget Sound and Columbia Gorge populations. The objective of the program is to increase the survival chances of young turtles in the wild by allowing hatchlings to grow to a size where they are too big to be preyed upon by bullfrogs. This method has been demonstrated to be effective (Haskell et al. 1996). Captive juvenile turtles attain the size of a 2-year old wild turtle in a single year. As of 1999, the Zoos had successfully reared and released 187 juvenile western pond turtles under this head-start program (Hays et al. 1999). Currently, over 1,000 turtles have been released back into the wild. The recovery objectives for the State of Washington are to establish at least 5 populations of >200 pond turtles, composed of no more than 70% adults, which occupy habitat that is secure from development or major disturbances. It is also necessary that the populations show evidence of being sustained by natural recruitment of juveniles. The core pond turtle sites should be wetland complexes that may be less susceptible to catastrophes than sites of a single water body (Hays et al 1999). The majority of the populations on National Forests or BLM Districts in Oregon and Washington occur within “Riparian Reserves”, a land management classification under the Northwest Forest Plan Standards and Guidelines (USDA and USDI 1994). Riparian reserves are designed to provide protection to riparian and aquatic systems and the species associated with those systems. Riparian reserve widths are generally determined by whether the water body is fish-bearing or not as well as the potential for tree growth within the area. However, Aquatic Conservation Strategy Objectives also encourage riparian widths to be designed to “maintain and restore habitat to support welldistributed populations of native plant, invertebrate, and vertebrate riparian-dependent species”. Where populations of pond turtles are present on FS and BLM managed lands, consider increasing riparian reserve widths to encompass known or potential overwintering habitat if monitoring indicates that not doing so would put these populations at risk. Consider protection of pond turtle nesting habitat in areas of high human use such as recreation or fishing sites. Turtle habitat adjacent to camping or fishing areas can be fenced and signed to reduce impacts from disturbance. Trails that impact turtle habitat can be relocated to avoid these sensitive areas. Interpretive signing in high use areas can also be considered. 3 Large is not defined nor are locations specified 12 Protect known turtle areas from prescribed fire to reduce mortality to overwintering adults and hatchlings still in the nest. In areas of pond turtle habitat, conduct burning after turtles have returned to ponds or streams. Where feasible, eradicate invasive species such as bullfrogs and bass to reduce juvenile mortality. Development of a “site management plan” may be considered where threats to the site/population may be numerous or complex. The format for the Site Management Plan can be found at: http://www.fs.fed.us/r6/sfpnw/issssp/documents/planning-docs/cp-smp-format-200704.doc Other pertinent information (references to Survey Protocols, etc): See Bury et al 2001 for Survey Protocol (57 pages). Bury, R.B., C. Barkhurst, R. Horn, L. Todd, S. Wray, R. Goggans, K. Beal, N. Sisk. February 2001. Western Pond Turtle: Survey Protocol and Monitoring Plan. Interagency Western Pond Turtle Working Group. Final Draft. ATTACHMENTS: Appendix 1. Key To Sexually Dimorphic Characters In The Western Pond Turtle (Clemmys marmorata) (Holland 1994). Preparer: Theresa Stone, Umpqua National Forest, Roseburg, OR Date Completed: 5 October 2007 Updated: May 2009 13 References: Ashton, D.T., A.J. Lind, and K.E. Schlick. 1997. Western Pond Turtle (Clemmys marmorata) Natural History. USDA Forest Service, Pacific Southwest Research Station, Redwood Sciences Laboratory, Arcata CA. 22 pp. Bettelheim, Matthew P. 2005. Marmorata: The Famed Mud Turtle of the San Francisco Market. California History. 82(4): pp 26-42. Bury, R. B. 1972. The effects of diesel fuel on a stream fauna. California Department of Fish & Game Bull. 58:291-295. Bury, R. B. 1979. Population ecology of freshwater turtles. Pages 571-602 in M. Harless and H. Morlock, editors. Turtles: perspectives and research. John Wiley & Sons, Inc. Reprinted in 1989 by Robert E. Krieger Publ. 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Master’s thesis, California State Polytechnic University, Pomona, CA. 81 pp. Haskell, A., T. E. Graham, C. R. Griffin, and J. B. Hestbeck. 1996. Size related survival of headstarted redbelly turtles (Pseudemys rubriventris) in Massachusetts. Journal of Herpetology 30:524-527. Hays, D. W., K. R. McAllister, S. A. Richardson, and D. W. Stinson. 1999. Draft Washington State recovery plan for the western pond turtle. Wash. Dept. Fish and Wild., Olympia, Wash. 53 pp. Holland, D. C. 1985a. An ecological and quantitative study of the western pond turtle (Clemmys marmorata) in San Luis Obispo County, California. M.S. Thesis, Calif. State Univ., Fresno Holland, D. C. 1988. Clemmys marmorata. Western pond turtle: Behavior. Herp. Review 19:87-88. Holland, D. C. 1991. A synopsis of the ecology and status of the western pond turtle (Clemmys marmorata) in 1991. Prepared for the USFWS, National Ecology Research Center, San Simeon Field Station. Holland, D. C. 1991b. 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University of California, Publications in Zoology 32(5):429441. Storm, R. M., W. P. Leonard, H. A. Brown, R. B. Bury, D. M. Darda, L. V. Diller, and C. R. Peterson. 1995. Reptiles of Washington and Oregon. Seattle Audubon Society. 176 pp. Ultsch, G.R. 2006. The Ecology of Overwintering Among Turtles: Where Turtles Overwinter And Its Consequences. Cambridge University Press; published online by Cambridge University Press. USDA Forest Service, and USDI Bureau of Land Management. 1994a. Record of decision for amendments to Forest Service and Bureau of Land Management planning documents within the range of the northern spotted owl; standards and guidelines for management of habitat for late-successional and old-growth forest related species within the range of the northern spotted owl. Portland, Oregon. USDA Forest Service, and USDI Bureau of Land Management. 1994b. Final supplemental environmental impact statement on management of habitat for latesuccessional and old-growth forest related species within the range of the northern spotted owl. Portland, Oregon. 2 volumes and appendices. Voight, V. F. 1963. Picta, the painted turtle. G. P. Putman’s Son’s. New York, N.Y. 63 pp. Zug, G. R. 1991. Age determination in reptiles. Society for the Study of Amphibians and Reptiles, Herpetological Circular No. 20. 17 Appendix 1. Key To Sexually Dimorphic Characters In The Western Pond Turtle (Clemmys Marmorata) (Holland 1994). CHARACTER a. neck FEMALE lateral and dorsal surfaces of head and neck usually mottled or ocellate MALE lateral and dorsal surfaces of head and neck often uniformly colored, especially in older animals b. nose nose relatively short nose relatively long c. maxilla maxilla often with fine dark vertical lines or “mustache” maxilla lightly marked or unmarked, especially in older animals d. snout angle of nose vertical or nearly vertical angle of nose usually 10-15 from vertical e. throat often flecked with numerous small dark flecks usually lightly marked or unmarked f. vent usually at or slightly posterior to posterior edge of carapace usually well posterior to posterior edge of carapace g. tail usually relatively long and thin usually relatively short and thick h. plastron area of femoral/anal seam junction usually flat area of femoral/anal seam junction usually slightly concave i. shell shell relatively high/deep in relation to length of carapace shell relatively low/shallow in relation to length of carapace 18