addendum a. species and habitat descriptions · lomatium utriculatum) and western buttercup...

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Addendum A. Species and Habitat Descriptions

3.1 Invertebrate Animals

For each of the five butterfly species that are addressed in the HCP, we include a description of its conservation status, population trends and distribution, life history and ecology, and habitat characteristics. When known, we also include species-specific threats and causes for decline. These causes are in addition to a common set of causes for decline for all five butterfly species, which include:

• Habitat Loss: o Habitat loss is the consistent, primary factor driving species extinctions and declines

world-wide (Groom et al. 2006), and the most common threat to butterfly populations (New et al. 1995). Prairies and oak woodlands in south Puget Sound have been converted to development, agriculture, gravel mines, and lost to forest succession resulting from elimination of fire and other beneficial sources of disturbance. In 1997, Crawford and Hall conservatively estimated that over 60,000 ha (>148,263 ac) of prairie existed historically in the south Puget Sound region, and that only 3% of that remained dominated by native vegetation. Prairie loss likely has continued since 1997, but current estimates are not available for this region. Refined the estimates of grassland habitat for the entire WPG ecosystem, and estimated the total amount of prairie, oak woodland, and grassland bluffs and balds prior to Euro-American settlement was over 72,000 ha (180,000 ac) (Chappell et al. 2001).

• Habitat Fragmentation: o Crawford and Hall (1997) found that historically in south Puget Sound there were

233 prairie sites, averaging 250 ha (618 ac) in size, including 18 large prairies (>405 ha), and contrasted that to 1997 conditions: 29 prairie sites, averaging 175 ha (432 ac) in size, with only 2 large prairies extant. Fragmentation of prairies directly threatens prairie butterflies by creating smaller and isolated populations, which increases the potential for population loss and inbreeding.

• Invasion of Prairie Communities: o Invasive plants have dramatically altered the ecological function of Pacific

Northwest prairies (Dunwiddie and Bakker 2011). Woody shrubs, including Scotch broom, and non-native grasses, especially tall oatgrass (Arrhenatherum elatius), bentgrasses (Agrostis), and sweet vernal grass (Anthoxanthum odoratum) have invaded most extant south Puget Sound prairies. Uncontrolled, these plants dominate native prairie vegetation, excluding butterfly host and nectar plants, and

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change vegetation structure and soil conditions. The invasion of native grasslands by exotic plants is a common threat to grassland butterflies (Warren 1993, Schultz 1998), and has occurred rapidly at historic and extant prairie butterfly sites (e.g., Hays et al. 2000). Many butterflies avoid areas with tall vegetation, including areas with Scotch broom and tall oatgrass (e.g., Hays et al. 2000, Potter and Olson 2012, Henry and Schultz 2012).

• Alteration of Natural Disturbance Regimes o Native Americans regularly burned prairies in the Pacific Northwest to support food

production and manage hunting sites (Norton 1979, Boyd 1986, Agee 1993), and this process supported open prairie and savannah. Soil disturbance also regularly occurred from both Native American harvest of bulbs and rhizome plant material (Turner 1999), and the activity of burrowing mammals, especially the Mazama Pocket Gopher (Huntly and Inouye 1988). Cultural practices changed when Euro-Americans began to settle the Pacific Northwest and the prairies; soil and vegetation disturbance from fire setting and prairie plant harvesting ceased. Encroachment by trees and shrubs, first native species and then non-native, combined with introductions of invasive grasses and herbaceous species, resulted in the loss of prairie due to forest encroachment, and dramatic alterations to the extant prairie.

• Prairie management. o Fire, herbicide use, mowing, and other prairie management techniques are

important tools for re-creating or simulating disturbance mechanisms that historically maintained prairies, reducing invasive species, and restoring endangered species habitat connectivity (Dunwiddie and Bakker 2011, Schultz et al. 2011). These prairie management practices, implemented to restore or enhance prairie vegetation and wildlife habitat, also can directly or indirectly harm butterflies (Schultz et al. 2011). Effects of these practices on butterflies are not completely understood. Prairie management in areas occupied by butterfly species of concern is necessary and must be undertaken with special methods and considerations to reduce or eliminate harm to these species.

3.1.1 Hoary Elfin (Callophrys polios) Puget Trough segregate

3.1.1.1 Conservation Status

The Hoary elfin is a butterfly in the Gossamer Wings Family (Lycaenidae). The Hoary Elfin is a south Puget Sound endemic and was recognized as a butterfly of conservation concern in the first Washington butterfly conservation status assessment (Pyle 1989) due to the small number of isolated populations, specialized and restricted habitat, and known threats to their habitat. Since that time, renewed taxonomic assessment, limited focal surveys by a few agencies and individuals, as well as limited life history observation have identified extant populations and confirmed the restricted nature of Hoary Elfin habitat and distribution.

Today, this butterfly remains a species of conservation concern (Schultz et al. 2011). In Washington, it is listed by the Department of Fish and Wildlife (WDFW) as one of 19 butterfly Species of Greatest

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Conservation Need in Washington’s Comprehensive Wildlife Conservation Strategy (WDFW 2005). The species is not listed by the U.S. Fish and Wildlife Service (USFWS).

3.1.1.2 Population Trends and Distribution

The Hoary Elfin Puget Trough segregate inhabits low-elevation grasslands (Pierce and Thurston counties) and open, heath woodlands (Mason and Kitsap counties) in the south Puget Sound region. The Puget Trough Hoary Elfin is a remarkably disjunct subspecies; neighboring polios subspecies occur hundreds of miles distant in far northeastern Washington, the southern central Oregon Coast, and in the Oregon Blue Mountains. Fewer than a dozen sites are known from Kitsap and Mason counties, and Hoary Elfin has not been confirmed on most for several decades (Hinchliff 1996). Four populations are documented from southern Pierce County (Hinchliff 1996; Gilbert pers. comm.); only one population has recently been confirmed extant and it inhabits the Artillery Impact Area of Joint Base Lewis-McChord (Gilbert pers. comm.). Ten populations are documented in Thurston County, including 2 on Joint Base Lewis-McChord; 5 are known extant (see Table 1).

Table 1. Known locations for Hoary Elfin in Thurston County, with current status as of 2014.

Location Last Record Status Bald Hill Natural Area Preserve 2014 Extant Johnson Prairie, Joint Base Lewis-McChord 2007 Unknown Mima Mounds Natural Area Preserve 2014 Extant Rocky Prairie Natural Area Preserve 2014 Extant Scatter Creek Wildlife Area North 2014 Extant Scatter Creek Wildlife Area South 2004 Unknown Tenalquot Prairie 2013 Extant Weir Prairie, Joint Base Lewis-McChord 2005 Unknown West Rocky Prairie Wildlife Area 1983 Unknown Wolf Haven 2006 Unknown

Most Hoary Elfin populations in Thurston County consist of a small number of individuals (<50), with a few sites likely supporting populations of over 200-300 individuals (A. Potter, WDFW, unpubl. data). Hoary Elfins venture little from their natal host Kinnikinnick (Arctostaphylos uva-ursi) patches, such that multiple separate populations and/or metpopulations likely occur within the same site.

3.1.1.3 Life History and Ecology

Hoary Elfins are highly sedentary butterflies; they do not migrate and spend their entire life cycle (egg, larva, pupa and adult) within or close to Kinnikinnick host plant patches, typically within 10-20 meters (A. Potter, WDFW, unpubl. data). Hoary Elfins are univoltine; which means they complete a single life cycle annually. One of the first butterflies to emerge in the spring on south Puget Sound prairies, adults are present between late-April and late-May. Males begin emerging first, followed by females; late-season individuals are primarily or solely females (A. Potter, WDFW, unpubl. data).

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Weather influences butterfly emergence and the flight period duration, with wet or cold conditions delaying emergence, and conversely, warm, dry conditions promoting earlier emergence. Weather conditions also affect the total flight period duration.

Hoary Elfins often perch motionless or while rubbing their hindwings together on Kinnikinnick (Arctostaphylos uva-ursi), a low-growing, prostrate, woody shrub in the Ericaceae (Heath or Heather) family. The butterfly’s deep brown, mauve, and frosty (hoary) wing coloring camouflages them superbly in Kinnikinnick. Adults court, mate, and nectar in or near Kinnikinnick; eggs are laid singly, larvae feed, and pupae overwinter on the host plant as well (Pyle 2002).

James & Nunnallee (2011) reared Hoary Elfins in captivity and describe their life cycle and life stages (egg, larva, pupa) (pp. 176-177). Eggs hatched in 5-9 days, larvae fed on kinnikinnick leaves and had 4 instar stages, and pupation occurred in protected locations approximately 25-29 days post egg-hatch (James and Nunnallee 2011).

Kinnikinnick is the sole host for this butterfly, and is also the primary nectar source for Hoary Elfin. Other spring-flowering species growing within or adjacent to Kinnikinnick are occasionally also used, they include spring gold (Lomatium utriculatum) and western buttercup (Ranunculus occidentalis) (A. Potter and L. Beyer, WDFW, unpubl. data).

Hoary Elfin population size and distribution are directly related to the size and location of their host plant patches. Large, multi-patch networks of Kinnikinnick can support large butterfly populations. Like many other native grassland butterflies (Severns 2008, Severns and Warren 2008, Henry and Schultz 2012), Hoary Elfin also requires low-growing vegetation; tall grasses, shrubs, or trees that invade Kinnikinnick patches reduce and can even eliminate habitat for this species.

3.1.2 Mardon Skipper (Polites mardon W.H. Edwards, 1881) -


The Mardon skipper is a butterfly in the Skipper Family (Hesperiidae). The Mardon Skipper was recognized as a butterfly of conservation concern in the first Washington butterfly conservation status assessment (Pyle 1989) due to its highly limited range, small number of populations, and known threats to their south Puget Sound prairie habitat. Since 1989, extensive surveys have assessed status of historically documented populations and searched for undiscovered sites across the butterfly’s range (Fleckenstein and Potter 1999, Harke and LaMarr 2000, Potter and Fleckenstein 2001, Harke 2001, Potter et al. 2002, Haggard 2003, Jepsen et al. 2007a, Jepsen et al. 2007b, Jepsen et al. 2007c, Jepsen et al. 2008, St. Hilaire et al. 2010, Kerwin 2011, unpubl. data: B. Bidwell, V. Harke, USFWS, T. Kogut, USFS, K. McAllister, WDFW, A. Potter, WDFW). Through these efforts, several populations located in the past, particularly in the south Puget Sound region, were determined to be extinct, many new populations were found in the Oregon and Washington Cascade Mountain Range, and a few populations were discovered in coastal California and Oregon.

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Today, the Mardon Skipper is a species of conservation concern throughout its range. In Washington, it is listed endangered by WDFW, and is 1 of 19 butterfly Species of Greatest Conservation Need (SGCN) in Washington’s Comprehensive Wildlife Conservation Strategy (CWCS) (WDFW 2005). The USFWS designated Mardon Skipper as a candidate for listing under the Endangered Species Act in 1999 (USFWS 1999). In a recent opinion, USFWS changed the species status to species of interest, based primarily on recent discovery of populations in the Oregon and Washington Cascade Mountains (USFWS 2012). The U.S. Forest Service and Bureau of Land Management list Mardon Skipper as a sensitive species (Kerwin 2011).


The Mardon Skipper occurs in 4 geographically distinct regions in Washington, Oregon, and California (Fig. 1) (Potter et al. 1999). It was originally described by W.H. Edwards in 1881 from material collected on “prairies of Washington Territory … near Tenino” (Morrison 1883, Brown and Miller 1981). At the southern most edge of their range, near the California and Oregon border, the butterfly exists on small, coastal grassland sites (Haggard 2003, Ross, lepidopterist, pers. comm.). Another group of Mardon Skipper populations inhabit meadows in the Cascade Mountain Range of southern Oregon, approximately 80-100 miles (129-161 km) northeast of the coastal populations (Hinchliff 1995). Despite searches by the U.S. Forest Service (Kerwin 2011) and others, no populations have been found further north in Oregon.

In the Washington Cascades, Mardon Skippers occur patchily in meadows and old clearcuts in the dry and transitional forest zones surrounding Mt. Adams, especially within the Gifford-Pinchot National Forest, Okanogan-Wenatchee National Forest, and Yakama Indian Reservation, from the communities of Trout Lake and Glenwood communities and east to Rimrock Lake (sites are in Klickitat, Skamania, and Yakima Counties) (Hinchliff 1996, Harke and LaMarr 2000, Harke 2001, Potter and Fleckenstein 2001, St. Hilaire et al. 2010, Kerwin 2011, G. King, YIN, unpubl. data). The U.S. Forest Service and USFWS identified approximately 40 Mardon Skipper populations in the southern Cascades of Washington (USFWS 2012).

The furthest north region inhabited by Mardon Skipper is south Puget Sound, where the butterfly inhabits glacial outwash prairies (Fig. 2) (Pyle 1989, Hinchliff 1996). Mardon Skipper was historically documented from 4 prairie sites (Hinchliff 1996) and was likely more widespread in this region. Recent intensive survey efforts determined the 4 historic sites no longer support Mardon Skipper populations, and located new populations on 4 prairie sites (Char and Boersma 1995, Potter et al. 1999, Fleckenstein and Potter 1999, Chramiec 2004, Wolford et al. 2007, unpubl. data: B. Bidwell, lepidopterist, V. Harke, USFWS, T. Lamarr, USFWS, A. Potter, WDFW; P. Dunn, TNC, pers. comm.). These 4 populations, 2 in southwestern Thurston County and 2 within Joint Base Lewis McChord (JBLM), are extant (Beyer 2012, D. Hays, WDFW pers. comm., A. Potter, WDFW, unpubl. data) and fall within the HCP area.

Mardon Skipper was first detected on JBLM, within the Artillery Impact Area (AIA) in 1997 during an incidental encounter (P. Dunn, TNC, pers. comm.). JBLM and WDFW biologists subsequently planned a comprehensive survey effort of the AIA, which JBLM undertook from 2004-2007

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(Chramiec 2004, Wolford 2006, Wolford et al. 2007). The JBLM Mardon Skipper populations are located on the outer edges of the AIA. Mardon Skipper numbers at AIA have been monitored incidentally during monitoring for the Taylor’s Checkerspot (Olson and Linders 2010, Linders 2012, M. Linders, WDFW, unpubl. data, A. Potter, WDFW, unpubl. data) and standardized butterfly counts were conducted on AIA sites by Wolford et al. (2007) and A. Potter (WDFW, unpbul. data). Daily Mardon Skipper counts from these efforts on JBLM sites have typically been low, except for high daily counts of 85 individuals on the Mortar Point 12 site and 65 individuals on the Range 76 site in 2007 (Wolford et al. 2007).

Systematic transect surveys conducted for Mardon Skipper on the Scatter Creek Wildlife Area in 2008 and 2009 yielded estimates of 100s of skippers occurring on the North Unit of Scatter Creek Wildlife Area, and likely over 1,000 butterflies inhabiting the Scatter Creek South Unit (Potter and Olson 2012). A. Potter and D. Hays (WDFW, pers. comm.) conducted Mardon Skipper surveys at Scatter Creek since 1997 and observed butterfly densities decrease noticeably in the primary Scatter Creek habitat areas during that time.


The Mardon Skipper is a small (20-24 mm; <1 in), tawny-orange butterfly with a stout, hairy body (Potter et al. 1999). The upper surface of both wings is orange with broad dark borders. The wings from below are light tan-orange with a subtly distinctive pattern of light-yellow to white rectangular spots. Males are smaller than females and have a small, dark streak (stigma) on the upper surface of the forewing. Like most members of the Hesperiinae, Mardon Skippers have a fast, skipping flight, bent antennae clubs, and a characteristic basking posture in which the forewings are held at a 45o angle and the hind wings are fully spread. On the south Puget Sound prairies, the Mardon Skipper look-alike that occurs during the same season is the Sonora Skipper (Polites sonora).

Mardon Skippers are univoltine; completing a single life cycle annually. They are a sedentary butterfly, they do not migrate; instead the species inhabits sites where they are found year-round (as an egg, larva, pupa and adult). In the south Puget Sound region, adults typically begin to emerge from chrysalids (pupae) in early-to-mid May, and the adult flight period often lasts into mid-June (Hinchliff 1996, A. Potter, WDFW, unpubl. data). Males emerge first, followed by females; late-season individuals are primarily or solely females (A. Potter, WDFW, unpubl. data). Weather influences butterfly emergence and the flight period duration, with wet or cold conditions delaying emergence, and warm, dry conditions promoting earlier emergence, and weather conditions also affect the total flight period (Potter et al. 2002).

Male Mardon Skippers seek mates by perching on low vegetation in specific places and then darting out to inspect passing butterflies to determine if they are female (A. Potter, WDFW, unpbul. data). Males detecting females commence courtship behavior; when males detect another male, a territory defensive behavior of tight, upward spiraling flight ensues (A. Potter, WDFW, pers. comm.). Females seek egg-laying sites by slowly flying and hovering just above the grassland vegetation, landing on their host plant grasses, and then quickly laying single eggs (A. Potter,

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WDFW, pers. comm.). Mardon Skipper egg-laying in the south Sound has been closely observed, and females have been found ovipositing (egg laying) mainly on the native prairie bunchgrass, Roemer’s Fescue (Festuca roemeri), but also occasionally on native California oatgrass (Danthonia californica) (Beyer 2012, Henry unpubl. data). On 2 sites in the south Sound, females selected egg-laying sites based on habitat structure: eggs were laid on small Roemer’s Fescue plants in sparsely vegetated prairie habitat (Henry and Schultz 2012).

Males and females feed by using their long proboscis to explore flowers and sip floral nectar. Hays et al. (2000) studied the Mardon Skipper’s use and preference of nectar plants for 2 years on a south Puget Sound prairie sites and found in the first year they strongly selected early blue violet (Viola adunca), and in the second year strongly selected common vetch (Vicia sativa). Other nectar sources regularly used in this region include common camas (Camassia quamash) and spring gold and where available wholeleaf saxifrage (Saxifraga integrifolia), seablush (Plectritis congesta), and fragrant popcorn flower (Plagiobothyrus figuratus) (Hays et al. 2000, Hays 2010, M. Linders, WDFW, unpubl. data, A. Potter, WDFW, unpubl. data).

The Mardon Skipper life cycle and life stages (egg, larva, pupa) were first described by Newcomer (1966) from butterflies he collected near Yakima, Washington and reared in captivity. Newcomer (1966) found that eggs hatch in 6-7 days and larvae fed for approximately 3 months. James & Nunnallee (2011) provide detailed descriptions and photographs for life stages of Mardon Skipper (pp. 388-389). James and Nunnallee (2011) identified 5 larval instar stages, and similar to others who have captive reared this skipper (Newcomer 1966, Grosboll, WDFW, unpubl. data), found the larvae create and inhabit grass shelters by webbing host plant blades together. Larval feeding is mostly nocturnal and larvae often rest in their grass nests during the day (James and Nunnallee 2011). Henry (2010) tracked Mardon Skipper phenology in the field (an almost impossible task) on a south Sound prairie and found larvae move little if at all from their natal host plant tuft, where they also overwinter. Whether or not larvae feed the following spring is not known. Prepupal larvae construct a strong shelter at the base of the host plant grass, in which pupation occurs (James and Nunnallee 2011).

3.1.2.4 Habitat Characteristics

Life history and habitat research has been conducted recently across the butterfly’s range (Hays et al. 2000, Haggard 2003, Imper 2003, Beyer and Black 2007, Beyer and Schultz 2010, Henry 2010, Henry and Schultz 2012). In the south Puget Sound region, Mardon Skipper inhabits glacial outwash prairies. Studies of habitat needs in this region include research on nectar plant selection and general habitat characteristics of Mardon Skipper (Hays et al. 2000), and 2 studies of oviposition habitat selection (Henry 2010, Henry and Schultz 2012, Beyer 2012).

Hays et al. (2000) found that Mardon Skippers use open grasslands with abundant Roemer’s Fescue interspersed with early blue violet. Adult Mardon Skippers selected for short, open, native fescue grasslands, which allowed access to nectar and oviposition plants. They selected areas with only limited cover of the invasive shrub Scotch broom (Cytisus scoparius).

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In south Puget Sound, Henry and Schultz (2012) found that Mardon Skippers lay eggs on Roemer’s fescue, and females select for small, mostly green Roemer’s fescue plants, in sparse, short-statured vegetation areas of the prairie. The importance of conducting region-specific habitat research was demonstrated with this study, the findings of which were contrary to Mardon Skipper oviposition habitat selection in the Cascade Mountains region (Henry and Schultz 2012, Beyer and Schultz 2009). Beyer (2012) also found Mardon Skippers almost solely laying eggs on Roemer’s Fescue. Henry and Schultz (2012) evaluated habitat at a restored, historic Mardon Skipper site and found that restored portions of the site now support Mardon Skipper oviposition habitat.

3.1.2.5 Threats/Reasons for Decline

Two of the 4 extant south Sound Mardon Skipper populations occur within the Artillery Impact Area (AIA) of JBLM. There are a variety of vegetation conditions within the several square mile AIA, most of which have been significantly affected by the frequent fires resulting from repeated ordnance explosion. The closed, undeveloped nature of the AIA, coupled with a low-intensity, high fire frequency, has in some areas supported significant patches of Mardon Skipper habitat. However, the frequency and type of use in the AIA (and JBLM) has changed over time. In recent years, development within the AIA has increased the footprint and intensity of use of roads and structures within Mardon Skipper occupied areas (M. Linders, WDFW, pers comm., R. Gilbert, JBLM, pers. comm.). Fire timing, frequency, and intensity also may have changed recently (R. Gilbert, JBLM, pers. comm.). Buildings and other structures, along with their intense accompanying use, affect Mardon Skippers directly, and reduce and fragment habitat. Vehicle traffic likely crushes eggs, larvae, pupae, and adults (Potter et al. 1999). Increased fire frequency and earlier fire dates are also likely threats to Mardon Skipper and their habitat.

Two key Mardon Skipper information needs identified by Schultz et al. (2011) were 1) determining the extent of genetic distinction between south Sound and other region’s populations, and 2) developing reintroduction methods to establish new populations on restored historic locales. Mardon Skipper researchers and managers have highlighted the importance of additional habitat study to test the single-year findings of Henry and Schultz (2012) in additional years and locations (Potter 2012). In general, prairies in the south Sound region have been well surveyed for this butterfly (A. Potter, WDFW, pers. comm.). For more detailed information on threats to Mardon Skipper, see the Factors Affecting Continued Existence section in Potter et al. (1999:9-15).

3.1.3 Oregon Branded Skipper (Hesperia colorado oregonia)

The Oregon Branded Skipper (Hesperia colorado oregonia) is a butterfly in the Skipper Family (Hesperiidae). It is not listed under the federal or state endangered species acts, but is identified as a Species of Greatest Conservation Need by the state of Washington.


The Oregon Branded Skipper was recognized as a butterfly of conservation concern in the first Washington butterfly conservation status assessment (Pyle 1989) due to the small number of isolated populations, specialized and restricted habitat, and known threats to their habitat. Since

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that time, renewed taxonomic assessment, limited focal surveys by a few agencies and individuals, as well as some life history research, have confirmed extant and extirpated populations, and the restricted nature of Oregon Branded Skipper habitat and distribution.

Today, this butterfly is recognized as a species of conservation concern throughout its range (Schultz et al. 2011). In Washington, it is listed by the Department of Fish and Wildlife as one of 19 butterfly Species of Greatest Conservation Need in Washington’s Comprehensive Wildlife Conservation Strategy (WDFW 2005) and is listed as an Endangered Species in Canada (COSEWIC 2013).

Many taxa currently considered Hesperia colorado, including subspecies oregonia were until recently named Hesperia comma (see Warren 2005). The Oregon Branded Skipper, a subspecies of Hesperia colorado which inhabits the Salish Sea region is often referred to as Hesperia colorado oregonia. However, the most recent taxonomic treatment of butterflies in the United States and Canada (Pelham 2008) limits that trinomial to a subspecies found in northern California and southwestern Oregon. Pelham (2014 pers. comm.) reports the new name for this subspecies will be updated to Hesperia colorado Salish Sea segregate.


The Oregon Branded Skipper inhabits low-elevation grasslands in south Puget Sound, the San Juan Islands and southern Vancouver Island. A small number of populations persist and most consist of a small number of individuals. Six populations are documented on prairies in the southern Puget Sound; 4 are known extant (see Table 1). The San Juan Islands do not currently support many areas of potential habitat and have not been extensively surveyed for this species. Single populations have been documented from San Juan and Orcas Island, with only the Orcas Island population known extant. On Vancouver Island, 17 sites have been identified and populations persist on only 4 (COSEWIC 2013).

Location Last Record Status Mima Mounds Natural Area Preserve, Thurston County

2013 Extant

Rocky Prairie Natural Area Preserve, Thurston County

2002 Likely extirpated

Scatter Creek Wildlife Area North, Thurston County

2013 Extant

Scatter Creek Wildlife Area South, Thurston County

2013 Extant

West Rocky Prairie Wildlife Area, Thurston County

2009 Unknown

Artillery Impact Area, Joint Base Lewis-McChord, Pierce County

2013 Extant

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Oregon Branded Skippers are univoltine, which means they complete a single life cycle annually. A sedentary butterfly, they do not migrate; instead, the species inhabits sites year-round (as egg, larva, pupa and adult). A late-summer butterfly, in the south Puget Sound region adults typically begin to emerge from their chrysalids (pupae) between late-July and mid-August, and the adult flight period lasts into early-September. Males begin emergence first, followed by females; late-season individuals are primarily or solely females (A. Potter, WDFW, unpubl. data). Weather influences butterfly emergence and the flight period duration, with wet or cold potentially conditions delaying emergence, and conversely, warm, dry conditions promoting earlier emergence. Weather conditions also affect the total flight period duration.

Male skippers seek mates by perching on low vegetation in select spots and then darting out to inspect passing butterflies (A. Potter, WDFW, unpubl. data). Males that detect females commence courtship behavior; when males detect another male they engage in a territory defense behavior of tight, upward spiraling flight. Males often seek out the tops of Mima Mounds, an example of “hilltopping” behavior and occasionally engage in “puddling”, using the proboscis to imbibe liquids and minerals from bare ground (or mammal excreta). Females search for egg-laying sites by slowly flying and hovering just above the grassland vegetation, landing on and exploring their host plant grasses and other short-stature vegetation (all of which are senesced), and then quickly attaching a single, white egg (A. Potter, WDFW, unpubl. data). Both males and females feed by using their long proboscis to explore flowers and sip floral nectar.

The Oregon Branded Skipper life cycle and life stages (egg, larva, pupa) were first described by Hardy (1954) from butterflies he collected on Vancouver Island, Canada and reared in captivity. Hardy (1954) found that the late-summer-laid eggs overwintered, hatching the following spring; subsequent larvae fed and developed in larval-made silken tents between grass blades for approximately four months; pupae developed for approximately one month and were also enclosed in silken tents. James & Nunnallee (2011) reared Hesperia colorado in captivity and documented their life stages with detailed descriptions and photographs (pp. 378-379); they identified 6 larval instar stages and observed predation from Minute Pirate Bug (Orios tristicolor).


Oregon Branded Skipper egg-laying has been observed in south Sound prairies on Roemer’s Fescue (Festuca roemeri) (Pyle 2002). In addition, during multiple years observing egg-laying females, including a modest focal study in 2013, oviposition was observed on Long-stoloned sedge (Carex inops) in multiple years, Roemer’s Fescue in a single year, and in addition, two non-natives, sweet vernal grass and, Hairy cat’s ear in 2013 (A. Potter and L. Beyer, WDFW unpubl. data). Eggs laid in 2013 were checked periodically until they disappeared, and only the eggs deposited 3 on Carex inops (n=3) persisted until spring. Despite these observations, the host plant species for this butterfly remain unconfirmed. It seems likely Carex inops is a host, due to repeated observations of its use for oviposition, and because this butterfly is a monocot-feeding skipper we know Hairy Cat’s Ear is not a host; however, egg-laying occurs in late-summer when all of these plants are senesced,

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and may occur on neighboring plants rather than directly on hosts. Observations of larval feeding in the wild are needed to confirm host plant species.

Oregon Branded Skipper relies on late-summer (August-September) floral nectar sources. In current condition, low-elevation grasslands and prairies support few late-summer flowering plants, either native or nonnative, leaving late-summer butterflies with a significant lack of food resources. They have regularly been found using the following plants: Tansy Ragwort (Tanacetum vulgare), nonnative Thistles (Cirsium ssp.), White-top Aster (Sericocarpus rigidus), Pearly Everlasting (Anaphilis margaritacea), and Hairy Cat’s Ear (A. Potter, WDFW, unpubl. data). Though nonnative, the first two species are key nectar resources for Oregon Branded Skipper, as well as for many other late-summer butterflies. There have been no formal studies of Oregon Branded Skipper nectar selection.

In the south Puget Sound region, Oregon Branded Skipper inhabits glacial outwash prairies. There have been no formal studies of their habitat requirements. Oregon Branded Skippers are found in areas dominated by short-stature, native grasses and sedges, especially Roemer’s Fescue and Long-stoloned Sedge, with open structure, and bare ground (or moss/lichen). All oviposition behavior observed in 2013 occurred in short, sparsely vegetated prairie (A. Potter, WDFW, unpubl. data).

3.1.4 Puget Blue (Plebejus (Icaricia) icarioides blackmorei, Barnes and McDunnough, 1919)


The Puget Blue was recognized as a butterfly of conservation concern in the first Washington butterfly conservation status assessment (Pyle 1989) due to the small number of populations, specialized and restricted habitat, and known threats to their habitat. Since that time, limited focal surveys by a few agencies and individuals, as well as some life history research, have confirmed extant and extirpated populations, and the restricted nature of Puget Blue habitat and distribution, especially within the south Puget Sound region.

Today, this butterfly is recognized as a species of conservation concern throughout its range. In Washington, it is 1 of 19 butterfly Species of Greatest Conservation Need (SGCN) in Washington’s Comprehensive Wildlife Conservation Strategy (CWCS) (WDFW 2005). The U.S. Forest Service and Bureau of Land Management list Puget Blue as a sensitive species (USFS/BLM 2012). In British Columbia, Canada, it is a species of Special Concern (Heron 2007).

All subspecies of Plebejus icarioides are strictly limited to lupine (Lupinus spp.) host plants. Two members of this species are federally endangered: Fender’s Blue (Plebejus icarioides fenderi), which occurs on scattered sites in the Willamette Valley, Oregon (USFWS 2010), and Mission Blue (Plebejus icarioides missionensis), now restricted to a few locales in northern California (BFCI 2006).

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The Puget blue is a Pacific Northwest endemic butterfly found in Washington State and British Columbia, Canada (Fig. 1). It was described originally by Barnes and McDunnough (1919) from material collected in the Victoria, British Columbia area by E.H. Blackmore, a well-known British Columbia lepidopterist (Guppy and Shepard 2001). In British Columbia, Puget Blue occurs on southern Vancouver Island, where it historically occupied low-elevation prairies and high-elevation meadows. Today, the low-elevation populations have been extirpated, and the butterfly is restricted to a few sub-alpine meadows (Guppy and Shepard 2001, J. Miskelly, lepidopterist, pers. comm.).

In Washington, it was documented historically from 20 sites; 1 in King County, 9 on south Puget Sound prairies, oak woodlands, and other open habitats (Pierce, Thurston, and Mason Counties), and 10 sites on subalpine meadows in the northeast Olympic Mountains (Hinchliff 1996). The Puget Blue was likely more widespread historically on south Sound prairies. It has been extirpated from King County, and likely from 1 known and many undocumented historic south Sound prairie locales. Recent survey efforts for this butterfly have found additional populations on 5 prairie sites (Char and Boersma 1995; unpubl. data from K. McAllister, WDFW and A. Potter, WDFW; D. Grosboll, TNC, pers. comm.), and on a few sub-alpine meadows in the southeastern Olympic Mountains (Yake 2005).

A total of 15 Puget Blue sites have been documented within the HCP planning area; populations are known to be extant on 7 sites, and the current occupancy status is unknown for 8 (Fig. 2). Documented sites range geographically from southwestern Thurston County to Joint Base Lewis-McChord lands in northeast Thurston and western Pierce Counties.

Knowledge of the distribution and site occupancy status of Puget Blue within the planning area is the result of detections made during general prairie butterfly surveys (Char and Boersma 1995, Hinchliff 1996, Fleckenstein and Potter 1999, Wolford et al. 2007, Fimbel 2008; unpubl. data B. Bidwell, K. McAllister, WDFW, A. Potter, WDFW), incidental observations (A. Potter, WDFW, unpubl. data; pers. comm. B. Bidwell, E. Delvin, UW, C. Fimbel, CNLM, D. Grosboll, TNC, K. McAllister, WDFW, W. Yake, lepidopterist), and focal research on this butterfly (Hays et al. 2000, Schultz et al. 2009, LaBar and Schultz 2012). There has been no comprehensive effort in the south Puget Sound region to revisit historic locales or identify potential habitat and survey for Puget Blue (A. Potter, WDFW, pers. comm.). The 8 Puget Blue sites documented within the planning area that are designated “occupancy status unknown” have not been surveyed recently to determine if the butterfly is extant.

Two Puget Blue populations (Scatter Creek Wildlife Area-South and JBLM-Johnson Prairie) have been estimated to support more than 500 adults (C. Schultz, WSU, unpubl. data). Small numbers of individuals have been reported from all other documented south Sound sites. Population monitoring, to assess butterfly abundance over time has not been conducted on any Puget Blue site.

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The icarioides butterflies are small, with a wingspan of <3.8 cm [1.5 inches] (Pyle 2002). They are classified within the subfamily Polyommatinae, commonly referred to as the blues. As this name implies, these butterflies, especially the males, are bright blue. Females are slightly larger than males and have an overall gray-brown color with blue highlights. Puget Blue can be difficult to distinguish from another, more common blue butterfly, the Silvery Blue (Glaucopsyche lygdamus).

Puget blues are univoltine; they complete 1 life cycle annually. They are sedentary insects, inhabiting their sites year-round as an egg, larva, pupa and adult. In the south Sound, adults (butterflies) typically begin to emerge from their chrysalids (pupae) in mid-to-late May. Although individual butterflies may live for only a few days, the entire adult flight period often lasts through June (A. Potter, WDFW, unpubl. data). Butterflies have been observed as early as early-May (B. Bidwell, lepidopterist, pers. comm.) and as late as early-July (Hinchliff 1996). Like other blue species, male Puget Blues seek out females using a behavior called “patrolling”, where the male flies back and forth in an area near host plants or vegetation edge (Scott 1986, A. Potter, WDFW, unpubl. data). Males also search for evaporated puddles and moist soil (a behavior called “puddling”), and animal urine and feces, from which they extract salts (Scott 1986, A. Potter, WDFW, unpubl. data). Female Puget Blues lay eggs individually on the underside of the lower leaves of Sickle-keeled Lupine (Lupinus albicaulis) (Hays et al. 2000) and possibly other perennial lupines.

Male and female butterflies feed by using their long proboscis to explore flowers and sip floral nectar. Hays et al. (2000) studied the Puget Blue’s use and preference of nectar plants for 2 years, on 2 south Sound prairie sites and found that they consistently, strongly preferred feeding from the unopened flowers of their host plant, Sickle-keeled Lupine. Two other native prairie plants, Manroot (Marah oreganus) and Graceful Cinquefoil (Potentilla gracilis) were frequently used in single, but not both years. Puget Blues avoided nectaring from several plants, including Scotch broom, clovers (Trifolium), and open Sickle-keeled Lupine flowers. In a single-year study, Dzurisin (2005) found Puget Blues at 1 south Sound site frequently obtaining nectar from 4 native prairie species: Sickle-keeled Lupine, Graceful Cinquefoil, Spring Gold (Lomatium utriculatum), and Western Buttercup (Ranunculus occidentalis). Annual variation in plant phenology and condition determines the availability of nectar resources and causes variation in availability and use among years.

Adults may not travel far from their lupine hosts; Downey and Fuller (1961) in a single year of study, visited many Puget Blue populations throughout the west, and never found butterflies further than 50 m from host plant patches. However, Puget Blues, primarily puddling males, have been observed regularly up to 500 m from the nearest host lupine patches (A. Potter, WDFW, unpubl. data).

The complete Puget Blue life cycle and life stages (egg, larva, pupa) were first described by Newcomer (1911). James & Nunnallee (2011) provide detailed descriptions and photographs of the species life stages (pp. 218-219). Eggs hatch in 5-10 days (James and Nunnallee 2011, A. Potter, WDFW, unpubl. data); the resulting caterpillars (larvae) feed on lupine and continue to grow,

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shedding their skins to expand in what are referred to as instar stages. Larvae are thought to be nocturnal, their concealment during the day, combined with camouflage, and ant tending (see below) are likely key features of their defense (James and Nunnallee 2011). In the second larval instar stage, young caterpillars seek a protected, concealed location at the base of their host plant, and enter a dormant phase (diapause) (James and Nunnallee 2011, LaBar and Schultz 2012), which for Puget Blues lasts roughly from late-July or early-August until the following spring. Upon breaking diapause in spring, larvae reinitiate lupine feeding on small, new growth; larvae grow to instar stage 4, and then become chrysalids (pupae) in late-April or early-May (LaBar and Schultz 2012), before finally emerging from pupae in 10-14 days as adults (butterflies) (James and Nunnallee 2011).

Many species in the Lycaenidae family engage in mutualistic relationships with ants, known as myrmecophily (Pierce et al. 2002). Interactions often consist of ants tending and milking butterfly larvae (Fig. 3), obtaining nutrition in the form of a nectar-like substance (honeydew) in the process, and also protecting larvae from predators and parasitoids; sometimes ants move butterfly larvae or pupae into ground chambers, including their nests (Downey 1962, Pierce et al. 2002).

Newcomer (1911) described an icarioides subspecies being tended by small, black ants. However, not all icarioides blues are tended by ants, as the Fender’s Blue has been intensively studied with no apparent interaction between ants and any life stage (Schultz, WSU, pers. comm.). Ant-tending has been observed in Puget Blues by Hays et al. (2000) and Potter (WDFW, unpubl. data), who observed Formica species ants milking and defending larvae. LaBar and Schultz (2012) and LaBar (2009) found a Puget Blue larva in an ant tunnel, Formica ants herding a larva into a small hole at the base of a lupine plant, and ants defending larvae.


In Washington, the Puget Blue inhabits low-elevation grasslands (prairies) in south Puget Sound, and sub-alpine meadows in the Olympic Mountains. In Puget Blue habitat and nectar use on 2 south Sound prairies, Hays et al. (2000) identified Sickle-keeled Lupine as the larval host and primary adult nectar source. The butterfly’s dependence on Sickle-keeled Lupine limits their habitat to areas and sites that support significant patches of this plant. Density of Sickle-keeled Lupine at 2 Puget Blue use areas varied between years and sites from 0.08-0.48 plants/m2 (Hays et al. 2000). Although no other Puget Blue larval host plants have been identified, Hays et al. (2000) determined other adult nectar plants that were preferred, including Manroot and Graceful Cinquefoil. Dzurisin (2005) found 4 species regularly used for nectar (Sickle-keeled Lupine, Graceful Cinquefoil, Spring Gold, and Western Buttercup. Because annual variation in plant phenology and condition determines availability of nectar resources and causes variation in availability and use among years, variety of nectar sources is an important habitat component. Another important habitat feature for Puget Blue is presence of bare ground depressions where water can collect and evaporate during the adult flight period. Males rely on these sites to obtain minerals. Puget Blues used prairie habitat high in forbs (broad-leafed herbaceous plants) and somewhat degraded, with Scotch Broom cover (total of all height classes) 60% in 1 Puget Blue use area, and cover of non-native grasses was 30% in another (Hays et al. 2000).

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Fire, herbicide use, mowing, and other prairie management techniques are important tools for re-creating or simulating disturbance mechanisms that historically maintained prairies, reducing invasive species, and restoring endangered species habitat connectivity (Dunwiddie and Bakker 2011, Schultz et al. 2011). Regarding herbicide use, a recent study of the effects of a grass-specific herbicide (sethoxydim) application to Puget Blue, found little to no impact on larval performance, or egg-laying, however, adult butterflies spent significantly less time in sprayed plots than controls (LaBar and Schultz 2012). These prairie management practices, implemented to restore or enhance prairie vegetation and wildlife habitat, also can directly or indirectly harm butterflies (Schultz et al. 2011). Effects of these practices on butterflies, including Puget Blue, are not completely understood. Prairie management in areas occupied by butterfly species of concern is necessary, and must be undertaken with special methods and considerations to reduce or eliminate harm to these species.

Few surveys have been conducted to understand distribution of Puget Blue in the south Puget Sound region, and populations have not been monitored to determine abundance, trends, or primary use areas. Knowledge of distribution, abundance, and habitat needs are essential elements to conserving and managing for Puget Blue, but have not been fully studied (Schultz et al. 2011). Information gaps specific to Puget Blue conservation also include the degree and extent of myremecophily and development of lupine propagation and transplant methods (Schultz et al. 2011).

A threat that is unique to Puget Blue is control and eradication of the lupine host plants on which they depend. Cattle and horses can be sickened by consuming lupine, which has caused eradication of lupine from many south Sound prairies historically used as pasture. This affects present day Puget Blue distribution and population size.

3.1.5 Taylor’s checkerspot (Euphydryas editha taylori, W.H. Edwards, 1888)


Once so numerous that Dornfeld (1980) described meadows near Corvallis, Oregon as “fairly swarming” with this butterfly, Taylor’s Checkerspot is recognized today, throughout its range, as imperiled. Taylor’s Checkerspot was identified as a butterfly of conservation concern in the first Washington butterfly conservation status assessment (Pyle 1989) due to its extensive loss of prairie habitat from development, agriculture, and forest succession, small number of populations, clearly important and largely unknown habitat requirements, and existence on unsecured private lands. Since 1989, extensive surveys have been conducted to determine the status of historically documented populations and search for undiscovered sites across the butterfly’s range (Fleckenstein and Potter 1999, Shepard 2000, Stinson 2005, Ross 2006, Holtrop 2010, COSEWIC 2011, Potter 2011, B. Bidwell, lepidopterist, unpubl. data, WDFW unpubl. data: K. McAllister, A. Potter, M. Walker, WDFW). Through these efforts, many populations located in the past were determined to be extinct; a few new populations were discovered of which some declined to extirpation and some persist. Some life history and habitat research is recent across the butterfly’s

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range (Hays et al. 2000, Severns and Warren 2008, Page et al. 2009, Severns and Grosboll 2011, Grosboll 2011).

Taylor’s Checkerspot was listed as endangered in 2013 (USFWS 2013) and is also listed as endangered by the state of Washington. Its NatureServe Global status is G5T1 (Critically Imperiled,) and its NatureServe State status is S1 (Critically Imperiled).

The butterfly is recognized as a butterfly of conservation concern throughout its range. In Washington, it is 1 of 19 butterfly Species of Greatest Conservation Need in Washington’s Comprehensive Wildlife Conservation Strategy (WDFW 2005). WDFW completed an extensive status review for this butterfly, which includes detailed accounts on Taylor’s Checkerspot taxonomy, natural history, habitat, and threats (Stinson 2005). The U.S. Forest Service and Bureau of Land Management list it as a sensitive species (USFS/BLM 2012). In British Columbia, Canada, it is classified as an endangered species under the Species at Risk Act (COSEWIC 2011).


Taylor’s Checkerspot is a Pacific Northwest endemic butterfly once found on over 80 sites in the Willamette Valley, Oregon, western Washington, and Vancouver Island, British Columbia, Canada (Fig. 1) (Stinson 2005, Ross 2006, Holtrop 2010, COSEWIC 2011, A. Potter, WDFW, unpubl. data, P. Severns, lepidopterist, pers. comm.). It was originally described in 1988 by W. H. Edwards from material collected in the Victoria, British Columbia area by a noted amateur lepidopterist, the Reverend George Taylor. In British Columbia, Taylor’s Checkerspot historically occupied at least 24 prairie-oak and coastal meadow sites in southern Vancouver Island, but today persists on only 1 site (COSEWIC 2011).

In Oregon, the butterfly occurs in the Willamette Valley, where over 14 sites were historically documented but only 2 are extant (Hinchliff 1995, Stinson 2005, Ross 2006, H. Rice, lepidopterist, pers. comm., P. Severns, lepidopterist, pers. comm.). Historically, the Taylor’s Checkerspot was likely more widespread throughout its range.

Washington. In Washington, Taylor’s Checkerspot was historically documented from 24 sites; 1 each in San Juan and Island Counties, 2 in coastal Clallam County, and 20 on south Puget Sound prairies, oak woodlands, and other open habitats (Lewis, Mason, Pierce, and Thurston Counties) (Hinchliff 1996, B. Bidwell, lepidopterist, pers. comm.). By 2004, it was documented extirpated (or likely extirpated) from all historic locales in Island, Lewis, Mason, Pierce, San Juan, and Thurston Counties (Stinson 2005). However, intensive survey efforts initiated in the 1990’s located additional populations of the butterfly on 5 south Puget Sound prairies (Char and Boersma 1995, Chramiec 2004; unpubl. data: B. Bidwell, J. Fleckenstein, DNR, and A. Potter, WDFW), forest balds in southeast Thurston County (unpubl. data: M. McCallum, DNR, K. McAllister, WDFW, A. Potter, WDFW, and M. Walker, WDFW), and a few forest balds and coastal sites in Clallam County (Holtrop 2010, A. Frost, entomologist, pers. comm., unpubl. data: A. McMillan, WDFW, A. Potter, WDFW, and T. Stuart, WDFW,).

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Thurston County HCP Area. At least 24 Taylor’s Checkerspot sites have been documented within the HCP area; the butterfly is extant on only 4, and 3 of those are recently reintroduced populations (Fig. 2) (Stinson 2005, Linders 2006, Linders 2012). Extant sites range geographically from southwestern Thurston County (2 populations) to Joint Base Lewis-McChord (JBLM) lands in western Pierce County (2 populations).

Knowledge of the distribution and site occupancy status of Taylor’s Checkerspot within the HCP area comes from observation data collected during general prairie butterfly surveys (Char and Boersma 1995, Hinchliff 1996, Fleckenstein and Potter 1999, Wolford et al. 2007, Fimbel 2008, unpubl. data: B. Bidwell, lepidopterist, K. McAllister, WDFW, A. Potter, WDFW), incidental observations (A. Potter, WDFW, unpubl. data, pers. comm.: B. Bidwell, lepidopterist, E. Delvin, UW, C. Fimbel, CNLM, D. Grosboll, TNC, K. McAllister, WDFW, W. Yake, lepidopterist), and focal research on this butterfly (Hays et al. 2000, Grosboll 2011, Potter 2011, Linders 2012). As part of the WDFW Taylor’s Checkerspot status review, the agency led a comprehensive effort in the south Puget Sound region to revisit historic locales and identify and survey potential habitat for the butterfly (A. Potter, WDFW, pers. comm.).

Taylor’s Checkerspot populations are closely monitored at the 4 extant south Sound sites (Olson and Linders 2010, Linders 2012). Three of these populations were recently established (or in 1 case perhaps augmented) by translocations of captive-reared butterflies (Linders 2006, Linders 2012). The 3 reintroduced populations occur on Scatter Creek Wildlife Area – South, Glacial Heritage County Park, and JBLM Artillery Impact Area – Range 51. During monitoring of the reintroduced populations, small numbers of butterflies have been observed at the first 2 sites, while large numbers of adults, 100s and perhaps 1,000s of individuals have been observed recently at the JBLM reintroduction site (Linders 2012). The sole extant south Sound population that is not the result of recent translocation is located on JBLM Artillery Impact Area – Range 76, and is also the single source population for the south Sound Taylor’s Checkerspot captive-rearing effort,. Close monitoring of this population has consistently detected 1000s of butterflies during recent years (Olson and Linders 2010, Linders 2012).


Description. Taylor’s checkerspot is a brightly colored, medium-sized butterfly with a striking checkered pattern of orange to brick red, black, and cream. On south Puget Sound prairies, no other butterfly resembles it. Females are larger than males, though both have the same checker-patterned wings.

Life cycle and behavior. Taylor’s Checkerspot is univoltine; it completes 1 life cycle annually. They are sedentary insects, inhabiting their sites year-round as an egg, larva, pupa, and adult. In the south Sound, adults (butterflies) typically begin to emerge from their chrysalids (pupae) in late-April, though this and all other life stage dates for this butterfly can vary significantly due to weather conditions (Linders 2006, A. Potter, WDFW, pers. comm.). Although individual butterflies may live only a few days, the entire adult flight period in the south Sound often lasts through late-May (Linders 2006, Olson and Linders 2010, Linders 2012, unpubl. data: D. Grosboll, TESC, K.

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McAllister, WDFW, A. Potter, WDFW). Butterflies in this region have been observed as early as late-March (A. Potter, WDFW, unpubl. data) and as late as early-June (Hinchliff 1996, Linders 2012, K. McAllister, WDFW, unpubl. data).

Males use 2 strategies for mate-finding: perching and patrolling (Bennett et al. 2011). In perching, males select specific sites to perch and then dart out at passing butterflies to determine if it is a female of its species. In patrolling, males search for females by almost constant flying, often along a regular route or territory. Females lay eggs in clusters, low on their host plants, which in the south Sound are the non-native English plantain (Plantago lanceolata) and native Harsh Paintbrush (Castilleja hispida) (Char and Boersma 1995, Hays et al. 2000, Severns and Grosboll 2011, Grosboll 2011, unpubl. data: D. Grosboll, TESC, M. Linders, WDFW, A. Potter, WDFW).

Male and female butterflies feed by using their long proboscis to explore flowers and sip floral nectar. Annual variation in plant phenology and condition affects availability of nectar resources thereby causing variation in plant species use among years. An early pollination study on south Puget Sound prairies (Jackson 1982) found Taylor’s Checkerspots nectaring solely on (Camassia quamash). Hays et al. (2000) observed (but did not quantitatively study) Taylor’s Checkerspot nectar habits on a south Sound prairie and found them primarily using common camas and Nine-leaved lomatium (Lomatium triternatum). Other nectar sources regularly used by Taylor’s Checkerspot in the south Sound region include: deltoid balsamroot (Balsamorhiza deltoidea), spring gold, Wholeleaf Saxifrage (Saxifraga integrifolia), and Seablush (Plectritis congesta) (Linders 2012, A. Potter, WDFW, unpubl. data).

Adult movement studies of the closely related E. editha bayensis and Melitaea cinxia have found these butterflies to be consistently sedentary, though a few individuals move some distance, most remain within a few 100 m (USFWS 1998, Nieminen et al. 2004). No research specific to Taylor’s Checkerspot has been conducted to determine their movement patterns or distance.

Several scientists have observed Taylor’s Checkerspot egg masses and larvae extensively in the south Sound, but their phenology in the wild has not been studied completely (Severns and Grosboll 2011; unpubl. data: D. Grosboll, TESC, M. Linders, WDFW, A. Potter, WDFW). Careful and detailed phenological data for Taylor’s Checkerspot larvae has been collected by the Oregon Zoo as part of a captive-rearing program (Barclay et al. 2010). James & Nunnallee (2011: pp. 286-287) provide detailed descriptions and photographs of the species life stages . Euphydryas editha eggs hatch in 8-9 days (James and Nunnallee 2011); eggs within a cluster typically hatching in synchrony (Barclay et al. 2010). The resulting caterpillars (larvae) create webbing and feed communally through the spring on the host plant species on which eggs were deposited, continuing to grow and shed their skins to expand, in what are referred to as instar stages. Larvae enter a dormant phase (diapause) in late-June or early-July (M. Linders, WDFW, unpubl. data, A. Potter, WDFW, unpubl. data) when host plants are senescing and no longer provide palatable vegetation. Larvae often diapause in a sheltered location under rocks, logs, or litter (Guppy and Shepard 2001). Diapausing Euphydryas editha larvae develop a thick exoskeleton that helps prevent dehydration (Scott 1986). The diapause phase lasts for many months, until early the following spring (January or February in the south Sound). Upon breaking diapause, Taylor’s Checkerspot larvae reinitiate feeding on a

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broader array of plant species. Plant species that held egg masses remain a major component of their diet, but additional post-diapause food sources (sea blush, Blue-eyed Mary (Collinsia parviflora), and Dwarf Owl-clover (Triphysaria pusilla) as available, also are used. Larvae pupate in March or April (M. Linders, WDFW, unpubl, data).


The Taylor’s Checkerspot inhabits grasslands in low-elevation prairies and meadows, coastal meadows and stabilized dunes, and montane meadows and balds. Balds are shallow-soiled, grass, herbaceous vegetation, or lichen and moss dominated sites, typically less than 5 ha (12.5 ac), that occur within forested lands (Chappell 2006). A few studies of Taylor’s Checkerspot habitat have been conducted outside of the south Puget Sound region, including in Oregon (Severns and Warren 2008), British Columbia (Page et al. 2009), and the north Olympic Peninsula (Severns and Grosboll 2011, Grosboll 2011). Egg-laying (oviposition) habitat is often studied with this and other butterflies because it is a limiting factor, determines the site of pre-diapause larvae, and influences the location of diapause, post-diapause, and pupation. Severns and Warren (2008) found that Taylor’s Checkerspots selected habitat for egg-laying that occurred within high cover of short-stature native bunchgrasses and adult nectar resources, indicating that females select egg-laying sites based on habitat condition. Page et al. (2009) found the most common activity of post-diapause larvae was basking and perching, demonstrating the importance of thermal habitats in this life stage. The British Columbia study population had multiple host plant species available and females’ selection of egg-laying sites in this environment was influenced by host plant phenology and condition (Page et al. 2009). A characteristic of egg-laying habitat consistently identified in the British Columbia and 3 Olympic Peninsula populations was the abundance of host plants (number or percent cover) (Page et al. 2009, Severns and Grosboll 2011, Grosboll 2011).

Within the south Sound region, the butterfly has been found on prairies and balds. Habitat selection by egg-laying females has been studied in 1 population, the sole extant south Sound site (JBLM Artillery Impact Area – Range 76) by Linders et al. (2009), Severns and Grosboll (2011), and Grosboll (2011). All researchers found that females selected habitat with high host plant density for oviposition. Grosboll (2011) determined that the butterfly selected for host plant patches with >10,000 cm3 volume. Severns and Grosboll (2011) found that the butterfly laid eggs more frequently along 2-track road edges than the open prairie, and explained this may be due to the strong association between the host plant at this site (English plantain) and the road beds.

Although there has been no quantitative study of Taylor’s Checkerspot nectar plant use or preference, several plants have been identified as key nectar sources in south Sound populations (common camas, deltoid balsamroot, sea blush, wholeleaf saxifrage, nine-leaved lomatium, and spring gold) (Jackson 1982, Hays et al. 2000, Linders 2012, M. Linders, WDFW, unpubl. data, A. Potter, WDFW, unpubl. data). Because annual variation in plant phenology and condition determines the availability of nectar resources and causes variation in availability (and therefore use) among years, variety of nectar sources is an important habitat component.

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Military training: The sole source population for Taylor’s Checkerspot captive rearing and translocation, along with the only other south Sound Taylor’s Checkerspot site that currently supports a robust population are located within the Artillery Impact Area (AIA) of JBLM. There are a variety of vegetation conditions within the AIA, most of which have been significantly affected by frequent fires that result from repeated ordnance explosion. The closed nature of the AIA, coupled with a low-intensity, high fire frequency, has in some areas supported significant patches of Taylor’s Checkerspot habitat. However, frequency and type of use in the AIA (and JBLM) has changed. In recent years, development within the AIA has increased the footprint and intensity of roads and structures within areas occupied by Taylor’s Checkerspot (M. Linders, WDFW, pers comm., T. Thomas, USFWS, pers. comm.). Fire timing, frequency, and intensity also may have changed (R. Gilbert, JBLM, pers. comm.). Buildings and other structures, along with their intense use affect Taylor’s Checkerspots directly and reduce the amount of habitat. Vehicle traffic likely crushes eggs, larvae, pupae, and adults (Stinson 2005). Increased fire frequency and earlier fire dates also are likely threats to Taylor’s Checkerspots and their habitat.

A recently identified potentially significant threat to Taylor’s Checkerspot is the widespread presence of a pathogen specific to the primary larval host English plantain (Stone et al. 2011). This fungal pathogen (Pyrenopeziza plantaginis), like the plant it specifically attacks, is native to Europe, and was first documented in the Pacific Northwest (and North America) in 2011; the length of time it has been present in these regions is unknown (Stone et al. 2011). The fungus has infected English Plantain at Taylor’s Checkerspot sites in Oregon (Stone et al. 2011), and Washington (P. Severns, lepidopterist, pers. comm.). Peak necrosis of plantain leaves resulting from infection occurs in late-winter and can overlap with the Taylor’s Checkerspot post-diapause larval period (Stone et al. 2011), a time when the plant is needed in abundance to feed larvae.

Taylor’s Checkerspot appears to be highly selective in its habitat requirements, however, habitat needs have not been fully studied. Knowledge of habitat needs for adults, larvae, and diapause are essential elements to conserving and managing for Taylor’s Checkerspot (Schultz et al. 2011). Severns and Grosboll (2011) and Grosboll (2011) studied egg-laying habitat selection, and both identified understanding larval survival in different environments and on different host plants as an important research topic. Methods to reliably develop and manage for Taylor’s Checkerspot habitat are needed. Grosboll (2011) identified the need to develop methods for enhancing host plant resources. Harsh Paintbrush and English plantain have been identified as Taylor’s Checkerspot host plants. On most recently known sites, only 1 of these species occurs; additional study is needed to determine the effects of multiple host species availability to short and long-term survival of checkerspot populations.

3.1.6 Valley Silverspot (Speyeria zerene bremnerii, W.H. Edwards, 1872)


Valley silverspot was first considered a rare butterfly in Oregon (Dornfeld 1980). Hammond and McCorkle (1984) thought the butterfly was likely extirpated from Oregon in the 1970’s, noting that

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Speyeria butterflies and their larval host plants (Viola sp.) are, across their range, among the first organisms to be eliminated from native grassland habitats by human-caused disturbance. Valley Silverspot was recognized as a butterfly of conservation concern in the first Washington butterfly conservation status assessment (Pyle 1989) due to its declining range, small number of populations, specialized and restricted habitat, and known threats to their habitat. Since that time, limited focal surveys by a few agencies and individuals, as well as some life history research, have confirmed extant and extirpated populations, population declines, and the restricted nature of Valley Silverspot habitat and distribution.

Today, this butterfly is recognized as a species of conservation concern throughout its range. In Washington, it is listed by the Department of Fish and Wildlife (WDFW) as a state candidate for listing, and as 1 of 19 butterfly Species of Greatest Conservation Need (SGCN) in Washington’s Comprehensive Wildlife Conservation Strategy (CWCS) (WDFW 2005). USFWS designates it a federal species of concern. The U.S. Forest Service and Bureau of Land Management list Valley Silverspot as a sensitive species (USFS/BLM 2011). In British Columbia, it is considered a species at risk and is red listed (British Columbia Data Centre 2012).

The closely related Oregon Silverspot (Speyereia zerene hippolyta), a Pacific Northwest coastal endemic subspecies, was listed as threatened by the USFWS in 1980 (USFWS 2001).

Taxonomic Note. A recent review of Willamette Valley, Oregon and south Puget Sound bremnerii specimens by noted butterfly taxonomist A. D. Warren (2005) found the Oregon butterfly previously considered bremnerii should be treated as a distinct subspecies. This unnamed Oregon subspecies (Speyeria zerene nr. bremnerii) is now extinct (Hammond and McCorkle 1984, Warren 2005). Hereafter, we follow Warren’s distinction, excluding Oregon from the range of bremnerii.


The Valley Silverspot is a Pacific Northwest endemic butterfly found in Washington State and the Canadian Province of British Columbia (Fig. 1). It was originally described by W.H. Edwards in 1872 from material collected on San Juan Island, Washington (Pelham 2008). In British Columbia, the butterfly occurred on approximately 19 southern Vancouver Island and 3 mainland sites (GOERT 2003, Guppy and Shepard 2001). Only a single British Columbia population on Salt Spring Island has been recently confirmed extant (GOERT 2003, J. Miskelly, lepidopterist, pers. comm.). Populations of a Speyeria zerene butterfly also have been found in northern Vancouver Island; however butterfly systematists disagree whether it is subspecies bremnerii (P. Hammond, OSU, pers. comm., J. Miskelly, lepidopterist, pers. comm.)

In Washington, Valley Silverspot was historically documented from a total of 26 sites; 2 in King County, 13 on south Puget Sound prairies and forest openings (Pierce, Thurston, and Lewis Counties), 3 in the San Juan Islands, 3 in far southwest Washington, and 5 sites on low-elevation and sub-alpine meadows in the northeast Olympic Mountains (Hinchliff 1996). Valley Silverspot was likely more widespread historically in southwest Washington, including on more south Sound prairie sites. Presently, it is known to be extirpated from King County, 3 Thurston County sites, and

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likely several additional known and undocumented Washington sites. Recent survey efforts have found additional populations on 8 prairie sites (Char and Boersma 1995, Yake 2007; unpubl. data: B. Bidwell, lepidopterist, K. McAllister, WDFW, A. Potter, WDFW).

A total of 15 Valley Silverspot sites have been documented within vicinity of the HCP planning area; populations are known to be extant on 8 sites, extirpated on 3, and the current occupancy status is unknown for 4 (Fig. 2). The butterfly’s distribution in Thurston County ranged geographically from a site documented in 1958 in the northwest corner of the county (“near Summit Lake”), to the far southeast (Bald Hill), several populations located in the southern portion of the county (Scatter Creek Wildlife Area, Mima Mounds Natural Area Preserve, etc.), to Joint Base Lewis-McChord lands near Yelm, and a site in Pierce County (Spanaway).

Knowledge of distribution and site occupancy status of Valley Silverspot in and near Thurston County results from butterfly detections made during general prairie butterfly surveys (Char and Boersma 1995, Hinchliff 1996, Fleckenstein and Potter 1999, Wolford et al. 2007, Yake 2007, Fimbel 2008; unpubl. data: B. Bidwell, lepidopterist, K. McAllister and A. Potter, WDFW), incidental observations (unpubl. data: B. Bidwell, lepidopterist, E. Delvin, UW, C. Fimbel, CNLM, D. Hays, WDFW, K. McAllister, WDFW, A. Potter, WDFW), and limited focal research on this butterfly (Hays et al. 2000). There has been no comprehensive effort in the south Puget Sound region to revisit historic locales or identify potential habitat and survey for Valley Silverspot (A. Potter, WDFW, pers. comm.). The 4 Valley Silverspot sites designated (Fig. 2) “occupancy status unknown” have not been revisited sufficiently to determine if populations are extant. Habitat was destroyed at the3 sites designated extirpated.

Intensive population monitoring, to assess annual butterfly abundance over time has not been conducted on any Valley Silverspot site. However, Valley Silverspot monitoring has been done sufficiently to determine that significant declines in population sizes have occurred since 1990 on 7 of the 8 extant south Sound prairie sites (Char and Boersma 1995, Hays et al. 2000, Wolford et al. 2007, Fimbel 2008, unpubl. data: B. Bidwell, lepidopterist, K. McAllister, WDFW, A. Potter, WDFW, The Nature Conservancy). Survey efforts on these 7 sites during the 1990’s typically located more than 100 butterflies on a single day, as compared to recent and more intensive searches, which have typically found fewer than 5-10 individuals. In recent years, Valley Silverspot has been reliably detected at only 4 of the extant sites (Mima Mounds NAP, Scatter Creek Wildlife Area-North, West Rocky Prairie Wildlife Area, and a privately owned site adjacent to Scatter Creek-North).


Description. The Valley Silverspot, a large, deep-orange butterfly with dorsal patterned black lines, is a striking butterfly, which flashes silver from large orb-like spots on the ventral hind wings. For butterflies, they are large, with a wingspan up to 6.4 cm [2.5 in] (Guppy and Shepard 2001, Pyle 2002); females are larger than males. Speyeria are classified within the subfamily Heliconiinae, whose members are commonly referred to as the longwings and fritillaries. In southwest

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Washington, Valley Silverspot can be difficult to distinguish from another, more common fritillary, the Great-Spangled Fritillary (Speyeria cybele pugetensis).

Life Cycle and behavior. The Valley Silverspot is univoltine; completing a single life cycle annually. The adults (butterflies) are strong fliers, capable and even prone to moving some distance; however, the species inhabits year-round (as an egg, larva, pupa, and adult) most sites where adults are found. In the south Sound, adults typically begin to emerge from their chrysalids (pupae) in early-to-mid July, with the entire adult flight period often lasting into September and even early-October (Hinchliff 1996, A. Potter, WDFW, unpubl. data). Males emerge first, followed by females; late-season individuals are primarily or solely females (A. Potter, WDFW, unpubl. data). Although most adult butterflies are short-lived, individuals in other species of Speyeria have survived for over a month (Scott 1986, James and Pelham 2011).

Male Valley Silverspots vigorously search for females by quickly flying above and around prairie vegetation (A. Potter, WDFW, unpubl. data). Females seek egg-laying sites by slowly flying across grassland habitat, landing on vegetation, and then crawling through ground-level plants and dead vegetation, while probing with their abdomen, until a suitable location is found (A. Potter, WDFW, unpubl. data). Similar oviposition behavior has been described by McCorkle and Hammond (1988) for the closely related Oregon silverspot. Speyeria butterflies lay eggs individually on species of violets. In the south Puget Sound, Valley Silverspot females have been observed laying eggs on and near Early blue violet, a native herbaceous plant (Hays et al. 2000, A. Potter, WDFW, unpubl. data).

Males and females feed by using their long proboscis to explore flowers and sip floral nectar. Hays et al. (2000) studied the Valley Silverspot’s use and preference of nectar plants for 2 years on a south Puget Sound prairie sites and found in the first year they nectared exclusively on the non-native Canada thistle (Cirsium arvense), and in the second year preferred native showy fleabane (Erigeron speciosus). Other regularly used nectar sources in this region include the native white-top aster (Aster curtus = Sericocarpus rigidus) and non-native tansy ragwort (Senecio jacobaea) (Hays et al. 2000, A. Potter, WDFW, unpubl. data). Availability of nectar resources has been found to be critical to female egg-laying in other Speyeria (Boggs 2003). Valley silverspots’ use of non-native nectar sources is not unusual; many other prairie butterflies use these species, especially on grasslands that are lacking native late-season nectar sources.

The valley silverspot life cycle and life stages (egg, larva, pupa) were first described by Hardy (1958). James & Nunnallee (2011) provide detailed descriptions and photographs for life stages of the closely related subspecies Speyeria zerene picta (pp. 236-237). James and Nunnallee (2011) found subspecies picta eggs hatched in 13-19 days and larvae entered a dormant condition (diapause) soon after for the winter, often gathering communally in plant litter. In spring, larvae become active again, feeding on young violet shoots and basking (Hardy 1958, James and Nunnallee 2011). Larvae continue to feed on Viola and grow throughout the spring and early-summer. Feeding is mostly at night and larvae often rest under leaves during the day (James and Nunnallee 2011). Hardy (1958) reared valley silverspot individuals from southern Vancouver Island, and found they went through 5 instar stages, entering pupation in mid-to-late June (total post-diapause larval period 73-84 days). James and Nunnallee (2011) rearing a related subspecies documented 6

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instar stages (L1-L6). Speyeria spp. larvae have several defense mechanisms, including possibly droplets of an unknown liquid on the hair-like setae of L1 larvae, and on the latter-stage (L2-L6) larvae, an eversible, ventral gland which produces a musky odor (McCorkle and Hammond 1988, James and Nunnallee 2011). Pupation sites were described by Hardy (1958) as above ground “in a slight hollow under a root or loose tuft of grass”; the pupal stage lasted 22-60 days.


In the south Puget Sound region, valley silverspot inhabits low-elevation grasslands (prairies) and forest openings. There have been no studies of valley silverspot oviposition or larval habitat selection or requirements. Early blue violet was identified as the larval host plant in the south Puget Sound (Hays et al. 2000, A. Potter, WDFW, unpubl. data). Early blue violet is also the larval host for the endangered and closely related Oregon Silverspot (McCorkle and Hammond 1988). Research on Oregon silverspot found it selects dense early blue violet patches (>20 plants/m2) for egg-laying (The Nature Conservancy 1990). It is likely that valley silverspot also requires patches of dense early blue violet.

In a 2-year study of valley silverspot habitat and nectar use on a south Puget Sound prairie, Hays et al. (2000) identified nectar plants preferred and regularly used by the butterfly (Canada Thistle, Showy Fleabane, White-top Aster, and Tansy Ragwort). Because nectar availability is so important to Speyeria fecundity (Boggs 2003), and annual weather conditions affect the availability of nectar species (Hays et al. 2000), abundant and diverse nectar species are key essential elements of Valley Silverspot habitat.

Hays et al. (2000) found valley silverspot nectar use areas characterized by 25-45% percent cover of bare ground, and were somewhat degraded, with 15-25% cover each of non-native grasses and woody shrubs, primarily scotch broom (Cytisus scoparius). Comparison of high-use nectar areas to the entire nectar use area found the high-use nectar areas had significantly greater percent cover of non-native grasses and forbs, and significantly lower cover of bare ground and woody shrubs (Hays et al. 2000).


Few surveys have been conducted to fully understand the current distribution of valley silverspot in the south Puget Sound region, and populations have not been closely monitored to determine abundance, trends, or primary use areas. This butterfly’s habitat requirements have not been fully studied. Knowledge of distribution, abundance, and habitat needs are essential elements to conserving and managing for valley silverspot (Schultz et al. 2011). Information gaps specific to valley vilverspot conservation also include development of successful Viola and native nectar source outplanting methods (Schultz et al. 2011).

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3.2 Vertebrate Animals

3.2.1 Mazama Pocket Gopher (Thomomys mazama Merriam, 1897)

3.2.1.1 Conservation status

The subspecies of the Mazama pocket gopher in Washington have been Candidates for listing under the federal Endangered Species Act since 2001 (USFWS 2001); 3 subspecies in Thurston County, and 1 in Pierce County were listed as Threatened in 2014 (USFWS 2014). The Mazama pocket gopher was listed as a state Threatened species by the Washington Fish and Wildlife Commission in 2006. The species had been listed as a candidate for state listing as threatened, endangered, or sensitive in Washington since 1996. Prior to that time, the Roy (T. m. glacialis), Tenino (T. m. tumuli), Tacoma (T. m. tacomensis), Shelton, (T. m. couchi), and Cathlamet (T. m. louiei) subspecies had been state candidates since 1991. As a state Threatened species, unlawful taking of Mazama pocket gophers is a misdemeanor under RCW 77.15.130. The western (Mazama) pocket gopher is a Species of Local Importance in the critical area ordinances of Thurston and Pierce counties. The Shelton pocket gopher (T. m. couchi) is a species of local importance in the critical area ordinance of Mason County.

3.2.1.2 Distribution and Population Trends

Mazama pocket gophers were historically more widespread and abundant on the glacial outwash prairies of the southern Puget Sound region. They also occur on subalpine meadows of the Olympic Mountains. Several populations are sufficiently distinct to be described as separate subspecies, particularly those that were geographically isolated. Other subspecies of Mazama pocket gophers are found in parts of western Oregon and in northern California. The species is currently represented in Washington by 6 extant subspecies: 1 in Clallam; 1 in Mason; 3 in Thurston, and 1in Pierce counties (Fig. 1). They were also historically found around Tacoma and in Wahkiakum County. The subspecies found in Thurston County are described here.

Gophers are seldom found in densely developed areas, or sites with very rocky soil. There are perhaps 3-4 large (i.e., 1,000s) Mazama pocket gopher populations in Thurston/Pierce counties. The largest populations appear to be found on the Olympia and Shelton Airports, Scatter Creek Wildlife Area, and Joint Base Lewis McChord. Many surviving gopher subpopulations are small (<50) and appear to be isolated from other subpopulations, although there are few data on dispersal to help delineate genetically connected populations. Small subpopulations are unlikely to persist for long without at least occasional demographic and genetic recharge by dispersing individuals from other nearby populations. Re-colonization becomes less likely as habitat is fragmented and populations isolated. Large populations, or clusters of subpopulations close enough and with land condition that permits exchange of dispersers, may be important for the persistence of each subspecies and the species.

Most of the Mazama pocket gophers in the southern Puget Sound region currently occur in ~10 general areas in Pierce, Thurston, and Mason counties. These concentrations of known gopher

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occurrences and prairie soil types are separated by distance or rivers and vary widely depending on soils present and the land-use history. What is known about abundance and distribution for the subspecies in Thurston County is summarized below.

• T. m. pugetensis. What is probably the largest population of Mazama pocket gophers is found in the loamy sand soils at the Olympia Airport and surroundings in Tumwater on the historical Bush Prairie. Gophers are scattered over several hundred acres of maintained grassland at the airport, where they are relatively unmolested by humans or domestic animals. Gophers are also found in vacant lots, yards, and pastures in nearby locations on both sides of Interstate 5. In 2005, McAllister and Schmidt (2005) derived a crude population estimate of 6,040 for the airport, but no trapping was done to determine how closely this approximated the number of actual gophers.

Chambers Prairie, extending from about Ward Lake to Lake St. Clair, is the largest area of Nisqually soil type (3,700 ac), and probably historically supported an extensive gopher population. Most of the area has residential development of various densities. Chambers Prairie has gophers scattered in vacant lots, roadsides, and rural and agricultural sites, but no large extensive populations like the airport are known. The northwestern half of the area is within the urban growth areas of Olympia and Lacey, and much is densely developed such that likelihood of extensive local extirpation is elevated. The southeastern half of this area also has turf, Christmas tree, and berry farms, and other smaller farms and pastures.

Little Chambers Prairie and Hawks Prairie contain substantial areas of loamy sand soils, but most of the suitable habitat is heavily developed, with dense residential neighborhoods, roads, and businesses. Small pockets of habitat with gophers exist on some less developed or undeveloped lands, but these appear to be small and isolated, and may not persist in the long-term.

• T. m. tumuli. Rocky Prairie, south of East Olympia and north of Tenino, totals about 2,200 ac. Within this area, WDFW West Rocky Prairie Wildlife Area (WLA) includes 270 ac of mounded and terraced prairie. No gopher population appeared to be present at West Rocky WLA until a translocation project established a gopher population using gophers captured at the Olympia Airport (Olson 2011b). A 750 ac area adjacent to West Rocky Prairie WLA is owned by a sand and gravel company. East of West Rocky Prairie WLA, Wolf Haven International maintains 38 ac of native mounded prairie with a small Mazama pocket gopher population established by translocation (Linders 2008). North of Wolf Haven International is a large area (~600 ac) of mounded prairie on private lands with Spanaway-Nisqually complex soil that was once a ranch. It supported a significant population of gophers in the early 1990s; current status of gophers at this site is unknown. West of this property is Rocky Prairie Natural Area Preserve (NAP) where very small numbers of gophers are detected occasionally. The translocation projects (2005-2008, 2009-2011) moved gophers from the Olympia Airport and 2 Tumwater sites, both within the range of T. m. pugetensis, and established populations in the range of T. m. tumuli. The population status of T. m. tumuli may have been tenuous, as Steinberg (1996) was unable to find any,

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and only very small numbers of gophers had been detected in the area since then. Any future translocations will maintain separation of subspecies, unless genetic analysis indicates taxonomic distinction is not warranted.

• T. m. yelmensis. Mound Prairie, near Grand Mound, is bisected by Interstate 5. West of I-5, north and south units of Scatter Creek WLA support significant gopher presence. After 2004, when scotch broom (Cytisus scoparius) control became widespread and intensive, gophers spread throughout the northern 2/3rds of the north unit, where they hadn’t been observed previously. Scatter Creek WLA contains about 600 ac of prairie, and is mostly Spanaway-Nisqually complex soils. The north unit has about 80 ac of Nisqually soil and the south unit has about 8 ac. Most of the land west of I-5 near Scatter Creek WLA is subdivided into 5 ac parcels, with some higher density, including the Grand Mound Urban Growth Area.

Rock Prairie, an area of >1,200 ac of private lands, is located southwest of Tenino. The area still supports Mazama pocket gophers on 2 large ranches (Steinberg 1996), and 1 ranch has a 500 ac Grassland Reserve Program easement with management guidelines that protect prairie vegetation and maintain conditions suitable for gophers. Some of the remaining private lands have not been surveyed for gophers.

The historical Tenalquot Prairie area includes Weir Prairie (Upper, Lower, and South Weir), and Johnson Prairie, which are in the Rainier Training Area of JBLM, and Tenalquot Prairie Preserve. Most of the area is Spanaway soil types. This area also includes private lands south of the Rainier Training Area. The Weir Prairie Research Natural Area consists of Upper Weir Prairie (547 ac) and Lower Weir Prairie (440 ac), and is protected from the most destructive forms of military training, such as off-road vehicle maneuvers and digging. A WDFW research team found a density of ~2 adult gophers/ac on Lower Weir Prairie during 2010 and 2011. Johnson Prairie is about 194 ac of native and semi-native grassland and is among the highest quality Puget prairies. It supports a substantial population of Mazama pocket gophers (Steinberg 1995, WDFW data), as well as a high diversity of plants, butterflies, Oregon vesper sparrows, and western toads (Remsburg 2000, Altman 2003). Past activities have primarily been foot maneuvers, parachuting, and limited vehicle use (Remsburg 2000). No tracked or wheeled vehicle use is allowed off established roads, because the site is designated a Secondary Research Natural Area. Civilian recreational impacts are an increasing concern on Johnson and Weir prairies because unauthorized off-road vehicle use has increased in recent years. These areas also are used frequently for hunting and horseback riding.

Tenalquot Prairie Preserve is a 125 ac preserve south of South Weir owned by The Nature Conservancy; WDFW has a conservation easement on the property. It is being restored to high quality prairie by Center for Natural Lands Management. Gophers are present in low numbers in the Spanaway soils of the area.

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Pocket gophers spend most of their time within their system of burrows. Gophers are believed to be generally solitary and exclude other gophers from their burrows except when breeding and when females have litters. When pocket gophers have established a territory, they generally remain there, although they will shift their home range in response to seasonally wet soils.

Thomomys pocket gophers adjust their annual cycle of activity to the seasonal changes of weather, soil, and plant growth where they occur (Cox and Hunt 1992). Pocket gopher territory (i.e., burrow systems) sizes vary with habitat quality and reproductive status. Using radio-telemetry, Witmer et al. (1996) estimated that the late winter-early spring home range of T. mazama on a fallow field averaged 108 m2 for 4 males (range 73–143 m2), and 97 m2 for 4 females (range 47–151 m2; 0.01–0.03 ac). WDFW personnel captured and average of 9 gophers/ac in a 22 ac plot at Olympia Airport, but some gophers remained in the plot (G. Olson, unpubl. data).

Mazama pocket gophers attain sexual maturity by the breeding season after their birth, when ~ 9 mo old and rear a single litter of ~5 (2-7) pups per year (Witmer et al. 1996, Verts and Carraway 2000). Gopher populations can increase dramatically in the summer after the dispersal of young of the year, and may increase to 3–4 times the spring adult population. In addition to this annual influx of young-of-the-year, gopher populations also fluctuate year-to-year due to environmental conditions. Pocket gopher populations are characterized by local extinction and recolonization (Baker et al. 2003). Territoriality and extreme weather may influence pocket gopher populations more than any other factors.

Pocket gophers have been called ‘keystone species’ and ‘ecosystem engineers’ because they affect the presence and abundance of plants and other animals (Vaughan 1961, 1974; Reichman and Seabloom 2002). Their extensive excavations affect soil structure and chemistry; food caches and latrines enrich the soil, affecting plant community composition and productivity. Mazama pocket gophers are an important prey species for many predators, including hawks, owls, coyotes, and weasels; their burrows provide retreats for salamanders, western toads, frogs, lizards, small mammals, and invertebrates (Stinson 2005).


Mazama pocket gophers live on open meadows, prairies and grassland habitats of the glacial outwash plain where there are porous, well-drained soils (Dalquest 1948). Mazama pocket gophers do not require high quality prairie, but can live in a wide range of grasslands, particularly if they include a significant component of forbs, such as clover, lupines, dandelions, false dandelions, and camas. In addition to remnant prairies, occupied sites in Washington include grassy fields at airports, pastures, fields, Christmas tree farms, and occasionally clearcuts. T. m. melanops is found in open parkland and subalpine meadows in the Olympic Mountains (Johnson and Cassidy 1997).

Although most of the populations are found in grasslands on land that historically was prairie, they will move into sites with well-drained soil where forest cover has been removed, including recent clearcuts. Gophers are known to populate sites after timber harvest and become common for a few

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years while grasses and forbs are available, but decline as the area regenerates to forest. This has been observed most frequently in Mason County. They are otherwise essentially absent from forest habitats in Washington. Mazama pocket gophers occur in woodland in Oregon, particularly in ponderosa pine communities, but they are absent from dense forest (Verts and Carraway 1999). Gophers also are rare where grassland has been taken over by dense Scotch broom (Steinberg 1996, Olson 2011b).

Perennial forbs are preferred for food over grasses, and fleshy roots and bulbs, such as camas are important when green vegetation is not available. The availability of forbs may provide nutrients important for gopher growth and reproduction. Gophers also eat fungi and disseminate the spores of species that have an important role in facilitating plant growth.

The distribution and abundance of pocket gophers are greatly affected by soils. Soil characteristics that affect gophers include depth and texture, particularly rock and clay content that affects burrowing ability, permeability that can result in periodic flooding of burrows, and water-holding capacity and fertility that affect growth of plant foods. In general, pocket gophers prefer deep, light-textured, porous, well-drained soils, and do not occur in peat or heavy clay soils (Chase et al. 1982, Baker et al. 2003).

Distribution of Mazama pocket gophers appears correlated with prairie soil types, but they are not found on all remnant prairie sites. They rarely occur where soil is very rocky (Steinberg 1996a, Olson 2011b). There are local populations in non-prairie loam, sandy, and gravelly soil types (e.g., Indianola loamy sand, Grove, Everett) that may have been unused by gophers historically due to forest cover. These occurrences often are adjacent to more typical prairie soils (e.g., Nisqually soils). They may be able to occupy any site that supports herbaceous vegetation, does not have significant tree cover, and is well-drained sandy, loamy, or gravelly soil. T. mazama in Washington have not been found in clay, and there are few records in silt soils. In sum, deep well-drained, sandy loam or loamy sand with sufficient fertility and water holding capacity to support desired forbs appears to provide optimal habitat (Baker et al. 2003).

3.2.1.5 Threats/Reason for Decline

Much of the Mazama pocket gopher habitat in the south Puget Sound has been lost to development, agriculture, and succession to forest, and what remains continues to be degraded by invasion of Scotch broom and other non-native plants. Residential development that becomes high density has been particularly destructive to prairie habitat, and probably led to extinction of T. m. tacomensis. Habitat loss has eliminated most of the prairie vegetation, though significant areas remain in grassland. Though Mazama pocket gophers are generally protected in recent years by state, county, and local regulation, development may result in some unavoidable habitat loss and additional fragmentation and isolation of habitat patches. Degraded sites may often represent habitat that can support young that have dispersed, but offer inadequate food to consistently support reproduction. Pocket gophers may not persist in high density residential areas due to effects of frequent mowing, herbicides, impervious surfaces, and perhaps elevated mortality rates resulting from predation by cats and dogs and illegal trapping or poisoning of gophers. Most occupied habitat on public lands is

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affected by non-conservation uses including military training and recreation. Gopher populations at airports can be affected by development of airport-related facilities and businesses and management of the vegetation around airport runways and taxiways. Gopher populations at airports benefit from mowing which prevents invasion of the extensive grassland by woody vegetation.

Trends in the human population suggest that amount and quality of habitat will continue to decline without protection and careful management of conflicting uses. Thurston County is projected to have 170,000 additional people and need an additional 50,000 detached single-family housing units, and >25,000 multi-family units by 2040 (Sustainable Thurston 2011:A11). As the habitat patches become smaller, fewer, and farther apart, the likelihood of each patch continuing to support grassland-dependent species declines as intervening habitat patches are lost. These trends generally affect gophers negatively.

The persistence of Mazama pocket gophers on roadsides, vacant lots, lightly grazed pastures, and within commercial timberland suggests that they are relatively resilient, and may be able to persist in rural and low density developed areas. However, recent extinction of the Tacoma pocket gopher indicates that life for gophers in high density residential and commercial areas is hazardous and recruitment and re-colonization is inadequate to maintain local populations. The last possible records of the Tacoma pocket gopher were animals that were killed by pet cats and identified as gophers by homeowners (Ramsey and Slipp 1974). It is not known if the mortalities from these sources have a significant effect on gopher populations, particularly in less densely settled areas. Dogs also are known to kill pocket gophers, but are probably less often free-roaming in unfenced areas. Pocket gophers can damage young trees and, like moles, their diggings can be an untidy nuisance to landowners desiring attractive lawns. They can also be a problem in vegetable gardens and at Christmas tree, berry, and vegetable farms in the area. Mazama pocket gophers are currently protected from killing without a permit; the frequency that they might be trapped or poisoned is unknown. When larger populations are suppressed by these methods, they readily recover if habitat remains suitable, but for small and isolated populations, mortality from persecution added to other hazards may lead to extirpation.

Livestock grazing. Gophers may survive in pastures in rural residential areas, but studies in California indicate that gopher density tends to decrease in heavily grazed pastures (Eviner and Chapin 2003). T. mazama has persisted on well-managed ranches in Thurston County.

Gravel mining. South Puget Sound prairies are located on glacial outwash gravels. Some of these glacial gravel deposits are very deep and valuable for use in construction and road-building, and prairie sites of significant size may be destroyed by gravel mining. One of the historic sites where Tacoma pocket gophers were collected became a large gravel pit, and 2 gravel pits have been opened on occupied gopher habitat in Pierce County south of Roy, and on historical Rock and Rocky prairies in Thurston County. These sites may be restorable to suitable condition for gophers when gravel removal operations have ceased if adequate layers of friable well-drained subsoil and topsoil are restored.

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Airport Management and Development. Pocket gophers occur in grasslands surrounding airport runways and adjoining lands at Olympia and Shelton. Airport safety considerations requires that the vegetation be mowed to maintain visibility, eliminate cover for large animals that might pose a hazard for aircraft, and provide a safety margin should aircraft overshoot or land short of the runway. This management benefits gophers by keeping out woody vegetation and maintaining the grassland. Development of aviation facilities and the surrounding port lands at the Shelton and Olympia Airports poses a potential of habitat loss for what may be the largest populations of T. mazama and T. m. couchi. The Olympia Airport designated 8.6 ac as a Mazama pocket gopher habitat conservation area in an interlocal agreement with WDFW as part of the Airport Five Year Development Plan, and any additional development would be subject to Tumwater critical area ordinances. The Port of Olympia is currently updating their master plan. The Plan projects significant future land developed for general aviation (~114 ac), aviation related/compatible industry (~245 ac), and additional area for parallel taxiways (Barnard Dunkelberg & Co. 2011).

The Port of Shelton had a habitat management plan prepared for the Shelton pocket gopher population on Sanderson Field to comply with Mason County regulations. The habitat plan was prepared in response to revisions in the Comprehensive Plan which identified several portions of the property for development (GeoEngineers Inc. 2003). The plan identifies an area of Port property where Scotch broom and other woody vegetation would be controlled to replace gopher habitat lost to development.

Military Training. The presence of Fort Lewis (part of Joint Base Lewis-McChord) has prevented the loss of habitat to agriculture and residential development for some of the largest remaining T. mazama populations. Mazama pocket gophers exist primarily on prairies where vehicular traffic is currently restricted to established roads, but there are no specific restrictions on training to protect gophers (J. Foster, pers. comm.). The number of Army personnel stationed at JBLM has increased and additional increase is planned (Ft. Lewis Directorate of Public Works 2010). Steinberg (1995) speculated that military training by mechanized units may have negatively affected some gopher populations by compacting the soil. The increase in training needs is likely to increase impacts on grasslands and pocket gophers, but the most damaging training has been concentrated on the same areas, so some less-used prairies have been maintained in good condition. Since gophers do not require native vegetation, the effect of degraded vegetation on gopher populations is uncertain. Changes that decreased the cover of perennial forbs would likely have a negative effect on gophers. Areas damaged by military training are repaired by the Land Rehabilitation and Maintenance program.

Fires that burn the vegetation, whether as part of restoration activities or as a side-effect of training during the summer, help reduce invasion by Douglas fir (Pseudotsuga menziesii) and Scotch broom and have maintained some of the highest quality prairie sites on JBLM. However, smaller portions of the AIA seem to burn too frequently, have a low percentage of native species, and a cover of mostly exotic annual grasses (Tveten and Fonda 1999).

Succession and invasive plants. The fire regime established and perpetuated by Native Americans maintained the south Puget Sound prairies for the past 4,000 years, or more. Fire suppression

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allows succession by native and exotic flora, and without vegetation management the native prairies would probably disappear. Fire suppression allows fire-sensitive species to invade and allows an unusual build-up of fuels that can lead to very hot fires that harm the normally fire-tolerant native species (Tveten 1997). The largest remaining prairie (91st Division) is maintained by prescribed and accidental fires, but large portions of these areas are also subject to disturbance during military training.

Fire suppression allows Douglas-fir to invade and overwhelm prairie. Disturbances such as grazing and vehicle traffic may accelerate colonization by Douglas-fir because Douglas-fir seed germination is enhanced by disturbance that increases mineral soil contact, while native plants may decline with the loss of the moss carpet. Prairie areas where Doug-fir control has been conducted in recent years include Johnson Prairie and Weir Prairie RNA on JBLM, Mima Mounds and Rocky Prairie NAP, Thurston County’s Glacial Heritage Preserve, and Scatter Creek WLA.

Scotch broom is the most visible invasive species that can cover prairies relatively rapidly. Olson (2011a) reported that Scotch broom negatively affected the probability of gopher site occupancy and plot use; the model suggested that plot use appears to decline as Scotch broom cover approached 10%. Parker (2002) reported that the glacial outwash prairie ecosystem is readily invaded by Scotch broom and that simply reducing soil disturbance and fires would not stop broom invasion (Parker 2002). Rook et al. (2011) noted that Scotch broom has long lasting effects on the soil that reduces germination and success of some native species. Scotch broom is killed through burning, hand pulling, or herbicide, but control requires an ongoing program because the plants produce abundant seeds that remain viable in the soil for several decades. Regular mowing can prevent additional Scotch broom seed production. Fire often stimulates germination of broom seeds in the soil, so a second burn, or herbicide is needed to kill the abundant seedlings. Portions of the Artillery Impact Area on JBLM are broom free, indicating that frequent burning prevents broom establishment, but this can also affect native species. All control methods can be detrimental to native species if not well planned.

There are numerous invasive exotic plants that degrade native prairies in the south Puget Sound region, in addition to Scotch broom. Techniques for restoration of the prairies and oak woodlands of the Willamette Valley-Puget Trough-Georgia Basin ecoregion are reviewed in Dennehy et al. (2011), Dunwiddie and Bakker (2011), Hamman et al. (2011), and Rook et al. (2011).

Implications of habitat loss for populations. Pocket gophers are vulnerable to local extinctions because of the small size of local breeding populations (Steinberg 1999). Low effective size of local populations and relatively large genetic differences between populations may be typical of gopher populations (Daly and Patton 1990). Pocket gophers have probably persisted by continually re-colonizing habitat after local extinctions; the loss of habitat patches and increases in hazards such as busy roads may have inhibited the re-colonization that historically occurred. Where additional habitat exists within a few hundred meters, some dispersal and resulting gene flow probably occurs between local populations, and vacant habitat is rapidly colonized. However, as habitat patches become smaller, fewer, and further apart, the likelihood of each patch continuing to support pocket gophers declines.

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3.2.2 Oregon Vesper Sparrow (Pooecetes gramineus affinis Miller, 1888)


The Washington Department of Fish and Wildlife (WDFW) includes this subspecies in the Priority Habitat and Species system, although management recommendations for it have not yet been developed. The American Bird Conservancy and Partners in Flight consider the subspecies and its habitat to be priorities for conservation in western Washington and western Oregon (Altman 2000, 2011). Prairie habitat for the Oregon Vesper Sparrows is a habitat of local importance in Thurston and Pierce counties.


Range and distribution. The breeding range of Oregon Vesper Sparrows extends from southwestern British Columbia through western Washington, western Oregon, and into the northwestern tip of California (Campbell et al. 2001, Jones and Cornely 2002, Altman 2003) (Fig. 1). The subspecies winters from central California west of the Sierra Nevadas to northwestern Baja California, Mexico (AOU 1957).

In Washington, Oregon Vesper Sparrows occur in lowland areas west of the Cascade Mountains (Jewett et al. 1953, Smith et al. 1997, Mlodinow 2005). Although nesting records are few, historical breeding range is believed to have extended from northern Skagit County, the San Juan Islands, and Clallam County (Dungeness and Sol Duc) south through southern Puget Sound, including Thurston County. Breeding also may have occurred in Clark County (Camas and Vancouver) (Fig. 1).

Based on records from 1992 to the present, the current breeding population in Washington is now limited almost entirely to remnant prairies in Thurston and Pierce Counties and grasslands on San Juan Island, though small numbers may still breed in eastern Clallam County and near Shelton in Mason County (Smith et al. 1997, Mlodinow 2005). Breeding season presence in Thurston County during the past 20 years has been recorded at Scatter Creek, Mima Mounds, West Rocky Prairie, Weir Prairie, Johnson Prairie, Tenalquot Prairie, the Olympia airport, Glacial Heritage Preserve, north of Bucoda, Goodard Road SW, and unspecified sites in Grand Mound, Rainier, Lacey, Tumwater, and Nisqually (Fig. 2; WDFW WSDM database). The vicinity of Yelm was once considered a prime area for the subspecies (Jewett et al. 1953), but is no longer occupied. Current breeding season records in Pierce County are focused around the prairie habitats of Joint Base Lewis-McChord (JBLM).

Migration season records since 1992 are scattered through all counties in western Washington except Pierce, Mason, Jefferson, and Wahkiakum counties (WDFW WSDM database). Some of these records likely represent vagrant occurrences of the subspecies P. g. confinis (Rogers 2000).

Population trends. Vesper Sparrow populations have been declining throughout North America since at least the 1960s (Jones and Cornely 2002). Abundance patterns of Oregon Vesper Sparrows reflect this trend, with declines evident across the breeding range (Beauchesne 2006, Altman 2011). In Washington, the subspecies was originally described as “fairly common” to “rather

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abundant” in localized areas of western Washington (Altman 2011), but apparently was never common over a widespread area. Larrison and Sonnenberg (1968) reported it as being of limited abundance and range by the mid-1960s. It was “rare and local….in remnant prairie areas” by the 1990s (Smith et al. 1997), with the exception of 91st Division Prairie on JBLM, where about 100 singing males were on established territories in 1998 (Rogers 2000). The current Washington population is threatened with extirpation (Mlodinow 2005), and is estimated at 250-300 birds in the Puget Lowlands and 50-100 birds on islands along the lower Columbia River (Altman 2011). Current numbers in Thurston County are unknown, but are apparently quite small (i.e., 0 to a few birds each) at Mima Mounds, Scatter Creek, and West Rocky Prairie (D. Canning, unaffiliated, unpubl. data).


Oregon Vesper Sparrows are present in western Washington mainly from early April through late September, with relatively few records during other months (Mlodinow 2005; WDFW, unpubl. data). Most spring migration occurs from early April to early May (Mlodinow 2005), with birds beginning to arrive at prairie sites in Thurston and Pierce counties after about April 10 (WDFW WSDM database). Males arrive up to a week earlier than females in other parts of North America (Best and Rodenhouse 1984). After nesting concludes, Vesper Sparrows typically gather in small groups until fall migration (Bailey and Niedrach 1965). Fall migration through western Washington is primarily from mid-August to late September, with fewer records extending into October (WDFW WSDM database). Migration usually occurs at night, with most individuals joining small flocks of up to 10 birds (Rising 1996, Jones and Cornely 2002). The species sometimes migrates with Horned Larks (Eremophila alpestris) and Savannah Sparrows (Passerculus sandwichensis) (Berger 1968, Hyde 1979). Individuals are rarely detected from November to March in western Washington (WDFW, unpubl. data).

Birds begin singing after arriving at their breeding sites (Altman 2003). Singing occurs most frequently early in the morning, subsides during the day, and then increases again from sunset to dusk (Jones and Cornely 2002). Singing is typically performed from elevated perches, such as fences, trees along the edges of fields, shrubs, grass, and the stalks of forbs, but may be conducted from the ground when perches are lacking (Berger 1968, Wiens 1969, Castrale 1983, Jones and Cornely 2002, Altman 2003). Territory sizes average 1.25 ha (range = 0.44 to 5.26 ha) (average = 3.1 ac, range = 1.1-13.0 ac) in western Oregon (Altman 1999, 2003), with larger sizes usually reflecting poorer food availability (Jones and Cornely 2002).

Vesper Sparrows become sexually mature a year after hatching and are seasonally monogamous (Jones and Cornely 2002). Females construct the nest alone (Rising 1996). Nests are made from grasses in the shape of a shallow bowl and have an outer diameter of 8-10 cm (3-4 in) (Berger 1968, Godfrey 1986, Peck and James 1987). Nests are placed on flat ground or in a shallow depression, and are usually located next to a clump of vegetation, crop residue, dirt clod, or at the base of a shrub or tree (Jones and Cornely 2002, Altman 2003).

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Oregon Vesper Sparrows nest from about late April to mid-July, with the few western Washington records reported from May 9 to July 7 (Bowles 1921, Altman 2003, Beauchesne 2006; WDFW, unpubl. data). It is unknown whether this subspecies produces second broods (Rogers 2000). Eggs measure 20 mm (0.8 in) long by 15 mm (0.6 in) wide on average (Jones and Cornely 2002). Clutch size for Vesper Sparrows (including P. g. affinis) is usually 3-5 eggs (range = 2-6 eggs). Incubation averages 12-13 days and is performed mostly by the female. Both parents feed the chicks, although primary responsibility of the first brood may fall to the male if the female begins a second brood (Berger 1968). Young fledge from the nest after 9-10 days on average and remain dependent on the parents for another 20-29 days (Perry and Perry 1918, Dawson and Evans 1960). Brown-headed cowbirds (Molothrus ater) parasitize Vesper Sparrow nests in parts of their range, including eastern Washington (Rising 1996, Campbell et al. 2001; M. Vander Haegen, pers. comm. in Altman 2003), but it is unknown if this occurs for Oregon Vesper Sparrows. Best and Rodenhouse (1984) reported that about half of breeding adults return to their nesting site the following year.

Diet is comprised of grass and forb seeds year-round, but is heavily supplemented with insects (especially grasshoppers, beetles, and caterpillars) and other arthropods during the breeding season (Berger 1968, Rotenberry 1980, Jones and Cornely 2002). Most foraging occurs on the ground, but birds will hop and hover to glean food from vegetation.

Average lifespan of Vesper Sparrows is unknown, but a maximum of 7.1 years has been recorded for a banded individual in the wild (Klimkiewicz 1997). Fledging rates average 3.0 young/successful nest and 1.0 young/active nest (Wray et al. 1982). Predation and farming activities are common causes of nest failure (Rodenhouse and Best 1983, Best and Rodenhouse 1984, Patterson and Best 1996, Best et al. 1997).


Vesper Sparrows inhabit a variety of grassland types, including shortgrass and tallgrass prairie, desert and semi-desert grasslands, shrub-steppe, croplands, hay fields, weedy fence rows and roadsides, and woodland edges (Campbell et al. 2001, Jones and Cornely 2002). Preferred areas for breeding territories typically have short sparse and patchy grassy and herbaceous cover, some bare ground, low to moderate shrub or tall forb cover, and low tree cover (Reed 1986, Campbell et al. 2001, Dechant et al. 2002, Jones and Cornely 2002). Some structural diversity of vegetation appears to be an important factor in site selection, with shorter vegetation chosen for foraging and scattered taller plants used for cover and singing perches (Davis and Duncan 1999, Beauchesne 2006).

Oregon Vesper Sparrows also show some variation in breeding habitat. In western Washington, the subspecies was originally widespread in prairies and pastures (Jewett et al. 1953), but had become restricted to the edges of open prairies by the 1990s (Rogers 2000, Mlodinow 2005). Breeding habitat in the state remains poorly quantified. Clegg (1998, 1999) reported that all breeding territories (n = 23) at JBLM were in areas of high quality prairie supporting intact Idaho fescue (Festuca idahoensis) located near prairie edge. Size of the prairie appears to be an important factor in current site selection, with only large prairies occupied now (S. Pearson, WDFW, pers. comm.).

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In strong contrast to western Washington, nearly all detections of Oregon Vesper Sparrows in Oregon’s Willamette Valley are in young Christmas tree farms (i.e., 2-5 years after planting) with extensive grass and weed cover, or in lightly grazed pastures with scattered shrubs and grass heights of less than 30-60 cm (1-2 ft) high (Altman 1999, 2003). Habitats avoided include cultivated grass fields, highly manicured Christmas tree farms, and fallow fields with grass heights exceeding 60 cm (2 ft) high. In southwestern British Columbia, the subspecies originally bred in pastures, agricultural land, and airport fields with patches of grasses and weeds (Campbell et al. 2001), but the few remaining territories are now present only in grasslands next to hayfields, which contain native and non-native plants (Beauchesne 2006).

The only study characterizing the microhabitat of nest locations of Oregon Vesper Sparrows reported that nests in the Willamette Valley were built in areas with relatively reduced grass cover (49%) and sizable amounts of bare ground (24%) and litter/ residue (21%) compared to other locations within territories (Altman 1999, 2000). Woody vegetation also was regularly present near many nests. Rogers (2000) reported reduced vegetation heights (average = 15-21 cm [6-8.5 in]) and densities at foraging locations compared to random sites in prairies in Pierce and Thurston counties, Washington.

Habitat selection during migration is poorly described. Migrating individuals have been observed in a variety of grassy habitats in western Washington (Mlodinow 2005).


Two major factors contributing to the declines of Vesper Sparrows in much of their North American range are habitat loss through conversion of native grasslands and shrublands to unsuitable types of agriculture, and the shift in farming practices to more intensive tillage and greater use of chemicals (Jones and Cornely 2002). Grazing impacts on Vesper Sparrows vary with grazing intensity and soil type, but locations exposed to heavy grazing typically support lower breeding densities than sites with moderate and light grazing (Kantrud and Kologiski 1982, Altman 1999). In addition to habitat modification, grazing can result in trampling of nests (Altman 1999).

Declining populations of Oregon Vesper Sparrows result primarily from habitat loss and degradation, and potentially from increased predation and human disturbance (Smith et al. 1997, Altman 1999, 2003, 2011, Rogers 2000, Beauchesne 2006). South Puget Sound prairies originally covered an estimated 60,470 ha (149,360 ac), but had declined in size by 90% by the mid-1990s, with only 3% remaining in intact prairies (Crawford and Hill 1997). During this period, the number of prairies in South Puget Sound fell from 233 to 29 sites and average size decreased from 260 to 175 ha (641 to 433 ac). This decline was driven by urban conversion, encroachment of Douglas-Fir (Pseudotsuga menziesii) forests caused by fire control, and conversion to farmland (Chappell and Kagan 2001). Many remaining prairies are degraded by the invasion of Scotch broom (Cytisus scoparius) and other non-native plants (Chappell and Kagan 2001).

Oregon Vesper Sparrows also may be experiencing increased predation from species associated with semi-urban and residential areas such as feral and domestic Cats (Felis catus), Raccoons

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(Procyon lotor), American Crows (Corvus brachyrhynchos), and Opossums (Didelphis virginiana) (Altman 1999, Rogers 2000, Pearson 2003, Stinson 2005, Beauchesne 2006). Human disturbance from various activities in prairies, such as military training, dog field trials, off-leash dog walking and training, horseback riding, bicycling, hiking, model airplane flying, school field trips, prescribed burning, Scotch broom control, and other habitat management activities also may be disruptive or harmful to the subspecies (e.g., Rogers 2000).

One other factor contributing to the vulnerability of populations is the Oregon Vesper Sparrow’s somewhat linear distribution through the lowlands of western Oregon to southwestern British Columbia (Altman 2011). This has helped cause substantial population fragmentation, resulting in a higher likelihood of reduced genetic variability and adaptability in surviving populations.

3.2.3 Oregon Spotted Frog (Rana pretiosa Baird and Girard, 1853)


The Oregon Spotted Frog is listed by the USFWS as Threatened (USFWS 2014b), and was listed as endangered in Washington in 1997. The species persists at only 6 Washington locations. In Thurston County, Oregon Spotted Frogs occur in the Black River drainage. The Oregon Spotted Frog population on Beaver Creek (a tributary of the Black River) occurs adjacent to West Rocky Prairie and is the only remaining population in south Puget Sound Lowlands associated with native prairie. Washington State status has been reported (McAllister and Leonard 1997, http://wdfw.wa.gov/publications/00382). The Draft Washington State Recovery Plan for the Oregon Spotted Frog (Hallock, in prep.) is currently under review; information herein relies heavily on information gathered for the recovery plan.


The Oregon Spotted Frog is a Pacific Northwest endemic historically distributed from southwestern British Columbia, Canada (Matsuda et al. 2006) to northeastern California, USA (Hayes 1997a), including the Puget Trough -Willamette Valley, and East Cascades-Modoc Plateau ecoregions. Oregon Spotted Frog populations have declined throughout the range and have been extirpated from large portions of their historical distribution. Range loss based on historical site analysis is estimated to be 79% but may approach 90% (Hayes 1997a). Available evidence indicates the species has been extirpated from the southern portion of its range in California and the lowland Willamette Valley in Oregon; the fate of populations at the northern extreme of the range in Canada is precarious (Hayes 1997a, Haycock 2000). Approximately 47 occupied sites are known to persist in Oregon (32), Washington (11) and British Columbia (4) (USFWS 2011, Gay and Bohannon 2011, Bohannon et al. 2012).

Locations of Oregon Spotted Frog populations in Washington went largely undocumented historically. McAllister and Leonard (1997) reviewed museum records from major herpetological collections of North America. These specimens reveal an historical distribution in the Puget Trough lowlands and southern Washington Cascades (McAllister 1995, McAllister and Leonard 1997) with nine widely separated populations verified by specimen records (McAllister and Leonard 1990,

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1991, McAllister et al. 1993). McAllister and Leonard (1997) identified 2 additional historical localities, Pattison Lake and Kent, based on reports by Professor James Slater and Warren Jones. In 2011and 2012, Oregon Spotted Frogs were found in the South Fork Nooksack River, Samish River, and Chilliwack River drainages (Gay and Bohannon 2011, Bohannon et al. 2012). Assuming that watersheds currently occupied were also occupied historically, Oregon Spotted Frogs occupied at least 14 watersheds in Washington. All Washington sites, historical and extant, are found below 634 m (2,080 ft.). Six extant occurrences persist in Washington including populations in the lower South Fork Nooksack River (Whatcom Co.), lower Chilliwack River (Whatcom Co.), upper Samish River (Whatcom & Skagit Cos.), upper Black River (Thurston Co.), lower Trout Lake Creek (Klickitat and Skamania Cos.) and Conboy Lake in Outlet Creek (Klickitat Co.) (Hallock, Recovery Plan, in prep.).

South Puget Sound lowlands and prairies (Pierce and Thurston Counties). The WDFW WSDM database contains 413 records of Oregon Spotted Frog in Pierce and Thurston Counties (Fig. 1). The vast majority of these records (403 or 97.5%) are from Thurston County where the species is extant. The 10 records for Pierce County include 8 museum specimens from 1937-1959 at Spanaway Lake, Little Spanaway Lake, and Spanaway Pond and a 1937 museum specimen collected 3 miles (4.8 km) west of Kapowsin. Oregon Spotted Frogs were last collected from Pierce County in 1959 at Spanaway Pond by J.N. Knudsen (Pacific Lutheran University museum specimens 40-43). The Spanaway sites were in the historical Nisqually Plains/Prairies. In 2008, an Oregon Spotted Frog reintroduction was started on Joint Base Lewis-McChord at Dailman Lake. The success of this project has yet to be determined.

In Thurston County, the only site documented historically was Patterson Lake (= Pattison Lake) where a museum specimen was collected in 1950. The rest of the 402 records come from various inventory and monitoring activities at sites along the upper Black River and its tributaries including Dempsey Creek, Salmon Creek, Blooms Ditch, Allen Creek, and Beaver Creek (Fig. 2). The population on Beaver Creek occurs adjacent to West Rocky Prairie and is the only population in the south Puget Sound Lowlands associated with native prairie. The rest of the sites are found in wetlands and creeks adjacent to pastures.


The Oregon Spotted Frog is a medium-sized, aquatic, ranid frog. They aggregate to breed following the coldest weeks of winter. Breeding frogs gather in seasonally flooded margins and shallows of emergent wetlands in areas that receive minimal shading from the surrounding vegetation. Frogs use the same breeding areas every year and depending on topography and site conditions, may lay eggs at the same site. Orientation to the breeding site is poorly understood but seems to involve a combination of non-vocal and vocal cues (Licht 1969, Risenhoover et al. 2001a). The male advertisement call sounds like faint, rapid, low-pitched tapping (Stebbins 2003). Calling occurs at the water surface and subsurface (Licht 1969, Bowerman 2010). The surface calls orient females to the egg deposition (oviposition) site (Licht 1969). Initiation dates of egg deposition vary by year depending on spring conditions (Licht 1969). In general, oviposition commences when subsurface

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waters are 7-9°C (45-48°F) and minimum water temperatures rarely fall below 5°C (41°F) (Licht 1971a, Hayes et al. 2000, McAllister and White 2001). Other cues also may be involved.

Oviposition start date has been tracked since 1996 at a study site on Dempsey Creek (elev. 60 ft [18 m], Thurston County). These data reveal a 3-week time span from mid-February to the first week of March when oviposition commences (K. McAllister, WA Department of Transportation, pers. comm.). Once initiated, breeding is “explosive” with many pairs breeding during a short time period (Licht 1969, Nussbaum et al. 1983, Briggs 1987). Most frogs spawn mid-day (Licht 1969) but nocturnal spawning also has been detected using wildlife cameras (J. Tyson & M. Hayes, WDFW, pers. comm.). Within a breeding area, multiple bursts of egg deposition may occur over a 2-3 week period.

Oregon Spotted Frogs may have a serially monogamous mating system with each female laying a single clutch per year that is fertilized by a single male, and each male breeds with only 1 female (Phillipsen et al. 2009). Fertilization is external. The male clasps the female around the upper body with his forearms in an embrace called amplexus. This embrace aligns the vents of the male and female in close proximity for spawning. The first pair of frogs to lay eggs selects the oviposition site. Each female lays a single globular egg mass that expands to the size of a softball. Additional females subsequently deposit their egg masses on top of or immediately adjacent to the initial egg mass. Eggs are deposited in shallow water typically ≤ 15 cm but up to 30 cm (up to 12 in.) deep (Licht 1969, Hayes et al. 2000, Lewis et al. 2001, McAllister and White 2001, Risenhoover et al. 2001a). Oregon Spotted Frogs occasionally lay egg masses on floating mats of prostrate Reed Canarygrass (Phalaris arundinacea) in waters that are deeper than typically used (> 30 cm, 12 in.) (McAllister and White 2001; M. Bailey, USFWS, pers. obs. and L. Hallock, WDFW, pers. obs.). When a communal egg mass cluster is established, males call from near it and on top of it (Licht 1969). Licht (1969) showed the significance of the egg mass clustering behavior by moving the initial egg mass. All subsequent females laid their eggs on the communal cluster at or near the new location and no females laid at the original location. At a low elevation site in British Columbia (Canada), females bred every year, averaging 643 eggs (range 249-935) in each mass (Licht1974).

Egg laying habits and certain aspects of the globular egg mass shape are adaptations for rapid development. The large egg mass retains more heat than smaller egg masses (Hassinger 1970, Duellman and Trueb 1986) and communal egg deposition produces higher daytime temperatures for the developing embryos (Licht 1971a, Duellman and Trueb 1986, McAllister and White 2001). The clustering of egg masses also may provide the majority of embryos protection from temporary stranding events, freeze damage, and egg predators. The placement of egg masses in the comparably warmer shallow waters and the selection of sites that receive minimal shading from the surrounding vegetation also speed development rates. Non-shaded habitat quickly warms on sunny days limiting potential freeze damage from cold nights. Embryos do not survive freezing (Licht 1971a). Non-shaded habitat also enhances development of algae that live symbiotically in the eggs and may be critical for oxygen delivery to and removal of nitrogenous waste from the innermost embryos in communal clusters (Pinder and Friet 1994).

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Embryo development to hatching can occur in as little as 14 days with 18-30 days being the typical development time (Lewis et al. 2001, McAllister and White 2001, Risenhoover et al. 2001a). The free-swimming larvae disperse from communal egg mass clusters a week or so after hatching. The tadpoles are primarily herbivorous feeding on algae, decaying vegetation, and detritus (Licht 1974); this life stage is dedicated to eating and growth. The tadpole stage lasts about 4 months (Licht 1974). In late summer, the tadpoles metamorphose into fully-formed, small frogs about 33 mm (1.3 in.) snout-vent length (Nussbaum et al. 1983).

Metamorphosed frogs prey primarily on invertebrates (Licht 1986b). Growth is rapid until adult sizes are achieved 1 to 2 years following metamorphosis (Licht 1975). At a low-elevation site in Thurston County, adult males continued to grow an average of 2.2 mm (0.09 in) per year and adult females grew 6.2 mm (0.24 in) per year (Watson et al. 2000). Longevity > 9 years was documented for a PIT-tagged Oregon Spotted Frog (K. McAllister, WA Department of Transportation, pers. comm.); longevity for most Oregon Spotted Frogs likely is shorter (Licht 1975, McAllister and Leonard 1997). Oregon Spotted Frogs do not have a prolonged period of hibernation (<1 month; Hayes et al. 2001, Hallock and Pearson 2001, Risenhoover et al. 2001b, Watson et al. 2003, Shovlain 2005) and they can be active under ice (Leonard et al. 1997, Hallock and Pearson 2001). Oregon Spotted Frogs rarely move long distances and have not been recorded moving > 2,360 m (7,750 ft.; Forbes and Peterson 1999, McAllister and Walker 2003).

Oregon Spotted Frogs suffer mortality mainly from predators and chance environmental events. Freezing temperatures and stranding of egg masses are the main threats to developing Oregon Spotted Frog embryos. An entire cohort can be lost in years when water retreats after breeding is underway. Freeze damage is a cause of embryonic mortality in years where temperatures drop below freezing after breeding is underway. The highest rates of embryo mortality are observed in years when the egg masses became temporarily stranded due to a period without precipitation that coincides with freezing night temperatures. Significant mortality also can result when tadpoles become isolated in breeding pools away from more permanent waters (Licht 1974, Watson et al. 2003).

In terms of predators, tadpoles are most vulnerable to predation when small (Licht 1974). In southwestern British Columbia, Licht (1974) found predators on Oregon Spotted Frog tadpoles to be mostly invertebrates. Fish also are likely predators on tadpoles (Hayes and Jennings 1986, McAllister and Leonard 1997, Hayes 1997a, Pearl 1999). The frogs are preyed on by a variety of vertebrate predators including native (Licht 1974) and non-native amphibians (e.g., American Bullfrogs, Lithobates catesbeianus formerly Rana catesbeiana; McAllister and Leonard 1997, Pearl et al. 2004), Common Garter Snake (Thamnophis sirtalis; Licht 1974, Hayes 1997a, McAllister and Leonard 1997, Forbes and Peterson 1999, Pearl and Hayes 2002, Watson et al. 2003), birds such as Sandhill Cranes (Grus canadensis; Hayes et al. 2006) and Great Blue Herons (Ardea herodias; Licht 1974), and mammals such as Mink (Neovison vison; Bowerman and Flowerree 2000; Watson et al. 2000, Hallock and Pearson 2001) and river Otters (Lontra canadensis; Hayes et al. 2005). Adult annual survival of a study population at Dempsey Creek (Thurston County) was 38% (Watson et al. 2000).

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Washington’s remaining populations of Oregon Spotted Frogs occupy still-water wetlands connected by riverine systems. The perennial creeks and associated network of intermittent tributaries provide aquatic connectivity between breeding sites, active season habitat, and overwintering habitat. Additionally, perennially flowing waters may provide the only suitable habitat during extreme summer drought or during winter when oxygen levels drop in still-waters under ice and snow. Associated wetlands have a mix of dominance types including aquatic bed, emergent, scrub-shrub, and forested wetlands. The seasonally inundated wetland margins are frequently hay fields and pasture. The less disturbed sites have wet meadows and prairie uplands. Some occupied sites are formed by American Beaver (Castor canadensis) activity. All the remaining Oregon Spotted Frog sites have moderate to severe habitat alteration including a history of cattle grazing and/or hay production as well as encroaching or established rural residential development. Hydrology has been altered to some extent at all sites.

Watson et al. (2003) stressed that the most important features for microhabitat use were water depth, flow characteristics (still water was used over flowing water), and a high degree of water surface exposure (i.e., 50-75% water) or conversely, a low to moderate degree of emergent vegetation (i.e., 25-50%). The predominant use of shallow water habitat by Oregon Spotted Frogs was illustrated by Watson et al. (2003), who found Oregon Spotted Frogs selected water depths of 10-30 cm (~4-11.7 in.) with less emergent vegetation and more submergent vegetation than adjacent habitats.

Oregon Spotted Frogs select breeding sites in seasonally flooded wetland margins adjacent and connected to perennial wetlands (Licht 1971a, Hayes et al. 2000, Pearl and Bury 2000, Watson et al. 2000, Hallock and Pearson 2001, Lewis et al. 2001, McAllister and White 2001, Risenhoover et al. 2001a, Watson et al. 2003, Pearl and Hayes 2004). Full solar exposure also seems to be a significant factor in breeding habitat selection (McAllister and White 2001, Pearl and Hayes 2004). Oviposition sites are in shallow waters with low vegetation structure that does not shade the eggs. Typically these locations are near shore but can also be in areas with extensive shallows. Low vegetation structure is typical of early successional vegetation stages but also can result from cattle grazing, haying and mowing. Heavy snow pack also can flatten emergent vegetation providing suitable oviposition conditions.

Post-breeding habitat use is the least studied of Oregon Spotted Frog habitat associations in Washington. During the summer drought (July to September), frogs in Thurston County were restricted to remnant pools that persisted during this time (Watson et al. 2003). At a site in Oregon, habitat use was primarily near-stream with frogs showing high micro-site fidelity (Shovlain 2005). During the coldest months, Oregon Spotted Frogs require well-oxygenated waters (Hallock and Pearson 2001, Hayes et al. 2001, Tattersall and Ultsch 2008) and sheltering locations protected from predators and freezing conditions (Risenhoover et al. 2001b, Watson et al. 2003). This is especially important during the coldest periods when activity of this ectotherm is expected to be the lowest.

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Slipp (1940) reported Oregon Spotted Frogs to be associated with prairie lakes and streams in the area between Tacoma and the Nisqually River (Tacoma Plateau/Nisqually Plains). Oregon Spotted Frogs require breeding habitat with low vegetation structure and full solar exposure (McAllister and White 2001, Pearl and Hayes 2004). Puget Sound prairies would have provided such habitat within an otherwise densely forested landscape.

3.2.3.5 Threats and Reasons for Decline

The decline of Oregon Spotted Frogs is attributable to several related factors. Among the most significant is the loss and alteration of wetland habitat. Oregon Spotted Frogs have life history traits, habitat requirements and population characteristics that make them vulnerable to such loss and limit their distribution. These include 1) a completely aquatic life history; 2) communal reproduction concentrated on the landscape with the same localized breeding areas used annually; 3) high levels of population fluctuation; 4) dispersal limited to aquatic corridors, 5) relatively large permanent wetlands (> 4 ha, 10 ac) that include shallow, warm-water habitats, 6) breeding habitats that have shallow water (≤ 30 cm, 12 in), short vegetation and full sun exposure with relatively stable hydrology and aquatic connectivity to permanent waters, and 7) overwintering habitats that provide adequately oxygenated water and shelter from freezing conditions and predators. Additional threats include geographic isolation of Oregon Spotted Frog populations, loss of natural processes that set back vegetation succession (e.g., beaver activity), invasion of exotic grasses into shallow wetland habitats, colonization of wetlands by non-native predators, and increase of water-borne pollutants and emerging diseases. This list of threats is neither exhaustive nor independent, as a number of factors are interconnected. Climate change is a further concern because it involves potential changes that are likely to increase effects of the above factors on Oregon Spotted Frog habitat.

Based on conservative estimates, Washington lost over 33% of its wetlands between pre-Euro-American settlement condition and the 1980s (Canning and Stevens 1990). This percentage accounts for complete loss from draining or filling, but does not account for alteration or degradation. Freshwater marshes and forested wetland experienced the greatest losses. Snohomish County estimated wetland losses of 180 acres (72 ha) per year during the 1990s. Assuming a similar rate, losses for the 8 urbanized counties with similar growth projections plus King and Pierce counties would be 1,800 acres (728 ha) per year (Canning and Stevens 1990). These counties are primarily in the Puget Sound Ecoregion where the majority of the historic distribution of Oregon Spotted Frogs in Washington State had been documented (McAllister and Leonard 1997). More specifically, case studies in Washington showed losses of freshwater wetland acreages reflected on U.S. Geological Survey quadrants to be 55% for Tenino and Yelm (south Thurston County), 82% for Tacoma South (Pierce County), and 70% for Lake Washington (King County) (Boule et al. 1983). Data on wetland changes in Washington since 1995 are lacking.

Invasive wetland species that alter wetland structure and function impact Oregon Spotted Frog habitat. Reed Canarygrass is present at all Washington sites and is the invasive plant of greatest concern due to the potential loss of Oregon Spotted Frog habitat from shading and impenetrable thatch. The grasses’ high rate of transpiration and ability to outcompete native plant species also

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are of concern for Spotted Frog habitat. In the Lower Puget Sound, Reed Canarygrass is especially problematic because there is no snow pack to compress it and the vertical structure shades breeding habitat.

South Puget Sound prairies. The south Puget Sound prairies were reduced to about 10% of their former abundance primarily due to agriculture and development (Crawford and Hall 1997). This likely affected the associated wetlands, especially seasonally flooded areas that would have been easily drained and converted to uplands. Historically, depressions and low areas of Thurston County, when drained, were better suited to hay and pasture than most of the well-drained upland soils and conversion to pasture was extensive. By 1947, pasture occupied more farmland than all other crops combined in Thurston County (Poulson et al. 1947).

3.2.4 Slender-Billed White-Breasted Nuthatch


The apparent decline in abundance of the Slender-billed White-breasted Nuthatch and range contraction from north to south in the Puget Lowlands substantiate its conservation rankings and concern for this species (USFWS 2011, WDFW 2012). The American Bird Conservancy and Partners in Flight consider the species and its oak habitat to be priorities for conservation in western Washington and Oregon (Altman 2000, ABC 2006, Altman 2011). Genetic analyses indicate that S. c. aculeata is genetically distinct from other regional populations in North America and demonstrates significant genetic differentiation among populations within this subspecies’ range (Spellman and Klicka 2007). This subspecies may now be extirpated from South Puget Sound with no evidence of breeding or sightings during the breeding season since the mid to late 1990s, respectively (Fig. 1; Chappell 2005, Altman 2011). This bird is a Species of Greatest Conservation Need (SGCN) in Washington’s Comprehensive Wildlife Conservation Strategy prepared by WDFW in 2005.


The Slender-billed White-breasted Nuthatch was historically found west of the Cascades in Washington (Kitchin 1934, Jewett et al. 1953). This species apparently did not breed historically in northwestern Washington north of Seattle or on the San Juan Islands (reviewed in Altman 2011). Christmas Bird Count data (1966-2006) indicate a 2.2% annual decline in this subspecies (Altman 2011).

Beginning in the early 1900s this species declined in abundance and experienced a contraction in its range in the Puget Lowlands. The species was “quite abundant” in the South Puget Lowlands in the 1850s and a “common resident, being especially numerous among the large oaks of the prairie country” around Tacoma in 1896 (Bowles 1929:53). However, the species had declined significantly by the early 1900s in the Tacoma area, with the species considered “not common” (Bowles 1906) and “practically unknown” (Bowles 1929), and in the Puget Sound region the species was considered “rare and local” (Dawson and Bowles 1909). By the early 1930s, it was “rare…almost accidentally in western Washington; formerly a common resident” (Kitchin 1934:16).

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In the 1940’s it was an “uncommon resident” in the Puget Sound area (Larrison 1952:88). By the early 1950s, Kitchin (1949:185) noted that although the species was “at one time fairly common in western Washington…they have been gone many years”, and Jewett et al. (1953:484) described it as “common in the oak region of the prairies in western Washington”. A synthesis of breeding records of the species during the 1960s by Altman (2011, original sources cited within) indicated that by the mid-1960s it was an ”irregular …breeder west of the Cascades” but still a “common bird in south Tacoma” and regularly occurring at Joint Base Lewis-McChord in oak and Ponderosa Pine habitats. By the late 1970s only 9 breeding sites were known in South Puget Sound, all with small populations (Chappell and Williamson 1984); these populations subsequently vanished over the next 25 years with the last known breeding in Lakewood south of Tacoma in 1995 (Fig. 1; Chappell 2005, Altman 2011). More recently, a small breeding population existed around Woodland and Kalama in Clark County up until 1995, but there is no indication that this population persists (Altman 2011).

Given the close association of this species with oak trees for nesting and foraging, historical and current distribution of oak habitats may provide insight into the historical and current distribution of the nuthatch within the planning area. Although information on historical distribution of prairies containing oak is lacking, the extent of prairie soils provide spatial limits to where oak habitat could have occurred historically. Locations of breeding locations provide additional information on where the species occurred historically. Current distribution for this species is based on recent efforts to map oak habitats in the Puget lowlands using aerial photography; no recent (since 1998) sightings of this species have been documented within the planning area (Fig. 2). In other parts of the subspecies’ range (Fig. 1) it is more common. In Oregon, the species is “common to uncommon…in western Oregon lowlands” and “common resident in oak and mixed forests, nut orchards, and suburban plantings in the Willamette Valley region” (Hagar 2006).


The Slender-billed White-breasted Nuthatch is a cavity user and year-round resident in western Washington (Anderson 1970, 1972). Pairs establish territories of about 10-15 ha (25-37 ac) and occupy the same territories year-round with more vigorous defense during the breeding season (Pravosudov and Grubb 1993, Hagar 2006). Because of weak cavity excavating capabilities, it relies mostly on naturally occurring cavities in living trees (Wilson et al. 1991, Viste-Sparkman 2006) and less so on cavities excavated by woodpeckers (Bent 1948, McEllin 1979, Viste-Sparkman 2006) for nesting. Tree cavities are also needed for roosting, and multiple cavities are used throughout the year (Gumtow-Farrior 1991). Nest building occurs in April and May (Hagar 2006) and nest cavities are lined with moss, fur, feathers, or similar soft materials (Jewett et al. 1953, Chappell and Williamson 1984). Nesting occurs early, with clutches typically completed no later than mid-April in Washington (Dawson and Bowles 1909) and the first week of May in Oregon (Griffee and Rapraeger 1937). Most nests are incubated in late April to early May (Griffee and Rapraeger 1937). Clutches contain 5-10 eggs (Griffee and Rapraeger 1937, Kitchin 1949, Matthysen 1998). Only the female incubates and the male brings food to her during this time (Pravosudov and Grubb 1993). After incubation for about 15 days the eggs hatch and both adults feed the young (Dawson and

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Bowles 1909, Pravosudov and Grubb 1993). Nestlings leave the nest after 25 to 27 days (Viste-Sparkman 2006) with most nests (71%) successfully fledging young (Viste-Sparkman 2006).

Foraging is concentrated on limbs and large branches of oak trees (Wagner 1981) where insects, including their eggs and larvae, are gleaned from tree surfaces (Jewett et al. 1953). In the Willamette Valley, weevils and earwigs were an important part of the diet during breeding and post-breeding periods with diversity of diet decreasing in winter months (Anderson 1976).


Across North America, the White-breasted Nuthatch is typically found in mature deciduous woodland and mixed deciduous and conifer forest (Pravosudov and Grubb 1993). In Washington and Oregon this species is associated with Oregon white oak (Quercus garryana) west of the Cascade Range and conifer forest, primarily ponderosa pine (Pinus ponderosa), east of the Cascade Range (Chappell 2005, Hagar 2006). The few studies that have investigated habitat characteristics of the Slender-billed White-breasted Nuthatch describe foraging (Wagner 1981) and nesting (Wilson 1980, Hagar and Stern 2001, Viste-Sparkman 2006) habitat associations; these observational studies limit interpretation of results.

Foraging. Foraging typically occurs in oak trees on the larger limbs and tree trunk, as documented in a California oak woodland (Wagner 1981), in the Willamette Valley (Anderson 1976), and in other parts of its range (Wilson 1970). Large branches are characteristic of open-grown oak trees with their associated “mushroom-like” crowns compared to slimmer branches and narrower tree crowns of closed-forest-grown oaks (Silen 1958, Burns et al. 1990). Fissured bark, which occurs more on larger oaks, supports a greater abundance and diversity of arthropods compared to smooth bark, and provides greater surface area for invertebrates (Jackson 1970, Nicolai 1986). More large limbs and greater surface area of fissured bark on the larger, open-grown oak trees may provide more optimal foraging surface area for nuthatches.

Nesting. Large decadent oak trees are important nest structures for the Slender-billed White-breasted Nuthatch (Wilson et al. 1991, Viste-Sparkman 2006). Most nests occur in cavities formed by natural decay processes. Further, in the Willamette Valley where this species nests mostly in natural cavities, the cavities occurred primarily in decadent portions of live trees rather than snags (Viste-Sparkman 2006). In addition to the presence of dead limbs and other forms of decadence, large diameter (>50 cm [19.7 in] dbh) and presence of multiple cavities also influence nest tree selection (Wilson et al. 1991, Viste-Sparkman 2006). In the Willamette Valley, nests in 50 oak trees averaging 70 cm (27.5 in) dbh (diameter at breast height), had a mean cavity height of 6.1 m (20.0 ft) (CI, 4.9 to 7.3 m), and mean diameter at the nest cavity was 49 cm (19.3 in) (CI, 40.7 to 87.5 cm) (Viste-Sparkman 2006). At home range scale (10 ha, 25 ac), pairs selected nest trees that were larger in diameter and likely to contain multiple cavities (Viste-Sparkman 2006). For oaks, abundance of cavities correlates positively with tree diameter (Gumtow-Farrior 1991, Wilson et al. 1991), and open-grown Oregon White Oaks are more prone to cavity development than oaks grown in dense stands (Gumtow-Farrior 1991). These findings highlight importance of large, open-grown oaks that depend on natural decay to produce most nest cavities.

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The Slender-billed White-breasted Nuthatch selects nest trees based on the local woodland structure. In the Willamette Valley, measures of tree size were important in determining where nuthatches nested. The positive influence of increases in tree diameter, canopy cover, and oak basal area and the negative influence of increases in small trees on nest tree use suggest that fewer larger trees surrounded nest trees (Viste-Sparkman 2006). In addition, density of White-Breasted Nuthatches was greater in areas with more large oaks and fewer small oaks (Viste-Sparkman 2006), results that are consistent with the Willamette Valley (Hagar and Stern 2001). These findings highlight the importance of large open-grown oaks with a sparse understory in woodlands as suitable nesting habitat.

Patch size and landscape configuration. Patch size and configuration of habitat influences nesting density of nuthatches. In the Willamette Valley, Slender-billed White-breasted Nuthatches were more abundant in smaller (<12 ha, 30 ac) than larger (>25 ha, 62 ac) woodland patches. The species responded positively to increased edge, particularly small woodlands with high edge-to-interior ratio compared to large woodlands (Viste-Sparkman 2006). Amount of oak and density of edge within an average home range and in the greater landscape (1 km, 0.6 mi) positively associated with nuthatch densities (Viste-Sparkman 2006). This species may respond positively to naturally occurring edges because they evolved in open conditions of oak savannas (Viste-Sparkman 2006).


Historically, oak habitats were widespread in South Puget Sound, comprising >40% of the region (Hanna and Dunn 1997). Currently, factors contributing to loss and degradation of prairie-oak habitats include urban, residential, and rural development; conversion to agriculture; loss of native species composition due to introduction of exotic species; fire suppression and associated encroachment by conifers; spread of exotic plant species; higher oak stand densities; conversion of oak woodland and forest to conifer stands for timber production; lack of oak recruitment; and harvest of oak trees (Hanna and Dunn 1997, Altman 2011). As a result, prairie-oak habitat was designated as a priority habitat for conservation (WDFW 2008) and is among the most threatened habitats in the Pacific Northwest (ABC 2006).

Loss and degradation of oak habitats has likely contributed to a range contraction from the Puget Lowlands (Altman 2011) and decline in numbers of this oak dependent bird beginning in the early 1900s (Bowles 1929). Lack of data on nuthatch numbers during the 1950s to 1990s, a period of great human population increase in the Puget Lowlands, restricts determining the timing and magnitude of decline for this nuthatch (Altman 2011). Strong association of the Slender-billed White-breasted Nuthatch with large diameter oaks makes it most threatened by dramatic loss and degradation of oak habitats in the Puget Lowlands (Altman et al. 2001, Altman 2011). In addition to loss of oak habitats, other factors that may have contributed to the species decline include scarcity of nesting cavities, competition from starlings for existing nest sites, insufficient oak mast to support overwintering populations, genetic drift due to small isolated populations, and unknown disease or natural disturbance (Chappell 2005). The nuthatch could be reintroduced in the Puget

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Lowlands from populations farther south (Altman 2011) if an assessment determined that threats had been addressed and a viable population could be supported by existing habitat.

3.2.5 Streaked Horned Lark (Eremophila alpestris strigata Henshaw, 1884)


The Streaked Horned Lark was listed as threatened under the Federal Endangered Species Act in 2013 (USFWS 2013). This Streaked Horned Lark is a Species of Greatest Conservation Need as recognized in Washington’s Comprehensive Wildlife Conservation Strategy.


The Streaked Horned Lark is a rare endemic subspecies found only in western Washington and Oregon (Fig. 1). It is perhaps the most distinct subspecies of the horned lark, a small common ground-dwelling passerine that prefers open grassland habitat (Beason 1995, Rogers 2000, Stinson 2005). The Streaked Horned Lark was once abundant on Puget Sound prairies, but has become increasingly rare with the decline in habitat and, in Washington, is now restricted to a few large open grassland sites. Rogers (1999) and MacLaren and Cummins (2000) conducted surveys to determine the status of Streaked Horned Larks in Washington, and visited locations on south Puget Sound prairie remnants, the San Juan Islands, northern Puget Sound sites (e.g., Skagit, Stillaguamish, Lummi Flats, Dungeness Spit), sites on the outer coast in Grays Harbor and Pacific counties, and along the lower Columbia River. No larks were detected at northern Puget Sound locations or in the San Juan Islands, and no new inland sites were found besides those already known at Fort Lewis, McChord AFB, Olympia Airport, and Shelton Airport.

Historically, Streaked Horned Larks bred from southern British Columbia, through the Puget Trough in Washington and in the Willamette and Rogue River Valleys in Oregon (Rogers 2000, Stinson 2005). The breeding range of the lark contracted over time with extirpation from former breeding sites in northern Puget trough, southern British Columbia, the Washington Coast north of Grays Harbor, and the Rogue River Valley of Oregon (Rogers 2000, Beauchesne and Cooper 2003, Stinson 2005). The Streaked Horned Lark is currently known to breed at about 13 locations in Washington: 6 inland sites (Table 1, Fig. 2), 3 coastal sites, and 4 Columbia River sites (additional Columbia River sites exist in Oregon).

The Streaked Horned Lark is by all indications very rare. Population estimates indicate that there are probably fewer than 2,000 Streaked Horned Larks remaining. Population estimates based on winter surveys produced estimates of about 500-600 in 2004-2005 (Pearson et al. 2005a). Pearson and Altman (2005) estimated about 330 birds breeding in Washington and 440 in Oregon; they cautioned that these estimates combined data from separate efforts over a time period of 8 years. Altman (2011) recently estimated a total population of 1,170-1,610. Camfield et al. (2010) reported that demographic data suggested an ongoing steep decline in the Washington population. McChord Field, which formerly had the highest number of nesting pairs has seen a marked decline (Fig. 3).

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Camfield et al. (2011) monitored Streaked Horned Lark nests on 7 sites in Washington and banded 58 adults (26 females, 32 males) and 88 juveniles. They developed a demographic model to estimate population trends and to identify the parameter and life stage that would be the most important targets for management. They reported that Streaked Horned Larks in Washington were declining rapidly and that local breeding sites were not sustainable without immigration. In addition, although there are no data on range-wide population trends for Streaked Horned Larks, territory mapping data from four sites in the Puget lowlands indicated that the number of territories had decreased 45% over 3 years from 77 territories in 2004, to 42 in 2007 (S. F. Pearson, WDFW, unpubl. data). They concluded that the highest priority for management was to increase adult survival, followed by improvement of juvenile survival and fecundity.


Horned larks forage on the ground, usually in short and sparse vegetation. Diet has not been studied in E. a. strigata, but horned larks are largely granivorous, both in winter (80-100% seeds) and in the breeding season (up to 73% seeds), while nestlings are fed insects exclusively (Beason 1995). Adults will dig up worms and insect larvae, and pry moth larvae from weed clumps to obtain food for chicks. Insects eaten include grasshoppers, beetles, and Lepidoptera larvae, and they also are adept at chasing and catching small insects (Beason 1995).

Territorial and courtship behavior. Streaked Horned Lark males begin to sing and establish territories after they arrive in Washington in the latter half of February and early March (Rogers 2000, Pearson 2003). Males sing from the ground and in flight. Ground singing functions in territorial defense and is often done from a post, rock, or dirt mound (Beason 1995). Aerial singing is part of an elaborate courtship display. Song flights last 0.5-8 minutes and are performed most frequently before nest-building, for a brief period after broods fledge, and when a nest is destroyed (Beason 1995).

Horned larks defend an ‘all purpose’ territory (Beason 1995). Territory sizes likely vary with habitat quality and lark density. Streaked Horned Lark territories in Oregon averaged 0.77 ha (1.9 ac; range 0.6-1 ha; n = 3) using the “repeat flush” territory mapping technique described by Wiens (1969) and (Altman 1999). In other subspecies, territories ranged from 0.3 – 5.1 ha (0.7-12.6 ac) (Beason 1995). Territories are defended until the last brood leaves the nest. There are no data on seasonal home ranges of broods after territories are abandoned, or on home ranges of winter flocks (Beason 1995). Bowles (1898) reported that some locations had high densities of nests, while large expanses of apparent habitat were vacant, suggesting that Streaked Horned Larks display aggregated nesting.

Nesting and brood rearing. Horned larks build a compact cup of dead grass, or other plant material that is usually placed in a depression scratched out to 5-7.5 cm (2-3 in) deep or a cavity from an upturned stone (Bowles 1900, Pickwell 1931, Campbell et al. 1997). Streaked Horned Larks have a long nesting season. Nest building in the south Puget Sound area was first observed mid-April to early May (Pearson and Hopey 2005). Clutch initiation dates vary with location; the first eggs are observed around the 1st of May (Pearson 2003, Pearson and Hopey 2004), though the early date for

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British Columbia is 5 April (Campbell et al. 1997). Bowles (1898) stated that one could confidently look for eggs at Washington locations between 1 May and the “last of July,” and perhaps earlier and later. Except at high elevations or high latitudes, horned larks typically raise 2 or more broods per season (Beason 1995). South Puget Sound birds seem to exhibit 2 peaks in clutch initiation, with the first peak from late April/early May and lasting until late May/early June; a peak of second clutches or renests after failures follows in late June to late July (Pearson and Hopey 2005). Nesting activity ended 8 August, 9 August, and 30 July in 2002, 2003, 2004, respectively (Pearson and Hopey 2004, 2005). The clutch size is most often 3; with a mean of 3.05 eggs ( 0.06 SE) for 135 clutches in Washington (Camfield et al. 2010). Clutch size may be affected by conditions, such as drought or a wet spring. Incubation lasts about 11 days, but occasionally up to 14 days during colder weather (Beason 1995).

The chicks attain 60% of the adult body weight in the first 8 days (Beason 1995). Kennedy (1913a in Jewett et al. 1953) noted horned lark chicks in eastern Washington leave the nest at 6-8 days; in British Columbia, chicks leave the nest at 7- 9 days (Campbell et al. 1997). The chicks can flutter and hop at departure, fly a few meters in a few days, and can walk and fly well by day 27 (Beason 1995). The parents provide food for a week or more after fledging. Chicks start to become independent by 3 weeks of age and are mostly independent at 4 weeks (Beason 1995).

Reproductive success. Pearson and Hopey (2005) reported that 63 of 167 (37%) active nests found on south Puget Sound study areas in 2002 - 2004 fledged at least 1 young. Overall nest success at 4 Puget lowland study sites calculated using the Mayfield method was 28%, 21%, and 28% in 2002, 2003, and 2004 (Pearson and Hopey 2005). Predation was the most frequent (69%) cause of nest failure at sites in south Puget Sound and caused 46% of failures at 2 coastal and 1 river island sites in 2004 (Pearson and Hopey 2005). Abandonment was the source of failure for 22% (23 of 106) of south Puget Sound and 46% (6 of 13) of coastal and river island nests. Some abandonment was human-related (e.g., tents erected next to nests on Gray Army Airfield). Failures directly caused by humans include 8 caused by mowing at south Puget Sound sites, and 1 that was crushed by a horse and rider on Midway Beach (Pearson and Hopey 2005). Recreational activities, including dog walking, beachcombing, vehicles, and horseback riding may increase predation and nest abandonment at coastal sites (Pearson and Hopey 2005).

In comparing the fecundity of the Streaked Horned Lark to an alpine subspecies, the pallid horned lark (E. a. articola), Camfield et al. (2010) found the replacement nest and multiple brood intervals for the Streaked Horned Lark to be almost 4 times longer than the pallid horned lark (22 vs. 6 days). This, combined with the Streaked Horned Lark’s smaller clutch size, lower hatchability of eggs, lower fledging success and high clutch depredation rates, resulted in higher annual fecundity for the pallid horned lark, despite the Streaked Horned Lark’s breeding season being over double the length of the pallid horned lark. Camfield et al. (2010) speculated that influences of anthropogenic habitat loss, habitat degradation, and increased nest predator populations on the vital rates of Streaked Horned Lark, may explain the mismatch between the authors’ predicted and observed life history strategy for Streaked Horned Lark.

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Breeding habitat. The Streaked Horned Lark nests on sparsely vegetated open habitats dominated by short grasses and forbs (Altman 1999, Rogers 2000, Pearson and Hopey 2005)and include airports, agricultural fields, sandy islands and coastal spits in Washington. Horned larks may select bare ground or short vegetation because adults normally walk rather than hop (Beason 1995). In agricultural areas in other parts of the country, horned larks often nest on bare ground, stubble fields, and pastures. Mowed fields adjacent to airport runways provide important nesting areas for Streaked Horned Larks in Washington (Rogers 2000, Pearson and Hopey 2005). When selecting territories, males on south Puget Sound sites seemed to avoid areas dominated by shrubs, perennial bunchgrasses, sod-forming perennial grasses, and non-native perennial forbs (Pearson 2003). They appear to select areas that are sparsely vegetated with short annual grasses and with a relatively high percent cover of rocks ( 9%) (Pearson and Hopey 2004, 2005).

Foraging sites. Streaked Horned Larks on Fort Lewis prairies selected foraging sites with a large percentage of bare ground (>40% of 1 m radius plots; included occasional mosses) and low vegetation (<30 cm, 12 in) (Rogers 2000). Rogers (2000) noted that larks seemed to select foraging sites that were atypical of the existing prairie landscape, but suggested that in historical prairies, “such sites would not have been hard to find.” Streaked Horned Larks in Oregon also used territories and nesting sites with a relatively high percentage of bare ground (Altman 1999). Given their selection for sparse, short vegetation and bare ground, Streaked Horned Larks may have historically been restricted to the driest parts of the south Puget Sound prairies. Larks may have selected areas where the vegetation was sparse because it burned frequently, had a poorly developed A horizon, had a high gravel/cobble content, or a combination of these factors (Pearson and Hopey 2004). In a 2004 experiment, burned plots on 13th Division Prairie received much higher use by post-breeding Streaked Horned Larks than unburned plots (Pearson and Hopey 2005).

Migration and winter habitat. Horned larks generally use the same open habitats during migration and winter, but perhaps with more frequent use of ocean beaches, dunes, and airports than during the breeding season (Beason 1995). All habitats where Streaked Horned Larks were detected in winter were large treeless/shrubless expanses with a high percentage of bare ground (Robinson and Moore 2004). Most birds were recorded on fallow ryegrass fields in the Willamette Valley and on dredged material along the lower Columbia River; smaller numbers were found on sandy Washington coastal sites (Robinson and Moore 2004, Pearson et al. 2005a).


Habitat loss, degradation, and fragmentation. Prairie habitat continues to be lost, particularly to residential development. In the south Puget Sound area, over 90% of the original grassland has been converted to other uses or succeeded to forest (Crawford and Hall 1997, Chappell et al. 2001). Olympia and Shelton Airports are planning for development of significant portions of their grasslands, which may affect nesting lark populations. As is typical of many grassland birds, horned larks seem to need rather large open areas, and habitat fragmentation is an important factor in their decline (Peterjohn and Sauer 1999, R. Rogers, pers. comm.). The smallest open area used for

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nesting by Streaked Horned Larks in the Puget lowlands is 131 ha (324 ac) (Sanderson Field/Shelton airport). The water and beaches surrounding coastal and Columbia River sites creates much larger open areas free of tall vegetation and obstructions and as a result, larks will use smaller expanses of open habitat under those conditions.

Loss of wintering habitat also is an issue. Sparsely vegetated dredged material along the lower Columbia River provides wintering habitat for Streaked Horned Larks. There are few specifics known about loss of this habitat type in Washington. Habitat is being lost at a North Portland, Multnomah County site in Oregon. Grading in preparation for development was apparently responsible for a decline from 150-200 wintering birds in 2002 and 2003 to 61 wintering birds in 2004 (Pearson and Altman 2005).

Fire suppression allows succession by native and exotic flora. Douglas-fir (Pseudotsuga menziesii) has invaded substantial portions of the historical prairies (Foster and Shaff 2003). Invasion by shrubs, tall vegetation, and turf-forming grasses would eliminate the short, open structure that larks seek for nesting and foraging. Nearly all the remaining prairie sites are degraded to some extent by exotic forbs and grasses, creating conditions that are not compatible with lark use. Pearson et al. (2005b) reported that late summer prescribed burn plots on 13th Division Prairie were selected by post-breeding adult and hatch-year larks, and by breeding birds the following spring; late summer prescribed burns created habitat conditions that were attractive to larks. Scotch broom (Cytisus scoparius) and other weedy plants are also invading some coastal (especially Damon Point) and Lower Columbia sites. Introduced beachgrasses (Ammophila spp.) reduce or eliminate unvegetated or sparsely vegetated sand used for nesting by Streaked Horned Larks at coastal sites.

Army Training on Joint-Base Lewis-McChord. Fort Lewis has generally been proactive in the conservation of prairie species, but larks are sometimes directly affected by Army training activities when they coincide with lark nesting (Pearson and Hopey 2004). Nest abandonment caused 20% of nest failures and some abandonment was likely caused by human disturbance during training activities. Military training activities may also affect horned lark nesting areas where disturbance of native vegetation leads to increases in exotic vegetation. Training activities on the Artillery Impact Area may result in a fire frequency that exceeds what is desirable for maintaining native prairie vegetation (Tveten and Fonda 1999); the potential effects on larks use is not known.

Control of Scotch Broom, Douglas-Fir and weedy forbs on Fort Lewis military bases is beneficial to larks by maintaining open prairie. The abundance and diversity of native forbs, mosses and lichens decline with disturbance. In heavily disturbed areas, mosses and lichens disappear and the soil surface is bare or covered with leaf litter (Clampitt 1993). Military training may benefit larks by maintaining lower vegetation density and higher bare ground than would exist without training activities or restoration of prairie. However, management that restores and maintains the sparse bunchgrass structure and abundant moss that existed historically may be optimal for lark nesting areas.

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Disturbance, mortality and development at airports and military airfields. Olympia Airport, Shelton Airport, Gray Army Airfield and McChord AFB contain most of the inland nesting Streaked Horned Lark population in Washington. Airports can be hazardous environments for nesting due to mowing, potential for collisions with aircraft, and special events hosted at military bases. Mowing of airports and military airfields likely benefits larks by keeping the vegetation short, but can cause mortalities to eggs, chicks, or adults during nesting unless it is timed to minimize impacts. Careful timing of mowing and adjustment of blade height can help minimize horned lark mortalities.

Gray Airfield adjusted its mowing schedule to minimize impacts to larks in 2003 and 2004. However, recently the paved area was expanded and the number of aircraft was increased and includes attack helicopters. This affected a portion of the habitat that was used by larks in recent years, and the hot downdraft produced by these aircraft may make some portion of the habitat unusable for lark nesting.

McChord AFB has not adjusted mowing schedules to minimize impacts to larks during the nesting season. Horned larks do not seem to be overly disturbed by the routine comings and goings of the large military cargo aircraft based there (S. Pearson, pers. comm.). However, McChord occasionally hosts large military training and civilian events that impact larks. The overall number of pairs detected on McChord Airfield has declined since 2004 (Anderson 2010a). Additionally, although the data have not been analyzed, anecdotal observations by surveyors indicate that there are fewer singing larks in recent than in previous survey years (Anderson 2010a).

Civilian airports. In recent years the Olympia Airport has hosted the highest number of nesting pairs of sites in Washington, and Shelton Airport has consistently hosted small numbers of birds. Olympia Airport has modified mowing schedules to minimize impacts to larks during nesting. The future loss of grassland to development at both airports may be significant.

Collisions with aircraft. Horned larks are particularly susceptible to being struck by aircraft, probably due to their affinity for the open, short-grass habitat surrounding runways. Horned larks are the most commonly reported species involved in collisions with Air Force aircraft, and represent almost 13% of all reported strikes (BASH 2009); very few horned larks were involved in bird strikes on civilian aircraft reported to the Federal Aviation Administration the difference between military and civilian aircraft is probably artifactual because Air Force personnel are required to report all bird strikes, while only 20% of bird strikes recorded at civilian airports are reported to the FAA (Cleary et al. 2003). Few are reported when little or no damage to the aircraft occurs. Dead larks have been found along the runways at McChord AFB and Gray Army Airfield (Pearson and Hopey 2005). It is not known how significant a source of mortality aircraft collisions are for the Streaked Horned Lark population in Washington, but 4 of 12 known nesting populations are at airports, and they include the sites with the highest nesting populations. Collisions may be more likely at airports with closely mowed vegetation concentrated next to runways, but less likely where mowed vegetation attracts larks to areas set back from active runways (S. Pearson, WDFW, pers. comm.). Assessments are needed to determine whether aircraft collisions are an important source of mortality of Streaked Horned Larks. Camfield et al. (2011) found that the nesting populations at airports were declining along with the coastal and Columbia River populations.

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Improving nesting habitat away from active runways may reduce collisions and improve adult survival if enough suitable habitat exists away from the runway

Management of Columbia River Islands. Pearson and Altman (2005) identified the dredged-material islands along the Columbia River as critical to persistence and recovery of this subspecies. To deepen and maintain the depth of the Columbia River shipping channel, the Army Corps of Engineers removes soil from the bottom of the shipping channel and deposits it on islands in the River. The dredged-material islands are publicly owned and larks breeding on these islands have higher reproductive success than at other breeding sites in Oregon and Washington. In addition, larks use these islands as over-wintering sites. Creation of dredged-material islands has provided nesting sites for Streaked Horned Larks and has potential to expand existing sites and create new ones. Deposition of new material at existing sites may help maintain the presence of bare ground and sparsely vegetated areas. However, depending on the timing, horned larks can be disturbed and nests destroyed if material is deposited on sites occupied during the breeding season. Center for Natural Land Management (CNLM 2011) reported encouraging results of experiments with tilling dredged-material sites to restore suitable conditions at several sites. The Army Corps of Engineers is developing a habitat management plan for these islands to minimize impacts to the population of breeding larks along the lower Columbia River.

Other human-related factors. Birds on prairie remnants within a matrix of suburbs may be subject to high rates of predation by American Crows (Corvus brachyrhynchos). Pearson (2003) observed crows depredating Streaked Horned Lark nests at airport nesting sites. Crow populations are high in urban habitats, perhaps due to a scarcity of predators and human associated food sources.

Population size and genetic health. Analysis of mitochondrial DNA indicates that Streaked Horned Larks probably have suffered a loss of genetic diversity (Drovetski et al. 2005). Diminished genetic diversity increases likelihood of populations suffering from inbreeding depression, reduced resistance to disease, and reduced adaptability to environmental change (Frankham et al. 2002). Inbreeding depression, in turn, can lead to reduced reproductive success. Streaked Horned Lark genetic health, represented by adequate genetic heterogeneity, is an important issue in populations in Washington, particularly in the Puget Trough. Anderson (2010b) reported that Streaked Horned Larks at 13th Division Prairie on Joint Base Lewis-McChord had significantly lower hatchability when compared to a guild of ground nesting birds and to Savannah Sparrows (Passerculus sandwichensis) at the site, suggesting the cause was not related to predation or other environmental factors at the site. The low hatching rate of Streaked Horned Lark eggs (44%), coupled with genetic data indicating a recent population bottleneck and low genetic diversity (Drovetski et al. 2005), suggested that inbreeding depression was playing a role in the decline of larks at 13th Division Prairie. A project was initiated in 2011 to address the issue of inbreeding and low hatching rate by moving eggs from Willamette Valley in Oregon to nests on 13th Divisions; the plan involved moving eggs from 5 lark nests in 2011, and again in 2012.

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3.2.6 Western Grey Squirrel (Sciurus griseus Ord, 1818)


In response to a petition to list the population under the federal Endangered Species Act, the USFWS conducted a status review of the Washington population of Western Gray Squirrels. They concluded that the Washington population did not represent a distinct population segment and therefore was not a listable entity, and that the population did not constitute a significant portion of the subspecies or its range (USFWS 2003). In 2004, the USFWS issued a 90-day finding on a separate petition stating that there was not substantial information to list the Washington population, the species, or any subspecies of Western Gray Squirrel (USFWS 2004). WDFW completed a state recovery plan in 2007 (Linders and Stinson 2007), and PHS Management Recommendations were recently updated (Linders et al. 2010).


Western gray squirrels range from north central Washington to the southern border of California, west to the coast in California, and east to the Nevada border (Hall 1981, Carraway and Verts 1994). In Washington, they currently occur in 3 isolated areas (Fig.1): Pierce and Thurston counties; foothills of the Cascades in Klickitat, Yakima, and Skamania counties; and in Chelan and Okanogan counties (Linders and Stinson 2007). The statewide population was estimated to total between 468 and 1,400 (Linders and Stinson 2007), but populations can fluctuate rather dramatically with disease and food supply.

Little information is available on historical population levels of the Western Gray Squirrel in Washington, but it’s generally believed that the species was more widespread. Western Gray Squirrels in the southern Puget Trough were considered uncommon during the late 1800s due to hunting intensity (Bowles 1921). Until 1933, county governments regulated hunting, and seasons were often long and rarely had bag limits. Bowles (1921) described an immense increase in Western Gray Squirrels in Pierce County, Washington, between 1896 and 1920 that he attributed to reduced hunting and an expansion of forests into Puget Sound prairies. Bowles (1921) and Couch (1926) described the species as common in the Pierce County area, and bark stripping by squirrels for food resulted in significant damage to trees. Western Gray Squirrels were frequently seen in Tacoma in 1941 (Flahaut 1941). By the late 1940s, Western Gray Squirrels had become scarce and were seldom seen across much of their Washington range (Booth 1947). Western Gray Squirrels were still present in the suburbs of Tacoma in the early 1950s, but declined with increasing development (Rodrick 1986).

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Of the 3 disjunct populations of Western Gray Squirrel in Washington, the Puget Trough population faces the greatest extinction risk (Linders and Stinson 2007). Rodrick (1986) conducted surveys in the Puget Trough in 1985–1986 and found Western Gray Squirrel sign on just 4 of 26 sites (15%); Fort Lewis appeared to harbor the only remaining squirrels in the Puget Trough. Western Gray Squirrels were found previously on adjacent private lands and in Thurston County, only 1 squirrel sighting was reported outside Fort Lewis (now Joint Base Lewis-McChord [JBLM]) since 1990 (WDFW data system; Fig.2). WDFW surveys in Thurston County on the Rainier Training Area and private lands nearby in 1996 failed to identify sign of Western Gray Squirrels during 36 hours of search effort. Available evidence suggested that the Puget Trough population might be dangerously low (Bayrakci et al. 2001).

In 2007, WDFW and the Department of Defense-JBLM, initiated a plan to augment the Western Gray Squirrel population on the Base with the goal of increasing genetic diversity and expanding the occupied area (Vander Haegen et al. 2007). From 2007–2011, a total of 83 Western Gray Squirrels were released on JBLM in Pierce County and the Rainier Training Area in Thurston County. Survival of translocated squirrels has been equivalent to that of resident animals and numerous females have produced young (Vander Haegen and Orth 2011). The JBLM augmentation project

Figure 1. Recent and historical range of Western Gray Squirrel in hydrologic units intersecting the planning area and elsewhere (inset) in the Northwest (modified from Northwest Regional Gap Analysis Project (NWGAP), Univ. of Idaho, September 2012). NWGAP methodology associates species range with an entire hydrologic unit that contains any records, thus depicting a broad generalization of occurrence rather than specific extent (Beauvais et al. 2012).

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and ecology study will be completed in 2012, but additional translocations to the Puget Trough population are planned as funds are available.


Arboreal and generally solitary in their habits, Western Gray Squirrels forage on the ground, but rarely stray far from trees. They use stick nests for resting and sleeping, and females frequently use cavity nests to birth and rear young. Tree seeds (pine, Douglas-fir, oak), green vegetation, hypogeous fungi (truffles and false truffles), and fruit and seeds of various shrubs are the main components of the Western Gray Squirrel diet.

Western Gray Squirrels mate over an extended period and lactating females have been captured from March to August in Klickitat County (Linders and Stinson 2007). Young are born after a gestation period of about 44 days and are weaned at about 10 weeks. The number of young surviving to emergence from natal dens (approx. 8 weeks of age) averaged 2.5 in Klickitat County (Vander Haegen et al. 2005).

Figure 2. Distribution of vegetation types that may contain suitable habitat (Linders and Stinson 1997), and current and historical (pre-1992) Western Gray Squirrel locations in Thurston and Pierce counties, Washington.

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Little is known about the population dynamics of Western Gray Squirrels. In general, population levels vary as a result of short-term factors including the food supply and environmental variation. Variation in the food supply, particularly mast (nuts, seeds, berries) production, has been cited as the most important factor affecting tree squirrel populations, although disease may have significant effects as well (Grinnell and Storer 1924, Carlson et al. 1982, Gurnell 1987).

Known and potential predators in the range of Western Gray Squirrels in the Puget Trough include the Red-tailed Hawk (Buteo jamaicensis), Northern Goshawk (Accipiter gentilis), Coyote (Canis latrans), Bobcat (Lynx rufus), House Cat (Felis silvestris), Barred Owl (Strix varia), Great Horned Owl (Bubo virginianus), Red Fox (Vulpes vulpes), , Domestic Dog (Canis familiaris) and Weasels (Mustela spp.) (Carraway and Verts 1994, Ryan and Carey 1995a, Bayrakçi 1999, Vander Haegen et al. 2005). Weasels likely killed several radio-collared Eastern Gray Squirrels (S. carolinensis) on Fort Lewis (Bayrakçi 1999). There are few data on the effect of predation on Western Gray Squirrel populations; predation may affect the magnitude of population fluctuations, but rarely cause them.

A study investigating potential competition between Western Gray Squirrels and non-native Eastern Gray Squirrels was initiated by a University of Washington graduate student in 2007. Preliminary data suggest that Eastern and Western Gray Squirrels occupied exclusive territories that differed in habitat structure and proximity to wetlands. This research is expected to be completed in 2012.


Western Gray Squirrels inhabit mixed forests of mature Douglas-fir (Pseudotsuga menziesii), Ponderosa Pine (Pinus ponderosa), and Oregon White Oak (Quercus garryana) , and various riparian tree species (Linders and Stinson 2007). Food supply is the most important factor regulating tree squirrel populations (Gurnell 1987), so optimal habitat will provide abundant pine and fir seeds, acorns, and hypogeous fungi. Presence of diverse other seeds and fruits, such as maples (Acer spp.), hazelnuts (Corylus cornuta), Oregon ash (Fraxinus latifolia), serviceberry (Amelanchier alnifolia), and Indian plum (Oemleria serasiformis), may help to provide a more stable food supply over time. Large diameter trees generally produce more seeds or acorns, while an interconnected canopy provides for arboreal travel and security for squirrels.

On JBLM, Western Gray Squirrel presence was positively correlated with mixed oak-conifer stands >8 ha (19.8 ac) in size that were <600 m (656 yd) from water. Squirrels favored stands containing more abundant and diverse food-bearing trees and shrubs, and mixed stands over pure oak stands (Ryan and Carey 1995a). High-use stands had significantly more basal area in Douglas-fir, more young oak trees, lower average ground cover, and more coarse woody debris. Desirable characteristics of habitat in the south Puget Sound region would include (Ryan and Carey 1995b):

• Mixed stands of Douglas-fir and oak (average dbh of Douglas-fir 19.1 inches (48.5 cm)) • Open understory with patches of shrubs. • A few scattered older oaks with cavities. • 6–10 tree and shrub species present that produce seeds or fruits eaten by Western Gray

Squirrels (including: snowberry [Symphoricarpos albus], hazelnut, Indian plum, Douglas-fir,

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Oregon White Oak, Salal [Gaultheria shallon], Serviceberry, Rosa spp., Blackberry [Rubus spp.] Red Huckleberry [Vaccinium parvifolium], Ribes spp., Bigleaf Maple [Acer macrophylum], Vine Maple [A. circinatum], Oregon Ash, Ponderosa Pine, Cascara [Rhamnus purshiana], Pacific Yew [Taxus brevifolia], Grand Fir [Abies grandis], Pacific Dogwood [Cornus nuttallii], Black Cottonwood [Populus balsamifera var. trichocarpa])

3.2.6.5 Threats /Reasons for Decline

Habitat destruction and degradation. Causes for the decline in the Puget Trough population likely include habitat loss, habitat alteration, and increased mortality related to vehicle traffic (Ryan and Carey 1995). Urban development poses a significant threat to Western Gray Squirrel habitat in the Puget Trough (Kessler 1990). Since the 19th century, oak woodlands have been subjected to logging, farming, and conversion to other land uses. Fire suppression, grazing, and removal of oak for firewood also affected the structure and quantity of these woodlands (Lang 1961, Kertis 1986, Franklin and Dyrness 1988). Fire suppression permitted Douglas-fir encroachment into oak woodlands resulting in the overtopping and death of oaks from competition and allowed Scotch broom (Cytisus scoparius) and other shrubs to invade the understory and compete with seedlings (Kertis 1986, Agee 1993). The cumulative effects of land conversion and fire suppression have caused a severe decline in oak woodlands throughout Washington (Larsen and Morgan 1998). Oaks are sensitive to surface disturbance such as grading and trenching because they have most of their roots within the top 2 feet of the soil surface (Ryan and Carey 1995a). Kessler (1990) estimated that there were about 4,130 ha (10,200 ac) of oak woodland in Thurston County in 1990. About 3,120 ha (7,700 ac) of this is on private lands (Kessler 1990, Ryan and Carey 1995a) and either exists in a matrix of suburban development where its habitat value for Western Gray Squirrel is severely compromised, or it is at risk of development.

As human populations continue to increase in the Puget Trough, development and land clearing will further reduce remaining Western Gray Squirrel habitat. A proposed southern extension of the primary runway on JBLM and an industrial park on the base could eliminate 103 ha (254) acres of oak and conifer woodlands (FHWA 2003). These military lands contain the largest tracts of publicly-owned oak woodlands in the Puget Trough region (Ryan and Carey 1995b).

Logging. Logging and land clearing may degrade Western Gray Squirrel habitat by disturbing the soil, destroying nests and potential nest sites, and fragmenting the tree canopy that squirrels use for travel and escape cover (Vander Haegen et al. 2004, Linders et al. 2010). Removal of the largest conifers reduces seeds available to squirrels, a critical food source. Soil disturbance, compaction, and reduction of canopy closure during logging affects the abundance of hypogeous fungi eaten by squirrels (Pederson et al. 1987, States and Gaud 1997). Overall, these activities may suppress squirrel populations by decreasing the food supply, reducing quality of nest sites, and increasing predation.

Along the lower Nisqually River in the south Puget Sound region, timber cutting began around 1890; by 1917, when Fort Lewis was established, most of the forests had been cut (Foster 1997). Between 1934 and 1952 the Army resumed clearcutting, so that by 1964, 90% of the forests on Fort Lewis were less than 70 years old. Most of the remaining Ponderosa Pine on Fort Lewis occurs in a

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500-ha forest that has been degraded by past management and fire suppression that allowed invasion by Douglas-fir, Scotch Broom, and exotic grasses. Portions of the area were lost to the construction of roads and training areas. Pine regeneration may be inhibited by lack of seed; Ponderosa Pines on Fort Lewis are not highly likely to bear cones until 50 cm (19.7 in) in dbh, and trees exceeding this size are uncommon (Foster 1997).

Roads. Vehicles contribute notably to Western Gray Squirrel mortality, especially when juveniles are dispersing (Ingles 1947, Gilman 1986, Verts and Carraway 1998, Weston 2005). With the continued expansion of human populations in the Pacific Northwest, road density and traffic volumes can be expected to increase across the landscape; this likely translates to an increased risk of death to squirrels on roads. Death by motor vehicle is a significant problem for the Puget Trough Western Gray Squirrel population; Ryan and Carey (1995b) reported that 16% (13 of 81) of the Western Gray Squirrels they observed died on roads; at least 4 squirrels were killed on Fort Lewis in 2005, and at least 4 in 2006 (Linders and Stinson 2007). Immature squirrels may suffer disproportionately from road-kill mortality (Gaulke and Gaulke 1984, Ryan and Carey 1995b). The actual amount of road-kill mortality may be underestimated because some squirrels are likely removed by scavengers.

Population size and isolation. Small population size and isolation is a potentially significant factor influencing the continued existence of Western Gray Squirrels in Washington. Western Gray Squirrel populations naturally fluctuate with mast production and disease. This natural variability puts smaller populations at greater risk of local extinction. The Puget Trough population is very small and cannot be expected to persist long without augmentation. An increasing number of studies indicate that goals to maintain viable populations of vertebrates need to be in the order of several thousands, rather than hundreds (Reed et al. 2003). The isolation of small populations typically results in a loss of genetic quality that may require introduction of individuals to counteract loss of fitness (Lacy 1987, Reed and Frankham 2003). Lack of genetic vigor may reduce the viability of populations and their ability to expand into adjacent habitat. Inbreeding depression has contributed to declines and extinctions of several species in the wild (Brook et al. 2002). Genetic health, represented by adequate genetic heterogeneity, may be an important issue in Western Gray Squirrel populations in Washington, particularly in the Puget Trough. Warheit (2003) reported that the Washington populations of Western Gray Squirrel showed reduced genetic diversity at all measures compared to populations in Oregon and California. Warheit (2003) noted that the reduction in genetic diversity may be a function of genetic drift resulting from the small population sizes in Washington.

Disease. Episodic outbreaks of disease, particularly Notoedric mange, have had a significant negative effect on populations of Western Grays Squirrels in Washington since at least the early 1930s (Carlson et al. 1982, Cornish et al. 2001). Local residents believe that Western Gray Squirrel populations in Klickitat County have never recovered to the numbers present prior to disease outbreaks in the 1930s and 1940s (Linders and Stinson 2007). Mange has not been reported in the Western Gray Squirrels in the Puget Trough, but other diseases may affect populations, including tularemia (Vander Haegen and Orth 2011). West Nile Virus has been confirmed in Western Gray

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Squirrels in California; it is not known if it will cause significant mortality in healthy Western Gray Squirrel populations.

3.3 Plants

3.3.1 Golden Paintbrush (Castilleja levisecta Greenm.)


Historically, golden paintbrush has been reported from more than 30 sites from British Columbia south through the Puget Trough of Washington to the Willamette Valley of Oregon. Throughout its range, populations have been extirpated due to agricultural, residential and commercial development. Fire suppression is also thought to have contributed to the decline of the species, as trees and shrubs invade its grassland habitat. The last known observation of the species in Oregon was in 1938 in Linn County (Consortium of Pacific Northwest Herbaria 2014). In British Columbia, golden paintbrush has apparently been extirpated from all but two sites. There are nine known extant natural occurrences in Washington. NOTE: As part of recovery efforts for the species, golden paintbrush has been planted at a number of sites from which it was not historically known. These planting are not included in references within this species profile to the number of occurrences of the species; all references in this profile are to naturally occurring populations.

The species was listed under the federal Endangered Species Act as threatened in 1997. It has a NatureServe global conservation status rank of G1 (critically imperiled) (NatureServe 2014). It is ranked S1 (critically imperiled) in both British Columbia and Washington. It is ranked SH (possibly extirpated) in Oregon. In Canada, golden paintbrush is included as Endangered on Schedule 1 of the Species at Risk Act (Species At Risk Public Registry 2014).

Golden paintbrush occurs in two ecosystem types that have a NatureServe conservation status rank of G1(critically imperiled): (1) Roemer’s fescue – white-top aster (Festuca roemeri – Sericocarpus rigidus) and (2) red fescue - (Oregon gumweed - great camas) (Festuca rubra (Grindelia stricta – Camassia leichtlinii).


Rangewide: Golden paintbrush was historically known from at least eleven sites in British Columbia, eight western Washington counties, and three counties in the Willamette Valley in Oregon. Today, there are just eleven known occurrences of golden paintbrush rangewide: two in British Columbia and nine in Washington. The last record of it occurring naturally in Oregon is from a specimen collected in 1938 from Linn County (Consortium of Pacific Northwest Herbaria 2014). The two extant occurrences in British Columbia reportedly have good viability; both are within Ecological Reserves, so they receive administrative protection. However, the species has been extirpated from nine of eleven historically known 11 sites, so the decline over the last 150 years has been significant. One occurrence within Victoria was known as recently as 1995, but has since been extirpated (B.C. Conservation Data Centre 2014a).

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Washington: Golden paintbrush was historically known from eight western Washington counties. It has apparently been extirpated from Pierce, Clark, King, Jefferson and Skagit counties. One occurrence (Davis Point in San Juan County) has been extirpated within the last 20 years. For the remaining extant sites in Thurston, Island and San Juan counties (Figure 1), the overall trend since population estimates began to be recorded in the early 1980s has been downward, although it has varied from year-to-year and from site-to-site (see Table 1). Drawing conclusions from the available data is also challenging, since counting/estimation methods have not been consistent. A notable exception to the downward trend has been the occurrence at Fort Casey State Park, where it appears that management focused on improving the habitat for golden paintbrush has been successful; clearing/mowing of areas that had been invaded by shrubs has resulted in a steady increase in the population over the last 10 years (Arnett 2011). Other populations have also shown at least a short term positive response to management actions, such as mowing or burning (USFWS 2007).

Figure 1. Distribution of extant occurrences of golden paintbrush in Washington (WNHP 2014b).

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Thurston County: There is one known, naturally occurring, extant occurrence of golden paintbrush in Thurston County (Figure 2). The Rocky Prairie Natural Area Preserve population has fluctuated over the last 15 years, but has remained within a range of approximately 6,000 to 9,500 individuals during that time. The most recent census counted 6,183 individuals. This is significantly less than an estimate of more than 15,000 from 1983, but it is not clear that comparable methods (i.e., comparable to those of today) were used (Arnett 2011).

TABLE 1.

Table 1. Trends for Washington Golden Paintbrush Occurrences (USFWS 2007) Site Name County Ownership Trend in 2005 Rocky Prairie Thurston State Stable/increasing Naas/Admiralty Inlet Island Private/State Increasing in the short term Fort Casey State Park Island State Increasing in the short term West Beach Island Private Stable? Forbes Point Island U.S. Navy Declining Ebey’s Landing Island Private Stable? False Bay San Juan Private Portions declining, others stable Long Island San Juan Private Unknown San Juan San Juan Private Unknown

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Figure 2. Known extant occurrence of golden paintbrush in Thurston County, Washington (WNHP 2014b).


Golden paintbrush is a short-lived perennial herbaceous plant; individual plants generally do not live longer than 5-6 years. It is thought to reproduce exclusively by seed; no vegetative spread has been reported. Shoots emerge from the ground and begin to grow in March. Flowering occurs primarily in April and May and continues into June. However, observations of flowering have occurred as early as February and as late as November. Fruits on earlier flowering individuals begin to form in May and June, and mature through mid-July. Plants generally are in senescence by mid-July. The capsules generally remain unopened on the senescent stems until late summer or early fall. Seeds are gradually shed throughout the fall. The seeds lack any adaptations to assist in long-distance dispersal; they are light and are most likely dispersed limited distances as wind shakes the dried, open capsules.

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Although Wentworth (1994) found that fruits can be produced in the absence of pollinator visitation, in a study of the breeding system of golden paintbrush, Kaye and Lawrence (2003) found that the species appears to require out-crossing for reliable seed set due to barriers to self-fertilization. Higher seed set was obtained between individuals from separate populations. They concluded that the vast majority of seeds are the result of insect-mediated crosses between different individuals.

According to the B.C. Conservation Data Centre (2014b), several insects have been observed visiting golden paintbrush flowers, including Bumblebees (Bombus californicus), a plume moth (Occidryas editha), a species of parasitic wasp, and others. Evans et al. (1984) also observed bumblebees (B. californicus) visiting flowers of golden paintbrush with pollen on their heads as they exited the inflorescence.

Golden paintbrush is a facultative parasite on the roots of other plant species. Its roots penetrate the roots of adjacent plants and thereby obtain water, nutrients and carbohydrates from these hosts. According to the British Columbia Conservation Data Centre, individuals grow more robustly in association with a host species, particularly with legumes (B.C. Conservation Data Centre 2014b). Wentworth (2000) demonstrated that in a greenhouse setting, golden paintbrush could grow to reproductive maturity in the absence of any host. Other researchers have demonstrated that both Roemer’s fescue (Festuca roemeri) and wooly sunflower (Eriophyllum lanatum) can be a host species (Kaye 2001, Lawrence and Kaye 2008, Sprenger 2008, Pearson and Dunwiddie 2006). Research also suggests that plantings are more successful when individuals are grown with a host (Pearson and Dunwiddie 2006).

Wentworth (1994) found that individual plants sometimes regressed from a larger size class to a smaller one. She suggested that this ability may provide individuals with the ability to survive in years in which resources are limited.


Historically, golden paintbrush occurred in open grassland habitats from the Williamette Valley to the southern end of Vancouver Island. At present, it is restricted to open grasslands at low elevations in the Puget Sound region, generally on glacial outwash or depositional material. Slopes can be flat to steep. Golden paintbrush does not tolerate a closed canopy. Historically, fire played an important role in the maintenance of its open prairie habitat.

Chappell and Caplow (2004) detailed site characteristics of the extant occurrences of golden paintbrush, including descriptions of both abiotic (physical) and biotic (primarily vegetation) parameters of each site. The most commonly associated species, across the full range of sites, include Nootka rose (Rosa nutkana), trailing blackberry (Rubus ursinus), red fescue (Festuca rubra), orchard grass (Dactylis glomerata), Pacific wood-rush (Luzula comosa), Kentucky bluegrass (Poa pratensis), yarrow (Achillea millefolium), field chickweed (Cerastium arvense), hairy cat’s-ear (Hypochaeris radicata), English plantain (Plantago lanceolata), bracken fern (Pteridium aquilinum),

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sheep sorrel (Rumex acetosella), Pacific sanicle (Sanicula crassicaulis), hairy vetch (Vicia hirsuta) and common vetch (Vicia sativa).

Chappell (2006) identified two plant associations in the Puget lowlands in which golden paintbrush occurs: (1) Roemer’s fescue – white-top aster (Festuca roemeri – Sericocarpus rigidus) and (2) red fescue - (Oregon gumweed - great camas) (Festuca rubra (Grindelia stricta – Camassia leichtlinii). Both of these plant associations have a NatureServe conservation status rank of G1S1 (critically imperiled).

The red fescue - (Oregon gumweed - great camas) association occurs in the northern portion of the species’ range in Washington on shallow soils over bedrock and on steep glacial bluffs. Sites are dominated by red fescue, with Oregon gumweed or great camas present. These sites are dry, occurring within the rain shadow of the Olympic Mountains. Douglas-fir, snowberry, and Nootka rose can encroach on this association and lead to its loss through successional change.

The Roemer’s fescue – white-top aster association represents most of what remains of native prairies in the southern Puget Sound area, including in Pierce and Thurston counties. These sites are dominated by Roemer’s fescue; white-top aster, houndstongue hawkweed, prairie lupine, Idaho blue-eyed grass, or sickle-keeled lupine are usually present. It occurs on level or mounded topography on glacial outwash. The soils are typically gravelly sandy loam in texture, deep and excessively drained. In the absence of fire, Douglas-fir commonly invades these open grasslands.

Thurston County: The following is adapted from Chappell and Caplow (2004): The Rocky Prairie site has mounded topography with a swale running through the site. Golden paintbrush occurs on all aspects. The intermound areas are gravelly; the mounds are much less gravelly and have deeper A horizons. Rocky Prairie soils are mapped as Spanaway-Nisqually complex (Pringle 1990). Roemer’s fescue (Festuca roemeri) is dominant or co-dominant throughout the site. Long-stolon sedge (Carex inops) and sweet vernalgrass (Anthoxanthum odoratum) are also common throughout the site. Oxeye daisy (Leucanthemum vulgare) and bracken fern (Pteridium aquilinum) are the most abundant non-graminoid herbaceous species. Sickle-keeled lupine is prominent within those areas with the highest density of golden paintbrush. Other frequent forbs include yarrow (Achillea millefolium), camas (Camassia quamash), wooly sunflower (Eriophyllum lanatum), strawberry (Fragaria virginiana), houndstongue hawkweed (Hieracium cynoglossoides), cut-leaf microseris (Microseris laciniata), graceful cinquefoil (Potentilla gracilis), western buttercup (Ranunculus occidentalis), Canadian goldenrod (Solidago canadensis), and Missouri goldenrod (Solidago missouriensis). Snowberry (Symphoricarpos albus) occurs locally and appears to be increasing. Moss cover, provided primarily by Dicranum scoparium, is consistently high. Non-natives present include common shepherd’s-cress (Teesdalia nudicaulis), common St. John’s-wort (Hypericum perforatum) and hairy cat’s-ear (Hypochaeris radicata).

3.3.1.5 Threats and Reasons for Decline

Rangewide: The decline of golden paintbrush has been attributed primarily to loss of prairie and grassland habitat resulting from conversion to agricultural, residential and commercial land uses

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(USFWS 2000). In Victoria, B.C., urbanization has resulted in the loss of several historical occurrences (B.C. Conservation Data Centre 2014). In Washington, at least one occurrence (Alki Point) was likely lost due to development. Development continues to pose a threat throughout the species historical range. Although no occupied sites have been directly lost to development in recent years, development immediately adjacent to sites has further isolated them from surrounding natural habitat.

Habitat modification due to succession in the absence of fire has also reduced the extent of grassland habitat throughout the range of the species, as woody species, including natives (such as Douglas-fir, Nootka rose and snowberry) and non-natives (such as Scotch broom), replace native forbs and grasses (NatureServe 2014; USFWS 2000). Prescribed fire has had positive results, at least in the short term, at both Rocky Prairie (Thurston County) and Forbes Point (Island County). However, accidental fires at Ebey’s Landing (Island County) during July of 2002 and 2007, when there were relatively heavy fuel loads and prior to seeds being fully developed, resulted in a significant decline in the population. Those fires also resulted in a significant shift to the site being dominated by non-native species (USFWS 2007).

Livestock and rabbit grazing have also been identified as a possible factor in the decline of populations (Sheehan and Sprague 1984, Gamon 1995). Grazing likely contributed directly (through herbivory and the resultant reduction in seed production) and indirectly (through a gradual change to sites being dominated by non-native grasses) to the extirpation of a number of historically known populations. More recently, herbivory by deer, rabbits and voles has been serious at two sites on Whidbey Island (USFWS 2007).

Recreational activities may have contributed to the decline, and ultimate loss, of populations within Victoria, B.C. (B.C. Conservation Data Centre 2014a) and at Deception Pass State Park in Skagit County, Washington. Soil compaction and direct trampling of individuals are thought to have contributed to the decline of these occurrences.

The two extant sites in British Columbia are within 1 to 5 meters of current sea level. A single catastrophic event, such as a major storm, oil spill or tsunami could potentially have a significant impact on these sites (B.C. Conservation Data Centre 2014a). Sea level rise as a result of global climate change could also impact these sites.

Thurston County: The primary threats to golden paintbrush at Rocky Prairie NAP (the lone extant Thurston County occurrence) are invasion of the grassland by trees and shrubs as a result of succession in the absence of fire, and competition from invasive, non-native species. Development of lands adjacent to Rocky Prairie will also further isolate the site from natural ecological processes which, over time, may have a negative effect on golden paintbrush.

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3.4 Prairie and Oak Habitats

The prairies and oak woodlands of the southern Puget Sound landscape are ecologically diverse, yet the diversity of plant community types was not recognized until Chappell (2006) developed a classification of upland plant associations in the Puget lowlands. Overall, more than 50 plant associations have been identified within the Puget lowlands, the majority of which are either forested ecosystems or do not occur within the prairie and oak woodland mosaic of the southern Puget Sound landscape.

Chappell’s classification is based primarily on plant species composition and physiognomy, and secondarily on environmental factors. His classification adheres to international and national standards for ecosystems classification (Federal Geographic Data Committee 1997, Grossman et al. 1998, Jennings et al. 2003). The ecosystem units in his classification are plant associations; they refer to existing vegetation rather than potential vegetation types.

3.4.1 Conservation Status and Threats

Most prairie and oak habitats and plant associations are considered imperiled or critically imperiled at either the state or global level, or both. Several of these plant associations also have rare species associated with them. Conservation of these plant associations will also contribute to the conservation of the rare and declining species that occur within them.

All of these associations are threatened to some extent by the cessation of the Native American practice of burning the prairies and woodlands. Without fire, the grasslands undergo succession to woodlands as trees and shrubs invade and become established, and the woodlands undergo succession to forests dominated by Douglas-fir. The grasslands and the more open woodland associations are also subject to Scot’s broom invasion. Not only does Scot’s broom directly out-compete native species, but by fixing nitrogen, it also creates conditions favorable to a different suite of species. The grasslands and more open woodlands are also subject to an increase in non-native grasses, often the result of past (or on-going) grazing practices.

Added to this mix of threats is the continuing pressure to develop land for residential, agricultural, commercial and industrial purposes. Most occurrences of the associations in Table 1 are now reduced in size, compromised in terms of their ecological condition, and isolated in the landscape as development has chipped away at their original extent. The result is that the remaining occurrences of these associations will take active management to retain them as features of the prairie and oak woodlands landscape.

3.4.2 Prairies

Roemer’s fescue – white-top aster: This plant association has a NatureServe conservation status rank of G1S1 (NatureServe 2014; WNHP 2014b), indicating that it is considered critically imperiled both within Washington and globally. According to Chappell (2006), most of what remains of native prairies in the southern Puget Sound region is this plant association. The Natural Heritage Program has 17 occurrences

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in its database (WNHP 2014a), of which eight are in Thurston County (see Table 1). More than half of the occurrences (9 of 17) are located on Joint Base Lewis McChord (JBLM), two are within Natural Area Preserves managed by the WA DNR (Mima Mounds and Rocky Prairie), one is managed by the WDFW (Scatter Creek Wildlife Area), and one is managed by Thurston County (Glacial Heritage).

A comparison of pre-2000 aerial photographs with aerial photographs from 2013 reveals that three areas where this association occurs on private land have undergone significant change. Two of these have undergone significant development and have likely been extirpated as a result: Cedarville (Grays Harbor County) and Nutmeg Prairie (Thurston County). The third, Roy Prairie (Thurston County), appears from the aerial photograph to be dominated by Scot’s broom, and whether it is even recognizable as this association is doubtful.

A number of rare plant and animal species are known to occur within this plant association, including six species listed under the federal Endangered Species Act: Taylor’s checkerspot (Euphydryas editha taylori) is listed as Endangered; Tenino pocket gopher (T. mazama tumuli), Yelm pocket gopher (T. mazama yelmensis), Roy Prairie pocket gopher (T. mazama glacialis), Olympia pocket gopher (T. mazama pugetensis), and golden paintbrush (Castilleja levisecta) are all listed as Threatened. Additionally, there are four animal species and two plant species of state-level concern. The Washington Department of Fish and Wildlife has listed Mardon skipper (Polites mardon) as Endangered, while Puget blue (Plebejus icarioides blcakmorei), valley silverspot (Speyeria zerene bremnerii), and Oregon vesper sparrow (Pooecetes gramineus affinis) are candidates for state listing (WDFW 2014). Rose checkermallow (Sidalcea virgata) and white-top aster (Sericocarpus rigidus) are considered Endangered and Sensitive, respectively, by the Natural Heritage Program (WNHP 2014b).

Table 1. Roemer’s fescue – white-top aster plant association occurrences. Site Name County Estimated Viability* Owner/Manager Mima Mounds NAP Thurston B WA DNR Rocky Prairie NAP Thurston B WA DNR Scatter Creek South Thurston BC WDFW Nutmeg Prairie Thurston BC Private? Glacial Heritage Thurston BC Thurston County Johnson Prairie Thurston BC JBLM Upper Weir Prairie Thurston BC JBLM Lower Weir Prairie Thurston C JBLM

* Estimated Viability Categories: A = Excellent, B = Good, C = Fair, D = Poor, E = extant (occurrence extant, but insufficient information to estimate viability), X = extirpated, H = historical (no recent information on the occurrence), and Blank = data field left blank (most of these should be E).

Roemer’s fescue – (field chickweed – prairie Junegrass): This plant association has a NatureServe conservation status rank of G2S1 (NatureServe 2014; WNHP 2014b), indicating that it is considered to be critically imperiled in Washington and imperiled globally. According to Chappell (2006), this plant association is the most frequently occurring native grassy bald association in the Puget lowlands. There are 21 occurrences of this association in the Natural Heritage Program database (WNHP 2014a) (Table 3).

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It occurs mostly in the northern portion of Puget Sound, but does occur as far south as the Bald Hill area in southeastern Thurston County. This plant association is represented in six DNR-managed natural areas, several state parks, national forest lands, and lands managed by local governments. All or portions of three occurrences are privately owned. Eight of the 21 occurrences have an estimated viability rating of good to excellent. No extirpated occurrences are included in the Natural Heritage database; however, it should be noted that Natural Heritage ecologists have traditionally focused attention on areas that are in good ecological condition, resulting in poor condition sites being under-represented in the database.

Rare species found within this plant association include Taylor’s checkerspot (Euphydryas editha taylori), a butterfly that is federally listed as an Endangered species (USFWS 2014), meconella (Meconella oregana), an annual plant species considered Endangered by the Natural Heritage Program, and common bluecup (Githopsis specularioides), another annual plant species considered Sensitive by the Natural Heritage Program (WNHP 2014b).

Table 3. Roemer’s fescue-field chickweed – prairie Junegrass plant association occurrences Site Name County Estimated Viability* Ownership Hamma Hamma Balds Mason A DNR (NAP) / USFS Burrows Island Skagit AB State Parks Mount Pickett San Juan B State Parks Burnt Hill Clallam B DNR Bald Hill Thurston B DNR (NAP) / Private Aldrich Butte Skamania B USFS McGlinn Island Skagit B Tribal Cypress Island NAP Skagit B DNR (NAP) Goose Rock Island BC State Parks Cypress Island South Skagit BC DNR (NRCA) Fidalgo Head Skagit BC Local gov’t Bowman Hill Skagit BC State Parks Hope Island Skagit BC State Parks Sugarloaf Skagit C Community Forest Pass Island Skagit C State Parks Hat Island Skagit C DNR (NRCA) Green Hill Thurston C Private Eden Valley Clallam C DNR Port Townsend Prairie Jefferson CD Local gov’t Smith Prairie Island CD Private Cypress Island South (2)

Skagit CD DNR (NAP)

* Estimated Viability Categories: A = Excellent, B = Good, C = Fair, D = Poor, E = extant (occurrence extant, but insufficient information to estimate viability), X = extirpated, H = historical (no recent information on the occurrence), and Blank = data field left blank (most of these should be E).

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Roemer’s fescue – rosy plectritis: This plant association has a NatureServe conservation status rank of GNRS1 (NatureServe 2014; WNHP 2014b), indicating that it is considered critically imperiled in Washington but that its global status has not yet been assessed. This association occurs on grassy balds. It occurs infrequently from the San Juan islands south to Clark County in Washington. Although the Natural Heritage Program has no occurrence records in the database (WNHP 2014a), Chappell (2006) collected data on a total of ten plots for this association in Clallam, Thurston, Mason, Skagit and Clark counties (Table 4). The ten plots were distributed among six sites, two of which are in the southeastern portion of Thurston County: Bald Hill and nearby Green Hill. Although the precise locations of the plots were not determined for this report, it is likely that they occur on a mix of federal, state, local government, tribal and private lands.

Rare species found within this plant association include Taylor’s checkerspot (Euphydryas editha taylori), a butterfly that is federally listed as an Endangered species (USFWS 2014) and common bluecup (Githopsis specularioides), considered Sensitive by the Natural Heritage Program (WNHP 2014b).

Table 4. Roemer’s fescue – rosy plectritis plant association: plot locations Bald Hill Thurston DNR Green Hill Thurston Private

California danthonia – wooly sunflower: This plant association has a NatureServe conservation status rank of GNRS1 (NatureServe 2014; WNHP 2014b), meaning that it is considered critically imperiled in Washington but that its global status has not yet been assessed. This association occurs on grassy balds; it occurs from southern British Columbia south to the western end of the Columbia River Gorge in Skamania County. Chappell (2006) describes it as being closely related to the Roemer’s fescue – field chickweed – prairie Junegrass association (see above), although it is dominated or co-dominated by california danthonia. Chappell identified seven plots as being this association (Table 5), although there are no occurrences in the Natural Heritage database (WNHP 2014a). The only plot location within Thurston County is in the Bald Hill area.

No rare species are known to occur within this association in Washington.

Table 5. California danthonia – wooly sunflower: plot locations Site Name County Ownership Bald Hill Thurston

3.4.3 Oak woodlands

Oregon white oak / long-stolon sedge – (common camas): This plant association has a NatureServe conservation status rank of G1S1 (NatureServe 2014; WNHP 2014b), indicating that it is considered critically imperiled both within Washington and globally. There are only five occurrences of this plant association in the Natural Heritage Program database (WNHP 2014a)(Table 7). The two best condition

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occurrences are both in Thurston County: one is within Bald Hill NAP (managed by DNR) and one is on JBLM.

Oregon white oak – (Oregon ash)/ snowberry: This plant association has a NatureServe conservation status rank of G2S1S2 (NatureServe 2014; WNHP 2014b), indicating that it is considered to be between critically imperiled and imperiled in Washington and considered imperiled globally. This association is limited to the southern half of the Puget lowlands and in the Willamette and Umpqua valleys in Oregon. Most locations are small and not of very high quality. There are only five occurrences included in the Natural Heritage database, three of which are in Thurston County (WNHP 2014a). The occurrence within Scatter Creek Wildlife Area has the best estimated viability; each of the components (size, condition and landscape context) of estimated viability were rated Good for this site (WNHP 2014a). The occurrence at Grand Mound is considered Fair; although it is intermediate in size, it has been influenced by past grazing and fire suppression. The Glacial Heritage occurrence is considered Fair-to-Poor because of changes resulting from fire suppression, the invasion of non-native species, and impacts from past grazing. Four of the five occurrences are on lands that are, to some degree, being managed for their conservation values. Oregon white oak / snowberry / long-stolon sedge: This plant association has a NatureServe conservation status rank of G2S2 (NatureServe 2014; WNHP 2014b), indicating that it is considered imperiled both within Washington and globally. This association essentially occurs throughout the range of Oregon white-oak within the Puget lowlands from San Juan County southward to Clark County. Despite its fairly wide geographic range, there are only five occurrences in the Natural Heritage Program database (WNHP 2014a), two of which are in Pierce County and one is in Thurston County. All three of these occurrences are within JBLM. The Island County occurrence is within Naval Air Station Whidbey Island and development for housing appears to have decreased its size considerably.

Oregon white oak – Douglas-fir / snowberry / sword fern: This plant association has a NatureServe conservation status rank of G4S3 (NatureServe 2014; WNHP 2014b), indicating that it is considered vulnerable in Washington but apparently secure globally. This association occurs throughout the range of oak within the Puget lowlands from British Columbia southward to the Willamette Valley in Oregon. This association is an intermediate successional stage between oak-dominated communities and various Douglas-fir forest communities. Douglas-fir, in the absence of disturbance or management intervention, is expected to increase over time and out-compete Oregon white oak. This association is more common on the landscape today than it would have been historically (Chappell 2006).

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addendum a. species and habitat descriptions · lomatium utriculatum) and western buttercup...

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