50226 Federal Register / Vol. 73, No. 166 / Tuesday, August 26, 2008 / Rules and Regulations 

of the June 25, 2008, final rule goes into 

effect on August 25, 2008, as scheduled. 

We will place the petitions we have 

received into the docket, and we will 

consider the arguments made in these 

petitions about the content of section 

40.67(b) along with other comments that 

we receive. On the basis of the 

comments we receive and any other 

information available to the Department, 

the Department will reconsider section 

40.67(b) and may retain, eliminate, or 

modify it. 

Because this action and the decision 

not to take similar action with respect 

to section 40.67(i) also completely 

respond to the parallel petitions to the 

Federal Railroad Administration (FRA) 

by some of the same parties, which raise 

the same issues about the same 

provisions of part 40, FRA is not taking 

any separate action on the petitions 

concerning the implementation of the 

amendments to 40.67 in the railroad 

industry. 

Issued this 21st day of August, 2008, at 

Washington, DC. 

Jim Swart, 

Director, Office of Drug and Alcohol Policy 

and Compliance. 

[FR Doc. E8–19816 Filed 8–22–08; 11:15 am] 

BILLING CODE 4910–9X–P 

DEPARTMENT OF THE INTERIOR 

Fish and Wildlife Service 

50 CFR Part 17 

[FWS–R5–ES–2008–0005; 92220–1113–

0000–C6] 

RIN 1018–AT37 

Endangered and Threatened Wildlife 

and Plants; Final Rule Removing the 

Virginia Northern Flying Squirrel 

(Glaucomys sabrinus fuscus) From the 

Federal List of Endangered and 

Threatened Wildlife 

AGENCY: Fish and Wildlife Service, 

Interior. 

ACTION: Final rule. 

SUMMARY: We, the U.S. Fish and 

Wildlife Service (Service), hereby 

remove the Virginia northern flying 

squirrel (Glaucomys sabrinus fuscus), 

now more commonly known as the 

West Virginia northern flying squirrel 

(WVNFS), from the List of Threatened 

and Endangered Wildlife due to 

recovery. This action is based on a 

review of the best available scientific 

and commercial data, which indicate 

that the subspecies is no longer 

endangered or threatened with 

extinction, or likely to become so within 

the foreseeable future. Habitat 

regeneration and recovery actions have 

resulted in a reduction in the threats, 

which has led to: (1) A significant 

increase in the number of known 

WVNFS captures and distinct capture 

locations; (2) verification of multiplegeneration reproduction and persistence 

throughout the range; (3) proven 

WVNFS resiliency; and (4) substantial 

improvement and continued expansion 

of suitable habitat rangewide. 

DATES: This rule becomes effective 

September 25, 2008. 

ADDRESSES: Comments and materials we 

received, as well as supporting 

documentation used in preparation of 

this final rule, are available for 

inspection, by appointment, during 

normal business hours, at our West 

Virginia Field Office, 694 Beverly Pike, 

Elkins, West Virginia 26241. Call (304) 

636–6586 to make arrangements. 

FOR FURTHER INFORMATION CONTACT: 

Diane Lynch, Regional Listing 

Coordinator, Northeast Regional Office, 

300 Westgate Center, Hadley, MA 01035 

(telephone: 413–253–8628); or Tom 

Chapman, Field Office Supervisor, or 

Laura Hill, Assistant Field Supervisor, 

West Virginia Field Office (see 

ADDRESSES). 

SUPPLEMENTARY INFORMATION: 

Background 

The northern flying squirrel, 

Glaucomys sabrinus, consists of 25 

subspecies, including the Virginia 

northern flying squirrel, G. s. fuscus. 

Miller (1936, p. 143) first described G. 

s. fuscus, based on specimens collected 

in the Appalachian Mountains of 

eastern West Virginia. The Virginia 

northern flying squirrel was listed as 

endangered under the Endangered 

Species Act (Act) of 1973, as amended 

(16 U.S.C. 1531 et seq.) effective on July 

31, 1985 (Service 1985 (50 FR 26999)). 

However, it was subsequently 

determined that a more suitable 

common name for G. s. fuscus is the 

West Virginia northern flying squirrel, 

due to the majority of the subspecies’

range occurring in West Virginia; thus, 

we refer to G. s. fuscus as West Virginia 

northern flying squirrel (WVNFS) 

throughout the rest of this document. 

Information about the WVNFS’ life 

history can be found in our final listing 

rule (50 FR 26999), the Appalachian 

Northern Flying Squirrels Recovery Plan 

(Service 1990, pp. 1–11), and the 

WVNFS 5-year review (Service 2006a, 

pp. 6–10). 

Previous Federal Actions 

On December 19, 2006, we published 

a proposed rule to delist the WVNFS (71 

FR 75924). Additional information 

regarding previous Federal actions for 

the WVNFS can be obtained by 

consulting the subspecies’ regulatory 

profile found at: http://ecos.fws.gov/ 

speciesProfile/ 

SpeciesReport.do?spcode=A09R. 

Recovery 

In 1990, the original recovery plan 

was approved, and at the time, the 

recovery criteria as they apply to the 

WVNFS were deemed objective, 

measurable, and adequate (Service 1990, 

p. 19). The original recovery criteria 

were not specifically reviewed or 

updated in the 2001 recovery plan 

amendment (Service 2001, pp. 1–6). 

Instead, the focus of the 2001 

amendment was an update to Appendix 

A, Guidelines for Habitat Identification 

and Management for the WVNFS. 

Implementation of the amended 

Appendix A guidelines by the 

Monongahela National Forest (MNF) 

effectively abated the main threat to the 

squirrel (i.e., habitat loss from timber 

management) throughout the majority of 

its range, by eliminating adverse 

impacts on all suitable habitat on the 

MNF without having to prove WVNFS 

presence (Service 2001, pp. 1–6; Service 

2006a, pp. 3–4). 

Recovery plans are not regulatory 

documents and are instead intended to 

provide guidance to the Service, States, 

and other partners on methods of 

minimizing threats to listed species and 

on criteria that may be used to 

determine when recovery is achieved. 

There are many paths to accomplishing 

recovery of a species, and recovery may 

be achieved without all criteria being 

fully met. For example, one or more 

criteria may have been exceeded while 

other criteria may not have been 

accomplished. In that instance, the 

Service may judge that, overall, the 

threats have been minimized 

sufficiently and the species is robust 

enough to reclassify the species from 

endangered to threatened or to delist the 

species. In other cases, recovery 

opportunities may have been recognized 

that were not known at the time the 

recovery plan was finalized. These 

opportunities may be used instead of 

methods identified in the recovery plan. 

Likewise, information on the species 

may be learned that was not known at 

the time the recovery plan was 

finalized. This new information may 

change the extent to which criteria need 

to be met for recognizing recovery of the 

species. Overall, recovery of species is 

a dynamic process requiring adaptive 

management, and judging the degree of 

recovery of a species is also an adaptive 

management process that may, or may 


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not, fully follow the guidance provided 

in a recovery plan. 

In the case of the WVNFS, new 

information on the subspecies has been 

learned that was not known at the time 

the recovery plan and the amendment 

were finalized. This new information 

includes habitat modeling efforts 

completed in 2006, completion of a 

forest plan amendment in 2006 with 

substantial provisions for protection of 

WVNFS and its habitat, our compilation 

in 2005 of the 20+ years of survey data, 

and our re-analysis of WVNFS 

persistence and geographic distribution 

based upon them. This new information 

changes the extent to which two of the 

four Recovery Plan criteria need to be 

met for recognizing recovery of the 

subspecies. Further details related to 

each recovery criterion are available in 

the Service-prepared document Analysis 

of Recovery Criteria for the West 

Virginia Northern Flying Squirrel 

(Service 2007a, pp. 1–16). An 

attachment to this document, ‘‘Table 3, 

Land use designations, restrictions, and 

primary management emphases in 

WVNFS habitat on the MNF,’’ provides 

supplementary information for 

downlisting criterion number 3. Based 

on our analysis of the best available 

data, we believe that the intents of the 

original recovery criteria have been met. 

In conjunction with the analysis of 

the recovery criteria, we analyzed the 

threats to the WVNFS under the 

framework of the five factors established 

in the Act. This analysis of the threats 

was based in part on the most recent 5- 

year review of the subspecies completed 

in 2006 (Service 2006a, pp. 1–20). This 

is available at http://www.fws.gov/ 

northeast/pdf/flysqrev.pdf. A further 

detailed discussion of the five factors is 

contained in the Summary of Factors 

Affecting the Species section of this rule 

below. 

Summary of Public Comments 

In our proposed rule (71 FR 75924), 

we requested that all interested parties 

submit information, data, and comments 

concerning: (1) Biological, commercial, 

trade, or other relevant data concerning 

any threat (or lack thereof) to the 

WVNFS; (2) additional information on 

the range, distribution, and population 

size of the WVNFS and its habitat; (3) 

the location of any additional 

populations of the WVNFS; and (4) data 

on population trends. The comment 

period was from December 19, 2006, 

through April 23, 2007 (71 FR 75924; 72 

FR 7852; 72 FR 9913). 

During the 120-day comment period, 

we received a total of 4,808 comments. 

Of these comments, we consider 18 (6 

from peer reviewers and 12 from other 

sources) to be substantive. The majority 

of comments received were form letters 

objecting to the proposed delisting rule 

but providing no new or supporting 

information. 

A. Distribution Concerns 

Issue 1—Some commenters asked us 

to quantify what portion of the 

historical range is currently occupied by 

WVNFS. 

Response—The historical range of 

WVNFS essentially corresponds to the 

distribution of old growth red sprucenorthern hardwood forest (500,000 to 

600,000 acres (ac)) prior to logging and 

fires at the turn of the 20th century. 

Much of the historical red spruce has 

been replaced by northern hardwoods. 

Current estimates of the amount of 

WVNFS habitat vary widely from 

242,000 ac (U.S. Department of 

Agriculture (USDA), Northern Research 

Station 2006, unpublished map) to 

600,000+ ac (Menzel et al. 2006b, p. 4), 

across which the WVNFS is widely 

dispersed. 

Historically, the red spruce-northern 

hardwood forest encompassed portions 

of eight counties, extending from the 

vicinity of Mount Storm (Grant County) 

in the north, to Cold Knob (Greenbrier 

County) in the south, east to the 

Allegheny Front (Pendleton and 

Highland Counties), and west to 

Webster County. Based upon monitoring 

from 1985 to the present, the WVNFS 

still occupies portions of these same 

eight counties, roughly corresponding to 

85 percent of the extent (breadth and 

width) of the historical range. With 

exception of the extreme northern 

portions of Grant County (roughly 5 

percent of the historical range), and the 

area from Briery Knob south to Cold 

Knob in Greenbrier County (collectively 

less than 10 percent of the historical 

range), the outer boundaries of the 

current distribution of the WVNFS 

closely match the extent of its historical 

range (Service 2007a, Figure 1). 

Additional information can be found on 

page 75926 of the proposed delisting 

rule (71 FR 75924). 

B. Population Concerns 

Issue 1—Some commenters expressed 

concern about an absence of population 

information and trend data. These 

commenters stated the Service had 

failed to consider population growth, 

population size, and linkages to other 

populations. Some commenters 

expressed concern about the use of 

persistence as an indicator of 

population health or stability and noted 

that the Service had not clearly defined 

the term ‘‘persistence.’’ The commenters 

stated that this approach is flawed 

because it is not tied to knowledge of 

the population, but merely to subspecies 

presence, which can be explained by 

immigration from other populations. 

Response—The Service has 

considered population dynamics when 

assessing the status of the WVNFS using 

the best available scientific data. The 

Service considers persistence to be the 

best indicator of successfully 

reproducing populations for this 

subspecies, given its poor detectability, 

its life history characteristics, and the 

20+ years of data from presence/absence 

surveys. 

We define persistence as continuing 

captures of WVNFS over multiple 

generations at previously documented 

sites throughout the historical range. 

Because WVNFS first reproduces at 1–

2 years, and has a relatively short life 

span, averaging approximately 3 years, 

persistence at a single monitoring site 

over 5 years indicates successful 

reproduction across multiple (three to 

five) generations (Service 2007c, p. 10). 

The Service has analyzed presence/ 

absence data to determine persistence of 

WVNFS across its range, taking into 

consideration detectability rates, life 

span, reproductive capacity, dispersal 

capability, linkages to other 

populations, and the naturally patchy 

habitat distribution of the subspecies 

(Service 2007c, pp. 5–6, 9–11). These 

data consistently indicate a relatively 

high degree of persistence (roughly 80 

percent) across the landscape, and are 

not indicative of a declining population 

of WVNFS. The data available for the 

remaining landscape (roughly 20 

percent) does not represent an absence 

or lack of persistence of the WVNFS, but 

rather is indicative of the WVNFS’ life 

history traits (i.e., elusive and hard to 

capture). Therefore, the data is simply 

less conclusive. This remaining 

landscape (roughly 20 percent) is still 

habitat for the WVNFS but success rates 

for capturing the WVNFS are lower. The 

persistence of WVNFS is likely 

facilitated by immigration. See Issues 2, 

3, and 4 and their responses under this 

section for additional information. 

Issue 2—Some commenters believe 

the Service must conduct a Population 

Viability Analysis (PVA) to identify a 

minimum viable population before a 

decision on delisting the WVNFS is 

made. These commenters noted that 

genetics-based computer models of 

minimum viable population sizes 

generally indicate that population sizes 

on the order of thousands of individuals 

(low thousands or higher) may be 

needed. In contrast, another commenter 

submitted a copy of a manuscript by 

Smith and Person (2007, pp. 626–636) 

that evaluated the estimated persistence 


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of a northern flying squirrel subspecies 

in fragmented habitats in southeast 

Alaska. This commenter concluded that 

dispersal likely will be the key to 

northern flying squirrel population 

viability, not total population size of 

individual patches. 

Response—A genetics-based 

computer model to identify minimum 

population sizes for WVNFS does not 

currently exist. In our view, there is 

insufficient information available to 

support an accurate or credible geneticsbased PVA model for WVNFS, and such 

an analysis would rely upon too many 

variables whose values would be 

speculative. Given the nature of the 

WVNFS life history and habitat 

information currently available, we 

believe that estimates of persistence, 

and an analysis of functional habitat 

connectivity, are the most credible form 

of PVA analysis. We therefore have 

done these analyses using the best 

available scientific data (for more detail, 

see Service 2007c, pp. 5–6, 9–11) 

resulting in evidence of persistence and 

a high degree of habitat connectivity. 

We also have considered the recent 

work by Smith and Person (2007, pp. 

626–636), who developed a birth-death 

process model to examine persistence of 

populations of a different northern 

flying squirrel subspecies in 

hypothetical, old-growth reserves 

isolated in managed landscapes in 

Alaska. We agree with these authors that 

functional habitat connectivity is more 

important to WVNFS population 

viability than total population size 

rangewide, or population sizes of 

individual habitat patches (See Issues 3 

and 4 and their responses below). 

Issue 3—Some commenters expressed 

concerns that habitat reserves may be 

too few, small, degraded, and isolated to 

support viable populations of WVNFS. 

These commenters emphasized the 

importance of functional habitat 

connectivity. 

Response—Within the range of the 

WVNFS in the central Appalachians, 

there are numerous patches of highquality, second-growth red spruce 

forest, with individual trees that are 

near maximum size and age, within an 

almost continuous matrix of more 

highly variable, second-growth red 

spruce and northern hardwood forest 

conditions. The habitat is still relatively 

well connected from the standpoint of 

WVNFS movement and does not 

significantly limit dispersal and 

movements (Service 2007c, Figure 1). 

Within the range of the WVNFS, above 

3,200 feet (ft), approximately 96 percent 

of the land is forested (627,237 ac) 

(USDA Forest Service 2007, unpubl. 

map). Patch sizes on the MNF also are 

fairly large and connected by numerous 

forested linkages, facilitating the 

likelihood of WVNFS dispersal (Service 

2007c, p. 6, Figure 1). For example, 

radio-tagged male WVNFS and other 

subspecies of northern flying squirrels 

have demonstrated an ability to make 

sudden, long-distance movements, 

presumably to find females. Some 

individuals have traveled up to 2 

kilometers (1.2 miles) in a night during 

the mating season, which is from late 

winter to early spring (Smith 2007a, p. 

871; Menzel 2003, p. 77, 117; Terry 

2004, p. 18; Weigl et al. 1999, pp. 59–

62; Weigl et al. 2002, p. 37, 145). 

Smith and Person (2007) modeled 

habitat reserve size for northern flying 

squirrels in Alaska. Habitat reserves 

must sustain individual insular 

populations, or the matrix of managed 

lands between reserves must allow 

dispersal among reserves to maintain 

wildlife populations within a 

metapopulation structure (Smith and 

Person 2007, p. 633). Out of an 

abundance of caution, Smith and Person 

(2007, p. 628) modeled the first scenario 

to estimate the persistence of northern 

flying squirrel populations occupying 

isolated fragments of habitat in a matrix 

of unsuitable habitat within a large 2- 

million-ac landscape in Alaska (p. 628). 

Lacking conclusive evidence of 

dispersal, the authors assumed their 

populations were closed (i.e., no 

immigration or emigration). They also 

assumed the habitat was static (i.e., 

patch size and patch quality are 

constant over as long as a 100-year 

period). Neither of these assumptions 

fits the situation in the central 

Appalachians where many, if not most, 

of the habitat patches containing 

WVNFS are connected by habitat, and 

through passive and active management, 

conditions are expected to continue 

improving. In addition, the authors 

relied heavily on 3 years of local 

demographic data and data from a 

longer-term study in Canada. These 

demographic data may be dissimilar to 

those of WVNFS in West Virginia and 

Virginia. For example, the authors used 

an estimated average litter size of 2, 

which is low compared to the WVNFS 

average litter size of 2.5–3.0 (Reynolds 

et al. 1999, p. 346; Stihler et al. 1998, 

p. 178). Estimated survival rates also 

may have been low because the value 

was based on recaptures of tagged 

individuals, and the lack of a recapture 

does not mean a squirrel has died. 

That said Smith and Person do 

provide a framework for judging the 

relative magnitude of patch sizes that 

may be needed for northern flying 

squirrel persistence in large forested 

landscapes. Smith and Person (2007, p. 

631, Table 5) estimated that the 

minimum area of an isolated patch of 

contiguous habitat to confidently 

sustain populations for at least 100 

years without immigration/emigration 

was 11,414 ac (4,621 hectares (ha)) 

(P=0.90). Furthermore, there was a high 

probability that G. sabrinus could 

persist in smaller (≥245-ac [99-ha]) 

isolated habitat patches for 25 years 

without migration (p. 631). Smith and 

Person (2007, p. 633) concluded that 

large reserves may not need to be 

contiguous, because interspersed lowerquality habitats can support northern 

flying squirrels for a short time and 

likely facilitate dispersal between 

patches of higher-quality habitat (Smith 

and Person 2007, p. 633). 

Because of the many assumptions, 

described above, of this model, which 

do not transfer well to the central 

Appalachians, we decided to do a 

coarse comparison of minimum patch 

sizes. Because the landscape for WVNFS 

appears to have a higher degree of 

functional connectivity than the study 

area in Alaska, we looked at the total 

acreages of contiguous and connected 

suitable habitat within each of seven 

core areas. [Five ‘‘core areas’’ were 

identified at the time the 1990 recovery 

plan was written (Service 1990. p. 16) 

as clusters of capture sites, and are 

referred to in the plan as Geographical 

Recovery Areas. Two more clusters were 

later identified when surveys found 

additional WVNFSs. Collectively these 

seven areas (hereafter called ‘‘core 

areas’’) encompass the entire extant 

distribution of WVNFS.] Out of an 

abundance of caution, we assumed 

these seven core areas were 

geographically separated (no 

immigration/emigration among them), 

although this likely is not the case. 

Using these conservative assumptions, 

the ‘‘minimum patch size of contiguous 

habitat’’ within each core area ranges 

from 9,353 ac (3,787 ha) for the smallest 

core area (Stuart Knob) to 120,484 ac 

(48,779 ha) for the largest core area 

(Cheat). Six of the seven core areas 

exceed the minimum patch size 

identified by Smith and Person (2007, p. 

631) as necessary to confidently sustain 

populations for at least 100 years 

without immigration/emigration (11,414 

ac or 4,621 ha). Thus we infer that there 

is adequate habitat for persistence of 

WVNFS populations within most, if not 

all, of the core areas. 

Whereas habitat conditions in Alaska 

(small, isolated, old-growth forest 

fragments in a matrix of unsuitable 

habitat) are quite dissimilar to those in 

the central Appalachians (large, wellconnected patches of predominantly 

second-growth forest in a matrix of 


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suitable habitat), it appears that habitat 

reserves of sufficient quantity, quality, 

and connectivity exist to sustain 

populations of WVNFS with influences 

of immigration and emigration. This 

habitat matrix provides a high degree of 

functional connectivity, as evidenced by 

persistence over multiple generations at 

monitoring sites across a range of forest 

conditions (Service 2007c, pp. 9–11). 

Issue 4—Some commenters asked the 

Service to analyze the viability of 

WVNFS metapopulations (multiple, 

relatively isolated breeding units). 

These commenters cited Weigl (2007, p. 

903), who claimed that ‘‘some second 

growth stands may well appear to 

support healthy densities of squirrels, 

but, in reality, are population sinks for 

migrants from neighboring old growth 

habitats and thus may not permanently 

maintain viable populations.’’ These 

commenters suggested the WVNFS may 

be undergoing a population decline that 

is influenced by source-sink dynamics 

of meta-population theory. 

Response—In response to this 

comment, the Service has conducted 

additional analyses to look for evidence 

of population sinks and sources in the 

central Appalachians. We found no 

evidence that the few remaining old 

growth patches of habitat in the central 

Appalachians, or other optimal habitat, 

are operating as potential sources of 

WVNFS recruits that disperse into 

suboptimal habitat (potential sinks) 

where populations are not sustained. 

Rather, our analysis of 21 years of 

monitoring shows no evidence of 

localized extirpation since the 

subspecies was listed. The WVNFS 

persists in or near all of the historical 

areas where it was originally known at 

the time of listing. Persistence of 

WVNFS across the range over multiple 

generations is consistently high, 

consistently distributed across habitat 

types (varying from 70 to 86 percent 

persistence) and geographic zones 

(varying from 80 to 85 percent 

persistence), and not significantly 

different from expected values (Smith 

2007a, p. 871; Service 2007c, p. 11, 

Table 1). Nestlings and juveniles are 

routinely documented at monitoring 

sites (76 percent of sites) (Service 2007c, 

p. 9). Because WVNFS has a relatively 

short life span (averaging approximately 

3 years), and first reproduces at age 1 or 

2, persistence at a single monitoring site 

over 5 years indicates successful 

reproduction across multiple (3+) 

generations. In addition, the observed 

roughly 1:1 sex ratio (492 males, 539 

females) is within the range needed for 

normal reproductive performance 

(Service 2007c, p. 11). Males are most 

likely to disperse, presumably to seek 

females (Ford 2007a). There is no 

indication of a predominance of 

dispersing males or juvenile males, 

which could be indicative of a metapopulation sink dynamic (such as an 

emigration front of individuals leaving 

former territory), or of a metapopulation source-dynamic (such as a 

colonizing front of individuals moving 

into former territory) (Ford 2007a). 

Collectively, these data show a 

relatively high degree of population 

stability and consistent habitat 

occupancy across multiple generations. 

Issue 5—Some commenters noted that 

the chance of capturing a WVNFS in a 

nest box is confounded by a very low 

rate of occupancy, plasticity in nest site 

selection, availability of nest sites, and 

relative abundance of WVNFS. These 

commenters state that it is as important 

to understand why an individual is 

present as to understand why it is not 

present. They state that a major caveat 

of relying on the nest box data as a 

measure of persistence is that it does not 

tell us anything about the habitat, and 

that it is impossible to infer what is 

optimal habitat and if it is available and 

can support the WVNFS. 

Response—The Service agrees that all 

of the factors mentioned above affect the 

chance of capturing a WVNFS; however, 

we disagree about inferences that can be 

drawn from persistence data. Continued 

persistence of WVNFS over the past 

century and occupation throughout 

most of its historical range tell us much 

about habitat and indicate that sufficient 

quality and quantity of habitat exists 

regardless of what may be perceived as 

‘‘optimal’’ habitat. Therefore, a strong 

inference can be made regarding habitat 

suitability based on the persistence, 

successful reproduction, and sex ratios 

that lack any indication of population 

sink dynamics (Service 2007c, pp 11, 

Table 1). 

Issue 6—Some commenters cited a 

paper by Weigl (2007, p. 900) as 

evidence that the WVNFS may have a 

longer life span than previously 

assumed. These commenters suggested 

that if this is true, then the Service may 

need to reanalyze reproductive data and 

conclusions about persistence. 

Response—Weigl (2007, p. 900) 

referred to a study of a different G. 

sabrinus subspecies in the Pacific 

Northwest as evidence that WVNFS may 

be relatively long lived. In this study, 

three squirrels were known to be at least 

7 years old at recapture; however, the 

majority of squirrels captured were not 

known to survive beyond 2–3 years 

(Villa et al. 1999, p.39). In the central 

Appalachians, recapture data for four 

WVNFS suggest the average lifespan is 

probably about 2 to 3 years (West 

Virginia Division of Natural Resources 

(WVDNR) and Service 2006, unpubl. 

data). Wells-Gosling and Heaney (1984, 

p. 4) also noted the average longevity of 

G. sabrinus was probably less than 4 

years. Our previous conclusions about 

persistence remain valid based upon an 

average lifespan of 2–3 years. 

C. Using the Best Available Science 

Issue 1—Some commenters were 

concerned about a lack of knowledge of 

the WVNFS life cycle and the 

consideration of science regarding the 

subspecies’ ecology. 

Response—The WVNFS life cycle and 

ecology is fairly well known from 

numerous studies in peer-reviewed 

journals, books, and technical 

publications. The Service has 

considered the best available scientific 

and commercial data regarding WVNFS 

life history and ecology. For a full list 

of the literature cited in this final rule, 

please contact the West Virginia Field 

Office (see ADDRESSES). 

Issue 2—Whereas four peer reviewers 

and some commenters were satisfied the 

best available science and data had been 

used in the development of the 

proposed rule, two peer reviewers and 

some commenters questioned the 

quality or interpretation of data used to 

support the proposed rule. These 

commenters offered manuscripts in 

press, or alternative literature citations 

or explanations of the data. 

Response—The Service has reviewed 

the manuscripts in press (now 

subsequently published) and literature 

citations provided by commenters. We 

have considered and incorporated the 

information provided in these 

documents where appropriate in this 

final rule. We have incorporated these 

documents into our administrative 

record and cited them in this rule where 

appropriate (including, but not limited 

to, sections of the rule dealing with 

WVNFS population dynamics; habitat 

use, quantity, quality, and connectivity; 

and climate change). The peer-reviewed 

scientific journal articles, peer-reviewed 

agency reports, and other literature cited 

in the final rule represent the best 

available science relevant to the 

decision. None of the alternative 

explanations of the data were as 

persuasive as the sources we have cited 

in the final rule. 

Issue 3—Some commenters disagreed 

with a choice of words in the summary 

sections of the proposed rule which 

referred to ‘‘an increase in the number 

of individual WVNFSs.’’ These 

commenters claimed that there is no 

evidence of an increase, noting that 

1,141 captures do not represent unique 

squirrels, because unknown portions 


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were recaptures. These commenters 

conducted an independent analysis of a 

WVNFS electronic database and field 

data reporting forms. They reported 

inconsistencies in the data base, and 

concluded there may have been as few 

as 654 unique captures. These 

commenters believe that such a low 

number of captures of unique 

individuals diminishes the credibility of 

conclusions reached by the Service 

about persistence. 

Some commenters also questioned 

whether an increase in WVNFS 

occupancy was simply a consequence of 

increased surveys or efficacy of survey 

efforts since listing. One commenter 

questioned our ability to detect a change 

in habitat occupancy. 

Response—Whereas the proposed rule 

did identify the total number of 

recaptures (71 FR 75926), the Service 

agrees that use of the phrase ‘‘increase 

in the number if individual WVNFSs’’

was not accurate, as we have not 

estimated the size of the WVNFS 

population. We have corrected this 

wording in the final rule. Based upon 

data collected through 2005, there has 

been an increase in the total number of 

known captures, from 10 at the time of 

listing, to 1,141 captures at the time of 

the proposed rule, of which there were 

78 total recaptures (6.8 percent). Due to 

multiple recaptures of some individuals, 

these 78 total recaptures represent 62 

individuals. 

Contrary to the commenters’ estimate 

of 654 unique captures, we calculate 

that there were a total of 908 unique 

captures (760 unique captures of adults 

and 148 unique captures of juveniles). 

These estimates take into account 

unique recaptures and unmarked 

individuals. About 8 percent of the 

adults escaped before they could be 

marked. Also, contrary to the 

commenters’ determination that 

‘‘several’’ nestlings were not tagged, 

nearly all of the 133 nestlings and about 

2 percent of the 154 juveniles captured 

were not marked as a precautionary 

measure. Researchers believe that 

marking small individuals with ear tags 

and/or pit tags is an unnecessary 

procedure that could increase mortality 

(Stihler 2007). The fact that these 

individuals were not marked is 

inconsequential when considering that 

there is less than a 5 percent probability 

of subsequent recapture. Rather, the 

capture of nestlings or juveniles is a 

good sign of reproduction (25 percent of 

all captures). 

The increase in the number of capture 

locations is useful in evaluating the 

distribution of WVNFS within its range, 

but cannot be used to estimate 

population sizes. The number of 

captures has increased with increased 

survey effort. While the area covered by 

surveys has increased over time, the 

efficacy of capturing WVNFS remains 

low. Based on original methodologies 

used at the time of listing, and still 

predominantly in use today, roughly 2 

percent of nest box or live trap checks 

result in detection of WVNFS (Terry 

2004, p. 46; Service 2006b, p. 13). This 

estimate of detectability is a simple 

calculation of the proportion of nest box 

or live trap checks that resulted in 

WVNFS capture. We have not used this 

simple estimate of detectability to 

calculate changes in habitat occupancy 

over time as suggested by one 

commenter. We evaluated whether the 

existing data set could be analyzed 

using more rigorous models for 

estimating detectability and changes in 

habitat occupancy (MacKenzie et al. 

2002, pp. 2248–2255; MacKenzie et al. 

2003, pp. 2200–2207; MacKenzie 2005, 

pp. 849–860; MacKenzie 2006, 1568–

1584); however, we felt it inappropriate 

given that model assumptions would 

have been violated and could not be 

validated. While there has been an 

increase overall in survey area, the 

techniques used were the same and the 

intensity of work at sites has not varied 

significantly in the past 20 years. 

As a result of these comments, the 

WVDNR has checked the data base 

against field forms and has made a few 

minor corrections. These changes do not 

substantially alter previous statistics 

reported by the Service or conclusions 

reached about persistence. 

Issue 4—Some commenters noted that 

monitoring sites were not randomly 

selected, which builds in bias. These 

commenters recommended that such 

data not be used for estimating 

population. 

Response—The Service acknowledges 

that monitoring sites were not selected 

randomly. The goal of the presence/ 

absence surveys was to find as many 

WVNFS as possible and to document 

their range and distribution. 

Consequently, few sites were placed in 

low-quality habitat, and many sites were 

placed in moderate or high-quality 

habitat. Because of this bias, the Service 

has not used these data to estimate 

population sizes, but rather to monitor 

presence/absence and persistence. 

D. Genetic Concerns 

Issue 1—Some public commenters 

were concerned about a lack of genetic 

research that might indicate risks due to 

isolation (e.g., genetic drift, inbreeding) 

or existence of discrete populations 

meriting ESA protection. 

Response—We considered 

information from several studies using a 

variety of genetic markers. Allozymic 

analyses by Browne et al. (1999, pp. 

205–214) found lower measures of 

polymorphism and heterozygosity in 

North Carolina, West Virginia, and 

Virginia populations of G. sabrinus 

compared with other northern flying 

squirrels, noting that population 

structure in the southeastern States is 

similar to that of other species that 

occupy habitat islands (Browne et al. 

1999, p. 212). Similarly, allozyme and 

mitochondrial DNA data examined by 

Arbogast et al. (2005, pp. 123–133) 

showed lower diversity of G. s. fuscus 

and G. s. coloratus compared with 

conspecifics (other flying squirrel 

species), but not relative to populations 

of the widespread southern flying 

squirrel. Sparks’ data from a small 

number of microsatellite loci showed 

moderate to high gene flow across 

populations of northern flying squirrels 

in West Virginia, Virginia, and North 

Carolina (Sparks 2005, pp. 16 and 23). 

In addition, the coefficient of inbreeding 

failed to differ between populations at 

Cheat Mountain, West Virginia, and at 

an unfragmented forested landscape in 

Washington State (Sparks 2005, p. 18). 

Also, no difference in levels of a 

parasitic helminth (a species of parasitic 

worm commonly found in the intestines 

of flying squirrels, the presence of 

which is often used as possible 

indicator of reduced fitness) was 

detected among G. sabrinus and two 

sympatric tree squirrels (Sparks 2005, 

pp. 19, 62). 

Arbogast et al. (2005, p. 130) and 

Weigl (2007, p. 902) speculate about 

potential future decreases in genetic 

diversity due to hypothetical habitat 

reductions. As discussed under Issue 3, 

Response to Comments, Section B—

Population Concerns, however, we 

believe that habitat is still relatively 

well connected from the standpoint of 

WVNFS movements. Interspersed 

lower-quality habitats that can support 

northern flying squirrels for a short time 

will also facilitate the low levels of 

dispersal necessary to maintain allelic 

diversity and heterozygosity while 

conserving local adaptations. 

Furthermore, Sparks (2005, p. 29) 

suggests that G. sabrinus may have a 

population structure adapted to some 

degree of inbreeding tolerance. 

In summary, after review of the 

genetic studies referenced above, we 

have not detected any genetic risk to the 

WVNFS due to isolation. Additionally, 

we are aware of no genetic, behavioral, 

ecological, morphological, 

physiological, physical, or other 

information supporting the existence of 

distinct population segments within the 

WVNFS. 


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E. Habitat Modeling Concerns 

Issue 1—One peer reviewer and some 

commenters thought the Service had 

applied the Menzel et al. (2006b, pp. 1–

10) model outside of its intended scope 

and for purposes not supported by the 

study the model is based upon. Some 

conclude that the Service is using the 

model to make a case that the agency 

can accurately predict habitat and 

WVNFS viability, by assuming that the 

model definitively predicts presence 

and absence. 

Response—Using logistic regression, 

Menzel et al. (2006b, pp. 1–10) 

developed a Geographic Information 

System (GIS)-based habitat model for 

WVNFS in West Virginia by 

synthesizing micro- and macro-habitat 

relationship data. The Service has 

applied this model appropriately to 

gauge the relative abundance and 

quality of habitat rangewide and to 

broadly estimate the predicted 

distribution of WVNFS on the 

landscape. We have not assumed that 

the model definitively predicts 

presence/absence of WVNFS. Nor have 

we argued that predicted habitat ensures 

WVNFS’ viability; predicted habitat is 

only one component. The model can 

give insights, albeit coarse, on habitat 

quality and its distribution across the 

landscape. As noted in the final listing 

rule (50 FR 26999) and recovery plans 

(Service 1990, pp. 12–16) for this 

subspecies, the abundance and quality 

of habitat are keys to the recovery of 

WVNFS because habitat loss and 

degradation were the main factors that 

led to the subspecies being listed as 

endangered. We have used the model at 

a landscape level to predict habitat 

quality and look for evidence of sinksource metapopulation dynamics. We 

have also used the model to highlight 

where managers should conduct followup site visits to determine actual 

squirrel habitat or where managers 

could reasonably assume no occupation 

without a site visit. A manager could 

use Ford et al. (2004, pp. 430–438) at 

the individual forest stand level to 

verify the quality of the habitat or what 

the probability level of occupation 

would be for that specific location. 

Issue 2—Some commenters criticized 

the Menzel et al. (2006b, pp. 1–10) 

habitat model for being unverified and 

untested. 

Response—The model has been 

verified and tested and proved to be 

quite accurate (81 percent) when the 

data were subjected to ground-truthing 

procedures to determine correct 

classification rates of occupiable and 

non-occupiable habitat (Menzel et al. 

2006b, p. 3–4). Staff from the WVDNR 

and MNF have used the model 

successfully to identify WVNFS habitat, 

corroborated by additional captures 

where the model had shown a high 

probability of occurrence. 

Issue 3—Some commenters stated that 

the Menzel model’s prediction of habitat 

from tracking data should have been 

verified in following years (different 

temporal frame) and on different areas 

of the range (different spatial frame). 

Response—The actual telemetry data 

used by the Menzel model did span 

several years and different areas. The 

model is based on actual data, which 

have been verified. 

Issue 4—Some commenters criticized 

the Menzel model for containing several 

untested assumptions: (a) There is a 

direct relationship between nest box use 

and preferred habitat; (b) quality of 

habitat is predicted by elevation and 

vegetative community; and (c) data from 

spring and summer tracking reveals 

information on habitat use the 

remainder of the year. 

Response—Addressing assumptions 

(a) and (b), Menzel et al. (2004, pp. 355–

368; 2006b, pp. 1–10; 2006a, pp. 204–

210) does not assert that probability of 

occurrence equates directly to preferred 

habitat; however, there is a clear 

correlation between high probability 

habitat (>75 percent probability of 

WVNFS occupancy) and habitat 

components such as red spruce and 

high elevation that were preferred by 

radio-collared individuals (Menzel et al. 

2006a, pp. 206–207). Addressing 

assumption (c), data from winter 

telemetry studies at Snowshoe 

Mountain Resort and Canaan Valley 

National Wildlife Refuge (Ford et al. 

2007 in press, pp. 6, 8) are similar to 

results from spring, summer and fall 

reported by Menzel et al. (2006a, pp. 

206–207). Winter data confirm that male 

home ranges are larger than female 

home ranges and both sexes key in on 

red spruce-dominated habitats for 

foraging (Ford et al. 2007, pp. 4, 6, 7). 

Issue 5—Some commenters stated that 

the Menzel model was based on limited 

spatial and temporal data from 4 sites 

and 13 animals; therefore, results can be 

generalized only with great caution. 

Response—The Menzel et al. (2006b, 

p. 3) model is not based on a limited 

subset of the data, but rather is based on 

most of the capture data through 1999 

and most of the telemetry data from 

WVNFS tracked in a variety of stand 

age-classes and compositions. All 

squirrels tracked for which home range 

sizes were calculated, had reached 

home range size asymptotes (the point 

on a graph indicating the minimum 

number of samples needed to calculate 

maximum home range size), indicating 

that sufficient location data exists to 

estimate home range size. Moreover, 

WVNFSs were tracked in a variety of 

poor to excellent habitat conditions. 

This methodology is consistent with 

similar examples of wildlife habitat data 

being collected from tagged individuals 

and then used in a modeling effort to 

extrapolate across a larger, but similar 

landscape (for example, Gibson et al. 

2004, pp. 75–89; Posillico et al. 2004, 

pp. 141–150). The Service believes it 

has interpreted these data appropriately. 

Issue 6—Some commenters stated that 

the Menzel model is a simplification of 

existing knowledge and does not 

account for important variables in 

WVNFS biology, such as forest age, 

structure, tree composition, and fungi. 

These commenters believe the model 

potentially overestimates optimal 

habitat by treating young forest the same 

as old forest, and by lumping other 

factors together (moist conditions, high 

rainfall, northern aspects, forest 

structure, suitable nest sites, food 

sources, etc.) based on elevation and 

spruce occurrence. 

Response—The Service concurs that 

Menzel et al. (2006b, pp. 1–10) is a 

simple habitat model that was meant to 

capture broad aspects of WVNFS 

distribution. The model tends to 

underestimate higher-quality habitat 

and to overestimate lesser-quality 

habitat, especially near the 50 percent 

predicted probability of occurrence 

threshold (Ford 2007b). However, we 

still think the model is useful and 

reasonably accurate for gauging the 

relative abundance and quality of 

habitat rangewide and for predicting the 

distribution of WVNFS on the 

landscape, and represents the best 

available scientific and commercial 

data. 

F. Ecosystem and Habitat Concerns 

Issue 1—Some commenters were 

concerned that delisting the WVNFS 

would jeopardize an entire ecosystem, 

especially when considering the critical 

role that WVNFS plays in dispersal and 

persistence of numerous fungi which 

have symbiotic relationships with trees. 

Response—The Service agrees that the 

WVNFS plays an important role in the 

red spruce-northern hardwood 

ecosystem (Smith 2007a, p. 862–863; 

Weigl 2007, pp.10–12). Habitat models 

for this subspecies implicitly recognize 

the symbiosis between WVNFS and tree 

fungus (Odom et al. 2001, pp. 245–252; 

Menzel et al. 2006b, pp. 1–10). The 

Service does not expect that delisting 

the WVNFS will have negative 

consequences for the ecosystem. The 

red spruce-northern hardwood 

ecosystem upon which the WVNFS 


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depends has substantially recovered and 

continues to improve (also see Issue 4 

in this same subsection below). The 

delisting process signifies elimination of 

endangerment of the WVNFS and 

elimination of the need for the Act’s 

protections. Delisting is a procedural 

acknowledgement of the recovered 

ecological status of this subspecies and 

the ecosystem upon which it depends. 

Issue 2—One commenter stated that 

protection of habitat is serving as a 

proxy for the status of the subspecies. 

Protection of habitat is critical to 

protection of the subspecies but does 

not ensure recovery. 

Response—While protection of 

habitat is important to the status of the 

subspecies, it is not serving as a 

substitute for other factors. In analyzing 

whether the WVNFS has recovered, the 

Service has considered the reduction of 

all threats to the subspecies, including 

the destruction, modification, or 

curtailment of its habitat or range; 

overutilization; disease or predation; 

inadequacy of existing regulatory 

mechanisms; and other factors. See the 

Summary of Factors Affecting the 

Species section below for additional 

information. 

Issue 3—Some commenters stated that 

there is a lack of a clear definition of 

habitat for WVNFS due to insufficient 

information on habitat needs. Factors 

comprising optimal habitat are complex 

and poorly understood. 

Response—The work of Menzel et al. 

(2004, pp. 355–368; 2006b, pp. 1–10), 

Ford et al. (2004, pp. 430–438; 2007 in 

press, pp. 4–7), and Mitchell (2001, pp. 

441–442) clearly define WVNFS habitat 

and its characteristics. 

Issue 4—Some commenters, including 

two peer reviewers, thought the Service 

had overemphasized spruce as a habitat 

component for WVNFS. These 

commenters note that the WVNFS 

inhabits deciduous forest at lower 

elevations without a spruce component, 

and therefore should not be considered 

an obligate to red spruce forest. These 

commenters state that additional 

hardwood forest needs to be protected. 

Some commenters also disputed that 

red spruce is preferred habitat of 

WVNFS, identifying biases in the work 

by Menzel. These commenters state that 

the Menzel habitat model is based on a 

small sample of nest boxes located in 

red spruce habitat, skewing this 

monitoring program toward a finding of 

red spruce as preferred squirrel habitat; 

however, actual squirrel capture data 

seem to refute the exclusive focus on 

red spruce (Menzel 2003, p. 93). 

Response—The Service never meant 

to imply that the squirrel is an obligate 

of the red spruce forest. However, the 

ecosystem in which WVNFS evolved 

consisted of a significant red spruce 

component, and it would be 

inappropriate to de-emphasize this 

important habitat feature. The WVNFS 

can be quite cosmopolitan, living within 

majority red spruce to nearly complete 

red spruce cover types, to majority 

hardwood to nearly complete hardwood 

cover types where the red spruce-fir 

component is minimal (Stihler et al. 

1995, p. 18; Menzel 2003, p. 68; Menzel 

et al. 2006a, pp. 207–208; Ford et al. 

2004, pp. 433–434; Reynolds et al. 1999, 

pp. 347–348). However, the 

preponderance of the data suggest a 

strong link to red spruce; there is a 

higher probability of WVNFS presence 

in areas with the most red spruce (as a 

percentage of the cover type) (Menzel 

2003, p. 68; Ford et al. 2004, pp. 433–

434, 2007 in press, pp. 12, 15–16; 

Menzel et al. 2006a, pp. 207–208). It is 

well documented that the entire range of 

the WVNFS was a red spruce dominated 

forest until heavily logged during the 

late 1800s and early 1900s (Mielke 1987, 

p. 219; Schuler et al. 2002, p. 89; 

Menzel et al. 2006b, p. 1; Rentch et al. 

2007, pp. 440–442). Home range sizes 

also are smaller in areas with more red 

spruce, suggesting that habitat quality is 

better in these areas because WVNFS do 

not have to travel as far to meet their 

ecological needs (Menzel 2003, pp. 77; 

Ford et al. 2007, p. 6). 

Additionally, no data in the central 

Appalachians show that WVNFS are 

heavily dependent upon pure 

hardwoods. Even so, protection of 

northern hardwood forest of 

considerable size is not a concern in the 

central Appalachians, since, within the 

range of WVNFS above 3,200 ft in 

elevation, approximately 96 percent 

(627,237 ac) of the land is forested 

(USDA Forest Service 2007, unpubl. 

map). At a coarser scale, within the 

more than 2 million ac of northern 

hardwoods in the high Allegheny 

landscape of West Virginia, Forest 

Inventory Analysis shows an 

approximately 15 percent increase in 

northern hardwoods from 1989 

(2,061,000 ac, SE = 4,400 ac) to 2000 

(2,393,600 ac, SE = 4,200 ac) (Griffith 

and Widmann 2003, pp. 30, 32). 

Finally, Menzel (2003, p. 93) does not 

support the commenters’ claims about 

bias. Sample bias was recognized and 

dealt with appropriately. The Menzel et 

al. (2006b, pp. 1–10) study used a 

sufficiently large sample of nest box and 

trap sites that produced WVNFS 

previous to 1999 in a statistical analysis. 

These occupied sites were then 

compared to 700+ locations that failed 

to produce WVNFS in a logistic 

regression analysis. Despite the fact that 

nest box and trap locations were skewed 

towards forest stands containing red 

spruce, captures occurred more 

frequently (in a greater proportion than 

habitat availability would suggest) in 

red spruce than in pure hardwood 

stands. 

Issue 5—Some commenters, including 

two of the six peer reviewers, expressed 

concern about the threat of extensive 

logging on Federal, State, and private 

lands within the range of the WVNFS. 

Some commenters claim the MNF 

proposes to log up to 40 percent of the 

area comprising Management 

Prescription (MP) 4.1, which focuses on 

red spruce and red spruce-northern 

hardwood restoration. 

Response—A substantial amount of 

WVNFS habitat is protected and 

managed consistently with the habitat 

needs of the WVNFS. Approximately 79 

percent of WVNFS habitat (189,785 ac) 

is protected from the threat of exploitive 

logging for the foreseeable future 

(Service 2007a, pp. 5–8). Privately 

owned lands potentially subject to 

continued timbering (50,997 ac or 21 

percent of WVNFS habitat) occur 

primarily at the edge of the subspecies’

range (Service 2007a, p. 8). These lands 

are not critical to the subspecies’

conservation, given the large amount of 

WVNFS potential habitat protected and 

managed on public lands in the core of 

the subspecies’ range. [For more details 

on the degree of land protection, see 

criterion # 3 in Service (2007a)]. 

The current MNF Forest Plan (USDA 

Forest Service 2006a, chapters II and 

III), protects WVNFS habitat primarily 

through land use designations, a 

predominantly passive management 

strategy, and binding standards that 

effectively remove the threat of habitat 

loss (via logging and other disturbances) 

on all WVNFS habitat on the forest 

(164,560 ac or 68 percent of the habitat 

rangewide). Standards TE 63–66 (USDA 

Forest Service 2006a, p, II–26–27) adopt 

and implement the provisions of 

appendix A of the recovery plan for the 

WVNFS, which severely limit 

vegetation management in all WVNFS 

habitat, including breeding, feeding, 

resting, and dispersal corridors (Service 

2001, appendix A). Only specific 

actions that have no adverse effect to 

WVNFS habitat, a discountable or very 

minor effect, or that demonstrate a 

beneficial effect (such as habitat 

restoration) are allowed in WVNFS 

habitat forest-wide. Based upon the 

Forest Service’s long-term (50+ years) 

desired conditions for the ecosystem 

(USDA Forest Service 2006a, p. III–12), 

the Forest Service’s intent shown in a 

Memorandum of Understanding signed 

by the MNF (Service et al. 2007, pp. 3 


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and 8), conversations with MNF staff, 

and the absence of any information to 

the contrary, we reasonably expect these 

standards to continue to apply 

regardless of the Act’s listing status of 

the WVNFS. This management strategy 

is also likely to continue post delisting, 

as the WVNFS would be managed by 

the Forest Service as a ‘‘sensitive 

species’’ (USDA Forest Service 2006c, p. 

18). 

The commenters’ reference to 

potential logging of 40 percent of the 

area of Management Prescription 4.1 

appears to stem from a 

misunderstanding of forest-wide 

standards TE63–66 and how they 

interplay with the other standards on 

specific prescription areas. Prescription 

area 4.1 encompasses 153,600 ac, of 

which 59 percent (roughly 91,332 ac) 

has been mapped as WVNFS habitat and 

is protected from commercial logging by 

standards TE63–66. The remaining 41 

percent of the area (62,268 ac) has not 

been mapped as WVNFS habitat. Within 

this 62,268-ac area, approximately 

27,300 ac (or 18 percent of the total 

acreage in prescription area 4.1) have 

been tentatively identified as suitable 

for timber production (USDA Forest 

Service 2006b, p. 3–354). These 27,300 

ac may be logged contingent on sitespecific project review and field checks 

to verify that these lands are not 

WVNFS habitat. Thus, at most, 18 

percent of the land in MP 4.1 could be 

logged over the life of the Forest Plan 

and all of this land would need to be 

demonstrated to not be suitable habitat 

for WVNFS, prior to logging. 

Logging of areas that are not WVNFS 

habitat will also need to comply with an 

array of other applicable standards in 

the management direction for 

prescription area 4.1 (USDA Forest 

Service 2006a, pp. III–14 to III–16), such 

as standards 4118 and 4119, which 

place limits on the amount and timing 

of disturbances within harvest units 

(USDA Forest Service 2006a, p. III–15). 

Standard 4118 states that no more than 

40 percent of forested National Forest 

System lands within each 4.1 

prescription area unit shall be harvested 

over a 10-year period. Standard 4119 

requires that unforeseen activities, such 

as timber salvage or pipeline 

installation, shall be counted toward the 

40 percent disturbance standard in 

4118. Thus there are additional limits 

on timbering, even in areas that are not 

WVNFS habitat, that further reduce 

forest disturbances. 

Limited logging in WVNFS habitat for 

purposes of restoration is also allowed 

in prescription area 4.1, consistent with 

standards TE 63–66, as long as it can be 

demonstrated to result in a minor/ 

discountable adverse effect or a 

beneficial effect to WVNFS. The Forest 

Service has an objective to restore 

approximately 1,000 to 5,000 ac of 

habitat over the next 10 years (USDA 

Forest Service 2006a, p. III–14, objective 

4107). Standard 4118 also applies to 

these restoration activities. Hence it 

places limits on the frequency of 

disturbances within stands. 

The Service is confident that these 

restoration efforts would benefit 

WVNFS in several ways, by: (1) 

Increasing amounts of coarse woody 

debris necessary for many fungal 

species; (2) increasing the size and 

importance of red spruce (an important 

fungal substrate); (3) increasing habitat 

patch size and connectivity; (4) 

increasing snags available as day dens 

for WVNFS; and (5) decreasing hardmast production, thereby lessening 

stand value to the southern flying 

squirrel competitor (Menzel et al. 2006a, 

p. 208). 

Issue 6—Some commenters expressed 

a view that all old growth forest across 

the range of WVNFS needs to be 

protected. These commenters cited 

Smith (2007a, pp. 864–865, 877) and 

Weigl (2007, p. 899, 902) as evidence of 

concerns about ongoing harvest of old 

growth forest, its replacement with 

plantations or regenerating stands, and 

the increasing fragmentation of much of 

the remaining habitat. 

Response—There is little to no 

harvesting occurring in old growth 

forests on public or private lands within 

the range of the WVNFS. There is very 

little old-growth remaining from the 

exploitive logging period in the late 

1800s/early 1900s. On the MNF, old 

growth currently comprises less than 1 

percent of the entire forest (USDA 

Forest Service 2006a, p. B–1). In 

addition, areas identified as old growth 

on the MNF are not suitable or allowed 

to be cut. The remaining known oldgrowth areas on the forest are protected 

by Botanical Area, National Natural 

Landmark, or Scenic Area designations, 

and are managed through specific Forest 

Plan direction and standards that 

prohibit timber removal and restrict 

other types of vegetation management in 

these areas (USDA Forest Service 2006a, 

p. B–4). Furthermore, ‘‘[t]imber harvest 

goals and objectives are based on 

achieving desired conditions for 

vegetation and habitat, not on regional 

economics’’ (USDA Forest Service 

2006b, Final Environmental Impact 

Statement (FEIS), Appendix I, p. I–152), 

so there is little risk of the MNF having 

adverse impacts on the WVNFS. 

Concerns about a significant increase in 

forest fragmentation throughout much of 

the remaining WVNFS habitat are 

unsubstantiated. There are no existing 

or predicted activities that are 

anticipated to significantly adversely 

affect forests within WVNFS range on a 

landscape level. 

Issue 7—Some public commenters 

cited a newspaper article as specific 

evidence that the impact of second 

home development in West Virginia is 

a significant threat to WVNFS. They 

requested that the Service reanalyze 

these impacts. 

Response—The Service has 

reanalyzed these impacts and come to 

the same conclusion as in its earlier 

analysis, that second home development 

is not currently a significant threat. The 

greatest development pressures in West 

Virginia are occurring, and are projected 

to continue to occur, outside of the 

range of the WVNFS, in the far eastern 

panhandle, and in and around the cities 

of Morgantown and Charleston (Stein et 

al. 2005, Figure 2). Second home 

development currently is occurring at 

the edge of the range of the WVNFS 

(primarily at Canaan Valley and 

Snowshoe Mountain). By 2030, housing 

density increases are projected to occur 

on private forests across 0 to 5 percent 

of the area corresponding to the core of 

the range of WVNFS (Stein et al. 2005, 

Figure 2). Such losses, if they occur, 

would be at the periphery of the range 

and minor in relation to the 242,000 ac 

of WVNFS habitat that exist within a 

larger landscape encompassing the 

range of WVNFS that is 96 percent 

forested (USDA Forest Service 2007, 

unpub. map). 

Issue 8—Some commenters thought 

that the impacts of roads had not been 

adequately considered. These 

commenters stated that roads create 

absolute barriers to flying squirrel 

movement. These commenters were 

concerned that construction of 

Appalachian Corridor H (a four lane 

divided highway running from Weston, 

WV, to the Virginia line), in particular, 

will open the region to further 

development and will isolate 

populations of WVNFS in Blackwater 

Canyon from populations and suitable 

habitat south of the highway. 

Commenters were concerned that 

populations of WVNFS in Blackwater 

Canyon north of the highway may not 

be able to survive on the remaining 

small island of habitat. They criticized 

the Service for not discussing these 

impacts in more detail in the proposed 

rule or 5-year review. 

Response—Construction of Corridor H 

through the extreme northern part of the 

WVNFS range is not expected to result 

in significant impacts to WVNFS or its 

habitat. As explained in the Land Use 

Planning section of the Factor A 


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analysis below, sufficient habitat will 

remain on both sides of the highway to 

support WVNFS (Service 2006b, pp. 4–

5, 16–29; 2007c, pp. 3–4, 14–26). 

Additionally, a cumulative effects 

assessment, conducted by the West 

Virginia Department of Transportation 

(2006, pp. 17–19) suggests there is an 

adequate amount of nonenvironmentally sensitive, lowelevation land, which is not WVNFS 

habitat, and is available to support all 

development reasonably expected to 

occur as a result of the highway 

construction. 

Issue 9—Some commenters were 

concerned that mining, drilling for gas, 

and construction of wind turbines in the 

habitat of WVNFS are increasing and 

therefore pose a threat to WVNFS. 

Response—There is no evidence that 

these activities have in the past, or will 

in the future, significantly threaten the 

WVNFS. This conclusion is based upon 

Service review of impacts to WVNFS 

from permit applications for coal 

mining, gas, and wind power projects. 

Surface mine projects in West 

Virginia average 302 ac in size, and 

underground mines average 34 ac of 

surface disturbance (Office of Surface 

Mining (OSM) 2005, p. 2). Most coal 

mining activity is concentrated in six 

counties (Boone, Kanawha, Mingo, 

Logan, Marshall, and Monongalia) 

outside the range of the WVNFS (OSM 

2005, p. 2). Within the range of WVNFS, 

small portions of Greenbrier, Randolph, 

Tucker, and Grant Counties have coal 

seams (OSM 2005, cover map); however, 

these areas were mined in the past and 

are not currently active. Given the cost 

of reopening a mine, it is unlikely that 

there would be a resurgence of active 

mining in these areas, considering that 

these sites require expensive acid-mine 

waste remediation (Fala 2007). In the 21 

years since the WVNFS’s listing, there 

have been only 2 or 3 projects out of 

thousands reviewed each year where the 

Service identified potential adverse 

effects from coal mining to WVNFS 

habitat, and each of these projects was 

in marginal habitat on the edge of the 

subspecies’ range. The Service has no 

information suggesting that coal mining 

activities will expand into WVNFS 

habitat. Given this lack of evidence of a 

threat and the above prior history and 

acreages involved, the potential for 

future impacts to WVNFS from this 

activity appears remote and 

insignificant. 

The Service has noticed a recent 

increase in gas drilling applications in 

West Virginia; however, the footprint of 

these projects typically is small, 

averaging approximately 1.5 ac per gas 

well. These projects also tend to use 

existing, short (<1 mile long) gravel 

access roads which do not pose a barrier 

to WVNFS dispersal. In the 21 years 

since the WVNFS’s listing, few if any 

gas projects have resulted in adverse 

impacts to WVNFS habitat, and none of 

these projects have resulted in take of 

WVNFS. The Service expects these 

trends to continue after the WVNFS is 

delisted. The minor impacts of these 

projects do not pose a substantial threat 

to WVNFS. 

There currently is one operating wind 

power project in West Virginia, two 

under construction, and one approved 

which will not be constructed. There 

also is one project in Virginia in the 

permitting application phase. These 

projects have ranged in size from 24 to 

372 ac of disturbance. Neither the 

presently operative project nor the two 

under construction have had impacts to 

WVNFS or its habitat. Although the 

Service has noticed an increase in 

prospecting for wind power projects in 

West Virginia, only a minority of these 

potential projects might adversely 

impact WVNFS or its habitat. Three of 

the 13 projects the Service has reviewed 

initially identified potential adverse 

impacts to WVNFS habitat (two projects 

in West Virginia and one project in 

Virginia). Two of these projects 

ultimately avoided WVNFS habitat 

because of the Act, and one of these 

projects was withdrawn due to 

difficulties seeking access from the 

Forest Service. Although prospecting is 

currently occurring, nearly half of all 

prospective wind energy applications 

filed for grid interconnection study 

within the mid-Atlantic region are 

withdrawn (Boone 2006, pp. 1–2). 

On national forest lands, project 

proponents currently must seek separate 

authorization for prospecting (surveys 

and setting up meteorological stations), 

as well as the construction and 

operation of wind towers. Even after the 

WVNFS is delisted, proposed wind 

farms in national forests within the 

range of WVNFS range would still need 

to be consistent with standards and 

guidelines in the forest plans. Therefore, 

we conclude that while prospecting in 

wind farms is increasing, only a 

minority may materialize, and fewer 

still might adversely affect the WVNFS. 

Based on these projections and the 

small acreage potentially involved, we 

conclude that wind power will not pose 

a significant threat to WVNFS or its 

habitat. 

G. Forest Pest Concerns 

Issue 1—Some commenters were 

concerned about the effects of beech 

bark disease and the hemlock woolly 

adelgid on the habitat of the WVNFS. 

Two peer reviewers noted that while 

these forest pests may have local 

impacts to WVNFS, they are not 

significant at the landscape level. Two 

peer reviewers discussed forest pests as 

potential threats but did not comment 

on their significance to WVNFS. 

Response—Any impacts to WVNFS 

habitat from beech bark disease or 

hemlock woolly adelgid are considered 

minor in the context of the subspecies’

range. A decline in American beech, as 

a result of beech bark disease, should 

provide additional snags and coarse 

woody debris for WVNFS. Additionally, 

a decline in beech nuts would also 

reduce the food supply of southern 

flying squirrels, a potential competitor 

of the WVNFS. 

Eastern hemlock currently comprises 

1 to 9 percent of forested land in 

counties within the range of WVNFS in 

West Virginia (Kish 2007, Figure 1). A 

predominantly eastern hemlock 

overstory is known to occur at 7 percent 

of WVNFS nest site locations (such as 

Blackwater Falls State Park), and its loss 

could affect the quality of riparian zone 

habitat useful for WVNFS dispersal 

between more isolated patches of red 

spruce–northern hardwood forest. 

Whether or not eastern hemlock is 

replaced by red spruce or northern 

hardwoods, thereby ameliorating losses, 

is unknown. However, research 

indicates that hardwood forests with 

little or no conifer component are not 

barriers to WVNFS movement (Menzel 

et al. 2006a, p. 207). Please refer to the 

5-year review (Service 2006a, pp. 17–18) 

and Factor A of the Summary of Factors 

Affecting the Species below for further 

information on both beach bark disease 

and the hemlock woolly adelgid. 

H. Acid Deposition Concerns 

Issue 1—Two commenters expressed 

concern about the effects of atmospheric 

acid deposition (also known as ‘‘acid 

rain’’) on WVNFS habitat, whereas one 

peer reviewer believed that such effects 

were largely speculative. 

Response—The Service agrees with 

the peer reviewer that such effects are 

largely speculative. Acid deposition is 

not a significant threat to the 

subspecies’ habitat. See Factor E under 

the Summary of Factors Affecting the 

Species section below for further 

details. 

I. Climate Change Concerns 

Issue 1—All peer reviewers agreed 

that the impacts of climate change on 

WVNFS are unclear. Whereas four peer 

reviewers concluded that measurable 

effects to WVNFS were not foreseeable, 

two concluded that the risk to WVNFS 

could not be discounted and requested 


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further analysis. Likewise, a coalition of 

commenters requested a more thorough 

analysis of the effects of global warming 

on WVNFS. They provided the Service 

a list of references to consider and 

submitted several unpublished maps of 

bioclimatic models for northern flying 

squirrels provided by Lawler (2007a, 

unpub. maps). This coalition of 

commenters believes that global 

warming is probably the greatest threat 

to WVNFS existence within the next 

100 years and likely will result in 

extinction of the WVNFS. 

Response—The Service has reviewed 

all evidence on climate change provided 

by the peer reviewers and members of 

the public, including references cited by 

the commenters, as well as others. 

While the Service acknowledges the 

general scientific consensus that global 

scale increases in temperature have 

occurred and are expected to continue 

into the future, we disagree with the 

commenters’ speculation that these 

changes will drive the WVNFS to 

extinction. Our ability to foresee 100 or 

more years into the future is limited by 

the current lack of reasonably accurate 

(Botkin et al. 2007, pp. 227–234; Meyers 

2008) climate change projection models 

localized for the range of the WVNFS, 

and simple stochastic events over such 

a long timeframe. 

Issue 2—Some commenters were 

concerned about the effect of climate 

change on interactions of WVNFS with 

the southern flying squirrel. They point 

to regional climate change studies 

projecting an increase in potential mast 

(nut producing) trees as evidence that 

the southern flying squirrel will 

outcompete WVNFS. In contrast, one 

peer reviewer noted that future climate 

conditions are unknown. He noted that 

hotter, drier summers but wetter, 

snowier winters might have little effect, 

or even a positive effect, on WVNFS if 

vegetation conditions remain 

unchanged, but wetter, snowier winters 

were less favorable for southern flying 

squirrels. 

Response—Bowman et al. (2005, pp. 

1486, 1490) speculated that southern 

flying squirrels in Canada had expanded 

their northern geographic range in 

response to climate warming between 

1994 and 2002, followed by a 

population crash in 2003 that resulted 

from an energetic bottleneck created by 

the combination of a cold winter that 

was preceded by a failed mast crop. 

They hypothesize that southern flying 

squirrels have the opportunity to 

expand their range northward during 

these warm periods, but acknowledge 

that there also is the possibility of large 

range contractions during cold spells 

(Bowman et al. 2005, p. 1491). They 

conclude that continued range 

expansions of southern flying squirrels 

are likely under continued global 

warming, although they expect that 

these expansions will be limited by the 

distribution of mast trees (Bowman et 

al. 2005, p. 1492). 

It is important to realize that 

projections about potential northward 

advance of oak forests in response to 

climate change relate to the potential 

distribution of suitable habitat wherein 

oaks could grow, not the actual 

distribution of the tree species. It is 

speculation that tree species will 

continue to move north because there 

are no barriers or constraints to 

migration (Hansen et al. 2001, p. 771). 

Iverson et al. (2004a, p. 787–799; 

2004b, pp. 209–219) investigated 

potential colonization of new suitable 

tree-species habitat under climate 

change for five eastern U.S species, 

including red oak. The results show the 

generally limited nature of likely 

migration over the first 100-year period 

following climatic change (Iverson et al. 

2004b, p. 216). They estimate that the 

proportion of new habitat that might be 

colonized within a century is low (15 

percent) for all five tree species, 

suggesting that there is a substantial lag 

between the potential movement of 

suitable habitat and the potential for 

tree species to migrate into the new 

habitat (Iverson et al. 2004a, p. 795). 

There is a relatively high probability of 

colonization within a zone of 10–20 km 

(depending on habitat quality and 

species abundance) of the current 

boundary, but a small probability of 

colonization as the distance from the 

current boundary exceeds about 20 km 

(Iverson et al. 2004b, p. 216). 

Looking at historical patterns, 

Schwartz et al. (2001, pp. 570, 574) and 

Iverson et al. (1999, Figure 7 on p. 89) 

predicted that migration rates of 1 to 10 

km/century might be the maximum 

future rates of tree colonization in 

fragmented habitats. Considering that 

the distribution of the WVNFS spans 

>170 km, it would take centuries for 

such potential shifts in oak species 

composition to materialize over a 

substantial portion of the range of 

WVNFS. Such slow colonization rates 

increase the likelihood that should red 

spruce decline significantly as a result 

of climate change, WVNFS would be 

able to survive in refugia of red sprucenorthern hardwood habitats (as 

projected by Delcourt and Delcourt 

1998, p. 927) and shift its range in 

response to similar slow, potential 

changes in southern flying squirrel 

distribution. 

Issue 3—Some commenters were 

concerned that the risk of wildfires 

would increase as a result of more 

frequent droughts, and thus would pose 

a threat to WVNFS. 

Response—Historically, natural fires 

in the Central Appalachians are 

believed to have been ‘‘relatively 

unimportant in the past, and to remain 

unimportant today, because of the wet 

weather that usually accompanies 

lightning’’ (Lafon et al. 2005, p. 129). 

Anthropogenic fires have played some 

role in the Central Appalachians for 

centuries as Native Americans used fire 

to drive game, improve wildlife habitat, 

maintain open meadows, and clear 

underbrush (Van Lear and Waldrop 

1989, pp. 1–2; Delcourt and Delcourt 

1997, p. 1013). European settlers also 

practiced widespread burning (Van Lear 

and Waldrop 1989, p. 3). As discussed 

by Weigl (2007, p. 898), wildfires 

ravaged the landscape during the period 

of industrial logging. Loggers set fires 

after clearcutting, and additional fires 

were ignited from sparks from the 

logging trains (Schuler et al. 2002, p. 

89). The fires associated with the 

logging practices of the early 1900s are 

not expected to reoccur, because the 

clearcutting is no longer taking place. 

While other parts of the Central 

Appalachians are currently considered 

to be especially fire-prone, the 

Allegheny Plateau, which contains most 

of the WVNFS habitat, is considered as 

‘‘having limited fire activity’’ (Lafon et 

al. 2005, p. 141). It is clear that fire has 

played some role in development of the 

current ecosystem for many centuries. 

Since climate appears to have a strong 

influence on fire regimes, potential 

climate changes will influence the 

number of fires, the area burned, and 

fire intensity (Lafon et al. 2005, p. 140). 

While there is the potential for 

occurrence of more frequent and intense 

fires during drought, there is also 

potential during wetter climatic periods 

for decreased fire activity. There are no 

scientific means, however, of accurately, 

or reasonably determining the net effect 

on WVNFS and its habitat of any 

potential change in the fire regime that 

may occur over the next century. While 

a long-term regime of intense, landscape 

level fires could significantly impact 

WVNFS habitat, those potential 

conditions are mere speculation given 

our present state of knowledge. 

Issue 4—Some commenters requested 

that the Service specifically review the 

potential contribution of global warming 

to the ‘‘recent’’ condition of red spruce, 

as described in several papers from the 

late 1980s [McLaughlin et al. 1987; 

Johnson et al. 1988; and Hamburg and 

Cogbill 1988 (miscited as Cogbill 1988 

by the commenters)]. They stated that 

the Service should fully examine all 


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studies of red spruce condition and 

factors contributing to that condition. 

Response—The Service has reviewed 

the three papers from the 1980s cited by 

the commenters, as well as other studies 

of red spruce condition. The three cited 

papers primarily focus on the northern 

and southern Appalachians, areas that 

are outside the range of the WVNFS. 

Although not directly applicable to 

WVNFS, papers covering areas outside 

the range of the WVNFS do provide a 

context for observed differences in 

regional trends of red spruce condition. 

The Service further examined potential 

impacts on the current and future 

condition of the red spruce-northern 

hardwood ecosystem in Factors A and E 

under the Summary of Factors Affecting 

the Species section of this final rule, as 

well as the responses to comments on 

these issues. 

Issue 5—Some commenters cited a 

paper by Delcourt and Delcourt (1998) 

as specific evidence that we can foresee: 

(1) The extirpation of red spruce-balsam 

fir and spruce-Fraser fir forests south of 

44 degrees north latitude (the White 

Mountains, New Hampshire); and (2) 

the movement of the southern range of 

these forests to northern New England 

in the next 100 years. 

Response—We have reviewed the 

paper by Delcourt and Delcourt (1998) 

cited by the commenters. We believe the 

projections made by these authors are 

likely overestimates of risk. First, the 

authors did not model the full range of 

possible climate extremes and did not 

use a full array of different climate 

models. They modeled possible future 

shifts in the spruce-fir ecotone from two 

climate models, assuming projected 

summer warming of 3.0 degrees or 6.4 

degrees Celsius (Delcourt and Delcourt 

1998, p. 926). The Intergovernmental 

Panel on Climate Change (IPCC) (2007c, 

Table 3.1) currently recognizes a 2 to 4 

degree Celsius increase by 2099 as the 

best estimate of warming under six 

climate models (with a likely range of 1 

to 6 degrees). Values substantially 

higher than 4 degrees cannot be 

excluded, but agreement of models with 

observations is not as good for those 

values (IPCC 2007c, part 2.3). Thus the 

3-degree warming scenario, identified 

by the Delcourts as their ‘‘most 

conservative’’ projection, falls within 

the middle range of the best estimate of 

temperature changes currently 

recognized by the IPCC. The Delcourts 

did not model the lower range (1–2 

degrees) of warming currently 

recognized by the IPCC. 

Second, the Delcourts provided little 

information about model assumptions 

and limitations, and did not attempt to 

validate their model, all of which 

greatly diminishes the usefulness of this 

paper. They did not quantify tree 

species extinction probabilities, or 

otherwise explain the basis for 

qualitative statements about their 

confidence in their predictions. Botkin 

et al. (2007, p. 231) notes that the type 

of niche-theory model used by the 

Delcourts is likely to overestimate the 

risk of tree species extinction. These 

types of models assume that observed 

distributions of trees are in equilibrium 

with their current environment, and that 

the tree species will become extinct 

outside of the regional values (Botkin et 

al. 2007, p. 231). However, local 

variation in climate due to topography 

or other factors could result in tree 

species being able to persist in suitable 

microhabitats even though the model 

projects no suitable habitat in these 

general regions (Hansen et al. 2001, p. 

765). The Delcourts focused on 

elevation and summer temperature as 

the primary factors controlling where 

spruce-fir could grow, but other factors 

would likely add considerable 

uncertainty, such as: the seasonality of 

precipitation, duration of cloud cover in 

the growing season, winter temperatures 

and frost-free chronologies, and sitespecific disturbances (White and Cogbill 

1992, pp. 4–16). 

The Delcourts suggested possible 

northern and upslope migration of red 

spruce under both a 3- and 6-degree 

warming scenario, with greater impacts 

occurring under the warmer scenario. 

For the moderate 3-degree warming 

scenario, the authors also suggested the 

possibility of spruce survival in refugia 

(Delcourt and Delcourt 1998, p. 928, 

Figure 4), similar to what happened 

during warmer and drier extremes of the 

post-glacial period 4000–5000 years 

before present (Delcourt and Delcourt 

1998, p. 927, Ware 1999, pp. 45–55) 

under similar temperature regimes. 

Although Delcourt and Delcourt 

(1989) modeled summer temperature 

changes through 2100, they provided no 

time frame for when vegetative 

responses would likely occur. They also 

did not provide any prediction of what 

the tree species composition would be 

in the forest that would succeed the 

spruce-fir forest in each of these 

scenarios. As discussed in climate 

change Response to Comments Issues 1–

3, future vegetative changes in response 

to such temperature changes could 

possibly occur over several hundred 

years. However, their possible impacts 

on WVNFS distribution and persistence 

are not reasonably foreseeable given the 

long time frames and high degree of 

uncertainty. Therefore, we do not find 

that the projections of Delcourt and 

Delcourt (1989) present a climate 

change threat to the WVNFS’ habitat 

that is likely to endanger the subspecies 

in the foreseeable future. 

J. Spruce Restoration Concerns 

Issue 1—Whereas one peer reviewer 

commented that restoration techniques 

have the ability to hasten improved 

overstory conditions and compositions 

favorable to WVNFS, some members of 

the public were concerned that spruce 

restoration efforts are misdirected and 

would not be successful. These 

commenters state there is only one 

master’s level study suggesting that such 

recovery may be feasible. 

Response—Forest management and 

silvicultural techniques, such as those 

being proposed, have long histories of 

implementation (Frank and Bjorkbom 

1973, pp. 1–29; Frank and Blum 1978, 

pp. 1–15; Carey et al. 1999, pp. 64–66). 

Several studies and modeling 

simulations indicate that restoration 

silviculture could be an effective tool for 

increasing the amount and quality of red 

spruce-northern hardwood forests in the 

central Appalachians (Rentch et al. 

2007, pp. 440–452; Schuler et al. 2002, 

pp. 88–98; Hornbeck and Kochenderfer 

1998, pp. 197–202). 

Efforts to restore or enhance red 

spruce-northern hardwood forests 

should in many cases enhance WVNFS 

habitat in the short-term, as well as the 

long term. For example, noncommercial 

efforts that involve red spruce release by 

girdling or stem-injection of herbicide 

will create snags suitable for day dens. 

In addition, removal of hard mast 

species such as northern red oak or 

American beech will lessen habitat 

suitability for the southern flying 

squirrel and therefore minimize any 

potential competition for dens and food, 

as well as lessen interspecific contact to 

spread the Stronglyoides parasite. See 

Factor C under the Summary of Factors 

Affecting the Species section below for 

additional information. 

And lastly, red spruce-northern 

hardwood restoration on the MNF is 

targeted at maximizing patch size and 

habitat connectivity for WVNFS (USDA 

Forest Service 2006a, p. III–14). These 

efforts are proceeding cautiously in 

unoccupied habitat, with monitoring to 

gauge success. These efforts are not 

clear cuts, but rather are light thinnings 

of northern hardwoods that open the 

canopy to provide additional light for 

growth of spruce. Spruce is naturally 

adapted to regeneration in small 

openings such as these. 

K. Overutilization Concerns 

Issue 1—One public commenter was 

concerned that once the WVNFS was 

delisted, its collection would no longer 


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be regulated and the subspecies would 

be threatened by overcollection. In 

contrast, two peer reviewers offered 

evidence that overutilization of WVNFS 

never has been a threat and would not 

become a threat should WVNFS be 

delisted. 

Response—Even for trained wildlife 

professionals, the WVNFS is an 

exceptionally difficult animal to catch. 

Thus the probability that a layman or 

commercial collector could capture or 

overcollect WVNFS is very remote given 

the subspecies’ low detectability, 

nocturnal and secretive habits, and 

remote localities where it occurs. Once 

delisted, WVNFS collection by hunting 

or trapping will still be illegal under 

West Virginia and Virginia state laws 

(West Virginia Code 20–2–5(26); Code of 

Virginia 29.1–521.A.10, 29.1–566 and 

29.1–530.A.), and its capture for 

scientific and educational purposes will 

still be regulated through collection 

permitting systems of the WVDNR (West 

Virginia Code 20–2–50) and the Virginia 

Department of Game and Inland 

Fisheries (Code of Virginia 29.1–568). 

For more information, see Factor B in 

the Summary of Factors Affecting the 

Species below. 

L. Adequacy of Regulatory Mechanisms 

Issue 1—Some commenters stated that 

much of the habitat believed to be 

important to the WVNFS is not fully 

protected in the long term. 

Response—There are no known 

rangewide threats to the subspecies’

forested habitat, thus full protection of 

this habitat is not required to maintain 

the WVNFS’s status as recovered. (See 

Factor A under the Summary of Factors 

Affecting the Species section for further 

details.) Seventy-nine percent of the 

modeled habitat of the WVNFS is being 

managed for the long term by provisions 

of forest plans, state management plans, 

wilderness and backcountry recreation 

designations, and conservation 

easements. For example, in the forest 

plan for the MNF (USDA Forest Service 

2006a, p. III–9, D–1), Management 

Prescription 4.1 focuses on protection, 

restoration and management of red 

spruce and red spruce-northern 

hardwood communities. This 

management prescription, as well as 

other management plans and 

agreements on state, Federal, and 

private lands, wilderness and 

backcountry recreation designations, 

and perpetual conservation easements 

will continue to apply following 

delisting of the WVNFS (Service 2007a, 

pp. 5–10). Collectively, all of these 

mechanisms provide reasonable 

certainty of protection and management 

of much of the habitat for WVNFS. 

Issue 2—Some commenters requested 

clarification on the status of forest plans 

for the Monongahela and the George 

Washington National Forests. These 

commenters were concerned that the 

Forest Service would not be able to 

implement these plans because of a 

lawsuit on the land management 

planning rule published in 2005. 

Response—In March 2007, a U.S. 

District court order enjoined the Forest 

Service from implementation and use of 

the land management planning rule 

published in 2005 until the Forest 

Service complied with the Court’s order 

for two combined cases (Citizens for 

Better Forestry et al. v. USDA and 

Defenders of Wildlife v. Johanns, C.A. 

C05–1144 (N.D. Cal.)). The Forest 

Service complied in 2008 by re-issuing 

its forest planning regulations. Forest 

plans currently in effect for the MNF 

and the George Washington National 

Forest (GWNF) were based on planning 

rules published prior to 2005; hence, 

their continued implementation and use 

in present form is not affected by the 

lawsuit or the new regulations. These 

existing plans provide guidance for 

management and monitoring of the 

WVNFS and its habitat, including 

prescriptions, goals, objectives, 

standards, and guidelines. Any 

subsequent revisions or amendments to 

these existing plans will require 

compliance with any planning 

regulations in effect at the time. Should 

the MNF choose to revise or amend 

their existing forest plan, we believe it 

is highly unlikely that the current 

WVNFS habitat would be affected (See 

the Factor A—Land Use Planning 

section under the Summary of Factors 

Affecting the Species below for further 

information). 

Issue 3—Some commenters noted the 

Service had entered into a 

Memorandum of Understanding (MOU) 

with the Forest Service, WVDNR, and 

others for continued management and 

protection of WVNFS. These 

commenters question whether the 

Forest Service would continue to 

protect a species without the force of 

law. 

Response—By signing the MOU, 

signatories demonstrate that they are 

committed to implementing the features 

within their discretion and authority 

(Service et al. 2007, pp. 1–8). The MOU 

affirms commitments made by the U.S. 

Forest Service MNF to implement 

standards and guidelines for the 

WVNFS and its habitat contained in the 

2006 Land and Resource Management 

Plan. This plan would not be 

invalidated by delisting the WVNFS. 

Also see response to section F,. 

Ecosystem and Habitat Concerns—Issue 

5 and section L,. Adequacy of 

Regulatory Mechanisms—Issue 2, above. 

Issue 4—Some commenters 

questioned the ability of the Forest 

Service, WVDNR, and others to fulfill 

their obligations in the MOU, given 

projected staff and budget cuts. 

Response—The Service is not relying 

upon the MOU as an enforceable 

regulatory mechanism under the Act. 

See Response to Issue 3 in this same 

subsection above. 

Issue 5—Some commenters were 

concerned that the MOU termination 

clause allows parties to opt out for any 

reason with 30 days’ notice. 

Response—MOUs commonly have 

early termination clauses. While some 

changes to the composition of the 

signatory parties to the MOU may occur 

over time, we expect that other parties 

will sign on and the MOU will continue 

to be implemented for the long-term by 

those participating at the time. See 

Response to Issues 3 and 4 in this same 

subsection above. 

M. Predator Concerns 

Issue 1—One commenter noted the 

Service had not discussed the impact on 

WVNFS of the reintroduction of the 

fisher, a potential predator on WVNFS. 

Response—Fishers (Martes pennanti) 

were reintroduced to West Virginia in 

the late 1960s or early 1970s, prior to 

the listing of WVNFS as endangered. 

Both animals have shown overlapping 

range expansions in the intervening 

decades, providing indirect evidence 

that fishers are not significant mortality 

agents for WVNFS. Most data from the 

eastern United States suggest that 

snowshoe hare, cottontails, voles, mice, 

and bird eggs comprise the majority of 

the fisher’s diet (Powell et al. 2003, p. 

643). Weigl (2007, p. 901) concluded 

that fishers probably can coexist with 

northern flying squirrels, with the 

exception of in small habitat islands, 

where there are fewer WVNFS and other 

prey is more limited. 

N. Other Natural Factors 

Issue 1—One peer reviewer and one 

public commenter thought the Service 

needed to give more consideration to 

the impact of parasites on WVNFS 

spread by southern flying squirrels 

(Glaucomys volans), given projections 

about climate change, acid deposition, 

oak decline in northern hardwood 

communities, and expansion of other 

seed- and nut-bearing hardwoods. 

Response—Recognizing that there are 

‘‘varying intensities’’ of parasitic 

infection of northern flying squirrels (G. 

sabrinus) in the wild, Weigl (2007, p. 

901) remains concerned about infection 

of G. sabrinus by the intestinal parasite 


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Stonglyoides robustus, based in part on 

his belief that there has never been 

stable sympatry of G. sabrinus and 

Stonglyoides robustus. While that may 

be true for the Carolina northern flying 

squirrel (G. s. coloratus) at the sites he 

studied in North Carolina and 

Tennessee, stable sympatric occurrences 

of WVNFS and the southern flying 

squirrel (G. volans) have been 

documented for decades at the Spruce 

Knob geographic recovery area in West 

Virginia (Wallace 2007, p. 2). The 

southern flying squirrel has been 

detected within all 7 of the generalized 

WVNFS core areas (or population 

centers), and at 20 percent of the 109 

WVNFS capture sites. Despite the 

presence of this competing species, 

there is no evidence of illness or 

mortality of WVNFS, and no evidence of 

local extirpation of WVNFS from any of 

these sites during 21 years of 

monitoring. Based on their documented 

co-occurrence in West Virginia and 

Virginia, and no documented lethal 

effects in the wild, we believe that 

speculation that impacts of climate 

change, acid deposition, or shifts in 

forest composition would decrease the 

fitness or survival of the WVNFS is 

unwarranted. The WVNFS has prevailed 

in repopulating its range in a habitat 

where the red spruce-northern 

hardwood compositions arguably favor 

the southern flying squirrel over the 

past 100 years. The Service does not 

believe that the WVNFS would have 

made this recovery if it suffered 

debilitating or lethal effects from 

sympatric relationships with parasitebearing species (See Factor E—

Competition with Southern Flying 

Squirrel under the Summary of Factors 

Affecting the Species section for further 

information). 

O. Miscellaneous 

Issue 1—Some commenters were 

concerned that we have ignored the 

WVNFS recovery plan criteria in 

determining that the subspecies has 

recovered. 

Response—As summarized above in 

the Recovery section of this final rule, 

our analysis shows that the intent of 

each criterion for downlisting and 

delisting has been satisfied and that 

most of the criteria have been achieved 

or substantially achieved. Although the 

recovery plan criteria are out-of-date, we 

conducted an analysis of how well these 

criteria have been met and summarized 

that analysis in the beginning of this 

final rule. New information has changed 

the extent to which these criteria need 

to be met for recognizing recovery of the 

subspecies. Species are listed or delisted 

under the Act based on whether they are 

threatened or endangered by one or 

more Factors (see Summary of Factors 

Affecting the Species section below). 

Up-to-date, threats-based recovery 

criteria can assist the Service in 

analyzing whether a species meets the 

definition of threatened or endangered. 

Recovery criteria are only one tool, 

however, the Service uses in making a 

classification determination. 

Issue 2—Some commenters expressed 

concern about not providing a postdelisting monitoring plan for public 

review, concurrently with the proposed 

rule. 

Response—The proposed and final 

delisting decisions are based firmly on 

an analysis of identified threats and 

changes in the subspecies’ status. They 

are not legally contingent upon future 

approval or implementation of the postdelisting monitoring plan. The Act 

contains no explicit requirements for 

either notifications or public comment 

opportunities relative to planning or 

implementation of post-delisting 

monitoring plans. Nevertheless, the 

Service sought input into these 

processes, as indicated by our request 

for public comment on the draft postdelisting monitoring plan (72 FR 57346), 

published in the Federal Register on 

October 9, 2007, prior to publication of 

this final rule, and by our finalization of 

the plan concurrent with this final 

decision on the delisting proposal. 

Issue 3—Some commenters expressed 

mistrust about the motivations behind 

delisting and accused the Service of 

catering to developers, the timber 

industry, and other extractive resource 

users. Some commenters also expressed 

value-based reasons as to why they 

opposed delisting, such as spiritual 

importance, animal rights, and need for 

humans to behave as caretakers and 

stewards of the WVNFS, not as pillagers 

of its habitat. The majority of comments 

received were one of three various form 

letters stating that the proposed rule was 

premature and based on inadequate 

scientific information, but provided no 

substantive information to support these 

statements. 

Response—Our decision to delist 

WVNFS is based solely on the best 

scientific and commercial data available 

and our five-factor analysis. This 

analysis indicates that the subspecies is 

neither threatened nor endangered. 

While we appreciate the values 

expressed by these commenters, such 

comments are either not relevant to the 

decision, or are outside the scope and 

authority of the final rule. 

Summary of Factors Affecting the 

Species 

Section 4 of the Act and its 

implementing regulations (50 CFR part 

424) set forth the procedures for listing 

species, reclassifying species, or 

removing species from listed status. 

‘‘Species’’ is defined by the Act as 

including any species or subspecies of 

fish or wildlife or plants, and any 

distinct vertebrate population segment 

of fish or wildlife that interbreeds when 

mature (16 U.S.C. 1532(16)). Once the 

‘‘species’’ is determined, we then 

evaluate whether that species may be 

endangered or threatened because of 

one or more of the five factors described 

in section 4(a)(1) of the Act. We must 

consider these same five factors in 

delisting a species. We may delist a 

species according to 50 CFR 424.11(d) if 

the best available scientific and 

commercial data indicate that the 

species is neither endangered nor 

threatened for the following reasons: (1) 

The species is extinct; (2) the species 

has recovered and is no longer 

endangered or threatened; and/or (3) the 

original scientific data used at the time 

the species was classified was in error. 

A recovered species is one that no 

longer meets the Act’s definition of 

threatened or endangered. The analysis 

for a delisting due to recovery must be 

based on the five factors outlined in 

section 4(a)(1) of the Act. This analysis 

must include an evaluation of threats 

that existed at the time of listing, those 

that currently exist, and those that could 

potentially affect the species once the 

protections of the Act are removed. 

The Act defines ‘‘species’’ to also 

include any subspecies or, for 

vertebrates, any distinct population 

segment. The Act defines ‘‘endangered 

species’’ as any species which is in 

danger of extinction throughout all or a 

significant portion of its range, and 

‘‘threatened species’’ as any species that 

is likely to become an endangered 

species within the foreseeable future 

throughout all or a significant portion of 

its range. 

For the purposes of this finding, the 

‘‘foreseeable future’’ is the period of 

time over which events or effects 

reasonably can or should be anticipated, 

or trends reasonably extrapolated, such 

that reliable predictions can be made 

concerning the status of the species. As 

discussed in the Summary of Factors 

section, we determined that any future 

threat from development will be 

localized and minimal, based on trends 

over the past 10 years. In addition, the 

Service has no indications that 

management of the forest for timber will 

have more than a minor impact on the 


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WVNFS based on the discussion in 

Factor A. WVNFS habitat has been 

improving steadily for the past 50–80 

years throughout its range and we 

expect this improvement to continue 

into the future. 

Climate change projection models are 

not reasonably accurate for the localized 

range of WVNFS, and therefore we 

cannot reliably predict that climate 

change will pose a threat in the future. 

All indications suggest that the squirrel 

is resilient enough to adapt to and 

survive gradual changes in the habitat, 

if there are any due to climate change. 

Therefore, we do not foresee any threats 

affecting the WVNFS into the future that 

would lead the species to become an 

endangered species. 

Section 4(a)(1) of the Act requires that 

we determine whether a species is 

endangered or threatened based on one 

or more of the five following factors: (A) 

The present or threatened destruction, 

modification, or curtailment of its 

habitat or range; (B) overutilization for 

commercial, recreational, scientific, or 

educational purposes; (C) disease or 

predation; (D) the inadequacy of 

existing regulatory mechanisms; and (E) 

other natural or manmade factors 

affecting its continued existence. Our 

evaluation of these factors is presented 

below. Following this threats analysis, 

we evaluate whether the WVNFS is 

threatened or endangered within any 

significant portion of its range. 

A. The Present or Threatened 

Destruction, Modification, or 

Curtailment of Its Habitat or Range 

WVNFS Distribution 

At the time of listing (1985), 10 

WVNFS individuals were known from 

Randolph and Pocahontas Counties, 

WV, and Highland County, VA (Service 

2006a, p. 8). It was thought that vast 

stretches of unsuitable habitat separated 

the four known population centers and 

that the WVNFS still existed but that it 

was very rare, and perhaps no longer 

present in much of its former range (50 

FR 26999). The final listing rule 

qualitatively described historic habitat 

losses and suggested that ‘‘in these last 

occupied zones, the squirrels [G. s. 

fuscus and G. s. coloratus] and their 

habitat may be coming under increasing 

pressure from human disturbances such 

as logging and development’’ (50 FR 

26999). 

The current known range of WVNFS 

follows the spine of the high Allegheny 

Plateau in a northeast to southwest 

alignment. Helmick Run (Grant County, 

WV) marks the northeast periphery and 

Briery Knob (Greenbrier County, WV) 

the southwest periphery, covering seven 

counties in West Virginia and Highland 

County, Virginia (Service 2006a, p. 25). 

As of 2006, there is a total of 109 

WVNFS capture sites, of which 107 are 

in West Virginia and 2 are in Highland 

County, Virginia (Service 2006a, pp. 8 

and Figure 2; WVDNR 2006a, pp. 

1–109). These capture sites are 

dispersed across seven general areas of 

habitat in the Allegheny Highlands 

region (Service 2006a, pp. 9 and Figure 

3). Distributed throughout the 109 

capture sites, there have been 1,198 

captures (including 85 recaptures) as of 

2006 (WVDNR 2006a, pp. 1–109). 

Collectively, the proportion of sites 

demonstrating persistence across 

multiple generations (83 percent), 

distributed among habitat quality types 

and within geographic zones; the 

routine documentation of nestlings and 

juveniles (76 percent of sites); and 

balanced to slightly skewed sex ratios 

demonstrate a relatively high degree of 

population stability and constant habitat 

occupancy (Service 2007c, pp. 9–11). 

Locally reproducing populations are the 

most likely factors for continuing to find 

WVNFS in numerous locations within 

their historical range over the last 

couple of decades, given their low 

detectability, relatively short life span, 

and relatively low reproductive 

capacity, and a naturally patchy nature 

of suitable forest habitat distribution 

(Service 2007c, p. 11). 

We now know that the WVNFS 

continues to occupy the areas identified 

in the 1985 final listing rule (50 FR 

26999) as well as numerous additional 

sites dispersed throughout its historical 

range, suggesting that its current range 

roughly approximates the extent of its 

historical range. Studies have confirmed 

the ability of the WVNFS to adjust its 

foraging and denning behavior (i.e., the 

ability to nest in a wide variety of trees) 

to persist in and around red sprucenorthern hardwood forest patches 

(Menzel et al. 2004, pp. 360, 363–364; 

Menzel et al. 2006a, pp. 1–3, 6, 7; 

Menzel et al. 2006b, p. 208; Ford et al. 

2004, p. 430). 

Habitat Quantity and Quality 

Prior to European settlement, there 

were in excess of 500,000 ac (some 

sources suggest 600,000+ ac) of oldgrowth red spruce–northern hardwood 

forests, the preferred habitat of the 

WVNFS, in the Allegheny Highlands. 

These forests (occupying ridges, slopes, 

and drainages) in West Virginia 

extended from the vicinity of Mount 

Storm (Grant County) in the north to 

Cold Knob (Greenbrier County) in the 

south, east to the Allegheny Front 

(Pendleton County), and west to 

Webster and Nicholas Counties. These 

red spruce–northern hardwood forests 

were more contiguous across the 

Allegheny Highlands than are the wellknown ‘‘sky-islands’’ of the Southern 

Appalachians, which support Carolina 

northern flying squirrels (G. s. 

coloratus) (Service 1990, pp. 16–17; 

USDA Forest Service–Northern 

Research Station 2006, unpublished 

data, pp. 2–3). 

Logging activity and associated 

widespread fires at the turn of the 20th 

century decimated the red spruce–

northern hardwood forests, resulting in 

younger forests with less red spruce 

and, in many areas, a mixed mesophytic 

(moderately moist environment), oakdominated forest (Menzel et al. 2006b, 

p. 6; Rollins 2005, pp. 12–13; Schuler et 

al. 2002, pp. 88–89). Loggers set fires 

after clearcutting, and additional fires 

were ignited from sparks from the 

logging trains (Schuler et al., 2002, p. 

89). The fires associated with the 

logging practices of the early 1900s are 

not expected to reoccur, because the 

clearcutting is no longer taking place. 

These fires did, however, consequently, 

result in less, and poorer quality, 

WVNFS habitat because younger forests 

with fewer red spruce provided reduced 

foraging and sheltering opportunities 

(Service 2006a, p. 6). Also, the presence 

of oak and its associated mast (i.e., 

acorns), provided a competitive 

advantage of food resources for the more 

aggressive southern flying squirrel. The 

WVNFS’ rarity was understood to be a 

consequence of its specialized use of a 

precipitously declining habitat type 

(Service 2006a, p. 11). 

Currently, it is estimated that there 

are approximately 242,000 ac of WVNFS 

habitat (USDA Forest Service–Northern 

Research Station 2006, unpublished 

data, p. 4). This estimate is based in part 

on the results of several habitat models, 

and includes all ‘‘optimal’’ habitat as 

well as ‘‘likely’’ habitat located in close 

proximity to red spruce–northern 

hardwood forests. ‘‘Likely’’ and 

‘‘optimal’’ are terms and definitions 

imparted by the Menzel model, with 

‘‘likely’’ areas having a greater than 50 

percent chance of being occupied by the 

WVNFS, and ‘‘optimal’’ areas having a 

greater than 75 percent probability of 

being occupied (Menzel 2003, pp. 84–

85, 87–89; Menzel et al. 2006b, pp. 4–

5). The models allow us to estimate the 

amount of potential and high-quality 

habitat in the Allegheny Highlands, 

prioritize areas for restoration and 

recovery (Menzel et al. 2006a, p. 7), 

assess anthropogenic (manmade) and 

geologic fragmentation of the red spruce 

forest, and analyze stewardship of the 

suitable habitat (Menzel et al. 2006b, p. 

7). 


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The forested areas used by the 

WVNFS across most of its range have 

continued to mature in the 20 years 

since listing. For example, about half of 

the rangewide areas modeled as optimal 

habitat are red spruce-northern 

hardwood forest stands on the MNF that 

are over 75 years old (Menzel et al. 

2006b, p. 4; Service 2006a, pp. 10–11; 

USDA Forest Service-Northern Research 

Station 2006, unpublished data, p. 2). 

Even though current habitat conditions 

are not as favorable for the WVNFS as 

historical conditions preceding the late 

1800s/early 1900s, current conditions 

are much improved compared to those 

at the time of listing. With the exception 

of localized habitat impacts, forest 

succession has resulted in older forest 

stands with improved forest structure, 

reflecting a continuing positive 

rangewide trend (Service 2006a, pp. 11–

14, 19–20). With regard to forest 

composition, the amount and extent of 

red spruce in the Central Appalachians 

also appears to be gradually increasing 

(Adams et al. 1995, p. 101; Schuler et 

al. 2002, p. 92–93; Rollins 2005, pp. 39–

51). Recent evidence also suggests 

improving trends in health and 

regeneration of red spruce-northern 

hardwood forests within the range of 

WVNFS (Adams et al. 1995, p. 101; 

Audley et al. 1999, pp. 179–199; 

Hornbeck and Kochenderfer 1998, pp. 

198–200; Schuler et al. 2002, p. 92–94; 

Rollins 2005, pp. 74–78). The forested 

landscape within the range of WVNFS 

provides a high degree of functional 

connectivity, as evidenced by large 

patch sizes, numerous linkages, and 

persistence over multiple generations at 

monitoring sites across a range of forest 

conditions (Service 2007c, pp. 5–6, 9–

11). 

We analyzed impacts that the balsam 

and hemlock woolly adelgids, insect 

parasites accidentally introduced from 

Europe (Service 1990, p. 13), may be 

having on the WVNFS’ habitat (Service 

2006a, p. 17). The balsam woolly 

adelgid infects balsam fir (Abies 

balsamea) trees, causing damage or 

mortality to the host trees (Service 1990, 

p. 13). However, we believe the effect of 

the balsam woolly adelgid on WVNFS 

habitat is discountable because balsam 

fir is limited to a minor component of 

the WVNFS habitat (Peart et al. 1992, p. 

149, 165). Red spruce occurs in or near 

stands of balsam fir, providing the 

WVNFS with alternative and higher 

value habitat where damage from the 

balsam woolly adelgid may have 

occurred. In addition, the impact of the 

balsam woolly adelgid on the small 

component of balsam fir within WVNFS 

habitat has already occurred (Service 

2006a, p. 17). 

The hemlock woolly adeglid has been 

in the United States since 1924. The 

insect damages eastern hemlock (Tsuga 

canadensis) trees by damaging new 

growth, which can cause defoliation and 

mortality (Service 2006a, p. 17). Only 7 

percent of the WVNFS capture sites are 

dominated by Eastern hemlock instead 

of red spruce (Service 2006a, p. 17). 

However, work conducted on the 

WVNFS indicates that hardwood forests 

with little or no conifer component are 

not barriers to movement (Menzel et al. 

2006a, p. 207). While hemlock woolly 

adelgid may remove the montane 

conifer component at less than 10 

percent of the known capture sites, 

most, if not all, of these areas are in 

close proximity to red spruce-northern 

hardwood forests, significantly reducing 

the occasions where loss of Eastern 

hemlock could be detrimental to the 

WVNFS (Service 2006a, p. 17). 

Additionally, the West Virginia 

Department of Agriculture has an active 

detection program for hemlock woolly 

adelgid and a treatment program that 

will remain in place regardless of the 

listing status of the WVNFS. Therefore, 

even though the hemlock woolly 

adelgid may impact a minor component 

of the squirrel’s habitat, we consider it 

to pose a negligible degree of risk to the 

WVNFS, because of the limited role of 

hemlock in the subspecies’ survival, and 

presence of red spruce in the majority 

of the areas (Service 2006a, p. 17). 

The potential impact of beech bark 

disease was also analyzed. Beech bark 

disease is caused by the beech scale 

insect (Cryptococcus fagisuga), followed 

by one of two fungi (Nectria coccinea 

var. faginata or N. galligena). The scale 

stresses and weakens the American 

beech tree (Fagus grandifolia) and the 

fungi then cause either localized lesions 

or decay and death of the entire tree 

(Service 2006a, pp. 17–18). Although 

American beech trees are common to 

the red spruce-northern hardwood 

forests of the Allegheny Highlands, in 

WVNFS habitat they usually occur in 

combination with red spruce and other 

hardwoods, particularly birch and 

maple. Therefore, despite having a 

devastating impact on the American 

beech component of the red sprucenorthern hardwood forest, beech bark 

disease is not thought to render WVNFS 

habitat unsuitable (Service 2006a, p. 

18). There is actually a potential shortterm benefit to the WVNFS due to the 

creation of new nest cavities in the 

holes of dead and decaying beeches. 

Foraging habitat for the WVNFS may 

also improve with increases in large 

woody debris on the forest floor from 

the dead beech trees, which could 

promote the growth of underground 

fungi, one of the WVNFS’ primary food 

sources (Carey et al. 1999, p. 54; Pyare 

and Longland 2001, p. 1008; Rosenberg 

and Anthony 1992, p.161; Waters et al. 

2000, p. 85). Additionally, the removal 

of beech nuts is thought to be more 

detrimental to the southern flying 

squirrel because it is a high-energy food 

source for that species, and, therefore, 

would counter any small amount of 

direct competition between the WVNFS 

and the southern flying squirrel. 

Therefore, while beech bark disease 

affects a minor component of WVNFS 

habitat rangewide, we consider it to 

pose an overall low-to-moderate degree 

of risk for WVNFS, and this risk may be 

offset by the potential benefits of 

creation of new nest cavities, increase in 

a primary food source, and potential 

harm to the food supply of the southern 

flying squirrel (Service 2006a, p. 18). 

We also analyzed the potential future 

impacts of climate change on the 

WVNFS’s habitat. While there is much 

speculation on potential future impacts 

of climate change on the WVNFS, it is 

important to recognize that there is no 

evidence that climate changes observed 

to date have had any adverse impact on 

WVNFS or its habitat. For example, 

within the range of the WVNFS, 

inexplicable crown dieback (Mielke 

1987, pp. 221–222) and declines in red 

spruce radial growth were reported in 

the 1980s (Adams et al. 1985, p. 315). 

Since the 1980s, there has been no 

evidence of widespread crown decline 

of red spruce throughout the range of 

WVNFS. By the late 1990s, Audley et al. 

(1998, pp. 177, 180, 190) noted that 

while a small percentage of individual 

trees sampled exhibited symptoms of 

reduced health and vigor, the majority 

of red spruce sampled in West Virginia 

appeared healthy. More recent 

dendrochronological surveys of red 

spruce stands in West Virginia detected 

this growth decline phenomenon 

occurring from about 1930 to 1990 

(Schuler et al. 2002, p. 93; Hornbeck 

and Kochenderfer 1998, pp. 199–200). 

Since this time period, the decline 

appears to have ended in the central 

Appalachians—growth rates have 

leveled or shown slight increases 

(Schuler et al. 2002, p. 93, figure 3; 

Hornbeck and Kochenderfer 1998, p. 

199–200) and regeneration remains 

unaffected (Schuler et al. 2002, pp. 92–

93). 

Red spruce is now recolonizing areas 

of hardwood forest near existing red 

spruce stands, areas that historically 

were red spruce until the logging and 

fires at the turn of the 20th century 

(Schuler et al. p. 2002, p. 89). There is 


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evidence that the red spruce-northern 

hardwood ecotone is either stabilizing 

or decreasing in elevation (expanding) 

to approximate its former extent (Adams 

et al. 1999, p. 235, Rollins 2005, p. 76). 

Rollins (2005, p. ii, 74–75) found that 

the amount and quality of red spruce at 

three study sites in the central 

Appalachians appeared to be gradually 

improving through natural regeneration. 

Since then, Rollins has studied 9 

additional sites, for a total of 12 

representative sites distributed in the 

northern, central, and southern portions 

of the range of WVNFS. Stand data on 

trees, saplings and seedlings, soil 

chemistry, red spruce foliar chemistry, 

and the percent of red spruce roots 

covered by symbiotic fungal 

mycorrhizae are currently being 

analyzed between two sampling periods 

(1985 vs. 2005). Although a final report 

is not yet available, preliminary results 

indicate a reversal of the crown dieback 

conditions observed in 1985 (Connolly 

2007). 

This pattern is contrary to general 

projections that climate changes in the 

next 100 years may shift the geographic 

ranges of flora and fauna upwards in 

elevation and northward (IPCC 2002, p. 

1). Considering the ecotone range 

expansion trends documented by 

Rollins (2005) and Adams et al. (1999), 

we expect that the extent and quality of 

the habitat for WVNFS is likely to 

continue to increase. 

We looked at the possible range of 

effects of climate change on the 

WVNFS. Under warmer scenarios, 

several regional models project that 

mixed (hardwood and conifer) forests in 

the northeastern United States 

(including West Virginia and Virginia) 

may decrease in potential area, as they 

gradually shift into Canada over the 

next 100 years or more. By some 

projections, this possible decrease in 

potential habitat could be as small as 

¥11 percent to ¥22 percent (Iverson et 

al. 2005, p. 34) or as large as ¥97 

percent over 100 years (Hansen et al. 

2001, p. 769). These models also project 

that northeast mixed hardwood and 

conifer forests may gradually be 

squeezed from the south by the advance 

of southeastern mixed forests to varying 

degrees (Inkley et al. 2004, p. 6). 

However, some models project that the 

biome remains intact under cooler 

scenarios (Hansen et al. 2001, p. 769; 

Inkley et al. 2004, p. 6, figure 3). As 

explained by Botkin et al. (2007, p. 230), 

‘‘the larger the scale of the primary units 

of the model, the simpler it is to 

estimate effects over large areas and 

times, but also the cruder the 

approximation is and the more likely 

that undesirable assumptions will 

prevail.’’ Given this caveat, as well as 

the huge variation in possible views of 

the future noted above, all with 

unknown likelihoods of occurrence, we 

conclude that it is not possible to 

translate these potential scenarios into 

potential effects on WVNFS or its 

habitat over any meaningful timeframe. 

We considered the map products 

provided by some of the public 

commenters (Lawler 2007a, unpub. 

maps). We spoke to Dr. Joshua Lawler 

(2007b), University of Washington, to 

gain a better understanding of the 

continentwide bioclimatic models he 

ran for all subspecies of northern flying 

squirrels. These models do not map 

vegetation directly, but attempt to do so 

indirectly by correlating the distribution 

of the various subspecies of northern 

flying squirrels to alternative scenarios 

of climate change. For the WVNFS, the 

models project that the future climate 

(2071–2100) within the range of the 

subspecies will be different from the 

baseline climate conditions of 1961–

1990. Contrary to the commenters’

speculation that these products project 

the extinction of WVNFS, the 

unpublished map products (Lawler 

2007a, unpub. maps) provided by the 

commenters indicate only an 

unquantified potential range contraction 

of WVNFS. Botkin et al. (2007, p. 231) 

notes that bioclimatic models vary 

greatly in their projections of extinction, 

and that Lawler et al. (2006) have not 

attempted to validate any of the models 

they are using. Lawler et al. (2006, p. 

1579) recognized that it would be 

difficult to translate these types of 

predictions into threats of extinction 

because actual range shifts would 

depend on dispersal, evolutionary 

flexibility, and species interactions. Dr. 

Lawler (2007b) stated that the model 

had a good degree of fit at the 

continentwide level, but the fit would 

be reduced, and the degree of 

uncertainty would be expected to be 

higher, at the State level. He indicated 

it is not possible to determine model 

error for the relatively small scale of the 

WVNFS’ range in West Virginia and 

Virginia. 

The WVNFS and other subspecies of 

G. sabrinus have demonstrated 

significant adaptability, resilience, 

mobility, and plasticity in habitat use by 

surviving landscape-level habitat 

changes during times of glacial retreat 

and advance during the Pleistocene, and 

by surviving intense landscape-level 

loss of forest during the late 1800s and 

early 1900s (Weigl 2007, p. 898). Over 

the past 100+ years, the WVNFS 

survived a change from a red sprucedominated forest to a loss of much of 

the forest habitat, to a transitional 

regeneration of a hardwood-dominated 

forest, and a more recent increase in the 

red spruce component. As several 

commenters point out, hardwood trees 

have always been an important 

component of WVNFS habitat and there 

is no evidence that a gradual increase in 

hardwoods would cause dramatic 

population declines for the WVNFS. In 

fact, Weigl (2007, p. 899), citing two 

other studies in the northeast, noted that 

‘‘the species is known to occupy 

hardwood habitat without spruce or 

fir.’’ The Service concludes that the 

WVNFS is expected to survive slow, 

gradual changes from long-term climate 

change. 

Based upon a review of the current 

scientific studies, peer-review 

comments, the unpublished maps 

provided by the commenters, and 

discussions with modelers, the Service 

concludes that there is no evidence that 

current changes in climate have had an 

adverse impact on WVNFS. Long-term 

projections about climate change and its 

possible effects on WVNFS are complex 

and best viewed as possible alternative 

views of the future that have unknown 

likelihoods of occurrence. Therefore, 

based on the above information, we 

have determined that we are unable to 

establish climate change as a threat to 

the WVNFS within the foreseeable 

future. 

Land Use Planning 

Available information indicates that 

the threat posed by past habitat loss has 

been largely abated across most of the 

WVNFS’ range. Implementation of the 

2001 recovery plan amendment (Service 

2001, p. 4) and the 2004 amendment to 

the MNF Land and Resource 

Management Plan (USDA Forest Service 

2004, pp. 84a–84c, 87, 234–234b) 

significantly removed the threat of 

habitat loss (via logging) across much of 

the WVNFS’ range. The recovery plan 

amendment recommended that suitable 

WVNFS habitat be considered during 

consultation with Federal agencies. The 

Forest Service reinforced this 

recommendation through an 

amendment to the MNF Land and 

Resource Management Plan, which 

limited vegetation management in all 

‘‘suitable habitat’’ (as determined 

collaboratively by the Forest Service, 

Service, and WVDNR) to: (1) Research 

activities covered under an Act section 

10 permit; (2) actions to improve or 

maintain WVNFS populations after 

research has demonstrated the 

beneficial effects of the proposed 

management; or (3) when project-level 

assessment results in no adverse effects. 

This conservation strategy has been 

carried forward into the MNF’s recent 


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Forest Plan Revision (USDA Forest 

Service 2006a, Management 

Prescriptions 4.1 and parts of 5.0, 5.1, 

6.2, and 8.0; USDA Forest Service 

2006c, pp. 12, 19–20, 27). 

It is important to note that section 7 

of the Act provides regulatory flexibility 

to Federal agencies to complete their 

missions. This process allows Federal 

agencies to incidentally ‘‘take’’

individuals of a listed species as long as 

they insure their actions are not likely 

to jeopardize the entire species or 

adversely modify critical habitat. This 

regulatory option provided the MNF the 

ability to harvest and manage timber 

even in occupied WVNFS habitat. 

However, the MNF has avoided impacts 

to the WVNFS altogether while still 

maintaining a viable timber harvest 

program, which continues under the 

revised plan (USDA Forest Service 

2006a). 

After the WVNFS is delisted, the MNF 

is likely to amend the Forest Plan to 

incorporate its latest forest planning 

regulations, and to formally recognize 

that the WVNFS is no longer an 

endangered or threatened species. But 

given the MNF’s desired future 

condition for Management Prescription 

4.1 (summarized below), and history of 

proactive recovery efforts directed 

toward WVNFS conservation, the 

Service believes that the MNF will 

continue management and monitoring 

the red spruce-northern hardwood 

ecosystem that supports the WVNFS. 

Furthermore, the MNF’s current timber 

management and harvest goals are based 

on achieving desired forest and habitat 

conditions and not on a regional 

economic or a supply/demand basis 

(USDA Forest Service 2006b, FEIS, 

Appendix I). The desired future 

condition for Management Prescription 

4.1 focuses on developing a late 

successional stage (>120 years) forest 

over time (50+ years) with the multi-age 

stand structure that likely existed prior 

to exploitive logging (USDA 2006a, pp. 

III–12). At the stand level, desired 

vegetation conditions include a mix of 

trees of different ages, complex vertical 

habitat structure, scattered small 

openings (<2 ac) dominated by shrubs 

and saplings, scattered over-mature 

trees, and an abundance of snags, den 

trees, and downed woody debris. 

Even if the MNF revises the current or 

subsequent Forest Plans to increase 

timber harvest, it is highly unlikely that 

the current WVNFS habitat would be 

impacted. About two-thirds of the MNF 

is fully stocked or overstocked timber. 

The MNF is growing nearly four times 

as much timber as is being harvested or 

dying from natural causes (USDA Forest 

Service 2006b, FEIS, Appendix I, p. I–

155). Therefore, with the current 

surplus of available timber and the 

relatively small portion of the available 

timber currently being harvested, the 

MNF could substantially increase its 

annual harvest rate within the 330,000 

available ac and still have no need to 

harvest in WVNFS habitat. The MNF’s 

FEIS for the Forest Plan Revision 

describes three forest management 

alternatives that would result in a 

greater acreage available for timber 

harvest than the selected alternative 

(from 900 to 17,300 ac more) (USDA 

Forest Service 2006b, FEIS, Summary). 

The alternative with the greatest acreage 

available for timber harvest also 

includes a greater total acreage withheld 

from timber harvest to protect WVNFS 

habitat, Indiana bat (federally listed as 

endangered) habitat, river corridors, 

scenic areas, and streams buffers 

(367,396 ac or 68,703 ac more) than the 

selected alternative (298,693 ac), 

providing supporting evidence that the 

MNF has sufficient timber reserves if it 

wanted to increase timber harvest and 

still can protect WVNFS habitat. 

The MNF is harvesting its timber 

outside of WVNFS habitat, at a 

sustainable rate. Alternatives have been 

identified that would provide additional 

acreage for timber harvesting without 

compromising WVNFS habitat. 

Therefore, the Service believes it is 

reasonable to expect the MNF will 

continue not to harvest timber in 

WVNFS habitat; a choice that would 

continue the agency’s previous 

contributions to improve the WVNFS’s 

status. We also believe that the MNF has 

the current and future capability to 

manage timber harvest in a way that 

does not harm the WVNFS after 

delisting and will do so. 

Looking beyond the MNF, there is no 

evidence of any new sources of habitat 

loss throughout the current range of the 

WVNFS. According to analyses using 

the Menzel model, approximately 68 

percent of areas modeled as habitat are 

now considered secured by public 

ownership and/or managed for the 

protection of the WVNFS (Menzel et al. 

2006b, p.4). These areas include Canaan 

Valley National Wildlife Refuge (NWR) 

(created in 1994), Blackwater Falls and 

Canaan Valley State parks, Handley 

Wildlife Management Area, Kumbrabow 

State Forest, and the MNF (Service 

2006a, pp. 12–14). An additional 5 

percent of habitat is considered secure 

in Virginia on the GWNF. 

Activities that have contributed to 

habitat loss and degradation since the 

time of listing occur only locally or 

occur on the periphery of the WVNFS’s 

range (Service 2006a, pp. 11, 14, 20). 

These activities include limited 

highway development, recreational 

development, mining and gas 

exploration, timber management, and 

wind farm development (see ‘‘Summary 

of Public Comments’’, part F, issue 9). 

With regard to activities that are 

reasonably foreseeable to occur, some 

low level of local impacts are likely to 

continue into the future; however, there 

is no indication that the activities would 

ever be likely to occur over a landscape 

level, or at such a magnitude as to pose 

a threat to the continued existence of 

WVNFS throughout its range or in any 

significant portion of its range (Service 

2006a, pp. 11, 14, 19–20). 

For example, construction of Corridor 

H through the extreme northern part of 

the range of WVNFS is not expected to 

result in significant impacts to WVNFS 

or its habitat. Roads can adversely affect 

WVNFS movement by fragmenting 

habitat, although not all roads create 

absolute barriers. WVNFS are capable of 

gliding up to 200 ft, with the majority 

of the glides ranging from 16 to 82 ft 

(Scheibe et al. 2007, p. 857; Vernes 

2001, pp. 1028–1029). WVNFS are 

known to have crossed logging roads, 

gravel roads, and ski slopes (Ford et al. 

2007, p. 8; Menzel et al. 2006a, p. 207; 

Terry 2004, pp. 18–19). Menzel et al. 

(2004, p. 358) noted that many WVNFS 

day dens were located along or near 

abandoned skidder trails. Weigl et al. 

(1999, p. 61) found that G. s. coloratus 

frequently crossed patches of nonforested habitat, and one crossed a 

paved road several times. However, 

telemetry studies conducted on G. s. 

coloratus near the 2-lane paved 

Cherohala Skyway in North Carolina 

failed to document any evidence of 

squirrels attempting to cross this 

highway, even though in many cases the 

home ranges of the tracked squirrels 

were located in close proximity to the 

highway right-of-way (Weigl et al. 1999, 

pp. 69–73). Mean distances between 

forest edges across both sides of the 

right-of-way for that study ranged from 

125 to 175 ft, and hence may have 

exceeded the normal gliding capability 

of a majority of G. s coloratus. 

Range-wide habitat modeling has 

estimated that more than 235,000 ac of 

suitable WVNFS habitat exists south of 

the proposed Corridor H alignment and 

an additional 4,400 ac of suitable 

WVNFS habitat exists in the Blackwater 

Canyon area to the north of the 

alignment (Service 2006b, p. 19). 

Construction of the proposed project 

could decrease habitat connectivity 

within the northern habitats, or even 

create a permanent barrier to dispersal 

of the WVNFS between northern and 

southern areas. However, the amount of 

suitable habitat north and northeast of 


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the Blackwater canyon (approximately 

4,400 ac) is considerable and we 

conclude that it is large enough that the 

current WVNFS population is likely to 

persist (Service 2006b, p. 23; Smith and 

Person 2007, p. 631). About 24,000 acres 

of suitable habitat exists in the 

Blackwater Canyon area south of the 

highway and this will remain connected 

by dispersal corridors to the remaining 

211,000 acres of suitable habitat. 

Although the 235,000 acres (this figure 

is comprised of 211,000 acres plus the 

24,000 acres in Blackwater Canyon) 

south of the proposed Corridor H 

alignment is not contiguous habitat, 

there are no sizeable gaps preventing 

squirrel dispersal, so we conclude that 

no portion of the population south of 

the alignment will be meaningfully 

affected by the road. This leaves only 

the question of the impact of the road 

footprint itself. A total of 745 ac of 

habitat for the WVNFS will be lost 

during construction of the proposed 

project (Service 2006b, p. 23; Service 

2008, p. 20). This equates to a total loss 

of only 0.1 percent of the available 

highly suitable and suitable habitat for 

the subspecies, and therefore does not 

represent a significant threat. 

The Service analyzed possible 

secondary impacts to WVNFS from the 

proposed Corridor H project from 

Parsons, WV to Davis, WV (Service 

2006b, pp. 1–39) and Davis, WV to 

Bismarck, WV (Service 2008, pp. 1–32). 

Construction of this four-lane divided 

highway is expected to increase human 

accessibility to surrounding lands and 

could spur increased development in 

the lands adjacent to the project. 

However, a cumulative effects 

assessment, conducted by the West 

Virginia Department of Transportation 

(WVDOT) (2006, pp. 17–19) suggests 

there is an adequate amount of nonenvironmentally sensitive, lowelevation land that is not WVNFS 

habitat and that is available to support 

all development reasonably expected to 

occur as a result of the highway 

construction. WVDOT (2006, p. 20) 

modeled the worst-case scenario for 

development that was reasonably 

certain to occur after the highway was 

built, taking into consideration 

development and traffic patterns, and 

trends in employment and population 

growth. They mapped the raw private 

land (currently undeveloped) that was 

available to accommodate projected 

development. This was defined as land 

that was located outside of the 100-year 

floodplain that did not have slopes 

greater than 25 percent, that did not 

have wetlands, and that did not have 

existing development or was not 

currently under public ownership. Thus 

is appears that the land identified as 

being available to accommodate 

development corresponds to those lands 

that have the greatest likelihood of being 

developed due to lack of constraints. 

As a general matter, because the 

majority of WVNFS habitat is publicly 

owned and managed, future 

development throughout the range of 

the WVNFS is expected to be minimal. 

The entire range of the WVNFS is 

within the Allegheny Mountains Valley 

Physiographic Region, an area of steep 

terrain and low human population 

density and growth. In 2005, the 

proportion of land use classified as low 

density and high density development 

within this physiographic region in 

West Virginia was 0.4 percent and 0.1 

percent, respectively (WVDNR 2006b, p. 

10). During 2000, population densities 

in the counties in West Virginia in 

which the WVNFS occurs were among 

the lowest in the State, ranging from 9.7 

to 40.4 persons per square mile 

(WVDNR 2006b, p. 17); and with the 

exception of Randolph County (0.3 

percent increase), the 10-year 

population trend (1990–2000) in all of 

these counties decreased (WVDNR 

2006b, p. 18). 

Summary of Factor A: Although the 

quantity and quality of WVNFS habitat 

is reduced from historical levels 

(preceding the logging and burning era 

of the late 1800s and early 1900s), we 

now know that the WVNFS is more 

resilient in its habitat use than formerly 

thought, probably because of its 

mobility and plasticity in nest tree 

selection. Additionally, the habitat is 

more connected than previously 

thought, and habitat trends are moving 

in a positive direction in terms of forest 

regeneration and conservation. Also, the 

subspecies continues to persist for 

multiple generations at many locations 

across its historical range. Impacts from 

proposed transportation projects and 

potential future housing development 

are localized and minimal. For the 

foreseeable future, any localized loss of 

habitat due to timber harvest or 

development on private lands will not 

reduce the overall quality of habitat for 

the WVNFS, rather it will just slightly 

reduce the amount of improvement in 

habitat conditions. For these reasons, 

and the lack of any rangewide threats to 

WVNFS habitat, the present or 

threatened destruction, modification, or 

curtailment of its habitat or range is no 

longer currently a threat to the WVNFS 

or likely to become so in the foreseeable 

future. 

B. Overutilization for Commercial, 

Recreational, Scientific, or Educational 

Purposes 

The final listing rule concluded that 

the WVNFS was not known to be 

threatened by human utilization but 

noted that flying squirrels are highly 

desirable as pets to some persons, and 

collecting for such purposes is at least 

a potential threat to the already rare 

WVNFS (50 FR 26999). The WVNFS has 

been captured only for scientific or 

educational purposes through nest box 

and live trap methods, and not for 

market collecting or commercial use. 

Capture for scientific or educational 

purposes has been very limited, is 

regulated by state permitting systems, 

and has not proven to be detrimental to 

the continued existence of the WVNFS. 

In the 21 years since listing, the 

Service has not received any evidence 

that commercial use in the pet trade or 

recreational use of the WVNFS is a 

threat. There are no law enforcement 

records of illegal harvesting or 

commercialization of the subspecies. 

Several factors indicate that delisting 

will not significantly change that. The 

WVNFS is a thinly dispersed, nocturnal 

mammal that is very difficult to catch. 

For example, Menzel captured the 

WVNFS at a rate of 0.227 captures per 

100 trap nights (Menzel 2003, p. 65), 

and the WVDNR’s nest box monitoring 

program has had only a 2 percent 

average success rate of squirrel 

occupancy per box checked (Service 

2006a, p. 7). Additionally, once the 

WVNFS is delisted, its collection by 

hunting or trapping will still be illegal 

under West Virginia and Virginia state 

laws (West Virginia Code 20–2-5(26); 

Code of Virginia 29.1–521.A.10; 29.1–

566 and 29.1–530A.). See further 

discussion in Factor D under the 

Summary of Factors Affecting the 

Species below. WVNFS is not currently 

defined as a game or furbearing animal 

that can be legally hunted or trapped in 

either state, and as such, there currently 

are no bag limits allowed for WVNFS 

(West Virginia Code 20–1-2; Code of 

Virginia 29.1–100, 29.1–530.A). 

Moreover, once the WVNFS is delisted, 

its capture for scientific and educational 

purposes will still be regulated through 

collection permitting systems of the 

WVDNR (West Virginia Code 20–2-50) 

and the Virginia Department of Game 

and Inland Fisheries (VDGIF) (Code of 

Virginia 29.1–568). 

Summary of Factor B: Overutilization 

for any purpose is not currently 

considered a threat and is not 

anticipated to emerge as a threat in the 

foreseeable future. 


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C. Disease or Predation 

The final listing rule (50 FR 26999) 

made no mention of disease as a threat 

to the WVNFS, and we are not aware of 

any evidence since the time of listing 

that suggests the health of WVNFS 

individuals is threatened by disease. Of 

the more than 1,000 squirrel captures 

since 1985, none have shown signs of 

disease (Service 2006a, p.15). 

The final listing rule predicted that 

increasing human recreational use of 

northern flying squirrel habitat might 

result in predation on the WVNFS by 

pets, especially cats (50 FR 26999). 

While natural predators of the WVNFS 

may include weasel, fox, mink, owls, 

hawks, bobcat, skunk, raccoon, snakes, 

and fisher, we are not aware of any 

scientific or commercial evidence since 

the time of listing to support pets 

preying upon WVNFS (Service 2006c, p. 

15), or to suggest that natural predation 

limits populations of WVNFS. As 

analyzed in our biological opinion for 

the Camp Wilderness Habitat 

Conservation Plan (HCP) (Service 2003, 

pp.12, 23), there are no documented 

deaths of northern flying squirrels, 

particularly the WVNFS, as a result of 

impacts of human recreational use or 

occupancy in, or near, its habitat, and 

pets are not predicted to be a substantial 

threat in the future (Service 2003, pp. 

12, 23–25). Since the majority of 

WVNFS habitat is found on the MNF, 

human encroachment into WVNFS 

habitat is uncommon and localized (e.g., 

Canaan Valley and Snowshoe 

Mountain) (Service 2003, pp. 12, 23–25; 

Service 2006c, p. 15; Service 2006a, pp. 

15, 20), and is therefore unlikely to 

become a threat to the WVNFS in the 

foreseeable future. 

Summary of Factor C: Disease and 

predation are not currently threats to the 

WVNFS and are not likely to become 

threats in the foreseeable future. 

D. Inadequacy of Existing Regulatory 

Mechanisms 

The final listing rule stated that this 

factor was not known to be applicable 

(50 FR 26999). Currently, all threats 

under Factors A-C, and E have been 

eliminated or abated, and no regulatory 

mechanisms are needed to delist the 

WVNFS. Therefore, the inadequacy of 

regulatory mechanisms is not 

considered a threat to the subspecies. 

Nevertheless, even though not 

considered necessary for delisting, the 

laws discussed below will continue to 

provide some level of benefits to the 

WVNFS. 

State Laws 

The State of West Virginia does not 

currently have any State laws protecting 

endangered species. However, for the 

reasons stated in the discussions of 

Factors A, B, C and E, there are no 

current threats to the subspecies as a 

whole that require additional regulation. 

Therefore, the lack of an endangered 

species State law in West Virginia is not 

expected to negatively impact the 

WVNFS. See Factor B above for 

additional information. 

In the Commonwealth of Virginia, the 

WVNFS has been listed as endangered 

under the Commonwealth’s endangered 

species act since its Federal listing in 

1985. This Commonwealth law, which 

is administered by the VDGIF, prohibits 

take of Commonwealth-listed species 

and is currently applicable to the 

WVNFS. The State has the authority to 

continue protection of the WVNFS 

under the State law once it is removed 

from the Federal List of Threatened and 

Endangered Wildlife (Virginia Code 

29.1–566) and intends to do so 

(Reynolds 2008). Lack of current threats, 

along with the Commonwealth’s 

endangered species act, ensures the 

WVNFS’ persistence in Virginia. See 

Factor B above for additional 

information. 

E. Other Natural or Manmade Factors 

Affecting the Continued Existence of the 

Species 

Competition With Southern Flying 

Squirrel 

The final listing rule (50 FR 26999) 

concluded that the WVNFS was 

threatened by competition with the 

southern flying squirrel for habitat and 

by the spread of a parasite from the 

southern flying squirrel to the WVNFS. 

However, evidence collected since the 

time of listing indicates that the 

occurrence and potential severity of the 

southern flying squirrel’s impacts are 

limited. The occurrence of the two 

subspecies has been documented at 20 

percent of the known occupied WVNFS 

sites with no evidence of local 

extirpation of WVNFS. Overcompetition by the southern flying 

squirrel for den sites does not appear to 

be affecting population persistence of 

the WVNFS. In addition, any 

competition between the two subspecies 

may be somewhat ameliorated by the 

spread of beech bark disease (see Factor 

A above for further information), which 

results in the reduced availability of 

beech nuts, an important food source for 

the southern flying squirrel (Service 

2006a, p. 18). 

The final listing rule cited evidence 

from a captive study in the 1960s that 

a nematode parasite, possibly carried by 

the southern flying squirrel, might be 

lethal to the WVNFS (50 FR 26999). The 

rule stated that while the southern 

flying squirrels appeared healthy, all the 

northern flying squirrels weakened and 

died within 3 months, and this 

mortality was associated with heavy 

infestations of the nematode parasite. 

All the southern flying squirrels also 

carried the parasite, but they remained 

in apparent good health and continued 

to breed (50 FR 26999). Based on review 

of the original dissertation, the cause of 

the northern flying squirrel mortality 

was never completely understood 

(Weigl 1968, pp. 129–150). Weigl et al. 

(1999, pp. 74–75, 2007 p. 902) 

hypothesized that survival and 

maturation rates of the parasite are 

limited by below-freezing temperatures 

that occur within the range of the 

WVNFS, but were not replicated in the 

1960s captive study. The conditions 

created in the captive study apparently 

do not closely relate to naturally 

occurring conditions, and observations 

of WVNFS individuals captured in the 

last 20 years (including areas also 

occupied by the southern flying 

squirrel) have revealed no signs of 

sickness, debilitation, or death due to 

parasitic infestation. 

Other Natural or Manmade Threats 

The 1985 final listing rule did not 

address additional threats under Factor 

E. However, the delisting criterion 

within the 1990 recovery plan 

addressed potential threats, such as 

forest pests (see Factor A) and acid rain, 

to the existence of the high elevation 

forests on which the squirrels (G. s. 

fuscus and G. s. coloratus) depend 

(Service 1990, p. 19). These potential 

threats were included in the overall 

analysis of the status of the WVNFS in 

the 5-year review (Service 2006a, pp. 4–

6) and are analyzed in more detail 

below. 

Acid rain (more appropriately referred 

to as acid deposition) has been cited as 

potentially damaging forest ecosystems, 

especially the spruce-fir forests in 

portions of the Appalachian Mountains 

(NAPAP 2005, p. 41). Although 

empirical data are lacking regarding 

specific effects on the WVNFS, the longterm potential exists for anthropogenic 

acid deposition to diminish the extent 

and quality of the boreal-like spruce 

forests that have survived on the high 

ridges and plateaus, by pushing them 

farther up the slopes, and, if warming 

continues, reducing and eventually 

eliminating habitat at higher elevations. 

However, there has been no evidence of 

acid deposition reducing the extent of 

red spruce-northern hardwood forests in 

the Central Appalachians since the 

WVNFS’ listing in 1985 (Service 2006a, 

p. 18, Adams 1999, p. 24) (See above 


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Response to Comments, I—Climate 

Change Concerns, Issues 4 and 5). 

Given the naturally acidic nature of 

soils in spruce forests, it is unlikely that 

acid deposition has contributed 

significantly to their further 

acidification (Johnson and Fernandez 

1992, p. 262; Johnson et al, 1992, pp. 

391, 396). These forests do not reach the 

very low winter temperatures observed 

farther north and have not exhibited the 

red spruce winter kill due to decreased 

cold tolerance that has been observed in 

the northern Appalachians and 

Adirondacks (Peart et al. 1992, p. 180; 

DeHayes 1992, p. 296; NAPAP 2005, p. 

41). Sulphate deposition in the Central 

Appalachians has dropped by at least 25 

percent in the last 10 years and pH of 

deposition has increased, making this 

runoff less acidic (Johnson et al. 1992, 

pp. 388, 391; Adams and Kochenderfer 

2007, p. 99–100, Adams et al. 2006, pp. 

4–6, 216–217). Deposition of nitrogen 

has either leveled off or may be slightly 

increasing, but the overall acid load is 

decreasing in high elevation red spruce 

forests of the Central Appalachians 

(Adams and Kochenderfer 2007, p. 100–

101; Johnson et al. 1992, p. 391; Adams 

et al. 2006, pp. 4–6, 266). Also, 

compared to many deciduous trees, red 

spruce also is more resistant to ozone, 

which is often found in combination 

with high levels of acid deposition 

(McLaughlin and Kohut 1992, pp. 360–

366; Adams 2007). Given the factors of 

naturally acidic soils, increasing pH of 

deposition, lack of extreme cold 

temperatures, resistance to ozone 

impacts, and lack of adverse impacts 

from nitrogen, there is no current 

information demonstrating a negative 

impact on these high elevation forests. 

Furthermore, the current trends of the 

decreasing overall acid load indicate 

that acid deposition is not a significant 

threat to the subspecies’ habitat in the 

foreseeable future. 

Thus, to the extent that the effect of 

acid deposition on G. s. fuscus and its 

habitat are reasonably predictable, we 

concluded that they are not a significant 

threat to the subspecies’ habitat in the 

foreseeable future. 

Summary of Factor E: Overall, our 

analysis of the other natural and 

manmade factors, either alone or in 

combination, indicates that the WVNFS 

is not in danger of extinction throughout 

all or a significant portion of its range, 

or likely to become endangered within 

the foreseeable future. 

Conclusion of the 5-Factor Analysis 

As demonstrated in our 5-factor 

analysis, threats to the WVNFS have 

been abated or sufficiently minimized 

over the entire range of the subspecies. 

Relative to the information available at 

the time of listing, recovery actions, 

forest regeneration, and a reduction or 

abatement of threats have led to: (1) A 

significant increase in the number of 

known WVNFS captures and distinct 

capture locations; (2) verification of 

multiple-generation reproduction and 

persistence throughout the range; (3) 

proof of resiliency of the squirrels; and 

(4) the substantial improvement and 

continued expansion of suitable habitat 

rangewide. 

The biological principles under which 

we evaluate the rangewide population 

status of the WVNFS relative to its longterm conservation are representation, 

redundancy, and resiliency (Groves 

2003, pp. 30–32). At the time of listing, 

the WVNFS was thought to be an 

extremely rare and declining taxon that 

had disappeared from most of its 

historical range. We now know that 

occupancy of available habitat has 

increased and is much more widespread 

and well connected than formerly 

thought, and that the geographic extent 

of the WVNFS’ range approximates 

historical range boundaries. The red 

spruce-northern hardwood forests have 

substantially recovered from the vast 

clear-cutting at the turn of the 20th 

century, and continue to improve. 

Additionally, we have learned that the 

WVNFS has adapted to changes in the 

spruce ecosystem over the past hundred 

years, and can successfully exploit the 

existing habitat conditions throughout 

the landscape. Habitat patch sizes 

within the core of the range of WVNFS 

are sufficiently large and well connected 

by numerous linkages to facilitate 

adequate WVNFS dispersal among 

population centers (Service 2007c, pp. 

9–12). Although there remains 

geographic separation (and likely has 

been since the end of the Pleistocene) 

between a few of the habitat areas 

supporting population centers at the 

edge of the range, this habit matrix 

overall provides a relatively high degree 

of functional connectivity, as evidenced 

by constant occupancy of habitat across 

a range of forest conditions over 

multiple generations. The WVNFS has 

demonstrated more flexibility in its 

habitat use than previously thought, 

including a capacity to move freely and 

become widely dispersed. Thus, there is 

adequate representation (i.e., occupancy 

of representative habitats formerly 

occupied by the WVNFS across its 

range) and redundancy (i.e., distribution 

of populations in a pattern that offsets 

unforeseen losses across a portion of the 

WVNFS’ range) (Service 2007c, pp. 6–

12). 

Compared to most other North 

American tree squirrels, G. sabrinus 

demonstrates resilience and behavioral 

plasticity (Weigl 2007, p. 898). The 

species survived glacial advances in the 

Pleistocene, as well as widespread loss 

of forest cover from logging and burning 

in the late 1800s/early 1900s (Weigl 

2007, p. 898; Rentch et al. 2007, p. 441). 

Studies have confirmed the ability of G. 

sabrinus to adjust its biology to survive 

a wide range of environmental 

conditions, such as: The ability to 

occupy forests of varying spruce and 

hardwood compositions (Weigl 2007, p. 

898–899); the ability to survive short 

cold snaps by dropping its body 

temperature without becoming torpid 

(Weigl 2007, p. 898); the ability to 

generally subsist on fungi and lichens, 

buds, berries, and staminant cones, but 

to occasionally use mast (Weigl 2007, p. 

898); the ability to delay reproduction in 

response to environmental variables 

(Weigl et al. 1999, p. 32, 79); the ability 

to nest in a wide variety of trees (Menzel 

et al. 2004, pp. 360, 363–364; Menzel et 

al. 2006b, pp. 1–3, 6, 7; Menzel et al. 

2006b, p. 208; Ford et al. 2004, p. 430); 

and the ability to recolonize new habitat 

areas over time by adjusting its activity 

patterns to meet ecological requirements 

in and around patches of forest (Menzel 

et al. 2006b, p. 208). 

Survey and monitoring efforts at 109 

sites over the past 21 years have shown 

a relatively high degree of population 

stability, as evidenced by a high degree 

of persistence and successful 

reproduction over multiple generations 

throughout the historical range (Service 

2007c, pp. 9–11). There is no evidence 

of extirpation of a local population or of 

a deleterious source-sink 

metapopulation dynamic (Service 

2007c, p. 11). As previously described, 

the current and future trend for habitat 

quantity, quality, and connectivity is 

expected to be favorable because of the 

continuing recovery of the red sprucenorthern hardwood ecosystem and the 

lack of rangewide threats to WVNFS 

habitat (see Factor A under the 

Summary of Factors Affecting the 

Species above, and Service (2007b, pp. 

3–8)). As habitat availability increases 

into the future, the carrying capacity of 

protected habitat should continue to 

ensure persistence of populations of the 

WVNFS. 

Recovery efforts have provided 

increased attention and focus on the 

WVNFS and the habitat upon which it 

depends. Numerous conservation 

actions have been implemented since 

1985 by land stewards, biologists, 

government agencies, and conservation 

groups. These include: Research and 

recovery actions specified in the 1990 

recovery plan and 2001 recovery plan 

update for the WVNFS; conservation 


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50246 Federal Register / Vol. 73, No. 166 / Tuesday, August 26, 2008 / Rules and Regulations 

provisions incorporated into future 

expansion of the Corridor H highway at 

the edge of subspecies’ range (Service 

2008, pp. 3–4, 22–26; 2006b, pp. 4–8, 

28–32); minimization and mitigation 

measures specified in two HCPs at 

Snowshoe Mountain, specifically the 

protection of approximately 200 ac of 

WVNFS habitat in perpetuity [BHE 

Environmental, Inc. (BHE) 2003, pp. 34–

42, Appendix F; BHE 2005, pp. 49–55]; 

red spruce plantings on public and 

private lands; and conservation 

provisions in the 1986 MNF Land and 

Resource Management Plan (USDA 

Forest Service 1986, pp. X–1–X–3), 2004 

Forest Plan Amendment (USDA Forest 

Service 2004, p. 84, 84a, 84c, 87, pp. 

234–234b), and 2006 Forest Plan 

Revision (USDA Forest Service 2006a, 

Management Prescription 4.1 and 

portions of 5.0, 5.1, 6.2, and 8.0). Of 

particular note are the habitat protection 

initiatives that have occurred on both 

public and private lands, the 

development of habitat models and 

research on red spruce-northern 

hardwood forest restoration, and the 

establishment of Canaan Valley NWR. 

In summary, all of the past, existing, 

or potential future threats to WVNFS, 

either alone or in combination, have 

either been eliminated or largely abated 

throughout all of its range. The major 

factor in listing the WVNFS was the loss 

of habitat due to the logging era at the 

turn of the 20th century. This threat has 

largely been abated as evidenced by the 

substantial recovery and continued 

improvement of the preferred habitat of 

the WVNFS, red spruce-northern 

hardwood forests. Therefore, we have 

determined that the WVNFS is not in 

danger of extinction or likely to become 

so throughout its range in the 

foreseeable future. 

Significant Portion of the Range 

Analysis 

Having determined the WVNFS is not 

in danger of extinction or likely to 

become so in the foreseeable future 

throughout all of its range, we must next 

consider whether the subspecies is in 

danger of extinction or is likely to 

become so in the foreseeable future in 

any significant portions of its range. 

A portion of a species’ range is 

significant if it is part of the current 

range of the species and if it is 

important to the conservation of the 

species because it contributes 

meaningfully to the representation, 

resiliency, or redundancy of the species. 

The contribution must be at a level such 

that its loss would result in a decrease 

in the ability to conserve the species. 

Applying the definition described 

above for determining whether a species 

is endangered or threatened in a 

significant portion of its range, we first 

addressed whether any portions of the 

range of WVNFS warranted further 

consideration. As discussed in Factor 

A—Land Use Planning, there is one 

small geographic area where localized 

habitat threats still exist due to a future 

road expansion. However, we 

concluded that this area did not warrant 

further consideration because this area 

is very small (in the context of the range 

of the WVNFS) and has no substantive 

effect on the viability of the subspecies, 

and thus there was no substantial 

information that this area is a significant 

portion of the range (see Service 

prepared document ‘‘Analysis of 

significant portion of the range for the 

West Virginia northern flying squirrel’’

(Service 2007b, pp. 1–6). Therefore, 

based on discussion of the threats 

above, we do not foresee the loss or 

destruction of any portions of the 

subspecies’ range such that our ability 

to conserve the subspecies would be 

decreased. Therefore, we find that the 

WVNFS is not in danger of extinction 

and is not likely to become endangered 

in the foreseeable future throughout all 

or a significant portion of its range. 

Effects of the Rule 

Promulgation of this final rule will 

affect the protections afforded the 

WVNFS under the Act. Taking, 

interstate commerce, import, and export 

of WVNFS are no longer prohibited 

under the Act. Removal of the WVNFS 

from the List of Endangered and 

Threatened Wildlife does not supersede 

any State regulations. Federal agencies 

are no longer required to consult with 

us under section 7 of the Act to ensure 

that any action authorized, funded, or 

carried out by them is not likely to 

jeopardize the subspecies’ continued 

existence. However, for the 

approximately 68 percent of the WVNFS 

habitat on the MNF, and the small area 

(5 percent) of habitat located within the 

GWNF, the activities impacting the 

WNVFS and its habitat must comply 

with appropriate Forest Service 

management plans. There is no critical 

habitat designated for the WVNFS. 

Post-Delisting Monitoring Plan 

Section 4(g)(1) of the Act requires us, 

in cooperation with the States, to 

implement a monitoring program for not 

less than 5 years for all species that have 

been recovered and delisted. The 

purpose of this requirement is to 

develop a program that detects the 

failure of any delisted species to sustain 

itself without the protective measures 

provided by the Act. If, at any time 

during the monitoring period, data 

indicate that protective status under the 

Act should be reinstated, we can initiate 

listing procedures, including, if 

appropriate, emergency listing. 

To further ensure the long-term 

conservation of the WVNFS, a postdelisting monitoring (PDM) plan has 

been developed that lays out a 10-year 

framework to monitor the status of the 

subspecies (Service 2007c, pp. 1–27). 

The Plan focuses primarily on 

monitoring of (1) Habitat status and 

trends and (2) implementation of habitat 

management plans and agreements. 

Habitat changes will be tracked 

rangewide by interpretation of remotesensed imagery obtained at or near the 

time of delisting (baseline), compared to 

the end of the PDM period. These data 

will be verified by a subsample of stand 

data and on the ground field checks. In 

addition, land managers will self-report 

annually on accomplishment of key 

components of land management plans 

or agreements for WVNFS, including the 

acreage of habitat modified (positively 

or negatively), as well as land 

management problems and solutions. 

The PDM plan also includes actions 

for monitoring of WVNFS distribution 

and persistence. The nest box and live 

trapping survey component will be 

largely a continuation of ongoing annual 

presence/absence surveys by the 

WVDNR, MNF, and other participants, 

but with an increased emphasis on 

covering as much of the extant 

distribution within core habitat areas as 

possible. This will help determine if 

WVNFS continue to be present in these 

areas over multiple generations. 

The PDM plan identifies measurable 

management thresholds and responses 

for detecting and reacting to significant 

changes in WVNFS habitat, distribution, 

and persistence. If declines are detected 

equaling or exceeding these thresholds, 

the Service, in combination with other 

PDM participants, will investigate 

causes of these declines, including 

consideration of habitat changes, low 

natality, deaths or emigration, weather, 

trap shyness, competition for nest sites, 

or any other significant evidence. The 

result of the investigation will be to 

determine if the WVNFS warrants 

expanded monitoring, additional 

research, additional habitat protection, 

and/or resumption of Federal protection 

under the Act. At the end of the 10-year 

monitoring program, the Service will 

conduct a final review. It is the intent 

of the Service to work with all of our 

partners towards maintaining the 

recovered status of the WVNFS. 

The final PDM plan is available on the 

Service’s northeast region Web site, 

http://www.fws.gov/northeast/ 

endangered. 


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Federal Register / Vol. 73, No. 166 / Tuesday, August 26, 2008 / Rules and Regulations 50247 

Paperwork Reduction Act 

This rule does not contain any new 

collections of information under the 

Paperwork Reduction Act (44 U.S.C. 

3501 et seq. ). An agency may not 

conduct or sponsor and a person is not 

required to respond to a collection of 

information unless it displays a 

currently valid OMB control number. 

National Environmental Policy Act 

We have determined that 

Environmental Assessments and 

Environmental Impact Statements, as 

defined under the authority of the 

National Environmental Policy Act of 

1969, need not be prepared in 

connection with regulations adopted 

pursuant to section 4(a) of the 

Endangered Species Act. We published 

a notice outlining our reasons for this 

determination in the Federal Register 

on October 25, 1983 (48 FR 49244). 

References Cited 

A complete list of all references cited 

herein is available upon request from 

the West Virginia Field Office (see FOR

FURTHER INFORMATION CONTACT above). 

Author 

The primary author of this final rule 

is Laura Hill, Endangered Species 

Biologist and species lead for the 

WVNFS in our West Virginia Field 

Office (see FOR FURTHER INFORMATION

CONTACT section). 

List of Subjects in 50 CFR Part 17 

Endangered and threatened species, 

Exports, Imports, Reporting and 

recordkeeping requirements, and 

Transportation. 

Regulation Promulgation 

■ Accordingly, we amend part 17, 

subchapter B of Chapter I, title 50 of the 

Code of Federal Regulations as set forth 

below: 

PART 17—[AMENDED] 

■ 1. The authority citation for part 17 

continues to read as follows: 

Authority: 16 U.S.C. 1361–1407; 16 U.S.C. 

1531–1544; 16 U.S.C. 4201–4245; Pub. L. 99–

625, 100 Stat. 3500; unless otherwise noted. 

■ 2. Amend § 17.11(h) is amended by 

removing the entry for ‘‘Squirrel, 

Virginia northern flying’’ under 

‘‘MAMMALS’’ from the List of 

Endangered and Threatened Wildlife. 

Dated: August 15, 2008. 

Bryan Arroyo, 

Acting Director, U.S. Fish and Wildlife 

Service. 

[FR Doc. E8–19607 Filed 8–25–08; 8:45 am] 

BILLING CODE 4310–55–P 


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