Bat Surveys on USPS Northern Region Lands in Montana: 2005 Prepared for: USDA Forest Service, Northern Region P.O. Box 7669 Missoula, Montana 59807 By: Paul Hendricks and Biyce A, Maxell Montana Natural Heritage Program Natural Resource Infoimation System Montana State Libraiy December 2005 MONTANA Natural Heritage Pfc^ram Bat Surveys on USPS Northern Region Lands in Montana: 2005 Prepai"ed for: USDA Forest Sei"vice_ Northern Region P.O. Box 6779 Missoula, Montana 59807 Agreement Number 05-CS-l 1015600-033 By: Paul Hendricks and Bryce A. Maxell JW MONTANA ^M Natural Heritage ^r^Ptt^iam ^^atate nilj^ Natural Resource f\ II Q^ T" N A Library © 2005 Montana Natural Heritage Program P.O. Box 20 1 800 • 1 51 5 East Sixth Avenue • Helena, MT 59620-1800 • 406-444-5354 i This document should be cited as follows: Hendricks, P. and B,A. Maxell. 2005. Bat Surveys on USPS Northern Region Lands in Montana: 2005, Report to the USDA Forest Service, Northern Region. Montana Natural Heritage Program, Helena, MT. 12 pp. plus appendices. Executive Summary The distribution and status of bats in Montana remain poorly documented on US Forest Service Noithern Region lands. This is of conservation interest because management activities on Forest Service lands (e.g., timber harvest, mine closures, closures of historic buildings) may have unintended consequences on habitats bats use for roosting and foraging and may therefore negatively impact bat populations. Additionally, the Noithern Region has designated Townsend's Big-eared Bat {Corynorhinus hmjisendii) a Species of Concern requiring special attention: it is widespread but infrequently captured in Montana, with few documented hibernacula and maternity roosts and a reputation for being highly susceptible to human distult^ance at roost sites. The Northern Region recognized the need for additional documentation of bats on Forest Service lands to address inventory and monitoring requirements, and initiated bat surveys in 2005 across the Region on selected National Forest (NF) Ranger Districts (RD). In Montana, these included Swan Lake RD-Flathead NF. Bozeman RD- Gallatin NF, Townsend RD-Helena NF, Libby RD- Kootenai NR and Judith RD-Lewis & Clark NF. Following a modified protocol based on the Oregon Bat Grid system, crews surveyed non-randomly chosen suitable habitats within randomly chosen 10 X 10 km" sample units in each RD; ten sites (often two/sample unit) on each District were sampled, for a total of 50 sites surveyed on Northern Region lands in Montana. Thus, this approach is primarily targeted at identifying species richness within grid cells; inferences on rates of occupancy are limited to the percent of !0 x 10 km- grid cells where a species was detected within each sampled RD. August 2005. Species captured included Townsend's Big-eared Bat at two sites, Big Brown Bat {EpTesicus fitsats) at 14 sites, Hoary Bat (Lasiuvus c'meveus) at 20 sites, Silver-haired Bat (Lasionycteris noctivagans) at 25 sites, California Myotis {Myotis californicus) at nine sites, Western Small-footed Myotis [M. ciiiolabrum) at eight sites. Western Long-eared Myotis (M. evotis) at 26 sites. Little Brown Myotis (M htcifiigus) at 32 sites. Fringed Myotis (M thysaiiodes) at four sites. Long-legged Myotis (M, volaus) at 23 sites, and Yuma Myotis (M. yumanensis) at two sites. No bats were captured at five of the sites sampled, although presence of bats was noted at each. The 2005 field survey filled important gaps in documented distributions in Montana, adding several new county records and underscoring the need for additional survey effort to define bat distributions on USFS landscapes more fully. A summary of all existing bat records across the region clearly shows large distribution gaps for all species, further underscoring the need for addition surveys. In particular, large portions of the Beaverhead -Dee rlodge NF. Custer NF. Flathead NF, Gallatin NF, and Lewis and Clark NF lack records for any bat species. We recommend that the USFS Northern Region continue with a grid- based random sampling scheme stratified by ecoregion or Ranger District, resulting in a site- occupancy approach that allows for valid inference of presence across the selected stratum, A grid- based sampling scheme is an important monitoring approach that should be extended beyond USFS lands and coordinated with other partner agencies and organizations to guide effective bat management across the state. Eleven species of bats, represented by 795 total individuals, were captured during late-June to mid- Acknowledgments We thank Fred Samson and Jenny Taylor (USPS) for initiating and promoting the project, through the USPS Regional Inventory and Monitoring (RIM) program, and overseeing its implementation. Jenny also organized the excellent training session, run by Pat Ormsbee (USPS) and Dr. Joe Szewczak (Humboldt State University); Kristi Dubois (MTPWP) helped coordinate the training by an"anging a home base in the PWP Region Office in Missoula. Pat and Jenny developed the sampling grid that guided surveys. Actual on-the-ground surveys on the Helena, Gallatin, and Lewis & Clark National Porests were conducted by Cori Lausen and Erin Baerwald. with the assistance of Shawna Campbell (Helena NP), Marion Cherry (Gallatin NP), Eric Tomasik (Lewis & Clark NP), Trixi Smith, and Bryce Maxell (MTNHP). Plathead and Kootenai National Porests sui"veys were conducted by Paul Hendricks (MTNHP). with the assistance of Jane Ingebretson (Plathead NP). Jenny Holifield (Kootenai NP), Henning Stabins (Plum Creek), and Scott Tomson (Lolo NP). Scott Blum (MTHNP) entered survey data into the Montana Natural Heritage Program's POD database, facilitating the production of new distribution maps and the updating of element occurrence data in the Montana Natural Heritage Program's Biotics database. IV Table of Contents Introduction 1 Methods 2 Results 4 Species and Numbers Captured 4 New County Records 5 Evidence ot Reproduction by Females 5 Adult Sex Ratios 5 Discussion 7 Overview 7 Analysis of Protocols 7 Conclusions and Recommendations 9 Need for a Montana Bat Grid 9 References Cited 11 Appendix A. Global/State Rank Definitions Appendix B. USPS Region I Montana Survey Sites for Bats: Summer 2005 Appendix C. Distribution Maps for Bats in Montana List of Figures Figure L Total sampling effort (net-hours) at each site (n = 50 sites) and the number of species captured 8 List OF Tables Table 1. Number of sites where bats were captured, and total number of individuals captured, on five Region I National ForestsinMontana. 24 June- 18 August, 2005 4 Table 2. Adult sex ratios (number of males: number of females) of bats captured on five Region 1 National Forests {one District each) in Montana. 24 June- 1 8 August. 2005 5 Introduction There hasbeen growing concern in recent decades regarding the status ot bats throughout North America, partly because of a general lack ot basic natural history information (Hayes 2003), and also because a variety of habitats traditionally used by bats for roosting and foraging have been subjected to widespread disturbance, alteration, reduced availability, orcomplete removal (Fenton 1997, Pierson 1998). As a result, six species or subspe- cies of bats in the continental United States cur- rently are classified as endangered under the United States Endangered Species Act of 1973 (C Shea el al, 2003); none of these bats occur in Montana, Conservation and protection of roosts are important long-term management activities for many North American bat species {Sheffield et al. 1992). Unfortunately. conser\'ation efforts for bats in Montana are often hampered by a lack of data on their habitat requirements. For example, the little data available from Montana on foraging and diet of bats have been obtained largely at water sources (Jones etal. 1973), with no knowledge of where the foraging bats are roosting. Conversely, studies of bat roosts in Montana {e.g.. Woithinglon 1991a. 1991b, Hendricks et al. 2000. 2004) lack informa- tion on where and how far the roost members go to feed and drink. Nor have patterns of roost selection and fidelity (e.g., Sherwin et al. 2003) been studied in Montana, even though it is understood that suitable summer and winter roosts may limit the local and regional distribution and abundance of many temperate-zone bats (Humphrey 1975, Dobkin et al. 1995). especially cave- and crevice- dwelling taxa. Most bat species use a variety of localized habitats for roosting, be they natural sites {e.g., caves, trees, rock crevices) or man-made sites (e.g.. buildings, mines, bridges). Sites may be used only during specific seasons of the year, and then for different purposes. Recent research on bat roosts in Mon- tana has followed the national pattern of inventory- ing and monitoring roosts in caves and abandoned mines (e.g.. Worthington 1991a, 1991b. Hendricks et al. 2000. 2004; Hendricks and Kampweith 200 1 ), and this remains an important activity for a state bat conservation plan. Nevertheless, sampling bats across the landscape at foraging sites continues to be critical for filling gaps in documented distri- bution, assessing relative abundance of local populations, and ultimately identifying roosts for these populations. Distribution and status of bats remain poorly documented on US Forest Service Northern Region lands in Montana. A summary of all existing bat records across the region clearly shows large distribution gaps for all species, further underscor- ing the need for addition surveys. In particular, large portions of the Beaverhead-Deerlodge NF, Custer NF, Flathead NF, Gallatin NF. and Lewis and Clark NF lack records for any bat species. This is of conservation interest because manage- ment activities on Forest Service landscapes may have unintended consequences affecting bat popula- tions and the habitats bats use for roosting and foraging. The Northern Region recognized the need for additional documentation of bats on Forest Service lands to address inventory and monitoring requirements, and therefore initiated bat surveys in 2005 across the Region on selected National Forest RangerDistricts. Additionally. Townsend's Big- eared Bat {Corytjorhinus toivnsendii). a 04 S2 state Species of Concern in Montana {see Appendix A for codes), is a Northern Region Species of Concern requiring special management attention because of few documented hibernacula and maternity roosts and a reputation for being highly susceptible to human disturbance at roost sites (Pierson etal. 1999). Methods The primary objective of the 2005 survey was to document bat species richness {number ot species) within sample units, with a longer-term objective of inferring sample unit occupancy by each species across entire Ranger Districts. Surveys for bats in Montana were conducted during summer (lale-June to mid-August) 2005 on one Ranger District (RD) in each of five National Forests (NF) of the North- ern Region: Flathead NF-Swan Lake RD. Gallatin NF-Bozeman RD. Helena NF-Townsend RD. Kootenai NF-Libby RD. Lewis & Clark NF-Judith RD. The sampled Flathead and Kootenai RDs are in northwestern Montana west of the Continental Divide. Sampled RDs for the other three Forests are in west-central and central Montana east of the Continental Divide. Survey sites spanned a range of elevations: l990-5030ft west of the Divide and 4533-7642 ft east of the Divide, The Oregon Bat Grid was used as a framework from which to draw randomly-selected grid cells for survey in the 2005 Northern Region sui"veys. A grid of square blocks, each 10 km on a side ( 100 km- in area), was created and overlaid on each RD, to create a target population of grid cells to infer occupancy rates. Each grid cell equaled a sample unit. Sample units were selected using randomly generated numbers; sample units to be surveyed were those with the lowest numbers and with reasonable access to potential sui"vey sites. Five sample units per RD were to be surveyed at two distinct locations for one night each. This differs dramatically in survey effort from the Oregon Bat Grid (draft protocols available from Pat Ormsbee), which calls for up to a total of 12 surveys in a sample unit, or until all species expected to occur in a sample unit have been detected. For the 2005 Northern Region Inventory, each RD received ten nights of survey eftbrt. resulting in a total of 50 sites sui"veyed on Northern Region lands (Appendix 2). Only one. two and three sample units received two nights of survey effort on the Gallatin, Helena, and Lewis & Clark, respectively; the remaining nights of sur\'ey effort on those Forests were spent in single surveys of additional sample units. Sites chosen for survey within a sample unit were determined in the field by survey crews or using information provided by other Forest Service personnel. Sites were usually features that might concentrate bat activity, most often water sources such as ponds and streams, less often bridges over streams, and caves and mines, and least often at abandoned buildings. Bats were captured using mist nets of various lengths and configurations, sometimes in conjunction with harp traps: number of nets deployed varied from site to site. Nets were to be deployed at twilight and left open for 3.5 hours, weather permitting, or until I hour passed with acoustic detections. Species identification was based on published keys and species accounts (van Zyll de Jong 1 985, Nagorsen and Brigham 1993. Adams 2003) and a key developed specifically for this project. Stan- dard measurements (weight, forearm length, ear length, sex. age. reproductive status} were obtained from each individual. Wing punch tissue samples were also collected from each captured bat, until five punches/species were accumulated from each site. Tissue was taken using sterile procedures and stored in biopsy tubes containing desiccant. Tis- sues are to be used for genetic identification of species pairs that are difficult to distinguish in the field (especially Myotis lucifitgus and M. yummieiisis). Genetic analysis had not been initialed at the time of this report. The survey protocol also called for acoustic moni- toring at each site using a Peltersson 240x detector or, in its absence, a Titley Anabat II detector. Calls were to be captured and analyzed using the appro- priate software (SONOBAT or ANABAT). Unfor- tunately, equipment often malfunctioned or was not available until late in the field season. The number of calls captured, analyzed, and used for species- specific determination were too few to include in this report. However, we feel acoustic sampling is an important component of all future inventory and monitoring efforts (see Hayes 1997 and O'Farrell and Gannon l999forexamples}. Data was recorded on standardized data sheets, and later transcribed to the Point Observation Database housed at the Montana Natural Heritage Program. Helena, where it is available for agency and public use. Statistical analyses follow procedures in Sokal and Rohlf( 1981). Results Species and Numbers Captured During summer 2005, 50 sites were sampled tor bats on USPS Northern Region lands in Montana: 20 sites west and 30 sites east ot the Continental Divide (see map in Appendix C). Bats were detected at all sites and captured at 90% of these (see Appendix B). Eleven species were captured, represented by 795 individuals (Table I); of these species, Townsend's Big-eared Bat {G4 S2) and Fringed Myolis (G4G5 S3) are Montana Animal Species of Concern (Montana Natural Heritage Program 2004). Nine species were captured at sites west and ten species at sites east of the Conti- nental Divide. The California Myotis was captured only west of the Divide. Townsend's Big-eared Bat and Western Small-footed Myotis were captured only east of the Divide. Most frequently captured in the total sample was the Little Brown Myotis, representing 35,8% of all captures from 64% of the survey sites (Table I ). Second most frequently captured was the Silver- haired Bat. representing an additional 22% of ali captures from half of the sui"vey sites. Hoary Bat. Western Long-eared Myotis. and Long-legged Myotis accounted for 8.8- 10.7% of the total individuals captured, at 40-52% of the survey sites. Big Brown Bat represented 6.7% of the total captures from 28% of the survey sites. The remain- ing five species each accounted for<3% of the tola! captures, and were found at 4% to 18% of the survey sites. Hoary Bat was visually identified at three sites west of the Continental Divide (one site on the Flathead NF. two sites on the Kootenai NF) where it was not captured; its large size and early emergence made this species identifiable in tTight. Table J. Nwiiber of sites where bats were captured, and total muiiber of individuals captured, on five Region 1 National Forests in Montana. 24 Jiine-18 August, 2005. Ten sites were sampled per Forest, each site for one night. Species and number of captures for each site are listed in Appendix 2. Species'" Forest Total Flathead Kootenai Helena Gallatin Lewis & Clark GOTO 0,0' 0,0 2,9 0,0 0,0 2,9 EPFU 0.0 2,5 3,4 4, 13 5,31 14. 53 LACI 0(lv).0 3(2v|,5 3,7 5,29 9,44 20 (3v). 85 LAND 0,0 5. 11 4,22 6, 33 10. 109 25. 175 MYCA 4.4 5. 10 0,0 0,0 0.0 9, 14 MYCI 0.0 0,0 3. 14 1, 1 4,6 8,21 MYEV 3,3 3.7 8, 17 5, 32 7. 16 26, 75 MYLU 4, 11 7. 33 4. 12 10, 176 7,53 32, 285 MYTH 0.0 2.2 2,4 0,0 0,0 4.6 MYVO 4,9 3.4 5, 13 4. 16 7.28 23. 70 MYYU 0, fl 1, 1 1, 1 n, (1 0,0 1 1 Total captures 27 78 103 300 287 795 Species Richness 4(+lv=5) 9 10 7 7 11 ^ Species codes: GOTO {Corynorhinus townsendii, Townsend's Big-eaied Bat), EPFU {Epresicus fuscus. Big Brown Bat). LACI {Lasiurus ciuereus. Homy Bat). LANO (Lasionycleris noctivagans)^ MYCA {Myotis caiiforiiicus, California Myotis). MYCI {Myotis ciiiolabruiu. Western Small-footed Myotis), MYEV (Myotis evotis^ Western Long-eared Myotis), MYLU (Myotis lucifitgus. Little Brown Myotis), MYTH (Myotis thysanodes^ Fringed Myolis), MYVO (Myotis volans. Long-legged Myotis), MYYU (Myotis yumanensis^ Yuma Myotis). '' Number of sites, number of individuals; v = sites with good visual observation but no capture. 4 New County Records The summer 2005 Northern Region survey resulted in new county records for nine species (see maps in Appendix C): Big Brown Bat (Gallatin). Hoary Bat (Judith Basin), Silver-haired Bat (Broadwater), California Myotis (Lake, Missoula). Western Small-footed Myotis (Gallatin). Western Long- eared Myotis (Gallatin). Fringed Myotis (Lincoln). Long-legged Myotis (Gallatin), and Yuma Myotis (Lincoln). Gallatin County (Gallatin NF) received the most new records, with the addition of four species. Additional bat inventory work conducted in 2003-2005 by the Montana Natural Heritage Program in eastern Montana resuhed in numerous additional county records for these and other species. Evidence of Reproduction by Females The best evidence of reproduction by a particular bat species on a Forest is the presence of pregnant and lactating females. Testicular males are repro- ductively active, but their presence in a sample does not necessarily indicate reproduction near the survey site, while post-lactaling females and volant juveniles may have dispersed from adjacent areas. Pregnant or lactating females were identified in seven of nine species captured west of the Conti- nental Divide; these included all Myotis species on the Flathead NF, with the addition of Yuma Myolis, Big Brown Bat, and Silver-haired Bat from the Kootenai NF. Only Hoary Bat and Fringed Myotis lacked evidence of reproduction on the two western Forests. East of the Divide, pregnant or lactating females were identified in seven often species captured; only Western Small-footed Myotis, Yuma Myotis. and Big Brown Bat lacked evidence of reproduction in samples of captured females. However, we expect reproductionby females of all species will be documented in the future with additional survey effort. Adult Sex Ratios Adult males were caught more often than adult females, as represented in pooled samples for the majority of species (six of seven) from east of Che Continental Divide, and where sample sizes ex- ceeded 20 individuals per species {Table 2). Of these, significantly more males were present in samples of Big Brown Bat, Hoary Bat. Western Small-footed Myotis, and Little Brown Myotis. Sample sizes of all species west of the Continental Divide, except Little Brown Myotis, were too small for statistical analysis. For this species, as with samples from east of the Divide, significantly more males were captured (chi squared = 4.85, F = 0,028). Table 2. Adutt sex ratios (number of mates: mmiber of females) of hats captured on five Region I National Forests {one District each) in Montana. 24 J ime- 18 August. 2005. Species' Forest /■ value Flathead Kootenai Helena Gallatin Lewis & Clark GOTO ___ ___ 1 8 ___ ___ — EPFU ___ 2:3 4 11:2 29:2 18.20*^ LAO ___ 5:0 4 23:4 37:7 19.92='-'^ LANO --- 5:6 10 12 22:11 53:56 NS MYCA 2:2 3:7 ___ ___ ___ ___ MYCI — --- 13:1 1:0 6:0 8.50^'^ MYEV 2:1 4:3 3:14 17:15 10:5 NS MYLU 5:6 28:5 12:0 147:26 51:1 81.11='-!^ MYTH --- 1:1 1:3 — --- -- MYVO 4:5 4:0 5:8 9:6 21:7 NS MYYU — (1:1 1:0 — — — ^ Species codes: COTO (Cor ynorh inns rownsendii. Townsend's Big-eared Bat). EPFU (Eptesicus fuscus. Big Brown Bal), LACI {Lasiurus cinereus. Hoary Bal), LANO [Lasionycterts nocrivagans). MYCA {Myoris califormciis. California Myotis), MYCI {Myotis cilio/abrum^ Western Small-footed Myotis), MYEV {Myotis evotis. Western Long-emed Myotis), MYLU {Myotis hfcifiigus. Little Brown Myotis). MYTH {Myotis thysanodes. Fringed Myotis). MYVO {Myotis volarts. Long-legged Myotis), MYYU {Myotis yumanensis^ Yuma Myotis). '' Tests the null assumption of equal numbers of males and females captured. Applies only to Forests (Helena, Gallatin, Lewis & CImk) where Districts surveyed are east of the Continental Divide and species sample size > 20 for the combined Districts. All chi squared values are Yates corrected: NS = not significant at a = 0.05, '^- P < 0.01,*^' P< 0.0001. Discussion Overview The summer 2005 survey helped fill distribution gaps, produced several new county records (Appen- dix C). and provided evidence that several species ot bats are using landscapes under Noit hern Region stewardship forreproductive activities, including Townsend's Big-eared Bat on the Townsend RD of the Helena NR Overall, adult males were captured more often than females where we sampled bats (Table 2), a bias that has been noted elsewhere in Montana (Worthington 1991a, 1991b. Hendricks et al. 2000, 2004). It is possible males outnumber females throughout the landscape, but the bias towards males in capture samples couid also be a result of differential habitat use by the sexes (Bogan et al. 1996, Ciyan et al. 2000} or differ- ences in capture probabilities. Because niche segregation of the sexes is a possibility, information on differences in habitat use by males and females isnecessary for monitoring bat populations, devel- oping conservation plans for Species of Concern (e.g.. Pierson et al. 1999), and effectively managing landscapes for bats. In particular, the 2005 survey identified numerous areas where bats are concentrating their activity while seeking food and water resources. Some of these sites may be useful in the future for monitor- ing efforts across Forest Districts, especially sites that were used by several bat species (see Appendix B). Thus, the 2005 survey was a valuable exeirise as a pilot effort in the development of a comprehen- sive survey and monitoring scheme, both for the Northern Region and all of Montana. Analysis of Protocols The largest failure of the 2005 survey was the inadequate use of. and facility with, acoustic technology and analysis. Acoustic technology has great potential to provide rapid assessment of species distributions over many sites and identify areas of significant concentrations of species and individuals. It also has an advantage over capture methods in requiring far less commitment of time and personnel. It is important, however, to have equipment available and field crews trained in the use ofthis technology well in advance of field surveys. Keeping electronic equipment functional in the field also requires special effort. We consider acoustic monitoring an impoilant component of future inventory and monitoring schemes, but it does not replace capture methods. There remains a need forrecordedcallsfrom individuals whose identity is definitive through morphologic and genetic measurement in order to build a library of reference calls for individual species from across the state. The three sets of data {acoustic, morpho- logic, genetic) will provide future workers using acoustic monitoring the reference tools they need to identify, and correct for, regional differences in calls. The protocol we used in 2005 called for deployment of mist-nets for 3.5 hours after twilight, weather permitting, or until I hour had passed without acoustic detections. The numbers of nets deployed, and the total duration of their deployment, were left to the discretion of field crews. The numbers of nets deployed at sites in 2005 ranged from 2- 16, and hoursof deployment after twilight ranged from 2 to 7.5. This resulted in large variation in survey effort among sites, and demands on crews varied correspondingly. What level of eftbrt is worth the expense in time and energy of the field crews, and does the extra level of effort result in additional speciesdocumented(theprimary objective of the 2005 survey}? The total amount of time all nets are deployed can be used to calculate net-hours, which is a measure of sampling effort. There was large variability in number of species captured and net-hours of effort in 2005, with up to seven species captured in as few as 15 net-hours or as many as 62.5 net-hours (Figure 1). In general, however, the number of species captured during a single night increased with increased sampling effort (Spearman r = 0.580, P < 0.000 1 , n = 50). The relationship appears to be curvilinear, with an asymptote at about 40-50 net-hours, suggesting this level of sampling intensity could represent the best balance between trapping effort and the time and energy commitment of the crew to capture new species. 8 n ♦ ♦ ♦ ♦ 6 - ♦ ♦♦ ♦ # « (A .9! u 0) D, 4 - 0) 4 ♦ « # 4 4 ♦ » ♦ » d ♦ ♦«» * # 2 - 4» 4»^ ♦♦ ♦ - 1 — 1 20 40 60 80 100 120 Net-hours Figure L Total sampling effort (uet-hours) at each site (n = 50 sites) artd the nwnber of species captured. Net-hours = {munber of nets deployed x minutes deployed) -^ 60. However, for 21 sites where 4-7 species were captured, the numberof nets deployed ranged from 3-16. with only 23-80% (mean = 48%) of deployed nets capturing bats: the Oregon Bal Grid protocols recommend use of up to three nets, using a muhi- visit approach for surveying sampling units. The maximum number of species captui"ed at 62% of these sites occurred by 40 net-hours, with a mean effort of 29.6 net-hours to reach the full species list at these 21 sites; the full species list of the netting session occurred a mean of 2.6 hours after nets were opened, A more useful sampling protocol for a broad-scale bat inventory, such as that initiated in 2005. would converge with thai used in the Oregon Bat Grid, which incorporates elements of a typical state bird atlas project to help guide the sampling effort applied to each sample unit. In the Oregon scheme, the primary objective is to document all species on a list of expected species generated foreach sample unit. Each sample unit is surveyed using muhiple detection methods, as we attempted to do in 2005, but also is visited as many times (up to 12) as it takes to achieve the species richness goal, rather than limiting the survey effort to two visits, as was done in 2005. Even for roost monitoring of a species like Townsend's Big-eared Bat. thei"e is so much detection variability during any single visit (due to a variety of environmental and sampling variables) that as many as nine visits to a site may be necessaiy to identify a non-roost (Sherwin et ah 2003). Although the 2005 survey filled distribution gaps and generated much useful data, limited human and monetary resources kept the survey from achieving the objective of determining species richness for most sample units visited. This failure gi"eatly limits or prohibits the ability to infer sample unit occupancy across Districts. Conclusions and Recommendations Knowledge of bats in Montana remains largely based on distribution records (e.g.. Nicholson 1950, Hoffmann et al. 1969. Swenson 1970, Swenson and Bent 1977, Swenson and Shanks 1979. Shryerand Flath 1980, Foresman 2001). although there are a few published studies focusing on other aspects of the biology of Montana bats (e.g.. Jones et al. 1 973, Hendricks et al. 2000). In addition, several agency- funded projects have addressed information gaps that help guide management activities at the BLM Field Office or USFS Ranger District landscape scale (e.g., Worthington 1991a, 1991b, Hendricks and Kampwerth 2001. Hendricks et al, 2004, 2005), Nevertheless, there are no studies from Montana addressing how bats use forested land- scapes of different stand types, ages, and structural complexity (e.g., Thomas 1988, Kalcouniset al. 1999). These significant gaps in our understanding of bat use of Montana landscapes remain a major barrierto implementing effective conservation measures. The modification of landscapes is often considered the major cause of population fluctuations of many bat species. Measui"es for the conservation of bats frequently rely on knowledge of species-habitat relationships, and the use of distribution maps are often used to assess a species status. Thus, a thorough knowledge of bat distributions as they relate to habitat features is critical for implementing management and conservation actions. However, intensive population surveys of bats are difficult to conduct because of the nocturnal behavior of bats, their large home ranges, and difficulty of species identification while in flight. The latter problem is being addressed with improved acoustic and radio transmitter technology, but intensive population surveys may remain beyond the scope of most conservation/management programs. Modeling landscape features used by bats offers an alternative approach to multi-species population monitoring that may be useful for bat conservation at a landscape scale (Jaberg and Guisan 200 1 ). To do modeling effectively, however, a large base of distribution records is necessary that sample all species and account for all activities (foraging. roosting, rearing young, mating, hibernating). This may be difficult to do across a landscape the size of Montana, but a systematic and standard method of data collection across the state will make habitat modeling a more realistic possibility. Need for a Montana Bat Grid Montana currently lacks any statewide scheme for bat inventory and monitoring. This will be a crucial component for effective implementation of the state bat conservation plan now under develop- ment. The objectives of a state bat grid are 1 ) to inventory the presence of bat species using a standardized survey effort and sampling unit across the survey region, 2) collect baseline data on acoustic, morphologic, and genetic characteristics that serve as reference for bat species identification, and 3) to provide a baseline assessment of site occupany rates in order to identify changes in status overtime. Inventorying and monitoring bat distri- butions and trends at this scale will place us in a better position to address conservation issues as they arise. To date, none of these objectives has been thoroughly addressed in Montana. We recommend the development of a bat grid that is applied to all of Montana. Simply determining whether or not a species is present or breeding in a grid cell is a valuable way of monitoring the distribution and status of species overtime and relative to a variety of associated variables {Hayek 1994; Olson et ah 1997). The Oregon Bat Grid offers a suitable model scheme from which to design a state bat grid, although modification of some protocols may be necessaiy for application to Montana, due to limitations in personnel and agency support, as well as land access issues. It is beyond the scope of this report to explore the details of what comprises a state bat grid. Never- theless, the scheme that is eventually developed should include a hierarchical scale of data collec- tion, stratified by ecoregion and/or Ranger District, that allows inference of grid cell occupancy rates. To infer occupany rales, it is necessary to determine detection probabilities, and to do this will require multiple surveys of a subset of sites in each stra- tum. Strength of inference is very limited with i"egard to the status of a population in any single grid cell using only the data on presence/non- detection, and may best be regarded as being informative relative to management actions at broader spatial scales. However, this is also an approach for raising red flags at individual habitat patches or local regions (and grid cells) so that these rapid assessment surveys can be followed up with more detailed studies of a populations' status. 10 References Cited Adams. R. A. 2003. Bats ot the Rocky Mountain West. University Press of Colorado, Boulder, CO. 289 pp, Bogan, M. A., J. G Osborne, and J. A. Clarke. 1996. Observations on the bats at Badlands National Park, South Dakota- Prairie Naturalist 28:115-123. Cryan, P. M.. M. A. Bogan. and J. S. Allenbach. 2000. Effect of elevation on distribution of female bats in the Black Hills, South Dakota. Journal of Mammalogy 81:719-725. Dobkin, D. S., R. D. Gettinger, and M. G. Gerdes, 1995. Springtime movements, roost use, and foraging activity of Townsend's Big-eared Bat [Fiecotus toivnsejidi'i) in central Oregon. Great Basin Naturalist 55:315-321. Hendricks. P, C. Currier, and J. Carlson. 2004. Bats of the BLM Billings Field Office in south- central Montana, with emphasis on the Pryor Mountains. Report to Bureau of Land Manage- ment Billings Field Office. Montana Natural Heritage Program, Helena. MT. 19 pp. + appendices. Hendricks, P. D. L. Center, and S. Martinez. 2000. Bats of Azure Cave and the Little Rocky Moun- tains, Montana. Canadian Field-Naturalist 114:89-97. Hendricks, P.. and D. Kampwerth. 2001. Roost environments forbats using abandoned mines in southwestern Montana: a preliminary assessment. Report to the U.S. Bureau of Land Management. Montana Natural Heritage Program. Helena. 19 pp. Fenton, M. B. 1997. Science and the conservation of bats. Journal of Mammalogy 78:1-14. Foresman. K. R. 2001. The wild mammals of Montana. American Society of Mammalogists, Special Publication No. 12. 278 pp. Hayek, L.C. 1994. Analysis of amphibian biodiversity data. Pp. 207-269. In: W. R. Heyer, M. A. Donnelly. R, W. McDiarmid, L. C. Hayek, and M. S. Foster (edsj. Measuring and monitor- ing biological diversity: standard methods for amphibians. Smithsonian Institution Press. Washington, D.C 364 pp. Hayes, J. P. 1997. Temporal variation in activity of bats and the design of echolocation-monitoring studies. Journal of Mammalogy 78:514-524. Hendricks, P., S. Lenard, C. Currier, and J. Johnson. 2005. Bat use of highway bridges in south-central Montana. Report to Montana Department of Transportation. Montana Natural Heritage Program, Helena. 31pp. Hoffmann, R. S., D. L. Pattie, and J. F. Bell. 1969. The distribution of some mammals in Montana, II. Bats. Journal of Mammalogy 50:737-741. Humphrey, S. R. 1975. Nursery roosts and community diversity of Nearctic bats. Journal of Mammalogy 56:321-346. Jaberg, C, and A. Guisan. 2001. Modelling the distribution of bats in relation to landscape structure in a temperate mountain environment. Journal of Applied Ecology ?,^\ 1 169- 1 1 8 1 . Hayes, J. P. 2003. Habitat ecology and conserva- Jones, J. K., Jr., R. P. Lampe. C. A. Spenrath, and tion of bats in western coniferous forests. Pp. 81-1 19 ///Mammal community dynamics: management and conservation in the coniferous forests of western North America (C. J. Zabel and R. G. Anthony, eds.). Cambridge University Pi^ss, Cambridge, UK. 709 pp. T. H. Kunz. 1973. Notes on the distribution and natural history of bats in southeastern Montana. Occasional Papers. The Museum, Texas Tech University 15:1-12, 11 Kalcounis. M, C, K. A. Hobsoti. R. M. Brigham. and K. R. Hecker. 1999. Bat activity in the boreal forest: importance of stand type and vertical strata. Journal of Mammalogy 80:673- 682. Nagorsen,D.W., and R.M. Brigham. 1993- Bats of British Columbia. UBC Press. Vancouver, BC. 164 p. Nicholson, A. J. 1950. Arecord of the Spotted Bat {Eudenjia macidata) forMontana. Journal of Mammalogy 31:197. 0'Farrell,M.J., and W.L.Gannon. 1999. A comparison of acoustic versus capture techniques for the inventory of bats. Journal of Mammalogy 80:2^30. Olson. D. H.. W. R Leonard, and R. B. Bury (eds), 1997. Sampling amphibians in lentic habitats: methods and approaches for the Pacific North- west. Northwest Fauna 4:1-134- O'Shea, T. J.. M. A. Bogan. and L. E. Ellison. 2003. Monitoring trends in bat populations of the United States and territories: status of the science and recommendations forthe future. Wildlife SocietyBulletin 31: 16-29. Pierson, E. D. 1998. Tall trees, deep holes, and scarred landscapes: conservation biology of North American bats. Pp. 309-325. In Bat biology and conservation (T. H. Kunz and P. A. Racey. eds.). Smithsonian Institution Press, Washington. DC. 365 pp. Pierson, E. D., and 14 others. 1999. Species conservation assessment and strategy for Townsend's Big-eared Bat { Coryjwrbuius town send ii townsendii and Corynorb'mus town send ii pall esc ens). Idaho Conservation Effort, Idaho Department of Fish and Game, Boise, ID. 68 pp. Sheffield, S. R.. J. H. Shaw, G. A. Heidt, and L. R. McClenaghan. 1992. Guidelines for the protec- tion of bat roosts. Journal of Mammalogy 73:707-710. Sherwin. R, E.. W. L. Gannon, and J. S. Altenbach. 2003. Managing complex systems simply: understanding inherent variation in the use of roosts by Townsend's Big-eared Bat. Wildlife Society Bulletin 3 1 :62-72. Shryer. J. and D. L. Flath. 1980. First record of the Pallid Bat {Antrozoas pallldus) from Mon- tana, Great Basin Naturalist 40:115. Sokal, R. R., andRLRohlf. 1981. Biometry, second edition. W.H. Freeman. San Francisco, CA. 859 pp. Swenson. J. E. 1970. Notes on the distribution of Myotis leib'u in eastern Montana, Blue Jay 28:173-174. Swenson, J, E., and J. C. Bent. 1977. The bats of Yellowstone County, soulhcentral Montana- Proceedings of the Montana Academy of Sci- ences 37:82-84. Swenson, J. E., and G.F Shanks, Jr. 1979. Note- worthy records of bats from northeastern Mon- tana. Journal of Mammalogy 60:650-652. Thomas.D.W. 1988. The distribution of bats in dift'erent ages of Douglas-fir forests. Journal of Wildlife Management 52:6 19-626. vanZylldeJong, C. G 1985. Handbook of Canadian mammals. 2. Bats. National Museum of Natural Sciences. Ottawa, ON. 212 p. Worthinglon. D. J. 1991a. Abundance and distri- bution of bats in the Pryor Mountains of south centra] Montana and northeastern Wyoming. Report for the Bureau of Land Management Billings Resource Area and Custer National Forest, Montana Natural Heritage Program, Helena, MT 23 pp. Worthington, D. J. 1991b. Abundance, distribu- tion, and sexual segregation of bats in the Pryor Mountains of south central Montana. Master's Thesis. University of Montana. Missoula. MT. 41 pp. 12 Appendix A. Global/State Rank Definitions Heritage Program Ranks The international network of Natural Heritage Programs employs a standardized ranking system to denote global (range-wide) and state status. Species are assigned numeric ranks ranging from 1 to 5. reflecting the relative degree to which they are ''at-risk'\ Rank definitions are given below. A number of factors are considered in assigning ranks — the number, size and distribution of known ''occurrences" or populations, population trends (if known), habitat sensitivity, and threat. Factors in a species' life history that make it especially vulnerable are also considered (e.g.. dependence on a specific pollinator). Global R\NK DEFINrnONS (NatureServe2003) Gl Critically imperiled because ofextreme rarity and/or other factors making it highly vulnerable to extinction G2 Imperiled because of rarity and/or other factors making it vulnerable to extinction G3 Vulnerable because of rarity or restricted range and/or other factors, even though it may be abundant at some of its locations G4 Apparently secure, though it may be quite rare in parts of its range, especially at the periphery G5 Demonstrably secure, though it may be quite rare in parts of its range, especially at the periphery Tl-5 Infraspecific Taxon (trinomial) — The status of infraspecific laxa (subspecies or varieties) are indicated by a "T-rank" following the species" global rank State Rank Definitions 51 At high risk because of extremely limited and potentially declining numbers, extent and/or habitat, making it highly vulnerable to extirpation in the state 52 At risk because of very limited and potentially decliningnumbers, extent and/or habitat, making it vulnerable to extirpation in the state 53 Potentially at risk because of limited and potentially declining numbers, extent and/or habitat, even though it may be abundant in some areas 54 Uncommon but not rare {although it may be rare in parts of its range), and usually widespread. Apparently not vulnerable in most of its range, but possibly cause for 1 o n g-term cone er n 55 Common, widespread, and abundant (although it may be rare in parts of its range). Not vulnerable in most of its range Combination Ranks G#G# or S#S# Range Rank — A numeric range rank (e.g., G2G3) used to indicate uncertainty about the exact status of a taxon Qualifiers NR Not ranked Q Questionable taxonomy that may reduce conservation priority — Distinctiveness of this entity as a taxon at the current level is questionable; resolution of this unceHainty may result in change from a species to a subspecies or hybrid, or inclusion of this taxon in another taxon. with the resulting taxon having a lower-priority (numerically higher) conservation status rank Appendix A - 1 X Presumed Extinct — Species believed to be extinct throughout its range. Not located despite intensive searches of historical sites and other appropriate habitat, and virtually no likelihood that it will be rediscovered H Possibly Extinct — Species known from only historical occurrences, but may never-the- less still be extant; further searching needed U Unraiikable — Species currently unrankable due to lack of information or due to substan- tially conflicting information about status ortrends HYB Hybrid — Entity not ranked because it represents an interspecific hybrid and not a species ? Inexact Numeric Rank — Denotes inexact numeric rank C Captive or Cultivated Only — Species at present is extant only in captivity or cultivation, or as a reintroduced population not yet established A Accidental — Species is accidental or casual in Montana, in other words, infrequent and outside usual range. Includes species (usually birds or butterflies) recorded once or only a few times at a location. A few of these species may have bred on the one or two occa- sions they were recorded Z Zero Occurrences — Species is present but lacking practical consei"vation concern in Montana because there are no definable occurrences, although the taxon is native and appears regularly in Montana P Potential — Potential that species occurs in Montana but no extant or historic occurrences are accepted R Reported — Species reported in Montana but without a basis for either accepting or rejecting the report, or the report not yet reviewed locally. Some of these are very recent discoveries for which the program has not yet received first-hand information: others are old, obscure reports SYN Synonym — Species reported as occurring in Montana, but the Montana Natural Heritage Program does not recognize the taxon; therefore the species is not assigned a rank * A rank has been assigned and is under review. Contact the Montana Natural Heritage Program for assigned rank B Breeding — Rank refers to the breeding population of the species in Montana N Nonbreeding — Rank refers to the non-breeding population of the species in Montana Appendix A - 2 Appendix B. USFS Region 1 Montana Survey Sites for Bats, Summer 2005 n £ a U w a oa OJ 1 — 1 U < -J jj a n o S o ^ > - >- - O > >- E < u >- E MYEV(I) MYCA(I),MYLU(1) O > in > >- u c > S rjj > >- >- EE _ rn < D >- >^ SE [J O 15 u Q. O c D -J >- E — -' < E -J , o c -J ;>- E o z < -J -J — >- '-^ r E u < -J \o -^ ^ Cu Z >- D -J -^ >- c E o ^ > >s pj >- E > PJ ^ >- — ^D '-^ -J O >- -E O z < -J 3 1 — 1 < 00 Dp < On < < 3 < < CO 3 3 3 1 — - CO 3 o 2 DJj < O m o o o r- — o O O O 00 o -t O O -t CO u-1 O CO ■n O CO -4-1 O -J n > U ■- r . ^1 _ DC u u a 13 ij c_ ■u IJ U 1- u O U 1-7 o u oj U •— ■5 o -J n O U o 3 o O u OJ 1- U t! U ij c Q. 3 fj 5 a. 00 O z ■n CO fN ■n PJ m ■rf CO r- CO OJ z O r- -rt — CO r^ oc o> — O Z On On r- 00 fi z ui O CpO CO 5 z oc "Ti Z O 00 o z r- a^ ■n PJ r- O rTi OJ O Z o "n z O r^i O ■O pj ■o ■r, '^ r- CO -^ z o o O o p:J O G^ Z CA CO n-^ O ui CO CO Z o O pi 00 z CO ■n o ■n a* m rTi a- '^ Z m O O ■n CO -^ ?7 U OJ -J n -J -J -J E o O in O n "a a a £ o IJ a. 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MYLU (104) o|£ Z S o < ^> -J f^ >- 3e > PJ >- E o z < > PJ >- E > E D - >- z < -J ^ ij-l fN ^iTd _] > -^ >- >- ;iM w SS E ^ > X S H gj H >- ^ ^EE C" 3- ^ :r;> X [X r ^ r-i Cl. i>^ >- w uj_E S go g >. r'D ^ H U -J R ^ ^ ^ E S ^ ^ Fj u > < ^ ^ 3ee fN — rJ tJ > Ph Z tJJ £3 < ^ > >- E u < > E 3 ^^ Cu fj 4> a < DO < DO < in 3 c 1— i 00 c 3 1 — 1 3 -TJ "3 — ^ 3 "3 1—1 m ■n O 1/1 c ■n -t "Tj 1/1 4> z J". 3JJ a 1 >> U a. 1— "5 O IS U B CO r- 3 n CD 1^ 1 c C 3 u X > U c a: y u en B r— 1 U p E OJ -:^ n -J n P n U as Z r- r- CO O ■O PJ o ■n O Z O O PJ o CpO ■n a^ z o PJ o Z CO O CO rj lo ui CO CO Z -^ ■n ■n O -^ ,3 z 00 z ■n m 00 \C m t Z 1 — 1 r7 z fN 00 rj O -2 O O ;^ < -S ^. o --- ' ^ S ' S CQ kJ ^ O ri o -J -2 ^ > O UJ > >- > o o E E Appendix B - 2 -4-1 Cl. a U » -4-1 eg CO -J i^ s ~" 2 ^ ^ S -^ a. z >_ LANO(l), MYCA(l), MYLU(I) < U >- < -3 00 -J >- p:. a. o ^ 3.3 < -J -J ;^ u < -J X >- 3 So ^> ^>- >^ >- LACI (seen), LAND (6), MYCA(2), MYEV(I), MYVO (2) n-, --^ '-' r < =J ^ y d - -] S S :^ ^ ■ ^ >- ^ 3~ ^ :^ o > ^ ^3S < ^ -J - >- z < -J MYLU(4). 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MYVO (1) D -J >- O z < -J eg P < 3 3 1 — - 3 1 — - •n 3 ■n 3 "5 CO DC < o 00 < 2 1 — J CO CpO -t O in in o o o r^. ■i- o o 1—1 o in o •-0 a-' ■n 01 B eg Z -4-1 1/) ^ 5b a a u p o6 o U o b n H o O N a U OJ 6 r- ^ 1 B - H O o a- -3 :^ O Lh U i U 1- > CO 'ij-. c c c in PJ CpO in CO Z Z in in H in o in -■o o CO in in in G^ in z o CO o r- in CO O in z ■n -t ■n ■n PJ CO CO o Z CO ■>^ P^ o 00 n o a -c qj <4^ Ui J .5 =3 u. vi? i^ F ^1 c := v^ T^ i:j c ^ C3 LA ^ ^ n ^ rr< O ■ c LA n < -J '-' Ed CO t- > \j >H :^ 5- o ■'-. ^ c ^ ^~ !> m ■^ LA V X an g 03 i^ U s Vj ^ s _^ E St. lA N 03 Vj LA O LU O 1 ^ Ed -J Ed e -I "« -> =0 >^ O ':^- 1^ 1^ LU u ^ r n Ed on ■^ ■3 -J "S ^ ^ ^ ^ D c/^ 00 ^ -1 i^ 1> [A ";^ S D J=5 T3 ^ LA >H Ed LA ^ >- U- n s , , c < J^ 1> ? ^ > ■- 4-1 >- 1 00 o P' 1^ ^ c: fl> c rin o DO n s fn -J DO P:h c c lU 1^ fli 00 Ed ^ U c: L^ 11 (J L-i "F -5 S: —1 <*— -» ' 1^ o ri >i_ >^ f3 lA h U Z l^ ■^ O H O i lA -t ^ U z < > O c U LA 1J LU -> o LA ■T3 >- pi- O rt S S S 1J LA 1J m LA lA 1J -J y Ed o O '^ ce: Cl. O ^ >> in LA ^ s Appendix B - 3 P. rr, O < -J > >- < -J > O > >- E 2 1— U < -J > >- o O > >- 00 1— 1 U < -J Co o o > >- O z < -J 1),MYLU(I), YVO (3) ■n O z < -J o > >- E LACl (7). LANO (56), MYEV(l), MYLU(5), MYVO(l) ! 3d 3 E^ ■n < -J >- >- s ^ ^ > S ~ ^ c "^ 3 --' -, E i> = i a ^ ^ "^ US' LA eg CO 1 — 1 U < -J U >- o z < -J -J >- Q- O z < -J _] >- 0- O z < -J > >- < -J u >- E < -J > pj ;^ E Cu Z PJ < o z < -J >- E o ^ ;S S OJ =[ 3 '3 r— 1 3 ^ ^ 3 "5 '5 F -^ E ** eg — p k— i k-l 1— » 1 — J 1 — 1 ►— i -TJ 1—1 ?3 f^i r^r ■^ iri r- CO o> o 1 — < C= 1— *- ' — 1 ' ' ' — ' — ' — ' rj ri T> CD £ '^ X &ft ;^ "z: > ^^ C^P t r- •n ■n a> o 00 r^ ^ O r- r- rj O "J". ^ . — ;; >- c ^ c U 1> -J c o >^ c C3 U c o 1- U o o DJj o U = o. u u u Lh o p:. o .£ 5 ^ i- pi ^^. cd s X >* Q _Dp CQ ■^ c -J LU ™ ^ t; CD :: >: ^ ,-3 5 S^S "? "O Z Z Z Z Z Z z z i>:r ^-£E ^ 00 -1- \o O' — o JN U-, t '^ "O =c C-i ' — 1 ■ — ' rj ^D o CO O 00 r^. a*i — ^— Ov ^N U-, ■n ^ ■n 00 \C rj r- -t r^ oc CO o o ■n y-i r- CO ?i c <■ >^ -^ r^ 00 :n c o CO DO u C on n CQ 13 O 3 c CQ 1> ■1—' c; i:; (> ^ -s ^ -1 = ^ ^' s = ^ r - U "5 i-Tp O. tu i-Tp 1 — J ^ 3 -^ 1 — 1 ^ -^ -= -i 00 5 £ -i^ ^ ^ -1^ ^ ^ ^ -1*: ^ ^5> c3 ^ B c3 J5 B k ^ J5 a VI U U U U U U U U U s ^ ^ ^ ^ ^ ^ =« ^ ^ ^ 3^s s O [/I •Ji on LA LA « LA LA LA ^ s ;£ =^ 1? ^ U. T> ?= S S ^ S ^ 5 ^ S 5 J ?^ -C — ' * ^5 -J 1> -J U U -J t-1 -J ?g Appendix B - 4 Appendix C. Distribution Maps for Bats in Montana u O CO □ o Q n QQ c To ■m D lU Hn r; u^ lO n ^ o O u^ na rj 1^ 4> +- ^ 6 3 O, Appendix C - 2 ^ —I I o o I o o ^^ CD m •D en m C ts> t/> c en O £^ g I is ^ ^ ^ ^ O 4^ ift Q « S 8 K 5 ^ ^ 8 Q t/> ^ Appendix C - 3 O i at m ■D o V) a: qs -ll- fTl 3 ^ ^ *0 n ir> ii^ n to X W iif? n ^M rj 4- 1^ LL + « Appendix C - 4 to to tn o 2 G Oh (3 6 ^ CJ Q. + m Appendix C - 7 Appendix C - 8 I o I o >■ E o □ o Appendix C - 9 Appendix C - 10 Appendix C - 1 1 I" I m m ti a ^ i t r HJ > c ^ R1 i^ u-t n o s \n ■r^ rM o M lb D- B-L CO ti ^ 5 « ■ Appendix C - 12 I c o Appendix C - 13 to o c Appendix C - 14 Appendix C - 15 .*2 i E E Appendix C - 16