Bat Diversity, Resource Use and Activity Patterns Along a Sonoran Desert Riparian Corridor

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Bat Diversity, Resource Use and Activity Patterns along a Sonoran Desert Riparian Corridor

Item Type text; Electronic Thesis

Authors Buecher, Debbie Jane Cramer

Publisher The University of Arizona.

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Link to Item http://hdl.handle.net/10150/193329 BAT DIVERSITY, RESOURCE USE AND ACTIVITY PATTERNS ALONG

A SONORAN DESERT RIPARIAN CORRIDOR

Deborah Jane Cramer Buecher

A Thesis Submitted to the Faculty of the

SCHOOL OF NATURAL RESOURCES

In Partial Fulfillment of the Requirements For the Degree of

MASTER OF SCIENCE WITH A MAJOR IN WILDLIFE AND FISHERIES SCIENCE

In the Graduate College

THE UNIVERSITY OF ARIZONA

2007 2

STATEMENT BY AUTHOR

This thesis has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library.

Brief quotations from this thesis are allowable without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the copyright holder.

Signed:____Debbie C. Buecher______

APPROVAL BY THESIS DIRECTOR

This thesis has been approved on the date shown below:

______12/04/07_____ John L. Koprowski Date Associate Professor of Wildlife and Fisheries Resources 3

ACKNOWLEDGMENTS This study would not have been possible without generous support and patience of my major professor, John L. Koprowski. He encouraged my interest in bat ecology (even though he clearly preferred squirrels) and guided me through the theoretical maze that is fieldwork in ecology. I also thank my graduate committee for their assistance, scientific expertise and tremendous patience during this process. I thank Coronado National Forest, Santa Catalina Ranger District (USDA) for allowing me to conduct research in Sabino Canyon. They were most generous in allowing me access to Sabino Canyon Recreational Area at all hours of the day and night. Forest Service personnel helped me through the FS Special Use Permit process and Forest Service biologist Josh Taiz was especially supportive of my study. The Nongame Branch of Arizona Game and Fish Department helped me acquire necessary Scientific Collecting Permit to work on bats in Arizona. This project was funded by an Arizona Game and Fish - Urban Heritage Grant, a grant from T & E, Inc., and a financial donation from The Friends of Sabino Canyon. A number of people made constructive editorial comments of earlier versions of this thesis, greatly improving it although they are not to blame for any errors or oversites that might appear, these are entirely my fault. My ‘editorial staff’ include: Annita Harlan, Scott Clemans, and members of the ‘Squirrel Lab’. Volunteers are often the unsung heroes of any large project. I have been most fortunate to have many fine people assist my mist-netting efforts along Sabino Creek. These friends have helped me in the dark and well past their bedtimes! My adventurous volunteers included: Scott Clemans, Shirley Cramer, Cathy Wasmann, Tania Yatskievych, Dale Mann, Natalie Furrey, Joan Martin, Kelly Hutton, Julia Tenen, Sue Clemans, Eric Martin, Bret Pasch, Brenna Larkin, Karen Munroe, Sid Schwartz, Sherry Barker, Gary Tenen, Leslie Pansing, Deputy Doug (Myrvold), Kari Bressmer, George Rallis, Alesha Williams, Barry Rosenbaum, Michelle Bezanson, Tovah Salazar, Bill & Jill Kellerman, Russell Davis, Geert-Henning & Ester Schauser (Germany), Joy & Steve Gurgevich and the 2005 Colorado College Ecology class. 4

DEDICATION

This thesis is dedicated to my husband Robert, who allowed me to pursue a lifelong dream by providing both emotional and financial support during this entire project. He was my engineering wizard in all things technical and without his encouragement this venture would have faltered in concept only.

To my Father who encouraged me when I began this journey but who was unfortunately not there to see its completion.

To my darling Mother who taught me that nature is wondrous and worthy of study but who insisted that I start my thesis…’It was a dark and stormy night…’

And last but not least, to my dear friend and mentor Ronnie Sidner, who spent many late nights in the field with me, sharing her tremendous knowledge of bats and encouraging me during endless phases of this endeavor.

“Scientific discovery consists in the interpretation for our own

convenience of a system of existence, which has been made with

no eye to our convenience at all.”

Norbert Wiener 5

TABLE OF CONTENTS

LIST OF FIGURES………………………………………………………………..……...8

LIST OF TABLES……………………………………………………………….………11

ABSTRACT……………………………………………………………………….……..12

INTRODUCTION………………………………………………….…….………….…..13

PRESENT STUDY……………………………………………………….………..…….15

REFERENCES………………………………………………………………….……….18

APPENDIX A: RIPARIAN AREAS AS HOT SPOTS OF BAT DIVERSITY IN ARID ENVIRONMENTS ………………………………………….……..….…………21

ABSTRACT……………………………………………………………….…....………..21

INTRODUCTION………………………………………………………….……………22

METHODS………………………………………………………………….….………..25

Study area…………………………………………………………………….….…….25

Capture methods…………………………………………………………….….….….28

Resource use…………………………………………………………………….….…30

Statistical analysis…………………………………………………………….….……30

RESULTS……………………………………………………………….……….………31

Species Diversity…………………………………………………..…………....…….31

Resource Use………………………………………………………….……….…..….32

Temporal Use………………………………………………………….…….…..….…32

Spatial Use………………………………………………………………….……..…..34

DISCUSSION…………………………………………………………….….…….....….34

Species diversity...... 34 6

TABLE OF CONTENTS - Continued

Resource use…………………………………………………………………………..36

Temporal Partitioning……………………………………………………..…………..36

Spatial Partitioning………………………………………………………...…………..37

CONSERVATION IMPLICATIONS……………………………………….…………..38

REFERENCES……………………………………………..……………………………41

APPENDIX B: FINDING THAT 4-STAR DINER OR HOW BATS MIGHT ‘ANTICIPATE’ PRODUCTIVE FORAGING AREAS………………………………...57

ABSTRACT………………………………………………………………..…….………57

INTRODUCTION……………………………………………………………………….58

STUDY AREA…………………………………………………………………………..59

METHODS………………………………………………………………..……………..61

Acoustic Sampling……………………………………………….……………..……..61

Data Analysis………………………………………………………………………….63

RESULTS………………………………………………………………………………..65

DISCUSSION……………………………………………………….…….……………..66

Proposed Model of Bat Distributions in Canyon Lands………………………………66

Physics and Meteorology……………………………………………….…………..…66

Hydrology……………………………………………………………….…………….67

Entomology…………………………………………………………….……….……..68

Mammalogy………………………………………………………………….………..69

Disciplines Combined…………………………………………………………………69 7

TABLE OF CONTENTS - Continued

MANAGEMENT RECOMMENDATIONS…………………………………………….70

REFERENCES…………………………………………………………………….…….71

APPENDIX C: SPECIES PRESENCE AND DISTRIBUTION....……………………..84

APPENDIX D: BAT CAPTURE DATA FOR SABINO CANYON 2002-2005……….89

APPENDIX E: COMPARATIVE ANALYSIS …………...…………………….…….128

APPENDIX F: SPECIES ACCOUNTS, SEASONAL DISTRIBUTIONS AND COMPARATIVE MEASUREMENTS OF THE SEVEN MOST FREQUENTLY CAPTURED SPECIES ALONG SABINO CREEK, 2002-2005……………………..134

REFERENCES……………………………………………………………………..…..151 8

LIST OF FIGURES

FIGURE 1.1, Species presence by month (2002-2005) along Sabino Canyon, documented with mist netting. The greatest species diversity occurs when there is little-to-no water available along the creek (May and June)……………….…..…...52

FIGURE 1.2, Seasonal distribution of a) number of bat species and b) numbers of individuals captured along Sabino Canyon during 2002-2005 showing ± one standard error……………………………………………………………….…….…53

FIGURE 1.3, Species presence at 12 pools along Sabino Canyon, documented with mist netting 2002-2005. Pool 1 is at the wildland-urban interface at the lower end of the canyon and Pool 12 is 5.1 km upstream from Pool 1. The seven most frequently captured species are well distributed along Sabino Canyon when the creek is flowing…….....……………………………..…..54

FIGURE 1.4, Number of bat species and individuals captured at pools along Sabino Creek as it changes from flowing water in the spring to isolated pools in early summer and then returning to flowing water after the onset of summer monsoon storms. High capture rate in early fall reflects migratory period of Sonoran Desert bats. Dashed line may reflect normal slope without migratory behavior……………………………………...... ………………………….………55

FIGURE 1.5, Linear regression of the number of bat species captured/night during summers 2002-2005. Negative slope reflects changes in species response to water availability along Sabino Creek. Early in the summer, when there is limited water, bat species are concentrated at the few remaining pools. However, once the summer monsoons begin, bats are able to disperse across the landscape to find drinking water…………………....………….56

FIGURE 2.1, Species presence along Sabino Creek documented with year-round mist netting 2002-2005 at 12 ephemeral pools along the canyon floor………....……….77

FIGURE 2.2, Map of Sabino Canyon showing sample sites used for the comparative acoustic sampling study. Sites 1 & 2 are located in riparian deciduous vegetation and Sites 3 & 4 are located in Sonoran desertscrub………….....…………………..78

FIGURE 2.3, Examples of passive acoustic sample site in deciduous riparian vegetation (top) and Sonoran desertscrub sample site (bottom)…….....………..….79

FIGURE 2.4, Plot of passive acoustic data by sample period along Sabino Creek – June-December 2004……………………………………………………...……..80 9

LIST OF FIGURES - Continued

FIGURE 2.5, Plot of passive acoustic data by sample period along Sabino Creek – January- December 2005…………………………………………..…….....……….81

FIGURE 2.6, A view of Acoustic Sample Site 3, where water flows from right to left. Arrow indicates the granite ridge extending from the west causing a 90o turn in Sabino Creek’s flowline, creating a possible eddy just upstream from the ridge…………………………………………………….....…….83

FIGURE 3.1, Map of Sabino Canyon Recreational Area, Santa Catalina Mountains – with netting locations highlighted………………………………….……….....……86

FIGURE 3.2, Species presence (2002-2005) documented with mist netting along Sabino Canyon shown by pool locations. Pool numbers refer to Table 3.2………………………………………………………………….....………87

FIGURE 3.3. Cumulative species curve with the number species documented along Sabino Creek plotted against total number of captured bats…………….....……….88

FIGURE 5.1, Sex ratios of bats captured along Sabino Creek during 2002-2003, prior to post-Aspen Fire sedimentation of pools………………………….....….…131

FIGURE 5.2, Sex ratios of bats captured along Sabino Creek during 2004-2005, after post-Aspen Fire sedimentation of pools…………………………….….…….132

FIGURE 6.1, Monthly distribution of sexes for big brown bats along Sabino Creek, Arizona 2002-2005, with limited netting during July……………………….….…135

FIGURE 6.2, Differential forearm lengths between male and female big brown bats...135

FIGURE 6.3, Monthly distribution of sexes for big free-tailed bats along Sabino Creek, Arizona 2002-2005……..……………….…………………………………………138

FIGURE 6.4, Differential forearm lengths between male and female big free-tailed bats, note reverse sexual dimorphism than usually seen between sexes in bats (n=24)...138

FIGURE 6.5, Monthly distribution of sexes for hoary bats, along Sabino Creek 2002- 2005, with limited netting during July…………………………..…………...……140

FIGURE 6.6, Differential forearm lengths between male and female hoary bats….…140

FIGURE 6.7, Monthly distribution of sexes for Mexican free-tailed bats along Sabino Creek 2002-2005, with limited netting during July………………….……………142 10

LIST OF FIGURES – Continued

FIGURE 6.8, Differential forearm lengths between male and female Mexican free- tailed bats……………………………………………………………………..……142

FIGURE 6.9, Monthly distribution of sexes for pocketed free-tailed bats along Sabino Creek 2002-2005, with limited netting during July……..……………….…...……145

FIGURE 6.10, Differential forearm lengths between male and female pocketed free- tailed bats, note that the females have shorter forearms than males (n = 251)….…145

FIGURE 6.11, Monthly distribution of sexes for western pipistrelle bats along Sabino Creek 2002-2005, with limited netting during July……………………..…………148

FIGURE 6.12, Differential forearm lengths between male and female western pipistrelles, females carry twins and may have a longer forearm to compensate (n = 192)……………………………………………………………………….…..148

FIGURE 6.13, Monthly distribution of sexes for Yuma myotis bats along Sabino Creek, Arizona 2002-2005………………………………………………………..….……150

FIGURE 6.14, Differential forearm lengths between male and female Yuma myotis...150 11

LIST OF TABLES

TABLE 1.1, Species abbreviation, scientific name, common name and numbers of bats of each species caught during 2002-2005 along Sabino Canyon with comparison of pre-fire vs. post fire capture rates. Differences in overall captures (2002-2003 vs. 2004-2005) reflect changes in netting effort and impacts from post-fire sedimentation……………………………………………………………………….51

TABLE 2.1, Statistical analysis of average number of hourly foraging call files at four sample sites, across 3 seasons (i.e., winter excluded) and between two habitats during June 2004 – December 2005……………………………………….82

TABLE 3.1, Compilation of bat species documented from Santa Catalina Mountains and Sabino Canyon Recreational Area, Pima County, Arizona…………...…….84

TABLE 3.2, Map locations and vegetation associations for mist net sites along Sabino Creek, 2002-2005……………………………………………………...…85

TABLE 4.1, Bat capture data from Sabino Canyon 2002-2005…….………….……….89

TABLE 5.1, To capture free-flying bats we used 2-ply 50 denier, 38 mm mesh nylon mist nets of either 6, 9, or 12 m stretched across flowing or pooled water along Sabino Creek (Kunz and Kurta 1988). We placed our nets perpendicular to the creek’s flow and recorded the direction that the bat was flying (upstream vs. downstream) when captured to evaluate any directional preference by bats (Adams and Simmons 2002). Of the seven most frequently captured species, N. femorosaccus was the only species that indicated any preference for flight direction during 2002-2005……....………………………………………...... ……129

TABLE 5.2, When netting we stacked two nets vertically with a single shelf overlap between upper and lower nets (covering a ~3.6 m vertical flight window) at each site and recorded the height a bat was flying upon capture. Of the seven most frequently captured species, only P. hesperus and M. yumanensis showed strong evidence for foraging just above the water’s surface along Sabino Creek during 2002-2005………………………………………………………………….....……130

TABLE 5.3, Results of comparative bat research across three Southwestern deserts, with hourly capture rates. These data suggest resource levels for bats are variable across different biotic communities and elevations ……...... ……..……...…..133 12

ABSTRACT

I quantified the bat assemblage associated with a Sonoran Desert riparian corridor at a wildland-urban interface using mist netting (2002-2005) to assess differential spatial and temporal resource use. My capture rate was high (17 species and 961 individuals) considering the aridity of the area; however, landscape complexity of this montane region undoubtedly contributes to foraging opportunities. I found that bats were distributed along the canyon when water was plentiful but their activity was concentrated at isolated pools during dry periods. I also found temporal variation in pool-use by the most frequently captured species. I conducted an acoustic study to measure bat-use between deciduous riparian and Sonoran desertscrub communities. I measured activity levels using number of acoustic call files. I found greater bat foraging in desertscrub and used a multidisciplinary approach to determine why bats might use the more arid environment.

All capture data and supporting analyses are included in appendices. 13

INTRODUCTION

The structural complexity of riparian landscapes provides critical habitat for many plant and animal species, supporting tremendous biodiversity (Ohmart and Anderson

1982). In the desert Southwest, these landscapes contribute to greater species diversity than their proportion of land area (Naiman et al. 1993, Neary et al. 2005). Increased bat diversity is often associated with riparian landscapes because of greater resource availability (Racey 1998, Grindal et al. 1999, Holloway and Barclay 2000), especially access to drinking water. However, riparian resources in arid landscapes can be temporally and/or spatially variable (Bell 1980, Gregory et al. 1991) and competition over resources may occur. Additionally, riparian environments often suffer from anthropogenic degradation resulting from increased urbanization, associated pollution and groundwater extraction, introduction of exotic species (Steiner et al. 2000), and most recently, impacts from catastrophic forest fires.

In hot, arid environments, multiple bat species often co-exist (Bell 1980,

Hoffmeister 1986, Schmidt 1999) where access to drinking water limits species distributions (Cutler 1996, Kuenzi and Morrison 2003, Rabe and Rosenstock 2005).

Consequently, competition over water and foraging resources likely occurs within these systems (Williams et al. 2006). Bats serve as an excellent model system to study resource competition (Findley and Black 1983) because they are long-lived K-strategists that live in stable, species-rich communities (Fenton 1997, Fleming et al. 1972, McNab

1971). We investigated resource partitioning by a Sonoran Desert bat assemblage along a riparian corridor at a wildland-urban interface to better understand how multiple bat 14

species co-exist within a streamside environment in an arid region. We used standard mist netting and passive acoustic sampling to quantify the community assemblage and to analyze impacts of seasonally ephemeral water resources on species’ presence. In any landscape, urbanization removes or alters native vegetation with buildings, roads and introduction of non-native landscaping (Radeloff et al. 2005). Although wildland habitat offers critical resources, its proximity to an urban setting results in anthropogenic degradation of habitat. Because this region is a hotspot of biodiversity (Mittermeier et al.

2004), it is important to understand the mechanisms that facilitate coexistence by bats

(Saunders and Barclay 1992) for long-term management and preservation of this unique environment. 15

PRESENT STUDY

I present four years of evidence for temporal and spatial resource use by bats along a Sonoran Desert riparian corridor at a wildland-urban interface, including detailed capture records in the appendices. Descriptions of bat diversity, resource use and activity patterns, determined from mist netting, are formatted for submission to Journal of

Biological Conservation (Elsevier Ltd). Results of passive acoustic sampling are formatted for Proceedings from the 6th Conference on Research and Resource

Management in the Southwestern Deserts: Borders, Boundaries and Time Scales – May

2006 (in press). Following is a summary of our major results.

We describe and quantify the bat assemblage associated with a Sonoran Desert

riparian corridor at a wildland-urban interface to assess differential spatial and temporal

resource use. We found tremendous bat diversity, 17 species representing three bat

families - Vespertilionidae, Molossidae and Phyllostomidae, along Sabino Canyon, Santa

Catalina Mountains, Arizona. This region has floristic and faunal ties to the Sierra Madre

Occidental, Mexico (McLaughlin 1995) and the proximity of our site to tropical bat

diversity (Nowak 1994) no doubt contributes to local diversity. Additionally, resources

available along arid Southwest riverine corridors in mountainous regions provide

tremendous landscape complexity for foraging by insectivorous bats.

We captured seven species in large enough numbers for statistical analysis, of

which western pipistrelles (Pipistrellus hesperus) and pocketed free-tailed bats

(Nyctinomops femorosaccus) were active year-round along the canyon. Mexican free-

tailed bats (Tadarida brasiliensis) were present all months except July but we did limited 16

netting during July due to weather conditions. Big brown bats (Eptesicus fuscus), hoary bats (Lasiurus cinereus), Yuma myotis (Myotis yumanensis) and big free-tailed bats (N. macrotis) were seasonal visitors. We found that bats distribute themselves spatially along the canyon when water is plentiful, partitioning water resources where possible.

However, when seasonal drying conditions reduce water resources to isolated pools, numbers of species and individuals increase at remaining pools as they compete for limited water. We also found distinct temporal variation in pool-use by the seven most frequently captured species, with P. hesperus the earliest species to arrive at pools and N. macrotis the latest.

The high species richness and numbers of animals captured along the stream corridor emphasizes the incredible importance of riparian systems to desert bats. SCRA represents only ~1.0% of the area of the Santa Catalina Mountains and yet reflects 82% of the range’s bat diversity. Additionally, the riparian ecotone supports 64% of the statewide bat diversity within only 567 hectares of land area.

We further investigated bat-use along a riparian system with passive acoustic monitoring. We sampled 21 nights (261 hours of passive acoustic sampling) in 2004 and

2005, recording 49,480 bat-call files at four sites along the stream corridor. Over 15 months, one bat detector recorded 12,260 call files, a second detector recorded 7,548 files, a third recorded 15,003 files and the fourth recorded 14,669 files. We expected that peak foraging activity, reflected by number of call files, would be along the lower portion of the canyon where dense deciduous riparian vegetation supports greater insect biomass

(Iwata et al. 2003, Mosley et al. 2006). However, we found that upper reaches of the 17

canyon often had greater bat activity and the detector at one site in Sonoran desertscrub recorded the greatest number of calls of all sites along the creek. We proposed a hypothetical model to investigate why bats might forage extensively at particular sites along an arid riparian corridor. Our proposed model incorporates an interdisciplinary approach to understand foraging behavior by bats, providing insight on how bats might perceive landscapes and food resources. 18

REFERENCES

Bell, G. P. 1980. Habitat use and response to patches of prey by desert insectivorous bats. Canadian Journal of Zoology. 58:1876-1883.

Cutler, P. L., 1996. Wildlife use of two artificial water developments on the Cabeza Prieta National Wildlife Refuge, Southwestern Arizona. M.S. thesis, School of Renewable Natural Resources, University of Arizona, Tucson.

Fenton, M. B. 1997. Science and the conservation of bats. Journal of Mammalogy 78:1- 14.

Findley, J. S., and H. Black. 1983. Morphological and dietary structuring of a Zambian insectivorous bat community. Ecology 64:625-630.

Fleming, T. H., E. T. Hooper, and D. E. Wilson. 1972. Three central American bat communities: structure, reproductive cycles, and movement patterns. Ecology 53:555-569.

Gregory, S. V., F. J. Swanson, W. A. McKee, and K. W. Cummins. 1991. An ecosystem perspective of riparian zones. BioScience 41:540-551.

Grindal, S. D., J. L. Morissette, and R. M. Barclay. 1999. Concentrations of bat activity in riparian habitats over an elevational gradient. Canadian Journal of Zoology 77:972-977.

Hoffmeister, D. F. 1986. Mammals of Arizona. University of Arizona Press, Tucson, Arizona, USA.

Holloway, G. L. and R. M. R. Barclay. 2000. Importance of prairie riparian zones to bats in southeastern Alberta. Ecoscience 7(2):115-122.

Iwata, T., S. Nakano, and M. Murakami. 2003. Stream meanders increase insectivorous bird abundance in riparian deciduous forests. Ecography 26:325-337.

Kuenzi, A. J., and M. L. Morrison. 2003. Temporal patterns of bat activity in southern Arizona. Journal of Wildlife Management 67:52-64.

McLaughlin, S. P. 1995. An overview of the flora of the Sky Islands of southeastern Arizona: diversity, affinities, and insularity. Pp. 60-70 in Biodiversity and management of the Madrean Archipelago: the Sky Islands of Southwestern United States and Northwestern Mexico. Gen. Tech. Rep. GTR-RM-264. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 19

McNab, B. K. 1971. The structure of tropical bat faunas. Ecology52:352-358.

Mittermeier, R. A., P. R. Gil, M. Hoffmann, and J. Pilgrim. 2004. Hotspots revisited: Earth’s biologically richest and most endangered terrestrial ecosystems.CEMEX, 205-216.

Mosley, E., S. B. Holmes, and E. Nol. 2006. Songbird diversity and movement in upland and riparian habitats in the boreal mixedwood forest of northeastern Ontario. Canadian Journal of Forest Research 36:1149-1164.

Naiman, R. J., H. Decamps, and M. Pollock. 1993. The role of riparian corridors in maintaining regional biodiversity. Ecological Applications 3:209-212.

Neary, D. G., P. F. Ffolliott, and F. DeBano 2005. Hydrology, ecology, and management of riparian areas in the Madrean Archipelago. Pp. 316-319 in: Connecting mountain islands and desert seas: biodiversity and management of the Madrean Archipelago II. Gen. Tech. Rep. RMRS-P-36. Fort Collins, CO; U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station.

Nowak, R. M. 1994. Walker’s bats of the world. Johns Hopkins University Press, Baltimore.

Ohmart, R. D., and B. W. Anderson. 1982. North American desert riparian ecosystems. Pp.433-466 In: Bender, G.L. (ed.), Reference handbook on the Deserts of North America Greenwood Press, Westport, Connecticut, pp. 433-479.

Rabe, M. J., and S. S. Rosenstock. 2005, Influence of water size and type on bat captures in the lower Sonoran Desert. Western North American Naturalist 65:87-90.

Racey, P.A., 1998. Ecology of European bats in relation to their conservation. In: Kunz, T. H., Racey, P.A., (Eds.), Bat biology and conservation. Smithsonian Institution Press, Washington, pp. 249-260.

Radeloff, V. C., R. B. Hammer, S. I. Stewart, J. S. Fried, S. S. Holcomb, and J. F. McKeefry, 2005. The wildland-urban interface in the United States. Ecological Applications 15, 799-805.

Schmidt, S. L. 1999. Activity patterns of California leaf-nosed and other bats at wildlife water developments in the Sonoran Desert. M.S. thesis, School of Renewable Natural Resources, University of Arizona, Tucson.

Saunders, M. B., and R. M. R. Barclay. 1992. Ecomorphology of insectivorous bats: a test of predictions using two morphologically similar species. Ecology 73:1335- 1345. 20

Steiner, R., J. Blair, L. McSherry, S. Guhathakurta, J. Marruffo, and M. Holm. 2000. A watershed at a watershed: the potential for environmentally sensitive area protection in the upper San Pedro Drainage Basin (Mexico and USA). Landscape and Urban Planning 49:129-148.

Williams, J. A., M. J. O’Farrell, and B. R. Riddle. 2006. Habitat use by bats in a riparian corridor of the Mojave desert in southern Nevada. Journal of Mammalogy 87:1145-1153. 21

APPENDIX A: RIPARIAN AREAS AS HOT SPOTS OF BAT DIVERSITY IN ARID ENVIRONMENTS

ABSTRACT

Riparian corridors in xeric environments act as contact zones between deciduous streamside and arid-adapted biotic communities, providing landscape complexity critical for wildlife. For bats foraging along riparian zones, this ecotone can supply food resources, day-roosts and access to drinking water. Yet, riverine areas are sensitive to anthropogenic alteration of habitat. We used standard mist netting to investigate the bat assemblage at a wildland-urban interface associated with a Sonoran Desert riparian corridor. We determined species richness and assessed differential spatial and temporal resource use. We documented 17 species, representing three bat families:

Vespertilionidae, Molossidae and Phyllostomidae. We found that bats distribute themselves spatially along the canyon when water is plentiful, partitioning water resources where possible. When seasonal drying reduces water availability to isolated pools, numbers of species and individuals increase at remaining pools as bats compete for limited water. We also found distinct temporal variation in pool-use by the most frequently captured species. Although this desert riparian corridor provides critical habitat for bats, increased urbanization, associated groundwater extraction and recent forest fires, aggravated by regional drought, threaten this biologically diverse system.

Riverine environments in arid landscapes are crucial for wildlife, both for foraging and as migratory corridors. Loss and degradation of riparian areas in arid regions put wildlife, 22

particularly bats, at risk and preservation of this ecologically important resource should be a management priority.

Keywords: Arizona, biodiversity, water availability, arid landscapes, resource partitioning, wildland-urban interface, fire

1. Introduction

Riparian systems are dynamic conduits of ecological function whose contributions to local and regional biodiversity are greater than their proportion of the landscape (Naiman et al., 1993; Peak and Thompson, 2006). These corridors are contact zones between streamside/aquatic floras, adapted to dependable water from shallow water tables, and hillslope vegetation adapted to local climate (Minckley and Brown, 1994; Neary et al.,

2005). Ecosystem function at the interface of these two very diverse biotic communities contributes to structural diversity and food web complexity (Gregory et al., 1991). Yet, many riparian systems are at risk from anthropogenic threats, including habitat fragmentation caused by urbanization and channelization of natural riparian systems

(Green and Baker, 2003). Urbanization alters or removes native vegetation with buildings, roads and introduction of non-native landscaping (Radeloff et al., 2005) with associated pesticide (Stansley et al., 2001; Haberl et al., 2007) and herbicide use (Boren et al., 1999). As a result, degradation and loss of riparian landscapes in North America and Europe in the last 200 y is estimated at 80% (Naiman et al., 1993). 23

Increased bat diversity is often associated with riparian landscapes because of access to drinking water (Racey, 1998; Grindal et al., 1999; Holloway and Barclay,

2000). Deciduous streamside vegetation also supports greater insect biomass contributing to foraging opportunities (Iwata et al., 2003; Mosley et al., 2006; Holloway and Barclay, 2000). Besides providing foraging habitat for bats (Carter, 2006), riverine landscapes provide corridors critical for seasonal migration (Williams et al., 2006). Bats are an incredibly successful group (~20% of extant mammals) whose species diversity

(>1000 species worldwide) reflects their ability to take advantage of foraging niches through different echolocation calls, wing morphologies and flight abilities (Altringham,

1996). Consequently, bats are excellent taxa to study the dynamics of complex environments. However, one of the greatest threats to bats is loss of habitat (Pierson,

1998; O’Shea and Vaughn, 1999) and bats foraging along a riparian landscape at a wildland-urban interface experience increased habitat degradation (Evelyn et al., 2004).

This impact on natural riparian systems increases the likelihood that wildlife, including bats, will compete for remaining resources (Menzel et al., 2001) within these species-rich systems (Naiman et al., 1993).

In competitive interactions, each species has an inhibitory effect on the fitness of a competitor, particularly if resources are limited (Gotelli, 1998; Ricklefs, 2001).

Energetic costs of competition negatively influence individual fitness, population sizes and species assemblages (Rosenzweig, 1995; Arlettaz et al., 2000). Species reduce competition by dietary specialization, exploiting different microhabitats, employing different foraging styles or efficiencies, or using temporal separation of habitat (Brown, 24

1971; Schoener, 1974; Vrezec and Tome, 2004; Herder and Freyhof, 2006; Luiselli,

2006; Davies et al., 2007). Bats serve as excellent models to study resource competition

(Findley and Black, 1983; Williams et al., 2006) because they are long-lived, K- strategists (i.e., low fecundity) that live in stable, species-rich communities (Fenton,

1997; Fleming et al., 1972; McNab, 1971). Mechanisms to reduce competition by bats include: differences in diet (Black, 1974; Barlow, 1997; Feldman et al., 2000), differential foraging behavior (Kunz, 1973; Aldridge and Rautenbach, 1987; Furlonger et al., 1987; Arlettaz, 1999; Nicholls and Racey, 2006) and different roosting ecology

(Kunz, 1982; Swift and Racey, 1983; Hill and Smith, 1986).

In arid environments, multiple bat species often co-exist in areas with limited resources (Bell, 1980; Hoffmeister, 1986; Kuenzi and Morrison, 2003) and it is important to understand those mechanisms that facilitate coexistence (Saunders and Barclay, 1992).

The distribution of bats in desert landscapes is particularly limited by their access to drinking water (Cutler, 1996; Kuenzi and Morrison, 2003; Rabe and Rosenstock, 2005).

Hot, dry summers thermally stress bats and flight increases moisture loss through elevated respiration rates (Chew and White, 1960; Thomas and Suthers, 1972). Although some bats concentrate urine with increased renal medullar structure to offset water loss

(Geluso, 1978; Bassett, 1982), most species must still have access to drinking water.

Additionally, pregnancy and lactation increase females’ water needs (Kurta and Bell,

1989). Lactating females must find an additional 20% of free water to offset water required for milk production (Kurta et al., 1990). For females rearing young in arid landscapes, this has implications for species’ distributions (Humphrey, 1975). Although 25

access to water is important to desert bats (Jones, 1966, Studier et al., 1970), few multi- year studies evaluated community dynamics of bats associated with a southwestern U.S. riparian area (Koprowski et al., 2005; Williams, et al. 2006). Although riparian resources may be temporally and/or spatially variable (Bell, 1980; Gregory et al., 1991), their importance suggests that, in theory, species packing might occur (MacArthur, 1970).

Increasing demands for water associated with human expansion into arid landscapes threaten riparian ecosystem function (Webb and Leake, 2006). To evaluate how an extraordinarily diverse bat assemblage responds to these impacts, we describe and quantify the community associated with a Sonoran Desert riparian corridor at an urban-wildland interface. We determined activity patterns of bats, revealing possible spatial and/or temporal resource partitioning and analyzed impacts of seasonally ephemeral water resources on species’ presence.

2. Methods

2.1 Study Area

We conducted this study in Sabino Canyon Recreational Area (SCRA), Santa Catalina

Mountains, Arizona, located on the northeast edge of the Arizona Upland Subdivision,

Sonoran Desert Biome (Turner and Brown, 1994). The Santa Catalinas encompass ~584 km2, rising over 2000 m above the 750 m Tucson Valley floor. SCRA is a rugged

canyon with steep cliffs and, because of its scenic beauty, is a popular riparian oasis (>1.5

million visitors/year) administered by Coronado National Forest (Bezy, 2004; Lazaroff et al., 2006). The Santa Catalinas are one of ~40 mountain ranges in the American 26

Southwest located between the Rocky Mountains and Sierra Madre Occidental in Mexico

(Warshall, 1995). These montane complexes occupy a unique location with tropical, subtropical and temperate influences (McLaughlin, 1995; Warshall, 1995), containing altitudinally stacked biotic communities that reflect tremendous biodiversity. Although riparian landscapes occupy <2% of this montane area (Neary et al., 2005), broad moisture and vegetation gradients along watersheds (Warshall, 1995; Whittaker and Niering, 1975) provide microclimate requirements appropriate for many bat species (Hoffmeister, 1986;

Adams, 2003). Unfortunately, past human attitudes and management policies concerning these montane complexes negatively impacted ~150 wildlife species now listed as vulnerable by federal agencies (Warshall, 1995). Despite being one of the biologically richest in the world, this region is now one of the most endangered biodiversity hotspots

(Mittermeier et al., 2004).

Mean annual winter temperature is 10o C and mean summer temperature is 30o C.

Annual precipitation is about 30 cm/y, with surrounding mountains getting up to 75 cm/y

(NOAA Weather Station, Tucson Airport). The Sonoran Desert is characterized by a

bimodal rain pattern (winter and mid-summer storms) and is the most lush of the four

North American desertlands (Turner and Brown, 1994). Sabino Creek begins in conifer

forest at the top of the Santa Catalina Mountains (2792 m elevation) and is the largest

watershed in the range (92 km2), resulting in surface flows approximately 8 months/y.

However, during dry seasons (spring and fall), the creek often sinks into the channel’s

sandy floor and the subsurface flow is inaccessible to wildlife. There is a dramatic

difference between riparian and desert-adapted vegetation along Sabino Creek. The 27

edges of the creek support riparian deciduous vegetation dominated by cottonwood

(Populus fremontii), willow (Salix gooddingii), and ash (Fraxinus velutina) (Minkley and

Brown, 1994). Adjacent to the riparian corridor, Sabino Canyon’s steep slopes are dominated by arid-adapted plants such as saguaro (Cereus giganteus), assorted other cacti

(Opuntia spp.) and palo verde (Parkinsonia spp.) (Turner and Brown, 1994). Due to the bimodal rainfall pattern, montane island effect and riparian influence, the Santa Catalina

Mountains provide considerable foraging opportunities for bats. Prior to this study, 21 bat species were documented from the range (Cockrum, 1960; Lange, 1960; Cox, 1962;

Cross, 1965; Reidinger, 1976; Hoffmeister, 1986; Sidner and Davis, 1995), representing three families: Molossidae, Vespertilionidae and Phyllostomidae.

Despite its tremendous biodiversity, Sabino Canyon is currently at risk from adjacent urban encroachment. Tucson’s population has grown ~469% since 1960 to 1

000 000 residents (http://www.pagnet/RegionalData/Population). This explosive growth removed native vegetation, introduced non-native landscaping (Green and Baker, 2003) and increased groundwater extraction (Webb and Leake, 2006). This has altered historic flows in nearby riverine corridors and negatively impacted native riparian vegetation, reducing foraging opportunities for bats. Although SCRA is surrounded on three sides by wilderness (Lazaroff et al., 2006), Tucson city limits are within 4.8 km of the southern boundary of SCRA and homes are located at the mouth of Sabino Canyon. Because bats associated with a wildland-urban interface experience fewer foraging resources

(Theobald et al., 1997), competition over remaining resources in adjacent wildlands is increased. 28

Besides loss of habitat from development, years of forest-wide fire-suppression

(Tune and Boyle, 2005) combined with regional drought (Swetnam and Betancourt,

1998) resulted in a major forest fire in 2003 (Aspen Fire), burning >34,000 h in

Coronado National Forest (http://www.fs.fed.us/r3/coronado/aspen). This crown fire occurred two years into our four-year study and negatively impacted the upper watershed of Sabino Creek. Subsequent post-fire sedimentation altered the flow regime of the creek, reducing water availability for bats during the hottest and driest months of May and June. With multiple negative impacts on habitat, it is critical that we understand not only which bat species are present, but also how each uses this landscape in order to inform management of those resources that will maintain populations and species diversity.

2.2 Capture methods

Between March 2002 and November 2005 we deployed mist nets at 12 sites along Sabino

Creek to evaluate bat assemblage composition. The Institutional Animal Care and Use

Committee (IACUC) of The University of Arizona (Protocol #05-062) approved our animal handling procedures. To capture free-flying bats, we used 2 ply 50 denier, 38 mm mesh nylon mist nets (Avinet Inc. Dryden, NY) stretched across flowing or pooled water

(Kunz and Kurta, 1988) along 5.1 km of lower Sabino Creek. This portion of the creek provides both areas of deciduous riparian vegetation and upper sections where Sonoran desertscrub extends to the canyon floor. We placed animals in individual cloth bags and recorded time of capture as hours after sunset (HAS). 29

We determined species, sex and reproductive condition (Racey, 1988) and evaluated tooth wear (Twente, 1955). For a short time after young became volant, we distinguished adult from sub-adult by determining amount of closure of cartilaginous epiphyseal plates in the wing bones (Anthony, 1988). We weighed each bat using a Pesola spring scale (±

0.25 g) and measured the right forearm length with a caliper (± 0.1 mm).

We quantified the bat assemblage along Sabino Creek to measure local and regional diversity (Shepherd and Kelt, 1999; Moreno and Halffter, 2001) and compared these values to previous work in the Santa Catalina Mountains. We defined α-diversity

(species richness) as the total number of species captured in SCRA and compared species richness to β- diversity (i.e., species turnover between SCRA and Santa Catalinas) using

Jaccard’s index (Sabo et al., 2005). A number of widely used indices to determine β- diversity use species abundance within each area (Magurran, 1988). However, mist netting does not equally reflect all species’ presence (i.e., netting bias) and is not suitable for determining species abundance (Kunz and Kurta, 1988). Because Jaccard’s index does not compare species’ abundances between two areas (Magurran, 1988) it is appropriate for quantifying bat diversity with netting data. We compared α- and β- diversity to broader regional diversity (γ-diversity). We evaluated species diversity using

Simpson’s (dominance) index because this metric has moderate discriminate ability, low sensitivity to sample size and is widely used (Thompson and Withers, 2003). To measure the degree of evenness in species composition, we used the Berger-Parker index

(Magurran, 1988). 30

2.3 Resource Use

We defined 4 seasons, reflecting general life history stages of temperate bats (Hill and

Smith, 1986; Hoffmeister, 1986) as: ‘winter’ (December to February), ‘spring’ (March to

May), ‘summer’ (June to August) and ‘fall’ (September to November). We identified three stream conditions to analyze impacts of seasonally ephemeral water on bats as: 1)

‘flowing water’, when the creek had moving water available throughout the study area; 2)

‘pools’ when the creek stopped flowing but water was available in pools of varying sizes along the canyon interspersed with short sections of dry streambed; and 3) ‘limited pools’ where drying conditions left long stretches of dry streambed with a few small isolated pools. We used these designations, correlated with numbers of species and total captures, to determine the influence of water availability on bat behavior and indications of resource allocation.

2.4 Statistical analysis

We used statistical software package JMP 4.0 (SAS 1996) to analyze capture data, evaluating only those species that we captured ≥ 24 individuals. We conducted the

Shapiro-Wilk test and performed transformations of all data not normally distributed to enable use of parametric tests (Ramsey and Schafer 2002). We used Pearson chi-squared

(χ2) goodness-of-fit tests for analysis of nominal data. We compared multiple means

using a Bonferroni corrected Tukey-Kramer HSD analysis. We used analysis of variance

(ANOVA) and multiple linear regression to investigate relationships of number of bat

species and number of individuals captured to: 1) water availability along Sabino Creek, 31

2) netting effort (hours nets deployed), 3) maximum daytime temperature and 4) percent moon illumination (Kunz and Kurta, 1988).

3. Results

3.1 Species Diversity

We set mist nets for 63 nights (385 net hours) between March 2002 and November 2005 and captured 961 bats representing 17 species (Table 1.1). The nectar bat, Leptonycteris curasoae, was the only species that we did not capture during our study that was historically documented from Sabino Canyon (Cockrum and Petryszyn 1991). Our overall capture rate of 2.5 bats/h (S.E. = 0.21) did not vary appreciably across seasons

(ANOVA F 3, 62 = 0.21, P = 0.89), although it was slightly higher (2.6 bats/h) in summer

and fall (S.E. = 0.41). Our capture rate in spring was 2.4 bats/h (S.E. = 0.41) and capture

rate for winter was 2.1 bats/h (S.E. = 0.55). Because water availability and netting effort

changed after the 2003 Aspen Fire, we tabulated percent total capture for each species

during pre- and post-fire periods (Table 1.1). Our cumulative species curve had a steep

initial slope with 15 bat species captured within the first 80 h of netting, but we did not

catch the 17th species until hour 238. The most frequently captured species (≥24

individuals) included, in the order of most-to-least: 316 Mexican free-tailed bats

(Tadarida brasiliensis), 253 pocketed free-tailed bats (Nyctinomops femorosaccus), 196

western pipistrelles (Pipistrellus hesperus), 70 big brown bats (Eptesicus fuscus), 29 hoary bats (Lasiurus cinereus), 26 Yuma myotis (Myotis yumanensis) and 24 big free- tailed bats (N. macrotis). 32

Sex ratios varied among the 17 species and we calculated male to female ratios, with a corresponding Pearson’s goodness-of-fit statistic, for the seven frequently captured species. The remaining 10 species were captured in numbers too low to conduct statistical analysis. We caught more females than males for N. femorosaccus (n = 251,

2 2 χ 1 = 6.06, p = 0.01) and P. hesperus (n = 185, χ 1 = 10.08, p = 0.01) but more males than

2 females for T. brasiliensis (n = 281, χ 1 = 4.87, p = 0.03). The numbers of females and

2 2 males did not differ for N. macrotis (n = 24, χ 1 = 1.50, p = 0.22), L. cinereus (n = 29, χ 1

2 2 = 2.79, p = 0.09), E. fuscus (n = 251, χ 1 = 0, p = 1.00) and M. yumanensis (n = 25, χ 1 =

0.15, p = 0.69).

Prior to this study, α-diversity for bats in SCRA was 13, but we captured five

additional species (Lasionycteris noctivagans, L. blossevillii, L. xanthinus, M. auriculus,

M. velifer, ), increasing α-diversity to 18. Previous γ-diversity for the Santa Catalina

Mountains was 21 species, which we increased to 22 by adding L. xanthinus. Jaccard’s

(similarity) index for quantifying β-diversity is 0.68, where complete species similarity

between α- and β-diversity equals 1. Simpson’s index (D = 0.77) for Sabino Canyon

suggests a 77% chance that any two bats captured will be different species. Analysis of

evenness, using the reciprocal of the Berger-Parker index (1/d = 0.33), indicates a low

evenness (Magurran, 1988) in Sabino Canyon.

3.2 Resource Use

Temporal Use

Netting results indicated variable seasonal presence for 17 bat species (Fig 1.1), with N.

femorosaccus and P. hesperus year-round residents. Tadarida brasiliensis were present 33

all months except July but we only mist net once in July (2003) because the initial onset of summer monsoons makes it difficult to mist net (i.e., winds, rain, lightning). Some species, like L. blossevillii, we captured multiple times but only in spring and fall, suggesting that they use Sabino Canyon as seasonal migrants. The highest species diversity occurred in spring, with 15 species documented along Sabino Creek and least diversity during winter. However, we captured more individuals in fall and the least number in winter (Fig. 1.2).

We found distinct temporal variation in pool-use by the seven frequently captured species (F 6, 361= 23.72, p < 0.0001). Pipistrellus hesperus typically visited water sources

first (n=193, ¯x = 1.41 HAS 95% C.I. 1.21-1.70) with a second peak in activity just prior

to sunrise. Tadarida brasiliensis often arrived within the first hour after sunset but

maximum activity occurred at hour 3 (n = 285, ¯x = 3.08 HAS 95% C.I. 2.85 – 3.33).

Nyctinomops femorosaccus arrived at pools after pipistrelle activity peaked, with greatest

activity between the third and fourth hours after sunset (n = 251, ¯x = 3.32 HAS 95% C.I.

3.07 – 3.57). Eptesicus fuscus activity peaked during the second hour (n = 70, ¯x = 2.71

HAS 95% C.I. 2.23 – 3.19). Lasiurus cinereus (n = 29, ¯x = 3.43 HAS 95% C.I. 2.69 –

4.18) and M. yumanensis (n = 26, ¯x = 3.57 HAS 95% C.I. 2.78 – 4.35) first arrived at

pools about the same time but were caught over a longer period in the evening.

Nyctinomops macrotis first arrived at pools the latest of the seven species (n = 24, ¯x =

4.90 HAS 95% C.I. 4.08 – 5.72). 34

Spatial Use:

When Sabino Creek was flowing, we rarely captured more that 3-4 species/night and the

7 most frequently captured species never segregated themselves at specific pools (Fig.

1.3). However, as the creek stopped flowing in late spring and water was isolated to a

few pools, capture rates increased to 7-9 species/night and total numbers of individuals/night also increased. Water availability (present vs. limited) influenced both number of species (t 21 = 2.37, p =0.03) and total individuals (t 21 = 2.15, p = 0.04)

captured each night (Fig. 1.4). This concentration of species and individuals generally occurred during late spring/early summer (May-June), prior to monsoon season. After the onset of summer storms (early to mid-July), when water was again available the length of Sabino Canyon, numbers of bats caught at each site declined.

Much of the variability seen in species numbers (ANOVA F 5, 24 = 6.74, p = 0.0005,

R2 = 0.58) was explained by 4 covariates. Percent moon illumination (F = 0.49, p = 0.49)

and high daytime temperature (F = 0.26, p = 0.62) did not strongly influence species

diversity. However, species numbers at pools increased with both number of netting

hours (F = 10.56, p = 0.003) and water availability (F = 5.48, p = 0.01). Amount of

available water was negatively correlated to number of bat species captured through

2 summer (F 2,43 = 15.65, p < 0.001, R = 0.43 – see Fig. 1.5).

4. Discussion

4.1 Species Diversity

The high species richness (18 species) and numbers of animals (961) captured along the

stream corridor emphasizes the incredible importance of riparian resources to desert bats. 35

SCRA represents only ~1.0% of the area of the Santa Catalina Mountains and yet reflects

82% of the range’s bat diversity. Additionally, the riparian ecotone supports 64% of the statewide bat diversity within only 567 h of land area. This diversity most likely reflects both riverine and montane influences, which allow bats to move across elevations (~2000 m) and between diverse biotic communities over short horizontal distances (~30 km).

The montane archipelago region has remarkable species richness in flora, birds, terrestrial mammals and herpetofauna (Warshall, 1995; Mittermeier et al., 2004) and our study shows high bat diversity.

Our capture rate was similar to recent studies in adjacent arid regions, including the Chihuahuan Desert (1.5 bats/h – Higginbotham, 1999) and Mojave Desert (1.3 bats/h

– Williams et al., 2006). Other researchers sampling manmade wildlife water developments in the Lower Colorado Subdivision, Sonoran Desert (Schmidt, 1999,

Kuenzi and Morrison, 2003) had lower capture rates (0.46-1.45 bats/h) but this may reflect different sampling methods, small size of the pools and/or their isolation within less complex habitat (Rabe and Rosenstock, 2005). Landscape diversity along mountainous riparian corridors contributes to resource complexity, supporting multiple species (Webb and Leake, 2006). Although seasonally ephemeral, riparian zones also supply critical water resources in arid environments (Bell, 1980; Gregory et al., 1991), often during the hottest months of the year. Additionally, landforms associated with rugged canyon systems provide requisite day-roosts across a spectrum of roost requirements, including a diversity of tree-roosts and crevices in rock outcrops with 36

appropriate microclimate conditions (Lewis, 1995; Webb et al., 1995; Holloway and

Barclay, 2000).

Because our study overlapped pre- and post-fire pool conditions along Sabino

Canyon, we documented impacts of altered water resources on bat behavior. The larger free-tailed bats, N. femorosaccus, N. macrotis and Eumops perotis, declined in capture numbers after post-fire sedimentation of pools. All three species have long narrow wings, are not highly maneuverable flyers (Kumirai and Jones, 1990; Milner and Jones,

1990; Best et al., 1996) and generally use larger pools to access water. Myotis yumanensis also declined in capture numbers after post-fire sedimentation of pools. This species is strongly associated with water (Duff and Morrell, 2007), generally foraging on emergent insects or taking insects trapped on the pool surface (Hoffmeister, 1986).

Historically, Sabino Creek has no doubt seen similar dramatic changes in flow patterns; however, these occurred prior to habitat loss in the adjacent basin, when bats could still get water along historic riverine corridors.

Resource Use

Temporal Partitioning

Distinct seasonal foraging patterns, including winter activity, occurred across variable

water conditions along Sabino Canyon. Bat species also visited pools at different times

after sunset, whether by active choice or passively, because transit times between day-

roosts and foraging areas delayed their arrival at pools (Cockrum and Cross, 1964;

Adams and Thibault, 2006). However, past competition may be influencing bat behavior

and molding present ecological patterns (Rosenzweig, 1978). 37

We had high capture rates on warm winter nights (¯x = 2.1 bats/h, S.E. = 0.55) and similar patterns are seen in other desert regions (O’Farrell et al., 1967; O’Farrell and

Bradley, 1970). Two bat species (N. femorosaccus and P. hesperus) from two families

(Molossidae and Vespertilionidae) were active year-round. We captured the greatest species diversity during spring and most individuals in fall. Greater species diversity in spring suggests that riparian corridors are important sites for water and foraging opportunities when bats migrate seasonally between winter and summer roosts. Netting more individuals during fall reflects 1) an increase in insect prey following summer monsoons and 2) both adults and newly volant young foraging on the landscape.

Although some bat species may only use riparian corridors on a seasonal basis, these landscapes remain a critical resource. Many birds have similar migratory behaviors but preserving only winter and summer destinations can negatively impact populations (Leu and Thompson, 2002; Weber et al., 1999). Migration is energetically costly and it is crucial that habitat be preserved at critical stopover sites where animals rest, forage and obtain water.

Spatial Partitioning

We found strong evidence for seasonal resource partitioning of pools through variation in spatial use. During seasons with water-flow, we observed individual bats and species distributed along the canyon, with no concentration at any particular location. This

suggests that bats reduce competition for water by accessing pools along the entire canyon. However, once drying conditions reduce water availability, resource partitioning 38

‘collapses’ and species must compete for water at the few remaining pools, reflected in higher species numbers and individuals during dry periods.

Water availability and number of net hours explained much of the variability in species diversity. However, percent moon illumination and high mean daytime temperatures did not. Because of temporal resource use by bats (Cockrum and Cross,

1964; Kuenzi and Morrison, 2003; Adams and Thibault, 2006), it is not surprising that netting for a longer period after sunset increases overall capture rate (Weller and Lee,

2007) and percent moon illumination was not significant because we avoided full moon

(Hecker and Brigham, 1999; Karlsson et al., 2002). Although high mean daytime temperatures should thermally stress bats day-roosting in desert landscapes, they did not strongly influence capture rates. However, the hottest temperatures in southern Arizona occur between May and August, both before and after the onset of summer monsoon storms. Therefore, bats are not responding specifically to hot temperatures but to availability of water resources (Kuenzi and Morrison, 2003; Rabe and Rosenstock, 2005).

5 Conservation implications

We sampled bats along a desert riparian corridor at an urban-wildland interface to understand seasonal habitat use, activity patterns, and resource partitioning. Previous researchers (Kurta and Teramino, 1992; Geggie and Fenton, 1985) investigated bat community structure in rural-urban settings in mesic environments but little work has been done in the arid American West (Koprowski et al., 2005; Williams et al., 2006).

The dramatic contrast between native desert vegetation and urban landscaping provides opportunities to investigate ecological questions regarding impacts of habitat alteration 39

on bats and possible changes in foraging behavior. Consequences of anthropogenic alteration of habitat on biotic and abiotic factors may have far-reaching implications for community ecology (Green and Baker, 2003; Webb and Leake, 2006). With a better understanding of how bats use the rural-urban interface, ecologists can anticipate how continuing urban sprawl will impact community assemblages (Kurta and Teramino,

1992; Evelyn et al., 2004).

Desert riparian ecosystems contribute tremendous function and biodiversity to the

American Southwest, despite the region’s aridity (Naiman et al., 1993; Neary et al., 2005;

Williams et al., 2006). An understanding of mammalian ecology and resource use within this region provides information essential to management agencies striving to protect resources and maintain wildlife populations. As nocturnal mammals, with an ability to forage long-distances over remote and rugged terrain, bats are often overlooked during species inventories (Adams, 2003). Although riparian corridors in arid landscapes provide critical habitat for bats (Williams et al., 2006), hot summer temperatures (>42°C) thermally stress mammals and potentially limit their distribution (Studier et al., 1970). In addition, because of the Sonoran Desert bimodal rainfall pattern, early summer months are the driest and water is not always available to canyon wildlife. Furthermore, western riparian landscapes are currently at risk due to excessive human use, impacts of regional drought (Swetnam and Betancourt, 1998) and potential climate change (Weiss and

Overpeck, 2005; Seager et al., 2007), reducing water availability for wildlife.

High bat diversity associated with hydric landscapes is not restricted to arid regions. Bats preferentially use riparian habitats because of complex habitat structure 40

and available resources at the ecotone between streamside (aquatic) and regional terrestrial systems (Grindal et al., 1999; Seidman and Zabel, 2001; Russo and Jones,

2003; Ford et al., 2005; Carter, 2006; Lloyd et al., 2006; Rogers et al., 2006; Duff and

Morrell, 2007). Unfortunately, humans also depend on riverine landscapes. Globally, water resources are becoming ‘endangered’ and <2% of contiguous U.S. rivers are pristine enough to warrant protection under the Wild and Scenic Rivers Act (Benke

1990). Because human populations worldwide are dependent on riverine landscapes for their water needs, it is not surprising that conflicts occur between human enterprises and the needs of wildlife. However, we must manage streamside landscapes for long-term sustainability. The potential loss of habitat for animals, such as bats, must be considered when managing the resource for biodiversity and long-term preservation.

Acknowledgements

This study was funded by Arizona Game and Fish Department and the Arizona Game and

Fish Commission with Heritage Urban Grant #U05012 and donations from non-profits T

& E, Inc. and Friends of Sabino Canyon. Fieldwork was supported by Coronado

National Forest, USDA, Sabino Canyon Ranger District. We would like to thank Bob

Buecher, Scott Clemans and many other field assistants for invaluable help and support during this project. C. Zugmeyer, M. Merrick, A. Harlan and S. Clemans reviewed earlier drafts of this manuscript. 41

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TABLE 1.1. Species abbreviation, scientific name, common name and numbers of bats of each species caught during 2002- 2005 along Sabino Canyon with comparison of pre-fire vs. post fire capture rates. Differences in overall captures (2002-2003 vs. 2004-2005) reflect changes in netting effort and impacts from post-fire sedimentation. Species 2002-03 2004-05 Grand Code Scientific Name Common Name 2002 2003 % Total 2004 2005 % Total Totals CHME Choeronycteris mexicana Mexican long-tongued bat 2 1 0.005 2 1 0.010 6 COTO Corynorhinus townsendii Townsend's big-eared bat 1 1 0.003 1 0 0.003 3 EPFU Eptesicus fuscus Big brown bat 30 18 0.073 18 4 0.073 70 EUPE Eumops perotis Western mastiff bat 4 4 0.012 0 1 0.003 9 LANO Lasionycteris noctivagans Silver-haired bat 0 1 0.002 0 0 0.000 1 LABL Lasiurus blossevillii Western red bat 1 1 0.003 1 4 0.017 7 LACI Lasiurus cinereus Hoary bat 8 11 0.029 8 2 0.033 29 LAXA Lasiurus xanthinus Western yellow bat 2 1 0.005 1 0 0.003 4 MYAU Myotis auriculus Southwestern myotis 1 0 0.002 2 0 0.007 3 MYCA Myotis californicus California myotis 3 0 0.005 1 4 0.017 8 MYTH Myotis thysanodes Fringed myotis 1 0 0.002 0 0 0.000 1 MYVE Myotis velifer Cave myotis 1 4 0.008 0 0 0.000 5 MYYU Myotis yumanensis Yuma myotis 7 14 0.032 2 3 0.017 26 NYFE Nyctinomops femorosaccus Pocketed free-tailed bat 106 104 0.318 24 19 0.143 253 NYMA Nyctinomops macrotis Big free-tailed bat 8 15 0.035 1 0 0.003 24 PIHE Pipistrellus hesperus Western pipistrelle 64 56 0.182 59 17 0.253 196 TABR Tadarida brasiliensis Mexican free-tailed bat 105 86 0.289 73 52 0.417 316

NET EFFORT: 2002 = 18 nights/121.75 h, 2003 = 22 nights/140.0 h, 2004 = 18 nights/96.25 h, 2005 = 5 nights/26.25 h. 52

FIGURE 1.1. Species presence by month (2002-2005) along Sabino Canyon, documented with mist netting. The greatest species diversity occurs when there is little-to-no water available along the creek (April to June). Leptonycteris curasoae was documented from Sabino Canyon in 1961 (Cockrum and Petryszyn 1991) but we did not capture it during our study. Species J F M A M J J A S O N D__ C. mexicana C. townsendii E. fuscus L. blossevillii L. cinereus L. noctivagans L. xanthinus M. auriculus M. californicus M. thysanodes M. velifer M. yumanensis P. hesperus E. perotis N. femorosaccus N. macrotis T. brasiliensis _____ J F M A M J J A S O N D__ 53

FIGURE 1.2 Seasonal distribution of number of bat species (a) and numbers of individuals (b) captured along Sabino Canyon during 2002-2005.

18 15 12 9 6 3 Number of Bat Species 0 Winter Spring Summer Fall

500

400

300

200

Number of Individuals 100

0 Winter Spring Summer Fall Season 54

FIGURE 1.3. Species presence at 12 pools along Sabino Canyon, documented with mist netting 2002-2005. Pool 1 is at the wildland-urban interface (lower end of the canyon) and Pool 12 is 5.1 km upstream from Pool 1. The seven most frequently captured species are well distributed along Sabino Canyon when the creek is flowing.

POOL NUMBER 1 2 3 4 5 6 7 8 9 10 11 12

Eptesicus fuscus

Lasiurus cinereus

Myotis yumanensis

Pipistrellus hesperus

Nyctinomops femorosaccus

Nyctinomops macrotis

Tadarida brasiliensis

1 2 3 4 5 6 7 8 9 10 11 12 55

FIGURE 1.4. Number of individuals and bat species captured at pools along Sabino Creek as it changes from flowing water in the spring to isolated pools in early summer and then returns to flowing water after the onset of summer monsoon storms. High capture rates in late summer reflect migratory period of Sonoran Desert bats and their dependence on riparian corridors. 60 10 # Indiv 9 # Species Migratory Period 50 8

7 40 6

30 5

4 Number of Species

Number of Individuals 20 3

2 10 1

0 0 Spring Early Summer Mid-summer Fall Water Available Limited Water Water Available Limited Water 56

FIGURE 1.5. Linear regression of the number of bat species captured/night during summers 2002-2005. Negative slope reflects changes in species response to water availability along Sabino Creek (R2 = 0.43, P < 0.001). Early in the summer, when there is limited water, bat species are concentrated at the few remaining pools. However, once the summer monsoons begin, bats are able to disperse across the landscape to find drinking water. 10 Monsoons Begin 9

Number of Species Captured/Night 2

LIMITED WATER WATER AVAILABLE 57

APPENDIX B: FINDING THAT 4-STAR DINER OR HOW BATS MIGHT ‘ANTICIPATE’ PRODUCTIVE FORAGING AREAS

ABSTRACT

Riparian corridors, particularly in the deserts of North America, support tremendous biodiversity. Unfortunately, anthropogenic pressures have caused historic riparian landscapes to dwindle until ≤1% remain. For proper conservation and management it is critical that we understand how wildlife exploit these streamside communities.

Historically, the behavior of nocturnal mammals has been difficult to monitor. In addition, for insectivorous bats, prey often occurs in heterogeneous patches, both temporally and spatially. However, bats often anticipate productive feeding areas, given their spatial memory and knowledge of the landscape. Using multiple disciplines, we investigated how rugged canyons in the desert Southwest could be modeled to predict where food resources for insectivorous bats might be concentrated. Hydrologic models can predict where water will be slowed or impounded by topographic irregularities.

These landscapes will affect currents and create eddies in the flow. Hypothetically, these same hydrologic models might also allow evaluation of other ‘fluids’, such as cold-air drainage, under similar topographic constraints. Because small insects move through air much like swimming through molasses, they may be at the mercy of these airflow patterns. An understanding that insects are ‘pooled’ by the fluid nature of air may allow bats to anticipate foraging opportunities on patchy food resources and may enable bat ecologists to more accurately track bat diversity. 58

INTRODUCTION

The structural complexity of riparian landscapes supports tremendous biodiversity, providing critical habitat for many plant and animal species (Ohmart and Anderson

1982). In the desert Southwest, these landscapes contribute to greater species diversity than their proportion of land area (Naiman et al. 1993, Neary et al. 2005). However, riparian environments are often at risk due to anthropogenic impacts from groundwater extraction, over-grazing, increasing urbanization, introduction of exotic species, mining and timber harvest (Steiner et al. 2000), reducing riverine landscapes to ≤ 1% of the area known prior to European settlement (Ffolliott and Thorud 1974). Because of concerns for native species due to these impacts, ecologists have evaluated resource use across several taxa (Greier and Best 1980, Hunter et al. 1988, Anderson 1994, Ellis et al. 2000).

However, bats are often overlooked in landscape studies due to nocturnal behavior, yet streamside landscapes are important foraging habitat for bats and may provide their only access to drinking water (Grindal et al. 1999, Holloway and Barclay 2000). Because limited work previously focused on bat-use of riparian areas in more mesic environments

(Hayes and Adams 1996, Lloyd et al. 2006), we investigated chiropteran community structure along a riparian corridor in a desert landscape (Koprowski et al. 2005).

Monitoring habitat-use by bats is challenging because of their ability to fly long distances over rugged terrain. In addition, most temperate bats weigh ≤15 g, requiring miniaturization of radio-transmitters for effective monitoring of bats during their nightly forage for food (Aldridge and Brigham 1988). Yet, radio-telemetry studies examining broad landscape questions across multiple bat species are still prohibitively expensive 59

due to their labor-intensive nature. However, bats use biosonar during foraging and for spatial orientation (Griffin 1958), providing an additional tool with which we can monitor habitat use. The recent availability of portable, field robust ultrasonic bat detectors makes evaluation of activity patterns easier to conduct (Murray et al. 1999, Johnson et al.

2002), increasing our knowledge of how bats use resources (Hayes 1997, Vaughn et al.

1997, Humes et al. 1999, Kalcounis et al. 1999). During 2004 and 2005, we conducted passive acoustic sampling to compare foraging efforts by bats between two biotic communities, deciduous riparian vegetation and arid Sonoran desertscrub. Although our study began as an investigation of bat-use along a riparian corridor, it ultimately provided greater insight into how bats perceive and perhaps even anticipate food resources in heterogeneous environments.

STUDY AREA

Sabino Canyon is a Sonoran Desert riparian corridor administered as a recreational area by Coronado National Forest. This rugged canyon system, adjacent to Tucson,

Arizona, is in the front range of the Santa Catalina Mountains. This range lies on the northeastern edge of the Arizona Upland subdivision (Turner and Brown 1994), Sonoran

Desertscrub Biome and rises over 2000 m above the 750 m desert floor. The Santa

Catalinas are composed of hard metamorphic granites and Sabino Canyon is known for its dramatic, vertical walls (Bezy 2004).

The proximity of Sabino Canyon to a large metropolitan area (~1,000,000 residents) ensures heavy visitation (> 1.5 million visitors/year) to this scenic attraction. Although the Forest Service closed the area to private vehicles, it provides areas to hike, picnic, and 60

bicycle. A tram system also transports visitors 5.6 km along the bottom of the canyon.

Therefore, without proper management both for preservation of wildlife and the visitors’ enjoyment, this tremendous human impact could have detrimental effects on the resource.

In addition, a major forest fire (Aspen Fire 2003) in the upper watershed of Sabino

Canyon (http://www.fs.fed.us/r3/coronado/aspen) applied further pressure on a resource already impacted by recent drought (Swetnam and Betancourt 1998).

Annual precipitation for Tucson is approximately 30 cm/year, with surrounding mountain ranges getting up to 75 cm/year (NOAA Weather Station, Tucson Airport).

The Sonoran Desert is characterized by a bimodal rainfall pattern, with about 45% of the annual precipitation occurring during winter storms, often producing snow above 1500 m. The remaining precipitation generally falls during intense summer monsoon storms

(Turner and Brown 1994). Sabino Creek begins high in the Santa Catalina Mountains

(2791 m elevation) in mixed conifer forest and is the largest watershed in the range (~ 92 km2), allowing Sabino Creek to flow for longer periods after storms. During snow melt or summer monsoons the canyon can flood, but normal precipitation patterns produce moderate-to-low flows for approximately 8 months a year. Early summer and late fall are the region’s driest periods and Sabino Creek often sinks into the channel’s sandy floor and is inaccessible to wildlife. The banks of Sabino Creek support deciduous riparian vegetation dominated by cottonwood (Populus fremontii), willow (Salix gooddingii), ash

(Fraxinus pennsylvanica), walnut (Juglans major), and sycamore (Platanus wrightii)

(Minckley and Brown 1994). The steep slopes of Sabino Canyon are dominated by 61

saguaro (Cereus giganteus), prickly-pear and cholla cacti (Opuntia spp.), palo verde

(Parkinsonia spp.), and brittle bush (Encelia farinosa) (Turner and Brown 1994).

Sabino Canyon offers critical habitat for bats, providing day-roosts, foraging opportunities and access to water (Cockrum 1960, Lange 1960, Cox 1962, Cross 1965,

Hoffmeister 1986, Reidinger 1976, Sidner and Davis 1995, Buecher 2002-2005, 2008).

The year-round presence of bats suggests that this riparian resource is important across multiple species, particularly during spring and summer months when 16 of the known 18 species are documented (Fig. 2.1).

METHODS

Acoustic Sampling

During 2004 and 2005 we conducted monthly passive acoustic sampling to compare bat-use between two landscapes along 5.1 km of Sabino Creek. We deployed 4 Anabat II frequency division bat detectors with compact flash (CF) capability (Titley Electronics,

Ballina, N.S.W. Australia) cabled to Zero Crossing Analysis Interface Modules

(ZCAIM). The latest version of Anabat II detector with CF card capability produces higher quality recordings and improves our ability to monitor habitat use (Milne et al

2004). Because pairwise acoustic sampling has more power during analysis (Hayes

1997), we placed two Anabat II detectors at sites in the lower canyon, representative of riparian deciduous streamside vegetation (Minckley and Brown 1994). We located Site 1

(elev. 829 m) at the lower end of Sabino Canyon at the wildland/urban interface (Fig.

2.2). We located Site 2 (elev. 855 m) 1.3 km above Site 1, at the upper end of the deciduous riparian landscape. We anticipated that this habitat, characterized by dense 62

vegetation (Fig. 2.3), could support greater insect biomass for foraging bats (Mosley et al.

2006). We chose two additional sites in the upper canyon in more arid, open habitat with less vegetation. We located Site 4 (elev. 899 m) at the upper end of our study area (5.1 km above Site 1) and Site 3 (elev. 960 m) was 1.9 km above Site 2 and 1.9 km below Site

4. We chose Sites 3 and 4 specifically at locations where Sonoran desertscrub extends from the hillsides to the canyon floor (Fig. 2.3). The distance between sampling sites varied because of constraints necessary to locate sites within these two biotic communities. Our null hypothesis assumed equal habitat use by bats with an alternative hypothesis reflecting differential use along the lower reach of Sabino Canyon due to additional foraging opportunities on insects associated with deciduous riparian vegetation.

In 2004, we sampled monthly June through August, in October and in December. In

2005, we sampled once each month (except February and October) and sampled twice in

January, July and September. We programmed each detector to turn on 30 minutes prior to sunset and off 30 minutes after sunrise the following morning. We set the sensitivity of all Anabat detectors to ‘6’ and calibrated the detectors twice during the study using the

‘Bat Chirp’ board designed and built by T. Messina

(http://[email protected]). This confirmed that each ultrasonic bat detector was properly calibrated, remaining within the 40 kHz test frequency. The equipment was secured at each site in a locked, waterproof housing. Because Anabat II microphones are sensitive to moisture, we placed each microphone at a 45 ° angle downward to a horizontal 11 x 11cm plexiglass reflector placed on top of a 2 m high PVC pole 63

(Livengood et al 2003). At all sites we aimed the microphone upstream along Sabino

Creek.

To minimize erroneous conclusions regarding habitat use from limited sampling, we articulated certain assumptions a priori (Sherwin et al. 2000, Gannon et al.2003). We assumed that (1) an acoustic ‘capture’ event correlated with the habitat in which recorded; (2) acoustic ‘captures’ are considered discrete (independent) events reflecting foraging behavior; (3) acoustic ‘captures’ used for analysis consist of ≥ 2 individual search phase calls; (4) all bats are randomly distributed in 3-dimensional space; (5) we eliminated pseudoreplication with simultaneous sampling at two localities in each of two habitats; and (6) conclusions should only apply to this or a similar locality and not extrapolated to all riparian landscapes.

Data Analysis

We analyzed echolocation calls recorded by the Anabat II bat detectors with

AnaBat 6.3 and AnaLook 4.8 software. We defined a ‘bat call’ as each individual vocalization by a bat and a ‘call sequence’ (bat pass) as any series of 2 or more individual calls. We removed all files containing only insect noise and any call files with <2 individual calls prior to analysis (O’Farrell et al. 1999, Gannon et al. 2003). Anabat II bat detectors sample continuously for 15 seconds or until there is a 5 second silence in ultrasonic bat calls. The detector then dumps the storage buffer and assigns a unique date/time stamp for each computer file. Within each computer call file there can be > 1 species or individual (O’Farrell et al. 1999) but because we were interested only in overall activity levels and not specific resource use between species, we used total 64

number of computer call files recorded in each habitat as our compartive measure of bat activity. With this system we simultaneously sampled at 4 sites/night/month, allowing us to evaluate and compare resource use and determine where bats are most active along the canyon across four seasons.

We analyzed acoustic data using statistical software package JMP 4.0 (SAS 1996) to investigate spatial patterns and examined differences in bat foraging between and among our four sample sites. Because sampling periods varied during the project, due to changing sunset and sunrise times, we used the average number of bat calls per night to measure activity during statistical analysis. Prior to analysis we performed Loge transformation of all data not normally distributed and compared multiple means using a

Bonferroni corrected Tukey-Kramer HSD analysis. We conducted a standard Student t- test between mean number of foraging call files in riparian vs. arid desertscrub to evaluate possible differences in how bats exploit the two environments. We used analysis of variance (ANOVA) to investigate relationships between the mean number of foraging calls/h to: 1) season 2) sample site and 3) habitat.

We defined four seasons, assigning months reflecting basic life history periods for temperate bats (Hill and Smith, 1986). ‘Winter’ included the three coldest months of the year (December to February), when many temperate bats are either hibernating or have migrated southward to find insects during months of reduced food resources. ‘Spring’

(March to May) can have variable weather, but defines the general emergence of bats from hibernacula and/or the movement of bats northward. ‘Summer’ (June to August) reflects a period when females are pregnant and lactating and, by late summer, when the 65

young are volant. During ‘fall’ (September to November) there is a general movement across the landscape as bats migrate from summer roosts to mating (swarming) sites and then on to hibernating or winter sites (Hill and Smith, 1986). Because there are fewer bat species active along the creek during winter, we excluded winter from our comparative analysis. To evaluate differences in foraging, as influenced by seasonal variation, we conducted a one-way Analysis of Variance (ANOVA) of three season means (spring, summer, fall).

RESULTS

We sampled 21 nights, (261 hours of passive acoustic sampling) in 2004 and 2005, recording 49,480 bat-call files at four sites along Sabino Creek. Plots of overall foraging activity show the distribution of bat calls by sampling period at each site in 2004 (Fig.

2.4) and 2005 (Fig. 2.5). Over 15 months, the bat detector placed at Site 1 recorded

12,260 call files, Site 2 detector recorded 7,548 files, Site 3 recorded 15,003 files and Site

4 recorded 14,669 files. We expected that peak foraging activity, reflected by number of call files, would be along the lower portion of Sabino Creek (Sites 1 & 2) where dense deciduous riparian vegetation exists, supporting greater insect biomass. However, we found that the upper reaches of the canyon (Sites 3 & 4) often had greater bat activity and the Anabat detector at Site 3 recorded the greatest number of calls of all sites along

Sabino Creek.

We analyzed bat foraging across three seasons (Table 2.1) but calls did not vary by season (ANOVA F 2, 65 = 1.18, P = 0.31). Comparing site means to each other (i.e., Site

1 to Site 2 and Site 3 to Site 4) suggest that our riparian sites (Sites 1 and 2) are fairly 66

different from each other (t = -1.88, df = 32, P = 0.07); however, the desertscrub sites

(Sites 3 and 4) are extremely similar (t = 0.12, df = 32, P = 0.91). Between habitats, there is a stronger relationship between the mean number of foraging bat call files in the

Sonoran desertscrub than in the deciduous riparian environment (t = 2.28, df = 66, P =

0.03).

DISCUSSION

Our results indicate that when 1.5 years of passive acoustic data are pooled for 3 seasons (spring, summer and fall), bat foraging differs considerably between habitat types, with more activity occurring in the Sonoran desertscrub than in deciduous riparian vegetation. Our results supported neither our null hypothesis nor our expectation that bats would preferentially forage within the deciduous riparian environment, requiring additional evaluation of how bats might perceive foraging landscapes along Sabino

Creek. However, we believe that the differences observed, particularly the high levels of foraging activity at Site 3, are real. We propose a hypothetical model to investigate why bats might forage extensively at particular sites along an arid desertscrub riparian corridor. Our proposed model incorporates an interdisciplinary approach to understand foraging behavior by bats, providing insight on how bats might perceive landscapes and food resources.

Proposed Model of Bat Distribution along Western Canyons

Physics and Meteorology

Cold air is denser than warm air, causing warm air to rise and cold air to sink (Geiger

1957, Halliday and Resnick 1966). Within mountainous regions this strongly influences 67

airflow patterns and local climate. Topography also has an enormous influence on the microclimate of valleys or canyons (Geiger 1957), particularly in the semi-arid

Southwest. During the day the basin floor heats, causing hot air to rise and move up- canyon, transporting small bits of dust and detritus up-slope. As warm air rises to higher elevations, it loses heat due to adiabatic cooling (Ricklefs 2001) and this cooler air mass sits on a ‘cushion’ of rising warmer air. However, after sunset this phenomenon reverses and cold air sitting in the upper watershed on the mountain, being heavier, settles and flows down-canyon much like a river of cold water (Geiger 1957). This is evident when crossing a dry arroyo or streambed at night, where air temperature at the flow-line of a canyon is often colder than the surrounding terrain.

Hydrology

Flowing water can be modeled using computer software (U.S. Army Corps of

Engineers 1990), predicting where water will be slowed or impounded by topographic irregularities. Rugged canyon morphology affects flow velocities and creates variable currents that produce side eddies in the channel. These circular currents capture leaves and detritus in small vortices and move debris upstream along the backside of a circular eddy until fluctuations in the current change the flow regime (Rouse 1978). Kayakers and river-runners benefit from this phenomenon, using eddies in the flow to maneuver safely through river rapids. These hydrologic models might also allow us to evaluate additional ‘fluids’, such as cold-air drainage under similar topographic constraints. We hypothesize that after sunset, cold air moves along canyon floors much like flowing water and creates eddies in the air column just as vortices occur in the water channel (Geiger 68

1957). This cool air is denser than a diurnal column of warm air (Denny 1993) and will capture dust, detritus and possibly insects in eddies created by canyon irregularities.

Entomology

Insect flight is an effort to counteract forces acting in opposition (i.e., gravity and drag) to desired motion (flight) and analysis of insect flight applies a proportional relationship of inertial forces to frictional forces (Chapman 1998). Forces acting on a flying insect include ‘lift’ created by wing movements, ‘drag’ or friction of the medium because of the insect’s mass and shape plus ‘thrust’, a function of the wings lifting the insect up and forward (Brodsky 1994). Drag acts in opposition to forward movement and depends on the shape of the insect (i.e., how streamlined its presence is to air resistance), the insect’s surface texture (smooth creating less drag than rough) and its Reynolds

Number. The Reynolds Number (Re) is a dimensionless value that measures the relative importance of the speed and size of an insect to the density and viscosity of the medium through which the insect is flying (Chapman 1998). Generally, the larger the insect, the larger the Reynolds number because fluid viscosity becomes less critical as the size of the insect increases (Brodsky 1994, Lehmann 2002). However, for prey-sized insects (≤ 5 mm), flight may be strongly influenced by air density and, for nocturnal insects flying along a cooler canyon floor, could be equated to swimming through a viscous fluid.

Extensive research in Europe modeling insect infestations between crop fields provides an example of how insects are distributed by winds (Lewis 1970, Landon et al.

1997). Lewis demonstrated that smaller insects (<3 mm2) drift with breezes, like inert particles, and are often retained in eddy zones. However, larger insects (> 4-5 mm2) will 69

actually use eddies as a refuge and will accumulate in shelter zones along the flow. In

England this often occurs along hedgerows, however any break or protection from wind will provide an opportunity for insects to accumulate (Lewis 1965). Epila (1988) also illustrated that insects are poor ‘aeronauts’ and, once airborne, are at the mercy of the prevailing winds until they encounter a shelter or wind shadow that affords protection. In this way insect movements may be influenced by down-canyon airflow patterns and the fluid nature of air (Lewis 1965), trapping them in eddies along the canyon bottom. This phenomenon may allow bats to predictably forage efficiently on patchy food resources and might provide an understanding of why bats congregate in larger numbers at particular sites along a rugged canyon.

Mammalogy

Insectivorous bats often cope with heterogeneous patches of prey, both temporally and spatially (Barclay 1991, Brigham 1991). Bell (1980) illustrated that bats often anticipate productive feeding areas, given their spatial memory and knowledge of the landscape. Thus, bats foraging along a canyon would be expected to anticipate pockets of prey that occur dependably due to topographic irregularities along the canyon floor.

These prey concentrations may be more predictable than natural emergence cycles of insects.

Disciplines Combined

Near Site 3, location of greatest bat foraging, a prominent rocky ridge extends into the canyon cross-section (Fig. 2.6), creating a topographic irregularity. As the creek flows towards this ridge of granite, it is forced to make an abrupt 90° bend. This likely 70

produces a large eddy in the airflow regime, essentially trapping insects in a protected column of upstream air. This could explain the high number of bat foraging calls, suggesting that bats are able to take advantage of rich food concentrations, particularly if these resources are more dependable than ephemeral insect swarms.

MANAGEMENT RECOMMENDATIONS

Three primary factors characterize the distribution of bats across a landscape: availability of appropriate roosts, accessibility to drinking water and adequate food resources (Racey and Entwistle 2003). Resource managers often focus on preservation of bat populations through protection of day-roosts, especially maternity colonies and hibernacula (Brady 1982, Brigham 1993, Thomas et al.1990, Tuttle 1977). Although this has proven critical in maintaining healthy bat populations, proper land management policies must also consider availability of other critical resources. When protecting bat populations, habitat associated with roosts must also guarantee continued access to drinking water and food resources. Our model investigated bat foraging patterns using a multi-disciplinary approach to evaluate how bats locate food at the landscape level, given the heterogeneous nature of those resources. This approach shows promise in predicting the presence of bats at the scale at which these animals must evaluate their habitat.

Hopefully, our data provide resource managers with additional insight and information critical to determining resource use by bats along an important riparian corridor in an arid region. We hope to conduct additional research directly investigating the environmental conditions (air flow, temperature gradients and insect concentrations) at Site 3 to test our model. Given the many pressures on this unique environment, this 71

study provides land managers with details necessary for making informed decisions regarding the protection and conservation of bat habitat in arid landscapes.

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FIGURE 2.1. Species presence by month (2002-2005) along Sabino Canyon, documented with mist netting. The greatest species diversity occurs when there is little-to-no water available along the creek (April to June). Leptonycteris curasoae was documented from Sabino Canyon in 1961 (Cockrum and Petryszyn 1991) but we did not capture it during our study. Species J F M A M J J A S O N D__ Choeronycteris mexicana Corynorhinus townsendii Eptesicus fuscus Lasiurus blossevillii Lasiurus cinereus Lasionycteris noctivagans Lasiurus xanthinus Myotis auriculus Myotis californicus Myotis thysanodes Myotis velifer Myotis yumanensis Pipistrellus hesperus Eumops perotis Nyctinomops femorosaccus Nyctinomops macrotis Tadarida brasiliensis _____ J F M A M J J A S O N D 78

FIGURE 2.2. Map of Sabino Canyon showing sample sites used for the comparative acoustic sampling study. Sites 1 & 2 are located in riparian deciduous vegetation and Sites 3 & 4 are located in Sonoran desertscrub.

SITE 4

SITE 3

SITE 2

SITE 1 Adapted from Coronado National Forest Map 2005 79

FIGURE 2.3. Examples of passive acoustic sample site in deciduous riparian vegetation (top) and Sonoran desertscrub sample site (bottom). 80

FIGURE 2.4. Plot of passive acoustic data by sample period along Sabino Creek – June- December 2004

2000

1800

1600

1400

1200

1000

800

Number of Call Files 600

400

200 Site 4 0 Site 3 J1 Site 2 J2 J3 J1 J2 Site 1 A O Sample Period D 81

FIGURE 2.5. Plot of passive acoustic data by sample period along Sabino Creek – January- December 2005

2500

2000

1500

1000 Number of Call Files

500

0 Site 4 J1 J2 F1 M1 Site 3 A1 M1 Site 2 J1 J1 J2 A1 Site 1 Sample Period S1 S2 N1 D1 82

TABLE 2.1. Statistical analysis of average number of hourly foraging call files at four sample sites, across 3 seasons (i.e., winter excluded) and between two habitats during June 2004 – December 2005

Variable R2 df F-Ratio P

Site 0.11 3 2.67 0.06

Season 0.04 2 1.18 0.31

Habitat 0.07 1 2.28* 0.03 * Student t-test 83

FIGURE 2.6. A view of Acoustic Sample Site 3, where water flows from right to left. Arrow indicates the granite ridge extending from the west causing a 90o turn in the flow of Sabino Creek, creating a possible eddy just upstream from the ridge. 84

APPENDIX C: SPECIES PRESENCE AND DISTRIBUTION TABLE 3.1. Bat species documented from Santa Catalina Mountains and Sabino Canyon Recreational Area, Pima County, Arizona (data include specimens and citations in literature. M. californicus/ciliolabrum identified as M. californicus using Bogan 1974, Constantine 1998, Gannon et al. 2001. Species Santa Catalinas (19) Sabino Canyon (18) PHYLLOSTOMIDAE Choeronycteris mexicana 4, 7, 9 3, 4, 7, 11 Leptonycteris curasoae 78 Macrotus californicus 2, 4 - VESPERTILIONIDAE Antrozous pallidus 4, 7, 10 - Corynorhinus townsendii 47, 11 Eptesicus fuscus 4, 7, 10 3, 4, 7, 11 Lasiurus blossevillii 4, 7 11 Lasiurus cinereus 4, 9 7, 11 Lasionycteris noctivagans 6, 9 11 Lasiurus xanthinus -11 Myotis auriculus 1, 2, 9 11 Myotis californicus 1, 4, 7 4, 11 Myotis ciliolabrum 1, 2, 4, 9 - Myotis thysanodes 4, 7, 9, 10 7, 11 Myotis velifer 411 Myotis volans 4, 7, 9 - Myotis yumanensis 4 2, 4, 11 Pipistrellus hesperus 4, 7 3, 4, 5, 7, 11 MOLOSSIDAE Eumops perotis - 3, 7, 11 Nyctinomops femorosaccus 4 3, 6, 7, 11 Nyctinomops macrotis - 2, 3, 7, 11 Tadarida brasiliensis 4 3, 4, 7, 11

1) Miller & Allen 1928 2) Cockrum 1960 3) Cox 1962 4) Lange 1960 5) Cross 1965 6) Hoffmeister 1986 7) Sidner 1984-1995 field notes 8) Cockrum and Petryszyn 1991 9) Sidner & Davis 1995 10) Buecher 2003 field notes 11) Buecher 2002, 2003, 2004, 2005 85

TABLE 3.2. Map locations and vegetation associations for mist net sites along Sabino Creek, 2002-2005 Elevation Vegetation Pool Name/Number Latitude Longitude (meters) Years Sampled Association

1. Below Sabino Dam N 32o 18' 51.8" W 110o 48' 41.2" 822 2002, 2004 Riparian

2. At Sabino Dam N 32o 18' 53.7" W 110o 48' 40.4" 824 2002, 2004 Riparian

3. 300' above Sabino Dam N 32o 18' 56.2" W 110o 48' 37.5" 829 2002, 2003, 2004, 2005 Riparian

4. Old Gaging Station N 32o 19' 01.1" W 110o 48' 38.8" 830 2002, 2003, 2004, 2005 Riparian

5. Shuttle Stop #1 Pool N 32o 19' 24.7" W 110o 48' 27.8" 855 2002, 2003 Riparian

6. Upstream Bridge #1 N 32o 19' 30.8" W 110o 48' 20.9" 861 2003 Riparian

7. Pool below Acropolis Wall N 32o 19' 39.1" W 110o 48' 07.9" 880 2002 Riparian

8. Upstream Bridge #4 N 32o 19' 40.7" W 110o 48' 04.3" 882 2002 Desertscrub

9. Upstream Bridge #5 N 32o 19' 44.9" W 110o 47' 54.4" 895 2002, 2003, 2004 Desertscrub

10. Upstream Bridge #7 N 32o 19' 48.7" W 110o 47' 39.1" 899 2004, 2005 Desertscrub

11. Above Bridge #8 N 32o 20' 00.4" W 110o 47' 24.4" 917 2005 Desertscrub

12. Above Anderson Dam N 32o 20' 24.9" W 110o 47' 06.4" 960 2002, 2003, 2004, 2005 Desertscrub

Note: 'Riparian' is Deciduous riparian vegetation associated with Arizona Upland streamside biotic community. 'Desertscrub' is Sonoran Desert vegetation associated with the Arizona Upland subdivision of the Sonoran Desertscrub biome (Turner and Brown 1996). 86

FIGURE 3.1. Map of Sabino Canyon Recreational Area, Santa Catalina Mountains - with netting locations highlighted.

9 8 7 10 6 5

4 3 Adapted from Coronado 1, 2 National Forest Map 2005 87

FIGURE 3.2. Species presence (2002-2005) documented with mist netting along Sabino Canyon shown by pool locations. Pool numbers refer to Table 3.2. Pool Number N 1 2 3 4 5 6 7 8 9 10 11 12__

Choeronycteris mexicana 6 Corynorhinus townsendii 3 Eptesicus fuscus 70 Eumops perotis 9 Lasionycteris noctivagans 1 Lasiurus blossevillii 7 Lasiurus cinereus 29 Lasiurus xanthinus 4 Myotis auriculus 3 Myotis californicus 8 Myotis thysanodes 1 Myotis velifer 5 Myotis yumanensis 26 Nyctinomops femorosaccus 253 Nyctinomops macrotis 24 Pipistrellus hesperus 196 Tadarida brasiliensis 316

Pool Number N 1 2 3 4 5 6 7 8 9 10 11 12__ 88

FIGURE 3.3. Cumulative species curve with the number species documented along Sabino Creek plotted against total number of captured bats.

4 Number of Species Captured

0 0 100 200 300 400 500 600 700 Number of Individuals Captured 89

APPENDIX D: TABLE 4.1. BAT CAPTURE DATA ALONG SABINO CANYON 2002-2005

Capture Genus ID No. Date Time Species Sex Repro FA Wt Netting Location 2159 06/30/02 2100 Chme f used 45.5 16.50 N 32o 19' 24.7" W 110o 48' 27.8" 2296 11/01/02 1200 Chme f used 45.9 18.75 N 32o 18' 53.7" W 110o 48' 40.4" 3138 06/09/03 203 Chme f parous 46.6 17.00 N 32o 19' 01.1" W 110o 48' 38.8" 4067 05/12/04 2340 Chme f preg 43.9 22.50 N 32o 18' 54.8" W 110o 48' 40.3" 5015 04/05/05 2130 Chme f preg 45.7 22.00 N 32o 18' 56.2" W 110o 48' 37.5" 4076 05/20/04 2156 Chme m not 46.2 17.00 N 32o 20' 24.9" W 110o 47' 06.4" 2111 05/16/02 310 Coto f parous 44.5 12.50 N 32o 18' 51.8" W 110o 48' 41.2" 3173 06/25/03 2120 Coto m ad 41.2 7.00 N 32o 19' 01.1" W 110o 48' 38.8" 4147 08/16/04 2000 Coto m ad not 41.2 7.00 N 32o 19' 48.7" W 110o 47' 39.1" 2030 05/13/02 2020 Epfu f early 48.3 13.00 N 32o 18' 53.7" W 110o 48' 40.4" 2032 05/13/02 2020 Epfu f early 47.1 12.00 N 32o 18' 53.7" W 110o 48' 40.4" 2034 05/13/02 2045 Epfu f early 46.3 14.00 N 32o 18' 53.7" W 110o 48' 40.4" 2078 05/16/02 2120 Epfu f null 46.0 11.25 N 32o 18' 51.8" W 110o 48' 41.2" 2087 05/16/02 2355 Epfu f null 48.5 13.00 N 32o 18' 51.8" W 110o 48' 41.2" 2135 06/11/02 2018 Epfu f not 48.4 11.50 N 32o 19' 01.1" W 110o 48' 38.8" 2150 06/11/02 2240 Epfu f preg 46.3 16.00 N 32o 19' 01.1" W 110o 48' 38.8" 2200 08/08/02 1132 Epfu f subad 47.0 11.00 N 32o 19' 40.7" W 110o 48' 04.3" 2224 09/04/02 2130 Epfu f parous 48.9 18.00 N 32o 18' 56.2" W 110o 48' 37.5" 3082 05/20/03 2035 Epfu f null 48.5 15.00 N 32o 19' 30.8" W 110o 48' 20.9" 3083 05/20/03 2110 Epfu f parous 47.0 16.00 N 32o 19' 30.8" W 110o 48' 20.9" 3085 05/20/03 2135 Epfu f null 48.1 14.25 N 32o 19' 30.8" W 110o 48' 20.9" 3107 05/25/03 2325 Epfu f parous 50.9 23.00 N 32o 18' 56.2" W 110o 48' 37.5" 3137 06/09/03 140 Epfu f preg 49.0 19.75 N 32o 19' 01.1" W 110o 48' 38.8" 90

TABLE 4.1. Continued 3156 06/18/03 2240 Epfu f preg 47.5 19.25 N 32o 19' 24.7" W 110o 48' 27.8" 3158 06/18/03 2318 Epfu f preg 48.0 18.75 N 32o 19' 24.7" W 110o 48' 27.8" 3160 06/18/03 2340 Epfu f preg 46.0 13.25 N 32o 19' 24.7" W 110o 48' 27.8" 3166 06/25/03 2015 Epfu f preg 46.9 17.00 N 32o 19' 01.1" W 110o 48' 38.8" 3192 08/04/03 1225 Epfu f parous 45.2 17.50 N 32o 18' 56.2" W 110o 48' 37.5" 3226 09/05/03 2150 Epfu f parous 48.6 17.50 N 32o 19' 44.9" W 110o 47' 54.4" 4068 05/12/04 2340 Epfu f null 47.8 15.50 N 32o 18' 54.8" W 110o 48' 40.3" 4077 05/20/04 2306 Epfu f parous 46.5 17.00 N 32o 20' 24.9" W 110o 47' 06.4" 4078 05/20/04 2309 Epfu f parous 50.0 18.25 N 32o 20' 24.9" W 110o 47' 06.4" 4080 06/18/04 2044 Epfu f parous 49.5 16.75 N 32o 18' 56.2" W 110o 48' 37.5" 4110 08/03/04 2125 Epfu f subad 48.2 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 4111 08/03/04 2420 Epfu f subad 46.2 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 4112 08/03/04 115 Epfu f subad 45.5 10.75 N 32o 19' 44.9" W 110o 47' 54.4" 4125 08/08/04 2025 Epfu f post lac 44.5 14.75 N 32o 18' 53.2" W 110o 48' 40.8" 4146 08/08/04 2250 Epfu f subad 51.8 15.75 N 32o 18' 53.2" W 110o 48' 40.8" 4155 09/05/04 1945 Epfu f parous 45.4 17.50 N 32o 18' 56.2" W 110o 48' 37.5" 4158 09/05/04 2100 Epfu f parous 45.4 15.75 N 32o 18' 56.2" W 110o 48' 37.5" 4160 09/05/04 2100 Epfu f parous 47.5 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 4161 09/05/04 2100 Epfu f regress 46.7 18.50 N 32o 18' 56.2" W 110o 48' 37.5" 4163 09/05/04 2200 Epfu f regress 48.0 18.00 N 32o 18' 56.2" W 110o 48' 37.5" 5019 04/05/05 2300 Epfu f null 48.2 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 2025 05/13/02 1940 Epfu m not 46.8 11.00 N 32o 18' 53.7" W 110o 48' 40.4" 2031 05/13/02 2025 Epfu m not 44.5 10.50 N 32o 18' 53.7" W 110o 48' 40.4" 2071 05/16/02 2017 Epfu m not 45.5 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 91

TABLE 4.1. Continued 2073 05/16/02 2027 Epfu m not 46.4 11.25 N 32o 18' 51.8" W 110o 48' 41.2" 2077 05/16/02 2037 Epfu m not 47.2 11.30 N 32o 18' 51.8" W 110o 48' 41.2" 2138 06/11/02 2043 Epfu m not 44.5 11.90 N 32o 19' 01.1" W 110o 48' 38.8" 2140 06/11/02 2055 Epfu m not 47.3 11.90 N 32o 19' 01.1" W 110o 48' 38.8" 2141 06/11/02 2055 Epfu m not 46.4 12.00 N 32o 19' 01.1" W 110o 48' 38.8" 2142 06/11/02 2059 Epfu m not 46.4 13.90 N 32o 19' 01.1" W 110o 48' 38.8" 2143 06/11/02 2124 Epfu m not 46.5 125.00 N 32o 19' 01.1" W 110o 48' 38.8" 2145 06/11/02 2126 Epfu m not 47.9 12.70 N 32o 19' 01.1" W 110o 48' 38.8" 2151 06/11/02 2350 Epfu m not 43.9 11.50 N 32o 19' 01.1" W 110o 48' 38.8" 2155 06/11/02 400 Epfu m not 47.5 11.50 N 32o 19' 01.1" W 110o 48' 38.8" 2160 06/30/02 2100 Epfu m not 46.0 14.50 N 32o 19' 24.7" W 110o 48' 27.8" 2166 06/30/02 2155 Epfu m not 48.9 15.00 N 32o 19' 24.7" W 110o 48' 27.8" 2173 06/30/02 2320 Epfu m not 47.5 14.00 N 32o 19' 24.7" W 110o 48' 27.8" 2195 08/06/02 2310 Epfu m not 47.2 13.75 N 32o 18' 56.2" W 110o 48' 37.5" 2198 08/08/02 2040 Epfu m scrotal 47.8 15.75 N 32o 19' 40.7" W 110o 48' 04.3" 2199 08/08/02 2040 Epfu m not 45.0 9.75 N 32o 19' 40.7" W 110o 48' 04.3" 2208 09/04/02 1950 Epfu m adnr 47.0 19.50 N 32o 18' 56.2" W 110o 48' 37.5" 2215 09/04/02 2025 Epfu m adnr 43.6 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 3064 05/01/03 2045 Epfu m not 49.4 18.50 N 32o 19' 44.9" W 110o 47' 54.4" 3066 05/01/03 2110 Epfu m not 46.0 14.00 N 32o 19' 44.9" W 110o 47' 54.4" 3099 05/25/03 2200 Epfu m not 46.1 14.25 N 32o 18' 56.2" W 110o 48' 37.5" 3116 06/09/03 2025 Epfu m not 48.8 14.50 N 32o 19' 01.1" W 110o 48' 38.8" 3153 06/18/03 2210 Epfu m not 44.0 12.75 N 32o 19' 24.7" W 110o 48' 27.8" 3167 06/25/03 2020 Epfu m ad 45.7 10.25 N 32o 19' 01.1" W 110o 48' 38.8" 3195 08/04/03 230 Epfu m not 49.5 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 92

TABLE 4.1. Continued 4057 05/12/04 2115 Epfu m not 43.5 11.25 N 32o 18' 54.8" W 110o 48' 40.3" 4107 08/03/04 2032 Epfu m subad 44.2 10.50 N 32o 19' 44.9" W 110o 47' 54.4" 4109 08/03/04 2125 Epfu m repro 47.8 16.00 N 32o 19' 44.9" W 110o 47' 54.4" 4152 08/16/04 2425 Epfu m ad 48.2 12.50 N 32o 19' 48.7" W 110o 47' 39.1" 5014 04/05/05 2100 Epfu m not 43.8 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 5029 09/03/05 1925 Epfu m not 48.3 15.50 N 32o 20' 24.9" W 110o 47' 06.4" 5030 09/03/05 1925 Epfu m subad 44.5 13.75 N 32o 20' 24.9" W 110o 47' 06.4" 2281 10/05/02 2455 Eupe f parous 77.0 63.50 N 32o 19' 44.9" W 110o 47' 54.4" 2284 10/05/02 120 Eupe f subad 76.6 59.00 N 32o 19' 44.9" W 110o 47' 54.4" 2289 10/05/02 430 Eupe f postlac 76.5 - N 32o 19' 44.9" W 110o 47' 54.4" 2324 12/28/02 1814 Eupe f parous 78.4 76.00 N 32o 19' 44.9" W 110o 47' 54.4" 3236 09/05/03 143 Eupe f subad 77.0 56.00 N 32o 19' 44.9" W 110o 47' 54.4" 3011 02/07/03 1900 Eupe m not 79.1 65.50 N 32o 19' 44.9" W 110o 47' 54.4" 3239 09/05/03 320 Eupe m subad 78.2 60.50 N 32o 19' 44.9" W 110o 47' 54.4" 3240 09/05/03 335 Eupe m subad 78.8 60.50 N 32o 19' 44.9" W 110o 47' 54.4" 5057 09/03/05 1244 Eupe m subad 79.4 62.00 N 32o 20' 24.9" W 110o 47' 06.4" 5073 11/04/05 1820 Labl m bk epi 38.0 8.00 N 32o 20' 00.4" W 110o 47' 24.4" 2013 3/22/02 2145 Labl m - 42.3 10.00 N 32o 18' 56.2" W 110o 48' 37.5" 4188 11/11/04 1845 Labl m blk epi 39.1 9.00 N 32o 18' 56.2" W 110o 48' 37.5" 5011 04/05/05 2005 Labl m - 39.5 9.25 N 32o 18' 56.2" W 110o 48' 37.5" 5034 09/03/05 2010 Labl m semi 40.3 9.75 N 32o 20' 24.9" W 110o 47' 06.4" 3320 11/15/03 1920 Labl m not 39.1 9.50 N 32o 18' 56.2" W 110o 48' 37.5" 5084 11/04/05 1955 Labl m bk epi 42.0 9.75 N 32o 20' 00.4" W 110o 47' 24.4" 93

TABLE 4.1. Continued 2054 05/13/02 2440 Laci f early 56.2 28.00 N 32o 18' 53.7" W 110o 48' 40.4" 2113 05/16/02 405 Laci f null 54.0 31.50 N 32o 18' 51.8" W 110o 48' 41.2" 2228 09/04/02 2140 Laci f not 55.2 25.50 N 32o 18' 56.2" W 110o 48' 37.5" 3070 05/01/03 2255 Laci f null 54.3 31.50 N 32o 19' 44.9" W 110o 47' 54.4" 3071 05/01/03 1000 Laci f parous 54.3 30.50 N 32o 19' 44.9" W 110o 47' 54.4" 3237 09/05/03 200 Laci f - 55.5 27.00 N 32o 19' 44.9" W 110o 47' 54.4" 4029 02/17/04 2250 Laci f parous 54.8 25.25 N 32o 19' 48.7" W 110o 47' 39.1" 4042 04/14/04 2035 Laci f parous 54.4 30.25 N 32o 19' 48.7" W 110o 47' 39.1" 4048 04/14/04 2200 Laci f null 53.8 26.00 N 32o 19' 48.7" W 110o 47' 39.1" 4054 04/19/04 2320 Laci f parous 56.9 31.00 N 32o 19' 01.1" W 110o 48' 38.8" 2104 05/16/02 140 Laci m not 62.0 25.50 N 32o 18' 51.8" W 110o 48' 41.2" 2212 09/04/02 2010 Laci m scrotal 52.6 20.50 N 32o 18' 56.2" W 110o 48' 37.5" 2297 11/05/02 1820 Laci m not 54.2 23.25 N 32o 20' 24.9" W 110o 47' 06.4" 2298 11/05/02 1830 Laci m not 53.7 22.50 N 32o 20' 24.9" W 110o 47' 06.4" 2299 11/05/02 1845 Laci m not 51.8 20.75 N 32o 20' 24.9" W 110o 47' 06.4" 3008 01/05/03 1900 Laci m not 54.4 25.00 N 32o 19' 44.9" W 110o 47' 54.4" 3031 03/22/03 1923 Laci m not 49.7 20.25 N 32o 18' 56.2" W 110o 48' 37.5" 3033 03/22/03 2003 Laci m not 54.2 19.50 N 32o 18' 56.2" W 110o 48' 37.5" 3106 05/25/03 2305 Laci m not 55.7 - N 32o 18' 56.2" W 110o 48' 37.5" 3248 09/12/03 315 Laci m semi 52.9 22.25 N 32o 19' 44.9" W 110o 47' 54.4" 3286 10/24/03 1840 Laci m not 52.4 21.25 N 32o 20' 24.9" W 110o 47' 06.4" 3287 10/24/03 1850 Laci m semi 49.5 22.00 N 32o 20' 24.9" W 110o 47' 06.4" 3291 10/24/03 2420 Laci m semi 52.2 20.75 N 32o 20' 24.9" W 110o 47' 06.4" 4018 02/17/04 1905 Laci m not 52.5 25.75 N 32o 19' 48.7" W 110o 47' 39.1" 94

TABLE 4.1. Continued 4156 09/05/04 2015 Laci m semi 54.0 24.25 N 32o 18' 56.2" W 110o 48' 37.5" 4182 10/23/04 1847 Laci m not 53.0 23.50 N 32o 20' 24.9" W 110o 47' 06.4" 4190 11/11/04 1950 Laci m not 53.6 24.00 N 32o 18' 56.2" W 110o 48' 37.5" 5063 09/03/05 245 Laci m bk epi 52.3 - N 32o 20' 24.9" W 110o 47' 06.4" 5079 11/04/05 1850 Laci m not 52.2 27.50 N 32o 20' 00.4" W 110o 47' 24.4" 3052 04/10/03 2005 Lano f null 42.1 8.25 N 32o 19' 44.9" W 110o 47' 54.4" 4052 04/19/04 2005 Laxa - - - - N 32o 19' 01.1" W 110o 48' 38.8" 2033 05/13/02 2025 Laxa f early 50.0 18.00 N 32o 18' 53.7" W 110o 48' 40.4" 3311 10/25/03 2320 Laxa f null 45.9 17.50 N 32o 18' 56.2" W 110o 48' 37.5" 2144 06/11/02 2124 Laxa m not 46.7 13.70 N 32o 19' 01.1" W 110o 48' 38.8" 2153 06/11/02 203 Myau m not 37.2 N 32o 19' 01.1" W 110o 48' 38.8" 4143 08/08/04 2220 Myau m ad 36.9 7..5 N 32o 18' 53.2" W 110o 48' 40.8" 4150 08/16/04 2205 Myau m ad 36.3 6.00 N 32o 19' 48.7" W 110o 47' 39.1" 5010 03/03/05 1940 Myca f null 33.1 3.50 N 32o 19' 48.7" W 110o 47' 39.1" 2050 05/13/02 2400 Myca m not 32.5 3.50 N 32o 18' 53.7" W 110o 48' 40.4" 2129 06/04/02 2126 Myca m not - - N 32o 19' 39.1" W 110o 48' 07.9" 4031 02/17/04 1207 Myca m not 31.4 2.75 N 32o 19' 48.7" W 110o 47' 39.1" 5006 01/16/05 1910 Myca m not 31.8 3.75 N 32o 18' 56.2" W 110o 48' 37.5" 5041 09/03/05 2305 Myca m scrotal 33.2 4.00 N 32o 20' 24.9" W 110o 47' 06.4" 5058 09/03/05 105 Myca m bk epi 32.7 3.75 N 32o 20' 24.9" W 110o 47' 06.4" 2147 06/11/02 2155 Mycilca m not - - N 32o 19' 01.1" W 110o 48' 38.8" 2029 05/13/02 2010 Myth f early 44.2 7.50 N 32o 18' 53.7" W 110o 48' 40.4" 2218 09/04/02 2040 Myve f null 42.4 7.50 N 32o 18' 56.2" W 110o 48' 37.5" 3065 05/01/03 2050 Myve m not 39.9 7.00 N 32o 19' 44.9" W 110o 47' 54.4" 3086 05/20/03 2135 Myve m not 43.6 8.50 N 32o 19' 30.8" W 110o 48' 20.9" 95

TABLE 4.1. Continued 3102 05/25/03 2230 Myve m not 41.8 7.75 N 32o 18' 56.2" W 110o 48' 37.5" 3128 06/09/03 2235 Myve m not 41.8 7.50 N 32o 19' 01.1" W 110o 48' 38.8" 2175 06/30/02 2415 Myyu f parous 43.7 4.75 N 32o 19' 24.7" W 110o 48' 27.8" 2201 08/08/02 1201 Myyu f null 33.5 4.75 N 32o 19' 40.7" W 110o 48' 04.3" 2283 10/05/02 115 Myyu f used 33.7 5.75 N 32o 19' 44.9" W 110o 47' 54.4" 3049 04/10/03 1945 Myyu f null 32.8 4.50 N 32o 19' 44.9" W 110o 47' 54.4" 3063 05/01/03 2005 Myyu f parous 31.9 5.00 N 32o 19' 44.9" W 110o 47' 54.4" 3097 05/25/03 2120 Myyu f parous 33.3 5.50 N 32o 18' 56.2" W 110o 48' 37.5" 3175 07/26/03 2000 Myyu f regress 35.1 - N 32o 19' 44.9" W 110o 47' 54.4" 3203 08/06/03 2220 Myyu f subad 32.5 5.25 N 32o 18' 56.2" W 110o 48' 37.5" 4073 05/20/04 2002 Myyu f preg 33.3 5.50 N 32o 20' 24.9" W 110o 47' 06.4" 4151 08/16/04 2340 Myyu f ad null 34.6 5.00 N 32o 19' 48.7" W 110o 47' 39.1" 5061 09/03/05 200 Myyu f - 35.8 4.75 N 32o 20' 24.9" W 110o 47' 06.4" 5065 09/03/05 307 Myyu f parous 35.4 6.00 N 32o 20' 24.9" W 110o 47' 06.4" 2012 3/22/02 2120 Myyu m - 33.5 5.50 N 32o 18' 56.2" W 110o 48' 37.5" 2189 08/06/02 2008 Myyu m - 34.6 - N 32o 18' 56.2" W 110o 48' 37.5" 2203 08/18/02 2203 Myyu m scrotal 32.6 4.50 N 32o 18' 56.2" W 110o 48' 37.5" 2294 11/01/02 2040 Myyu m semi 33.8 5.25 N 32o 18' 53.7" W 110o 48' 40.4" 3058 04/10/03 2205 Myyu m not 34.8 4.75 N 32o 19' 44.9" W 110o 47' 54.4" 3098 05/25/03 2200 Myyu m not 36.3 5.00 N 32o 18' 56.2" W 110o 48' 37.5" 3176 07/26/03 2008 Myyu m not 32.8 5.00 N 32o 19' 44.9" W 110o 47' 54.4" 3180 07/26/03 2310 Myyu m subad 32.4 4.20 N 32o 19' 44.9" W 110o 47' 54.4" 3184 08/04/03 2200 Myyu m semi 39.8 7.00 N 32o 18' 56.2" W 110o 48' 37.5" 3241 09/05/03 355 Myyu m ad,not 33.1 4.00 N 32o 19' 44.9" W 110o 47' 54.4" 96

TABLE 4.1. Continued 3243 09/05/03 455 Myyu m ad,not 33.2 4.75 N 32o 19' 44.9" W 110o 47' 54.4" 3249 09/28/03 1919 Myyu m scrotal 33.2 5.75 N 32o 19' 44.9" W 110o 47' 54.4" 3299 10/25/03 1840 Myyu m not 32.4 5.00 N 32o 18' 56.2" W 110o 48' 37.5" 5037 09/03/05 2200 Myyu m not 34.8 4.75 N 32o 20' 24.9" W 110o 47' 06.4" 3193 08/04/03 1225 Nyfe - - - - N 32o 18' 56.2" W 110o 48' 37.5" 4145 08/08/04 2245 Nyfe f lac 48.3 15.00 N 32o 18' 53.2" W 110o 48' 40.8" 2016 04/12/02 2137 Nyfe f early 47.7 17.00 N 32o 18' 53.7" W 110o 48' 40.4" 2018 04/12/02 2157 Nyfe f early 47.0 15.00 N 32o 18' 53.7" W 110o 48' 40.4" 2026 05/13/02 1950 Nyfe f early 47.5 13.25 N 32o 18' 53.7" W 110o 48' 40.4" 2036 05/13/02 2055 Nyfe f early - - N 32o 18' 53.7" W 110o 48' 40.4" 2038 05/13/02 2112 Nyfe f early 46.0 13.25 N 32o 18' 53.7" W 110o 48' 40.4" 2089 05/16/02 2410 Nyfe f parous 47.5 12.50 N 32o 18' 51.8" W 110o 48' 41.2" 2090 05/16/02 2422 Nyfe f parous 48.0 15.25 N 32o 18' 51.8" W 110o 48' 41.2" 2102 05/16/02 129 Nyfe f parous 46.5 16.00 N 32o 18' 51.8" W 110o 48' 41.2" 2103 05/16/02 130 Nyfe f parous 47.0 14.50 N 32o 18' 51.8" W 110o 48' 41.2" 2106 05/16/02 210 Nyfe f null 46.0 13.50 N 32o 18' 51.8" W 110o 48' 41.2" 2107 05/16/02 239 Nyfe f parous 46.0 15.00 N 32o 18' 51.8" W 110o 48' 41.2" 2115 05/16/02 405 Nyfe f parous 47.0 16.75 N 32o 18' 51.8" W 110o 48' 41.2" 2130 06/04/02 2250 Nyfe f preg 47.4 16.50 N 32o 19' 39.1" W 110o 48' 07.9" 2152 06/11/02 2350 Nyfe f null 46.6 13.50 N 32o 19' 01.1" W 110o 48' 38.8" 2165 06/30/02 2155 Nyfe f preg 46.9 15.00 N 32o 19' 24.7" W 110o 48' 27.8" 2170 06/30/02 2245 Nyfe f null 46.3 11.50 N 32o 19' 24.7" W 110o 48' 27.8" 2171 06/30/02 2305 Nyfe f null 46.4 13.75 N 32o 19' 24.7" W 110o 48' 27.8" 97

TABLE 4.1. Continued 2172 06/30/02 2305 Nyfe f null 47.1 12.00 N 32o 19' 24.7" W 110o 48' 27.8" 2180 06/30/02 215 Nyfe f null 46.2 12.50 N 32o 19' 24.7" W 110o 48' 27.8" 2190 08/06/02 2023 Nyfe f postlac 48.0 14.75 N 32o 18' 56.2" W 110o 48' 37.5" 2191 08/06/02 2050 Nyfe f parous 45.6 15.00 N 32o 18' 56.2" W 110o 48' 37.5" 2206 09/04/02 1930 Nyfe f juv 45.5 12.75 N 32o 18' 56.2" W 110o 48' 37.5" 2207 09/04/02 1930 Nyfe f juv 46.2 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 2209 09/04/02 1950 Nyfe f juv 46.2 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 2210 09/04/02 2010 Nyfe f juv 45.5 13.75 N 32o 18' 56.2" W 110o 48' 37.5" 2211 09/04/02 2010 Nyfe f used 47.8 16.50 N 32o 18' 56.2" W 110o 48' 37.5" 2213 09/04/02 2010 Nyfe f juv 44.6 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 2219b 09/04/02 2040 Nyfe f regress 45.5 15.00 N 32o 18' 56.2" W 110o 48' 37.5" 2221 09/04/02 2110 Nyfe f juv 45.2 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 2222 09/04/02 2110 Nyfe f juv 44.4 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 2225 09/04/02 2130 Nyfe f juv 43.4 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 2226 09/04/02 2135 Nyfe f juv 46.4 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 2227 09/04/02 2135 Nyfe f used 46.4 15.30 N 32o 18' 56.2" W 110o 48' 37.5" 2229 09/04/02 2140 Nyfe f parous 46.6 14.75 N 32o 18' 56.2" W 110o 48' 37.5" 2230 09/04/02 2145 Nyfe f used 48.0 14.75 N 32o 18' 56.2" W 110o 48' 37.5" 2232 09/04/02 2245 Nyfe f juv 45.2 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 2233 09/04/02 2305 Nyfe f parous 46.9 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 2240 09/04/02 2150 Nyfe f used 47.9 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 2241 09/04/02 2150 Nyfe f juv 45.5 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 2245 09/04/02 430 Nyfe f regress 47.5 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 2250 10/05/02 1852 Nyfe f subad 45.0 11.00 N 32o 19' 44.9" W 110o 47' 54.4" 2252 10/05/02 1903 Nyfe f subad 46.4 10.50 N 32o 19' 44.9" W 110o 47' 54.4" 98

TABLE 4.1. Continued 2255 10/05/02 1925 Nyfe f subad 46.3 11.75 N 32o 19' 44.9" W 110o 47' 54.4" 2256 10/05/02 1955 Nyfe f null 46.3 14.00 N 32o 19' 44.9" W 110o 47' 54.4" 2261 10/05/02 2145 Nyfe f parous 46.2 15.00 N 32o 19' 44.9" W 110o 47' 54.4" 2263 10/05/02 2145 Nyfe f subad 45.6 11.00 N 32o 19' 44.9" W 110o 47' 54.4" 2266 10/05/02 2157 Nyfe f subad 45.8 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 2268 10/05/02 2229 Nyfe f used 45.6 15.50 N 32o 19' 44.9" W 110o 47' 54.4" 2269 10/05/02 2235 Nyfe f subad 45.5 12.25 N 32o 19' 44.9" W 110o 47' 54.4" 2275 10/05/02 2350 Nyfe f used 45.3 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 2285 10/05/02 320 Nyfe f regress 45.5 12.00 N 32o 19' 44.9" W 110o 47' 54.4" 2290 10/05/02 440 Nyfe f subad 46.3 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 2291 10/05/02 530 Nyfe f used 47.0 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 2302 12/07/02 1800 Nyfe f parous 46.3 15.50 N 32o 18' 53.7" W 110o 48' 40.4" 2305 12/07/02 1800 Nyfe f parous 46.5 15.25 N 32o 18' 53.7" W 110o 48' 40.4" 2310 12/07/02 1845 Nyfe f parous 47.5 16.00 N 32o 18' 53.7" W 110o 48' 40.4" 2325 12/28/02 1818 Nyfe f - 46.2 12.75 N 32o 19' 44.9" W 110o 47' 54.4" 2328 12/28/02 1830 Nyfe f - 46.9 14.75 N 32o 19' 44.9" W 110o 47' 54.4" 2333 12/28/02 2030 Nyfe f parous 48.0 16.50 N 32o 19' 44.9" W 110o 47' 54.4" 2334 12/28/02 2103 Nyfe f null 47.4 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 3006 01/05/03 1810 Nyfe f null 45.3 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 3025 03/10/03 1930 Nyfe f parous 47.1 12.75 N 32o 19' 44.9" W 110o 47' 54.4" 3029 03/10/03 2155 Nyfe f parous 45.3 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 3030 03/10/03 2155 Nyfe f parous 47.1 13.75 N 32o 19' 44.9" W 110o 47' 54.4" 3036 03/22/03 2150 Nyfe f parous 47.7 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3037 03/22/03 2150 Nyfe f parous 46.6 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 99

TABLE 4.1. Continued 3042 03/22/03 2215 Nyfe f null 46.9 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 3043 03/22/03 2245 Nyfe f null 46.4 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 3056 04/10/03 2045 Nyfe f parous 44.2 13.25 N 32o 19' 44.9" W 110o 47' 54.4" 3081 05/20/03 2023 Nyfe f early 45.6 14.25 N 32o 19' 30.8" W 110o 48' 20.9" 3087 05/20/03 2155 Nyfe f parous 46.7 17.00 N 32o 19' 30.8" W 110o 48' 20.9" 3088 05/20/03 2200 Nyfe f parous 47.0 18.25 N 32o 19' 30.8" W 110o 48' 20.9" 3089 05/20/03 2210 Nyfe f parous 47.2 17.75 N 32o 19' 30.8" W 110o 48' 20.9" 3091 05/20/03 2220 Nyfe f parous 46.5 16.50 N 32o 19' 30.8" W 110o 48' 20.9" 3096 05/25/03 2110 Nyfe f parous 45.5 17.50 N 32o 18' 56.2" W 110o 48' 37.5" 3104 05/25/03 2235 Nyfe f parous 46.4 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3124 06/09/03 2105 Nyfe f preg 46.8 16.25 N 32o 19' 01.1" W 110o 48' 38.8" 3152 06/18/03 2145 Nyfe f preg 47.6 16.00 N 32o 19' 24.7" W 110o 48' 27.8" 3155 06/18/03 2216 Nyfe f preg 47.0 20.25 N 32o 19' 24.7" W 110o 48' 27.8" 3179 07/26/03 2215 Nyfe f postlac 46.8 15.75 N 32o 19' 44.9" W 110o 47' 54.4" 3183 08/04/03 2120 Nyfe f ad,null 45.2 14.75 N 32o 18' 56.2" W 110o 48' 37.5" 3187b 08/04/03 2309 Nyfe f parous 46.0 14.75 N 32o 18' 56.2" W 110o 48' 37.5" 3191 08/04/03 2340 Nyfe f parous 46.2 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3196 08/04/03 250 Nyfe f postlac 46.8 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 3201 08/06/03 2120 Nyfe f postlac 46.4 15.60 N 32o 18' 56.2" W 110o 48' 37.5" 3204 08/06/03 2340 Nyfe f postlac 46.5 16.50 N 32o 18' 56.2" W 110o 48' 37.5" 3207 08/06/03 2400 Nyfe f ad,null 44.8 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3210 09/05/03 1935 Nyfe f subad 46.2 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 3213 09/05/03 1945 Nyfe f subad 45.7 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3214 09/05/03 1945 Nyfe f subad 48.2 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 100

TABLE 4.1. Continued 3217 09/05/03 2005 Nyfe f subad 48.6 13.75 N 32o 19' 44.9" W 110o 47' 54.4" 3218 09/05/03 2005 Nyfe f subad 46.3 13.25 N 32o 19' 44.9" W 110o 47' 54.4" 3219 09/05/03 2025 Nyfe f subad 46.7 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 3220 09/05/03 2035 Nyfe f subad 47.8 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 3223 09/05/03 2120 Nyfe f subad 46.6 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 3228 09/05/03 2150 Nyfe f subad 45.7 12.25 N 32o 19' 44.9" W 110o 47' 54.4" 3231 09/05/03 2240 Nyfe f subad 45.4 13.25 N 32o 19' 44.9" W 110o 47' 54.4" 3232 09/05/03 2240 Nyfe f subad 47.1 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 3233 09/05/03 2325 Nyfe f parous 47.6 15.75 N 32o 19' 44.9" W 110o 47' 54.4" 3238 09/05/03 140 Nyfe f subad 45.8 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 3242 09/05/03 413 Nyfe f postlac 46.2 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 3247 09/12/03 215 Nyfe f parous 46.6 16.00 N 32o 19' 44.9" W 110o 47' 54.4" 3250 10/03/03 1842 Nyfe f subad 46.7 9.75 N 32o 18' 56.2" W 110o 48' 37.5" 3254 10/03/03 1900 Nyfe f subad 46.8 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3255 10/03/03 1926 Nyfe f subad 45.3 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3256 10/03/03 1926 Nyfe f subad 44.2 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 3257 10/03/03 1926 Nyfe f parous 47.2 12.80 N 32o 18' 56.2" W 110o 48' 37.5" 3260 10/03/03 1940 Nyfe f subad 47.2 13.75 N 32o 18' 56.2" W 110o 48' 37.5" 3262 10/03/03 1955 Nyfe f subad 45.0 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 3263 10/03/03 1955 Nyfe f subad 47.1 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 3266 10/03/03 2020 Nyfe f subad 47.6 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3267 10/03/03 2100 Nyfe f null 45.5 13.40 N 32o 18' 56.2" W 110o 48' 37.5" 3268 10/03/03 2100 Nyfe f subad 46.3 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3269 10/03/03 2125 Nyfe f subad 45.0 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 101

TABLE 4.1. Continued 3270 10/03/03 2140 Nyfe f regress 46.6 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3275 10/03/03 2230 Nyfe f subad 45.9 14.75 N 32o 18' 56.2" W 110o 48' 37.5" 3278 10/03/03 2300 Nyfe f subad 45.6 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 3281 10/03/03 2415 Nyfe f regress 47.3 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 3300 10/25/03 1840 Nyfe f null 48.0 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3304 10/25/03 1910 Nyfe f null 46.9 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 3305 10/25/03 1920 Nyfe f parous 46.4 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3310 10/25/03 2150 Nyfe f regress 47.2 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3313 10/25/03 2415 Nyfe f parous 46.5 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 4001 01/11/04 1820 Nyfe f parous 46.5 12.75 N 32o 18' 56.2" W 110o 48' 37.5" 4003 01/11/04 1820 Nyfe f null 46.4 14.25 N 32o 18' 56.2" W 110o 48' 37.5" 4004 01/11/04 1830 Nyfe f null 46.5 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 4005 01/11/04 1845 Nyfe f null 46.2 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 4010 01/11/04 2155 Nyfe f null 46.8 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 4011 01/11/04 2230 Nyfe f parous 47.0 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 4013 01/11/04 2305 Nyfe f - 46.2 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 4015 01/11/04 2350 Nyfe f parous 47.0 15.75 N 32o 18' 56.2" W 110o 48' 37.5" 4019 02/17/04 2050 Nyfe f parous 47.0 14.25 N 32o 19' 48.7" W 110o 47' 39.1" 4032 02/17/04 1207 Nyfe f null 46.8 13.00 N 32o 19' 48.7" W 110o 47' 39.1" 4046 04/14/04 2130 Nyfe f parous 47.2 14.75 N 32o 19' 48.7" W 110o 47' 39.1" 4081 06/18/04 2410 Nyfe f preg 46.5 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 4154 09/03/04 2020 Nyfe f subad 45.9 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 4159 09/05/04 2100 Nyfe f subad 44.3 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 4165 09/05/04 2205 Nyfe f subad 46.4 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 102

TABLE 4.1. Continued 4171 09/05/04 2315 Nyfe f regress 47.6 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 4175 09/05/04 2355 Nyfe f subad - - N 32o 18' 56.2" W 110o 48' 37.5" 5003 01/16/05 1855 Nyfe f parous 47.1 12.25 N 32o 18' 56.2" W 110o 48' 37.5" 5054 09/03/05 1230 Nyfe f subad 45.4 13.00 N 32o 20' 24.9" W 110o 47' 06.4" 5056 09/03/05 1230 Nyfe f parous 46.6 14.50 N 32o 20' 24.9" W 110o 47' 06.4" 5064 09/03/05 305 Nyfe f subad 47.1 14.50 N 32o 20' 24.9" W 110o 47' 06.4" 5075 11/04/05 1830 Nyfe f subad 45.1 12.50 N 32o 20' 00.4" W 110o 47' 24.4" 2002 3/22/02 1910 Nyfe m - 49.0 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 2010 3/22/02 2050 Nyfe m - 46.6 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 2017 04/12/02 2146 Nyfe m no 47.6 15.00 N 32o 18' 53.7" W 110o 48' 40.4" 2040 05/13/02 2150 Nyfe m not 47.4 13.50 N 32o 18' 53.7" W 110o 48' 40.4" 2041 05/13/02 2200 Nyfe m not 46.4 16.50 N 32o 18' 53.7" W 110o 48' 40.4" 2042 05/13/02 2208 Nyfe m not 46.4 14.00 N 32o 18' 53.7" W 110o 48' 40.4" 2045 05/13/02 2330 Nyfe m not 47.8 14.50 N 32o 18' 53.7" W 110o 48' 40.4" 2049 05/13/02 2400 Nyfe m not 48.6 15.25 N 32o 18' 53.7" W 110o 48' 40.4" 2053 05/13/02 2430 Nyfe m not 46.9 14.50 N 32o 18' 53.7" W 110o 48' 40.4" 2091 05/16/02 2425 Nyfe m not 48.5 14.50 N 32o 18' 51.8" W 110o 48' 41.2" 2096 05/16/02 2455 Nyfe m not 48.0 15.00 N 32o 18' 51.8" W 110o 48' 41.2" 2131 06/04/02 2315 Nyfe m not 44.4 10.75 N 32o 19' 39.1" W 110o 48' 07.9" 2148 06/11/02 2215 Nyfe m not 46.6 13.00 N 32o 19' 01.1" W 110o 48' 38.8" 2168 06/30/02 2225 Nyfe m not 45.5 11.75 N 32o 19' 24.7" W 110o 48' 27.8" 2205 09/04/02 1915 Nyfe m adnr 47.6 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 2216 09/04/02 2025 Nyfe m juv 45.8 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 2217 09/04/02 2030 Nyfe m juv 45.8 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 103

TABLE 4.1. Continued 2219 09/04/02 2040 Nyfe m juv 46.1 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 2231 09/04/02 2150 Nyfe m adnr 48.0 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 2248 10/05/02 1846 Nyfe m adnr 46.7 12.25 N 32o 19' 44.9" W 110o 47' 54.4" 2249 10/05/02 1846 Nyfe m adnr 46.1 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 2251 10/05/02 1857 Nyfe m subad 45.4 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 2253 10/05/02 1915 Nyfe m subad 47.0 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 2257 10/05/02 1958 Nyfe m adnr 47.0 15.25 N 32o 19' 44.9" W 110o 47' 54.4" 2259 10/05/02 1958 Nyfe m adnr 48.3 15.25 N 32o 19' 44.9" W 110o 47' 54.4" 2260 10/05/02 2145 Nyfe m subad 45.0 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 2264 10/05/02 2145 Nyfe m subad 45.1 11.00 N 32o 19' 44.9" W 110o 47' 54.4" 2265 10/05/02 2150 Nyfe m adnr 47.6 14.75 N 32o 19' 44.9" W 110o 47' 54.4" 2267 10/05/02 2157 Nyfe m subad 46.8 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 2270 10/05/02 2305 Nyfe m adnr 46.3 14.25 N 32o 19' 44.9" W 110o 47' 54.4" 2271 10/05/02 2327 Nyfe m subad 46.1 14.00 N 32o 19' 44.9" W 110o 47' 54.4" 2272 10/05/02 2343 Nyfe m subad 45.5 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 2273 10/05/02 2350 Nyfe m subad 46.1 14.00 N 32o 19' 44.9" W 110o 47' 54.4" 2274 10/05/02 2350 Nyfe m subad 46.2 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 2276 10/05/02 2350 Nyfe m subad 47.0 12.25 N 32o 19' 44.9" W 110o 47' 54.4" 2277 10/05/02 2430 Nyfe m subad 47.9 14.00 N 32o 19' 44.9" W 110o 47' 54.4" 2278 10/05/02 2430 Nyfe m subad 46.9 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 2279 10/05/02 2430 Nyfe m subad 46.8 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 2280 10/05/02 2445 Nyfe m subad 45.6 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 2287 10/05/02 350 Nyfe m subad 44.9 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 104

TABLE 4.1. Continued 2288 10/05/02 425 Nyfe m adnr 46.2 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 2307 12/07/02 1820 Nyfe m not 46.5 12.50 N 32o 18' 53.7" W 110o 48' 40.4" 2308 12/07/02 1820 Nyfe m not 46.7 15.50 N 32o 18' 53.7" W 110o 48' 40.4" 2317 12/15/02 1800 Nyfe m not 47.8 15.50 N 32o 20' 24.9" W 110o 47' 06.4" 2327 12/28/02 1825 Nyfe m not 47.6 14.25 N 32o 19' 44.9" W 110o 47' 54.4" 3003 01/05/03 1750 Nyfe m not 47.1 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 3010 02/07/03 1900 Nyfe m not 47.4 13.25 N 32o 19' 44.9" W 110o 47' 54.4" 3017 03/10/03 1850 Nyfe m not 46.5 12.00 N 32o 19' 44.9" W 110o 47' 54.4" 3032 03/22/03 1950 Nyfe m not 46.2 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3034 03/22/03 2003 Nyfe m not 47.5 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3041 03/22/03 2215 Nyfe m not 48.0 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3062 05/01/03 1955 Nyfe m not 47.8 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 3084 05/20/03 2125 Nyfe m not 47.4 14.50 N 32o 19' 30.8" W 110o 48' 20.9" 3090 05/20/03 2210 Nyfe m not 46.8 15.00 N 32o 19' 30.8" W 110o 48' 20.9" 3101 05/25/03 2215 Nyfe m not 47.9 17.00 N 32o 18' 56.2" W 110o 48' 37.5" 3108 05/25/03 2330 Nyfe m not 46.4 15.25 N 32o 18' 56.2" W 110o 48' 37.5" 3109 05/25/03 2335 Nyfe m not 47.3 14.25 N 32o 18' 56.2" W 110o 48' 37.5" 3187 08/04/03 2309 Nyfe m ad,not 47.5 15.25 N 32o 18' 56.2" W 110o 48' 37.5" 3199 08/06/03 2100 Nyfe m ad,not 47.0 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3211 09/05/03 1935 Nyfe m subad 46.1 12.25 N 32o 19' 44.9" W 110o 47' 54.4" 3212 09/05/03 1945 Nyfe m subad 45.5 14.00 N 32o 19' 44.9" W 110o 47' 54.4" 3215 09/05/03 1945 Nyfe m subad 46.5 14.50 N 32o 19' 44.9" W 110o 47' 54.4" 3216 09/05/03 1955 Nyfe m subad 47.6 13.50 N 32o 19' 44.9" W 110o 47' 54.4" 3221 09/05/03 2055 Nyfe m subad 47.2 11.00 N 32o 19' 44.9" W 110o 47' 54.4" 3224 09/05/03 2135 Nyfe m subad 46.9 14.25 N 32o 19' 44.9" W 110o 47' 54.4" 105

TABLE 4.1. Continued 3227 09/05/03 2150 Nyfe m subad 46.8 13.75 N 32o 19' 44.9" W 110o 47' 54.4" 3229 09/05/03 2150 Nyfe m subad 46.7 12.00 N 32o 19' 44.9" W 110o 47' 54.4" 3230 09/05/03 2150 Nyfe m subad 47.5 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 3251 10/03/03 1900 Nyfe m sub not 46.2 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3252 10/03/03 1900 Nyfe m sub not 48.7 - N 32o 18' 56.2" W 110o 48' 37.5" 3258 10/03/03 1926 Nyfe m sub not 48.0 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3259 10/03/03 1940 Nyfe m sub not 48.1 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3261 10/03/03 1950 Nyfe m sub not 47.0 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 3264 10/03/03 2010 Nyfe m sub not 47.2 14.25 N 32o 18' 56.2" W 110o 48' 37.5" 3265 10/03/03 2020 Nyfe m sub not 47.4 14.30 N 32o 18' 56.2" W 110o 48' 37.5" 3271 10/03/03 2145 Nyfe m sub not 47.7 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3272 10/03/03 2200 Nyfe m ad not 48.4 15.50 N 32o 18' 56.2" W 110o 48' 37.5" 3273 10/03/03 2200 Nyfe m sub not 47.9 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3274 10/03/03 2215 Nyfe m sub not 46.5 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3280 10/03/03 2320 Nyfe m sub not 46.8 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3282 10/03/03 2430 Nyfe m sub not 45.6 12.75 N 32o 18' 56.2" W 110o 48' 37.5" 3295 10/24/03 120 Nyfe m not 47.5 14.00 N 32o 20' 24.9" W 110o 47' 06.4" 3303 10/25/03 1910 Nyfe m not 46.7 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3309 10/25/03 2040 Nyfe m not 48.0 15.00 N 32o 18' 56.2" W 110o 48' 37.5" 3312 10/25/03 2320 Nyfe m not 47.1 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3315 10/25/03 215 Nyfe m ad not 47.8 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 4009 01/11/04 2025 Nyfe m ad not 47.5 15.00 N 32o 18' 56.2" W 110o 48' 37.5" 4045 04/14/04 2035 Nyfe m not 46.8 13.00 N 32o 19' 48.7" W 110o 47' 39.1" 4047 04/14/04 2130 Nyfe m not 47.2 12.75 N 32o 19' 48.7" W 110o 47' 39.1" 106

TABLE 4.1. Continued 4164 09/05/04 2205 Nyfe m subad 46.2 12.75 N 32o 18' 56.2" W 110o 48' 37.5" 4180 10/23/04 1840 Nyfe m subad 46.1 11.00 N 32o 20' 24.9" W 110o 47' 06.4" 4184 10/23/04 1930 Nyfe m subad 49.1 11.75 N 32o 20' 24.9" W 110o 47' 06.4" 5004 01/16/05 1905 Nyfe m not 48.4 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 5036 09/03/05 2145 Nyfe m ad not 46.5 14.75 N 32o 20' 24.9" W 110o 47' 06.4" 5038 09/03/05 2240 Nyfe m not 47.0 14.00 N 32o 20' 24.9" W 110o 47' 06.4" 5039 09/03/05 2255 Nyfe m ad not 46.7 14.50 N 32o 20' 24.9" W 110o 47' 06.4" 5040 09/03/05 2255 Nyfe m not 47.8 15.00 N 32o 20' 24.9" W 110o 47' 06.4" 5048 09/03/05 2340 Nyfe m subad 48.1 15.50 N 32o 20' 24.9" W 110o 47' 06.4" 5051 09/03/05 2350 Nyfe m subad 47.4 14.60 N 32o 20' 24.9" W 110o 47' 06.4" 5052 09/03/05 2350 Nyfe m ad 48.5 15.50 N 32o 20' 24.9" W 110o 47' 06.4" 5055 09/03/05 1230 Nyfe m subad 47.8 14.00 N 32o 20' 24.9" W 110o 47' 06.4" 5059 09/03/05 150 Nyfe m subad 45.7 13.50 N 32o 20' 24.9" W 110o 47' 06.4" 5062 09/03/05 210 Nyfe m subad 46.3 14.00 N 32o 20' 24.9" W 110o 47' 06.4" 5080 11/04/05 1855 Nyfe m not 47.4 15.25 N 32o 20' 00.4" W 110o 47' 24.4" 5085 11/04/05 2020 Nyfe m subad 45.5 11.50 N 32o 20' 00.4" W 110o 47' 24.4" 5086 11/04/05 2020 Nyfe m subad 46.3 14.50 N 32o 20' 00.4" W 110o 47' 24.4" 2015 04/12/02 2127 Nyma f early 61.3 25.00 N 32o 18' 53.7" W 110o 48' 40.4" 2019 04/12/02 2202 Nyma f early 61.1 26.00 N 32o 18' 53.7" W 110o 48' 40.4" 2043 05/13/02 2208 Nyma f early 61.7 28.25 N 32o 18' 53.7" W 110o 48' 40.4" 2110 05/16/02 300 Nyma f parous 62.0 30.50 N 32o 18' 51.8" W 110o 48' 41.2" 2114 05/16/02 405 Nyma f parous 60.0 29.00 N 32o 18' 51.8" W 110o 48' 41.2" 2154 06/11/02 335 Nyma f early 61.5 25.50 N 32o 19' 01.1" W 110o 48' 38.8" 107

TABLE 4.1. Continued 2262 10/05/02 2145 Nyma f regress 59.5 26.00 N 32o 19' 44.9" W 110o 47' 54.4" 2282 10/05/02 105 Nyma f parous 61.3 24.75 N 32o 19' 44.9" W 110o 47' 54.4" 3054 04/10/03 2045 Nyma f parous 60.7 24.25 N 32o 19' 44.9" W 110o 47' 54.4" 3277 10/03/03 2244 Nyma f regress 63.4 24.90 N 32o 18' 56.2" W 110o 48' 37.5" 3289 10/24/03 2130 Nyma f subad 58.5 20.50 N 32o 20' 24.9" W 110o 47' 06.4" 3292 10/24/03 2420 Nyma f regress 60.0 24.75 N 32o 20' 24.9" W 110o 47' 06.4" 3297 10/24/03 125 Nyma f null 59.2 24.50 N 32o 20' 24.9" W 110o 47' 06.4" 3302 10/25/03 1855 Nyma f parous 60.2 21.50 N 32o 18' 56.2" W 110o 48' 37.5" 4050 04/14/04 2330 Nyma f parous 63.4 26.25 N 32o 19' 48.7" W 110o 47' 39.1" 3288 10/24/03 1921 Nyma m sub not 62.0 21.75 N 32o 20' 24.9" W 110o 47' 06.4" 3290 10/24/03 2135 Nyma m sub not 62.2 23.50 N 32o 20' 24.9" W 110o 47' 06.4" 3293 10/24/03 1902 Nyma m not 63.4 26.00 N 32o 20' 24.9" W 110o 47' 06.4" 3294 10/24/03 112 Nyma m not 59.6 25.50 N 32o 20' 24.9" W 110o 47' 06.4" 3296 10/24/03 125 Nyma m not 62.5 24.40 N 32o 20' 24.9" W 110o 47' 06.4" 3298 10/24/03 245 Nyma m subad 59.0 24.75 N 32o 20' 24.9" W 110o 47' 06.4" 3301 10/25/03 1848 Nyma m not 63.5 25.25 N 32o 18' 56.2" W 110o 48' 37.5" 3306 10/25/03 1925 Nyma m not 63.6 26.75 N 32o 18' 56.2" W 110o 48' 37.5" 3316 10/25/03 215 Nyma m not 63.5 25.75 N 32o 18' 56.2" W 110o 48' 37.5" 2158 06/30/02 2050 Pihe - juv 30.2 2.50 N 32o 19' 24.7" W 110o 48' 27.8" 2184 06/30/02 420 Pihe f lac 30.4 3.50 N 32o 19' 24.7" W 110o 48' 27.8" 2062 05/16/02 1942 Pihe f parous 30.4 3.40 N 32o 18' 51.8" W 110o 48' 41.2" 2075 05/16/02 2037 Pihe f parous 30.3 5.10 N 32o 18' 51.8" W 110o 48' 41.2" 2081 05/16/02 2215 Pihe f null 31.4 4.50 N 32o 18' 51.8" W 110o 48' 41.2" 2084 05/16/02 2318 Pihe f parous 30.0 4.00 N 32o 18' 51.8" W 110o 48' 41.2" 108

TABLE 4.1. Continued 2092 05/16/02 2425 Pihe f null 30.5 5.00 N 32o 18' 51.8" W 110o 48' 41.2" 2093 05/16/02 2430 Pihe f null 31.3 4.75 N 32o 18' 51.8" W 110o 48' 41.2" 2117 06/04/02 1950 Pihe f preg 31.4 4.25 N 32o 19' 39.1" W 110o 48' 07.9" 2119 06/04/02 1958 Pihe f preg 31.6 4.25 N 32o 19' 39.1" W 110o 48' 07.9" 2120 06/04/02 1958 Pihe f preg 31.3 4.75 N 32o 19' 39.1" W 110o 48' 07.9" 2121 06/04/02 1958 Pihe f preg 32.2 4.70 N 32o 19' 39.1" W 110o 48' 07.9" 2122 06/04/02 2005 Pihe f preg 30.2 4.70 N 32o 19' 39.1" W 110o 48' 07.9" 2124 06/04/02 2005 Pihe f preg 31.8 4.80 N 32o 19' 39.1" W 110o 48' 07.9" 2126 06/04/02 2015 Pihe f preg 31..5 5.25 N 32o 19' 39.1" W 110o 48' 07.9" 2127 06/04/02 2015 Pihe f preg 31.3 5.25 N 32o 19' 39.1" W 110o 48' 07.9" 2128 06/04/02 2095 Pihe f preg 30.5 5.20 N 32o 19' 39.1" W 110o 48' 07.9" 2132 06/11/02 2013 Pihe f null 30.5 4.00 N 32o 19' 01.1" W 110o 48' 38.8" 2133 06/11/02 2013 Pihe f preg 29.8 4.75 N 32o 19' 01.1" W 110o 48' 38.8" 2136 06/11/02 2025 Pihe f preg 29.1 5.25 N 32o 19' 01.1" W 110o 48' 38.8" 2137 06/11/02 2040 Pihe f preg 29.4 6.00 N 32o 19' 01.1" W 110o 48' 38.8" 2149 06/11/02 2215 Pihe f preg 30.2 4.75 N 32o 19' 01.1" W 110o 48' 38.8" 2156 06/30/02 1930 Pihe f used 31.3 3.50 N 32o 19' 24.7" W 110o 48' 27.8" 2157 06/30/02 2045 Pihe f used 31.4 4.25 N 32o 19' 24.7" W 110o 48' 27.8" 2161 06/30/02 2112 Pihe f used 30.4 4.50 N 32o 19' 24.7" W 110o 48' 27.8" 2167 06/30/02 2223 Pihe f postlac 30.7 4.50 N 32o 19' 24.7" W 110o 48' 27.8" 2169 06/30/02 2240 Pihe f postlac 32.6 3.25 N 32o 19' 24.7" W 110o 48' 27.8" 2176 06/30/02 2430 Pihe f lac 31.7 4.50 N 32o 19' 24.7" W 110o 48' 27.8" 2178 06/30/02 2450 Pihe f postlac 30.5 3.75 N 32o 19' 24.7" W 110o 48' 27.8" 2179 06/30/02 2450 Pihe f lac 31.4 4.50 N 32o 19' 24.7" W 110o 48' 27.8" 109

TABLE 4.1. Continued 2183 06/30/02 405 Pihe f lac 29.0 4.00 N 32o 19' 24.7" W 110o 48' 27.8" 2185 06/30/02 420 Pihe f lac - - N 32o 19' 24.7" W 110o 48' 27.8" 2186 06/30/02 430 Pihe f postlac - - N 32o 19' 24.7" W 110o 48' 27.8" 2187 06/30/02 430 Pihe f - - - N 32o 19' 24.7" W 110o 48' 27.8" 2188 06/30/02 430 Pihe f lac - - N 32o 19' 24.7" W 110o 48' 27.8" 2196 08/08/02 1910 Pihe f parous 32.2 3.25 N 32o 19' 40.7" W 110o 48' 04.3" 2202 08/18/02 1854 Pihe f subad 30.4 3.00 N 32o 18' 56.2" W 110o 48' 37.5" 2246 10/05/02 1835 Pihe f regress 30.0 4.75 N 32o 19' 44.9" W 110o 47' 54.4" 2316 12/15/02 1745 Pihe f parous 28.9 3.25 N 32o 20' 24.9" W 110o 47' 06.4" 2321 12/28/02 1800 Pihe f - 31.1 3.00 N 32o 19' 44.9" W 110o 47' 54.4" 3004 01/05/03 1805 Pihe f null 29.8 3.50 N 32o 19' 44.9" W 110o 47' 54.4" 3014 03/10/03 1842 Pihe f null 32.0 3.75 N 32o 19' 44.9" W 110o 47' 54.4" 3016 03/10/03 1848 Pihe f null 29.4 3.00 N 32o 19' 44.9" W 110o 47' 54.4" 3018 03/10/03 1853 Pihe f parous 30.3 3.25 N 32o 19' 44.9" W 110o 47' 54.4" 3019 03/10/03 1855 Pihe f parous 30.6 3.25 N 32o 19' 44.9" W 110o 47' 54.4" 3021 03/10/03 1855 Pihe f null 30.6 3.00 N 32o 19' 44.9" W 110o 47' 54.4" 3110 06/09/03 1930 Pihe f used 31.3 4.00 N 32o 19' 01.1" W 110o 48' 38.8" 3115 06/09/03 2020 Pihe f preg 31.3 5.50 N 32o 19' 01.1" W 110o 48' 38.8" 3117 06/09/03 2025 Pihe f preg 31.2 5.25 N 32o 19' 01.1" W 110o 48' 38.8" 3118 06/09/03 2040 Pihe f preg 29.9 5.50 N 32o 19' 01.1" W 110o 48' 38.8" 3122 06/09/03 2050 Pihe f preg 30.0 6.25 N 32o 19' 01.1" W 110o 48' 38.8" 3123 06/09/03 2050 Pihe f preg 30.8 5.75 N 32o 19' 01.1" W 110o 48' 38.8" 3125 06/09/03 2110 Pihe f preg 31.7 6.00 N 32o 19' 01.1" W 110o 48' 38.8" 3126 06/09/03 2120 Pihe f preg 31.0 5.75 N 32o 19' 01.1" W 110o 48' 38.8" 3140 06/18/03 2005 Pihe f lac 30.9 4.50 N 32o 19' 24.7" W 110o 48' 27.8" 110

TABLE 4.1. Continued 3143 06/18/03 2021 Pihe f lac 31.8 4.00 N 32o 19' 24.7" W 110o 48' 27.8" 3144 06/18/03 2030 Pihe f preg 30.4 5.75 N 32o 19' 24.7" W 110o 48' 27.8" 3146 06/18/03 2030 Pihe f preg 30.6 6.00 N 32o 19' 24.7" W 110o 48' 27.8" 3147 06/18/03 2030 Pihe f preg 31.4 5.25 N 32o 19' 24.7" W 110o 48' 27.8" 3148 06/18/03 2042 Pihe f preg 29.0 5.25 N 32o 19' 24.7" W 110o 48' 27.8" 3149 06/18/03 2044 Pihe f preg 30.8 5.00 N 32o 19' 24.7" W 110o 48' 27.8" 3150 06/18/03 2044 Pihe f preg 31.4 5.00 N 32o 19' 24.7" W 110o 48' 27.8" 3151 06/18/03 2109 Pihe f preg 31.3 6.00 N 32o 19' 24.7" W 110o 48' 27.8" 3154 06/18/03 2216 Pihe f lac 30.2 3.50 N 32o 19' 24.7" W 110o 48' 27.8" 3165 06/25/03 2007 Pihe f preg 29.9 4.50 N 32o 19' 01.1" W 110o 48' 38.8" 3168 06/25/03 2040 Pihe f lac 30.3 5.00 N 32o 19' 01.1" W 110o 48' 38.8" 3169 06/25/03 2040 Pihe f lac 31.1 5.00 N 32o 19' 01.1" W 110o 48' 38.8" 3170 06/25/03 2050 Pihe f lac 30.3 4.50 N 32o 19' 01.1" W 110o 48' 38.8" 3171 06/25/03 2055 Pihe f lac 31.3 4.50 N 32o 19' 01.1" W 110o 48' 38.8" 3174 06/25/03 2120 Pihe f lac 30.8 4.00 N 32o 19' 01.1" W 110o 48' 38.8" 3178 07/26/03 2125 Pihe f postlac 29.9 4.50 N 32o 19' 44.9" W 110o 47' 54.4" 3181 08/04/03 2000 Pihe f postlac 29.9 4.50 N 32o 18' 56.2" W 110o 48' 37.5" 3197 08/06/03 1955 Pihe f subad 31.4 3.75 N 32o 18' 56.2" W 110o 48' 37.5" 3198 08/06/03 2010 Pihe f subad 30.8 3.50 N 32o 18' 56.2" W 110o 48' 37.5" 3225 09/05/03 2139 Pihe f subad 30.8 3.50 N 32o 19' 44.9" W 110o 47' 54.4" 4035 04/14/04 1930 Pihe f null 29.3 3.50 N 32o 19' 48.7" W 110o 47' 39.1" 4041 04/14/04 2020 Pihe f parous 30.5 4.50 N 32o 19' 48.7" W 110o 47' 39.1" 4049 04/14/04 2230 Pihe f null 31.2 4.00 N 32o 19' 48.7" W 110o 47' 39.1" 4056 05/12/04 1945 Pihe f preg 30.3 5.00 N 32o 18' 54.8" W 110o 48' 40.3" 111

TABLE 4.1. Continued 4074 05/20/04 2036 Pihe f preg 31.4 5.00 N 32o 20' 24.9" W 110o 47' 06.4" 4082 07/08/04 1920 Pihe f lac 31.2 4.25 N 32o 19' 07.8" W 110o 47' 01.8" 4083 07/08/04 1929 Pihe f sub 28.1 2.50 N 32o 19' 07.8" W 110o 47' 01.8" 4084 07/08/04 1930 Pihe f lac 32.1 3.75 N 32o 19' 07.8" W 110o 47' 01.8" 4087 07/08/04 2030 Pihe f lac 31.4 3.75 N 32o 19' 07.8" W 110o 47' 01.8" 4089 07/08/04 2055 Pihe f regress 31.2 3.75 N 32o 19' 07.8" W 110o 47' 01.8" 4091 07/08/04 2105 Pihe f lac 29.1 4.50 N 32o 19' 07.8" W 110o 47' 01.8" 4092 07/08/04 2105 Pihe f lac 31.4 4.25 N 32o 19' 07.8" W 110o 47' 01.8" 4093 07/08/04 2105 Pihe f lac 31.0 4.50 N 32o 19' 07.8" W 110o 47' 01.8" 4094 07/08/04 2110 Pihe f post lac 30.8 4.20 N 32o 19' 07.8" W 110o 47' 01.8" 4095 07/08/04 2125 Pihe f lac 31.0 4.50 N 32o 19' 07.8" W 110o 47' 01.8" 4096 07/08/04 2150 Pihe f lac 30.4 3.75 N 32o 19' 07.8" W 110o 47' 01.8" 4097 07/08/04 2150 Pihe f lac 32.1 4.75 N 32o 19' 07.8" W 110o 47' 01.8" 4099 07/08/04 2150 Pihe f subad 30.5 2.75 N 32o 19' 07.8" W 110o 47' 01.8" 4101 08/03/04 1957 Pihe f subad 30.5 3.75 N 32o 19' 44.9" W 110o 47' 54.4" 4104 08/03/04 2013 Pihe f subad 31.3 4.00 N 32o 19' 44.9" W 110o 47' 54.4" 4105 08/03/04 2013 Pihe f subad 31.3 4.00 N 32o 19' 44.9" W 110o 47' 54.4" 4113 08/08/04 1934 Pihe f subad 30.7 3.25 N 32o 18' 53.2" W 110o 48' 40.8" 4115 08/08/04 1945 Pihe f subad 31.3 3.20 N 32o 18' 53.2" W 110o 48' 40.8" 4116 08/08/04 1955 Pihe f subad 28.4 3.25 N 32o 18' 53.2" W 110o 48' 40.8" 4117 08/08/04 1955 Pihe f parous 31.3 3.75 N 32o 18' 53.2" W 110o 48' 40.8" 4118 08/08/04 2005 Pihe f subad 30.9 3.50 N 32o 18' 53.2" W 110o 48' 40.8" 4121 08/08/04 2020 Pihe f subad 31.4 3.50 N 32o 18' 53.2" W 110o 48' 40.8" 4123 08/08/04 2020 Pihe f subad 30.7 3.50 N 32o 18' 53.2" W 110o 48' 40.8" 112

TABLE 4.1. Continued 4128 08/08/04 2043 Pihe f subad 32.5 4.25 N 32o 18' 53.2" W 110o 48' 40.8" 4153 09/03/04 1935 Pihe f subad 29.6 3.50 N 32o 19' 44.9" W 110o 47' 54.4" 4176 10/23/04 1800 Pihe f null 30.5 3.75 N 32o 20' 24.9" W 110o 47' 06.4" 4177 10/23/04 1815 Pihe f null 31.5 4.25 N 32o 20' 24.9" W 110o 47' 06.4" 4192 12/13/04 1750 Pihe f null 32.6 3.75 N 32o 19' 48.7" W 110o 47' 39.1" 5012 04/05/05 2030 Pihe f parous 30.4 5.25 N 32o 18' 56.2" W 110o 48' 37.5" 5026 09/03/05 1920 Pihe f regress 30.1 4.00 N 32o 20' 24.9" W 110o 47' 06.4" 5028 09/03/05 1925 Pihe f null 30.9 3.75 N 32o 20' 24.9" W 110o 47' 06.4" 5032 09/03/05 2000 Pihe f subad 31.5 3.25 N 32o 20' 24.9" W 110o 47' 06.4" 5033 09/03/05 2010 Pihe f subad 28.2 3.50 N 32o 20' 24.9" W 110o 47' 06.4" 5067 11/04/05 1805 Pihe f null 30.7 4.25 N 32o 20' 00.4" W 110o 47' 24.4" 5068 11/04/05 1805 Pihe f used 29.6 4.50 N 32o 20' 00.4" W 110o 47' 24.4" 5069 11/04/05 1810 Pihe f used 31.4 4.50 N 32o 20' 00.4" W 110o 47' 24.4" 5070 11/04/05 1815 Pihe f null 31.8 4.50 N 32o 20' 00.4" W 110o 47' 24.4" 5071 11/04/05 1815 Pihe f used 28.4 4.00 N 32o 20' 00.4" W 110o 47' 24.4" 5083 11/04/05 1940 Pihe f used 27.5 4.50 N 32o 20' 00.4" W 110o 47' 24.4" 2020 05/06/02 1846 Pihe m no 29.1 2.50 N 32o 19' 24.7" W 110o 48' 27.8" 2021 05/06/02 1910 Pihe m no 28.8 2.50 N 32o 19' 24.7" W 110o 48' 27.8" 2022 05/06/02 1929 Pihe m no 29.8 2.50 N 32o 19' 24.7" W 110o 48' 27.8" 2023 05/06/02 2010 Pihe m no 30.5 3.25 N 32o 19' 24.7" W 110o 48' 27.8" 2028 05/13/02 2000 Pihe m not 29.3 3.00 N 32o 18' 53.7" W 110o 48' 40.4" 2064 05/16/02 1958 Pihe m not 29.4 2.75 N 32o 18' 51.8" W 110o 48' 41.2" 2065 05/16/02 2000 Pihe m not 30.0 3.50 N 32o 18' 51.8" W 110o 48' 41.2" 2080 05/16/02 2135 Pihe m not 28.5 3.30 N 32o 18' 51.8" W 110o 48' 41.2" 113

TABLE 4.1. Continued 2116 05/16/02 505 Pihe m not 30.5 3.35 N 32o 18' 51.8" W 110o 48' 41.2" 2118 06/04/02 1951 Pihe m not 30.0 3.25 N 32o 19' 39.1" W 110o 48' 07.9" 2123 06/04/02 2005 Pihe m not 29.4 3.50 N 32o 19' 39.1" W 110o 48' 07.9" 2125 06/04/02 2005 Pihe m preg 28.1 3.90 N 32o 19' 39.1" W 110o 48' 07.9" 2134 06/11/02 2014 Pihe m not 30.4 3.00 N 32o 19' 01.1" W 110o 48' 38.8" 2146 06/11/02 2130 Pihe m not 29.4 3.60 N 32o 19' 01.1" W 110o 48' 38.8" 2162 06/30/02 2140 Pihe m - - - N 32o 19' 24.7" W 110o 48' 27.8" 2163 06/30/02 2140 Pihe m not 29.1 3.25 N 32o 19' 24.7" W 110o 48' 27.8" 2174 06/30/02 2415 Pihe m not 29.5 3.50 N 32o 19' 24.7" W 110o 48' 27.8" 2194 08/06/02 2200 Pihe m not 29.7 3.75 N 32o 18' 56.2" W 110o 48' 37.5" 2197 08/08/02 1915 Pihe m not 31.8 3.75 N 32o 19' 40.7" W 110o 48' 04.3" 2204 09/04/02 1905 Pihe m semi 29.5 3.00 N 32o 18' 56.2" W 110o 48' 37.5" 2247 10/05/02 1840 Pihe m semi 29.0 3.00 N 32o 19' 44.9" W 110o 47' 54.4" 2254 10/05/02 1925 Pihe m scrotal 29.6 3.75 N 32o 19' 44.9" W 110o 47' 54.4" 02296b 11/05/02 1805 Pihe m semi 28.5 3.25 N 32o 20' 24.9" W 110o 47' 06.4" 3001 01/05/03 1742 Pihe m not 29.4 2.75 N 32o 19' 44.9" W 110o 47' 54.4" 3012 03/10/03 1835 Pihe m not 29.9 3.25 N 32o 19' 44.9" W 110o 47' 54.4" 3013 03/10/03 1840 Pihe m not 29.7 2.75 N 32o 19' 44.9" W 110o 47' 54.4" 3015 03/10/03 1845 Pihe m not 39.1 3.00 N 32o 19' 44.9" W 110o 47' 54.4" 3020 03/10/03 1855 Pihe m not 29.1 2.50 N 32o 19' 44.9" W 110o 47' 54.4" 3095 05/25/03 1950 Pihe m not 30.5 3.40 N 32o 18' 56.2" W 110o 48' 37.5" 3111 06/09/03 1940 Pihe m not 27.4 2.75 N 32o 19' 01.1" W 110o 48' 38.8" 3112 06/09/03 1943 Pihe m not 27.3 3.00 N 32o 19' 01.1" W 110o 48' 38.8" 3113 06/09/03 2006 Pihe m not 29.2 3.00 N 32o 19' 01.1" W 110o 48' 38.8" 114

TABLE 4.1. Continued 3114 06/09/03 2006 Pihe m not 29.8 3.25 N 32o 19' 01.1" W 110o 48' 38.8" 3119 06/09/03 2043 Pihe m not 30.0 3.50 N 32o 19' 01.1" W 110o 48' 38.8" 3141 06/18/03 2015 Pihe m not 30.1 3.50 N 32o 19' 24.7" W 110o 48' 27.8" 3142 06/18/03 2015 Pihe m not 30.5 3.25 N 32o 19' 24.7" W 110o 48' 27.8" 3172 06/25/03 2105 Pihe m ad 29.1 3.00 N 32o 19' 01.1" W 110o 48' 38.8" 3177 07/26/03 2055 Pihe m subad 30.6 3.50 N 32o 19' 44.9" W 110o 47' 54.4" 3283 10/24/03 1755 Pihe m ad not 29.6 3.25 N 32o 20' 24.9" W 110o 47' 06.4" 3284 10/24/03 1810 Pihe m scrotal 29.3 3.75 N 32o 20' 24.9" W 110o 47' 06.4" 3285 10/24/03 1830 Pihe m semi 30.5 4.25 N 32o 20' 24.9" W 110o 47' 06.4" 3317 11/15/03 1750 Pihe m epi 28.3 3.25 N 32o 18' 56.2" W 110o 48' 37.5" 3324 12/21/03 1750 Pihe m not 27.8 3.25 N 32o 19' 44.9" W 110o 47' 54.4" 4016 02/17/04 1830 Pihe m bk epi 29.6 3.25 N 32o 19' 48.7" W 110o 47' 39.1" 4017 02/17/04 1845 Pihe m not 29.1 2.75 N 32o 19' 48.7" W 110o 47' 39.1" 4036 04/14/04 1930 Pihe m not 28.8 3.00 N 32o 19' 48.7" W 110o 47' 39.1" 4038 04/14/04 1940 Pihe m not 29.6 3.00 N 32o 19' 48.7" W 110o 47' 39.1" 4039 04/14/04 1951 Pihe m - - - N 32o 19' 48.7" W 110o 47' 39.1" 4072 05/20/04 1952 Pihe m not 30.7 4.00 N 32o 20' 24.9" W 110o 47' 06.4" 4085 07/08/04 1934 Pihe m subad 28.8 2.50 N 32o 19' 07.8" W 110o 47' 01.8" 4086 07/08/04 2000 Pihe m ad not 28.6 3.25 N 32o 19' 07.8" W 110o 47' 01.8" 4088 07/08/04 2048 Pihe m subad 27.7 2.75 N 32o 19' 07.8" W 110o 47' 01.8" 4090 07/08/04 2055 Pihe m subad 29.9 2.75 N 32o 19' 07.8" W 110o 47' 01.8" 4098 07/08/04 2150 Pihe m subad 30.0 2.75 N 32o 19' 07.8" W 110o 47' 01.8" 4100 08/03/04 1957 Pihe m ad not 29.9 3.75 N 32o 19' 44.9" W 110o 47' 54.4" 4102 08/03/04 1957 Pihe m subad 29.7 3.50 N 32o 19' 44.9" W 110o 47' 54.4" 115

TABLE 4.1. Continued 4103 08/03/04 1957 Pihe m ad semi 30.0 3.75 N 32o 19' 44.9" W 110o 47' 54.4" 4106 08/03/04 2018 Pihe m subad 29.7 3.25 N 32o 19' 44.9" W 110o 47' 54.4" 4114 08/08/04 1934 Pihe m ad semi 30.8 3.25 N 32o 18' 53.2" W 110o 48' 40.8" 4119 08/08/04 2005 Pihe m ad semi 30.2 3.25 N 32o 18' 53.2" W 110o 48' 40.8" 4122 08/08/04 2020 Pihe m subad 30.6 3.25 N 32o 18' 53.2" W 110o 48' 40.8" 4131 08/08/04 2025 Pihe m ad semi 30.2 3.50 N 32o 18' 53.2" W 110o 48' 40.8" 4133 08/08/04 2025 Pihe m subad 29.7 3.25 N 32o 18' 53.2" W 110o 48' 40.8" 4148 08/16/04 2020 Pihe m subad 29.3 3.00 N 32o 19' 48.7" W 110o 47' 39.1" 4149 08/16/04 2044 Pihe m subad 29.6 3.20 N 32o 19' 48.7" W 110o 47' 39.1" 4178 10/23/04 1825 Pihe m bk epi 30.0 4.00 N 32o 20' 24.9" W 110o 47' 06.4" 4181 10/23/04 1840 Pihe m bk epi 29.2 3.75 N 32o 20' 24.9" W 110o 47' 06.4" 4185 10/23/04 2015 Pihe m bk epi 28.9 3.50 N 32o 20' 24.9" W 110o 47' 06.4" 4191 12/13/04 1740 Pihe m not 28.9 3.50 N 32o 19' 48.7" W 110o 47' 39.1" 5025 09/03/05 1912 Pihe m bk epi 28.7 3.50 N 32o 20' 24.9" W 110o 47' 06.4" 5027 09/03/05 1920 Pihe m semi 27.7 3.50 N 32o 20' 24.9" W 110o 47' 06.4" 5031 09/03/05 1950 Pihe m bk epi 30.0 4.00 N 32o 20' 24.9" W 110o 47' 06.4" 5072 11/04/05 1815 Pihe m bk epi 28.2 3.75 N 32o 20' 00.4" W 110o 47' 24.4" 5081 11/04/05 1900 Pihe m not 30.1 4.00 N 32o 20' 00.4" W 110o 47' 24.4" 5082 11/04/05 1908 Pihe m bk epi 26.3 3.00 N 32o 20' 00.4" W 110o 47' 24.4" 2094 05/16/02 2430 Tabr - - - - N 32o 18' 51.8" W 110o 48' 41.2" 2099 05/16/02 100 Tabr - - - - N 32o 18' 51.8" W 110o 48' 41.2" 2329 12/28/02 1830 Tabr - - - - N 32o 19' 44.9" W 110o 47' 54.4" 4130 08/08/04 2043 Tabr - - - - N 32o 18' 53.2" W 110o 48' 40.8" 4172 09/05/04 2315 Tabr - - - - N 32o 18' 56.2" W 110o 48' 37.5" 116

TABLE 4.1. Continued 4174 09/05/04 2355 Tabr - - - - N 32o 18' 56.2" W 110o 48' 37.5" 5053 09/03/05 2310 Tabr - - 42.7 10.25 N 32o 20' 24.9" W 110o 47' 06.4" 2001 3/22/02 1905 Tabr f early 41.6 10.00 N 32o 18' 56.2" W 110o 48' 37.5" 2007 3/22/02 1930 Tabr f early 43.0 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 2008 3/22/02 1950 Tabr f early 43.3 10.75 N 32o 18' 56.2" W 110o 48' 37.5" 2011 3/22/02 2120 Tabr f early 43.0 10.25 N 32o 18' 56.2" W 110o 48' 37.5" 2024 05/06/02 2010 Tabr f early 43.8 10.00 N 32o 19' 24.7" W 110o 48' 27.8" 2047 05/13/02 2355 Tabr f early 42.5 10.00 N 32o 18' 53.7" W 110o 48' 40.4" 2048 05/13/02 2355 Tabr f early 42.2 10.50 N 32o 18' 53.7" W 110o 48' 40.4" 2051 05/13/02 2430 Tabr f early 42.5 12.50 N 32o 18' 53.7" W 110o 48' 40.4" 2063 05/16/02 1942 Tabr f early 44.5 8.50 N 32o 18' 51.8" W 110o 48' 41.2" 2069 05/16/02 2002 Tabr f parous 41.9 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 2070 05/16/02 2010 Tabr f parous 42.4 10.30 N 32o 18' 51.8" W 110o 48' 41.2" 2072 05/16/02 2021 Tabr f parous 43.1 10.80 N 32o 18' 51.8" W 110o 48' 41.2" 2079 05/16/02 2121 Tabr f parous 42.7 10.00 N 32o 18' 51.8" W 110o 48' 41.2" 2082 05/16/02 2225 Tabr f parous 43.9 11.50 N 32o 18' 51.8" W 110o 48' 41.2" 2085 05/16/02 2318 Tabr f parous 42.5 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 2086 05/16/02 2323 Tabr f parous 44.0 12.50 N 32o 18' 51.8" W 110o 48' 41.2" 2088 05/16/02 2400 Tabr f null 43.5 10.75 N 32o 18' 51.8" W 110o 48' 41.2" 2095 05/16/02 2445 Tabr f parous 43.0 9.50 N 32o 18' 51.8" W 110o 48' 41.2" 2097 05/16/02 2458 Tabr f parous 43.0 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 2100 05/16/02 100 Tabr f - 43.0 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 2105 05/16/02 155 Tabr f parous 41.0 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 2164 06/30/02 2145 Tabr f preg 42.8 12.00 N 32o 19' 24.7" W 110o 48' 27.8" 117

TABLE 4.1. Continued 2214 09/04/02 2020 Tabr f parous 43.1 9.50 N 32o 18' 56.2" W 110o 48' 37.5" 2220 09/04/02 2110 Tabr f used 42.4 11.30 N 32o 18' 56.2" W 110o 48' 37.5" 2223 09/04/02 2115 Tabr f used 42.0 9.80 N 32o 18' 56.2" W 110o 48' 37.5" 2234 09/04/02 2305 Tabr f null 42.4 10.30 N 32o 18' 56.2" W 110o 48' 37.5" 2235 09/04/02 2330 Tabr f parous 42.5 9.00 N 32o 18' 56.2" W 110o 48' 37.5" 2242 09/04/02 345 Tabr f juv 41.6 8.50 N 32o 18' 56.2" W 110o 48' 37.5" 2243 09/04/02 345 Tabr f regress 42.5 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 2244 09/04/02 2150 Tabr f used 44.3 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 2292 11/01/02 1835 Tabr f parous 43.9 14.50 N 32o 18' 53.7" W 110o 48' 40.4" 2293 11/01/02 1855 Tabr f not 43.3 14.00 N 32o 18' 53.7" W 110o 48' 40.4" 2295 11/01/02 2040 Tabr f null 42.1 12.75 N 32o 18' 53.7" W 110o 48' 40.4" 2309 12/07/02 1845 Tabr f - 41.0 10.25 N 32o 18' 53.7" W 110o 48' 40.4" 2311 12/07/02 1915 Tabr f - 43.5 10.25 N 32o 18' 53.7" W 110o 48' 40.4" 2312 12/07/02 1915 Tabr f parous 43.2 11.50 N 32o 18' 53.7" W 110o 48' 40.4" 2313 12/07/02 1915 Tabr f parous 43.3 13.50 N 32o 18' 53.7" W 110o 48' 40.4" 2314 12/07/02 2000 Tabr f parous 43.8 13.00 N 32o 18' 53.7" W 110o 48' 40.4" 2318 12/15/02 1800 Tabr f parous 41.9 9.00 N 32o 20' 24.9" W 110o 47' 06.4" 2320 12/28/02 1800 Tabr f - 44.6 12.00 N 32o 19' 44.9" W 110o 47' 54.4" 2326 12/28/02 1825 Tabr f parous 43.9 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 2330 12/28/02 1840 Tabr f parous 43.9 12.50 N 32o 19' 44.9" W 110o 47' 54.4" 2331 12/28/02 1920 Tabr f parous 43.7 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 2332 12/28/02 2015 Tabr f regress 42.6 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 3002 01/05/03 1750 Tabr f parous 42.5 9.75 N 32o 19' 44.9" W 110o 47' 54.4" 118

TABLE 4.1. Continued 3005 01/05/03 1810 Tabr f parous 42.7 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 3009 01/05/03 1920 Tabr f parous 43.7 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 3022 03/10/03 1908 Tabr f parous 42.2 9.50 N 32o 19' 44.9" W 110o 47' 54.4" 3024 03/10/03 1920 Tabr f parous 44.0 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 3026 03/10/03 1951 Tabr f parous 42.6 10.00 N 32o 19' 44.9" W 110o 47' 54.4" 3027 03/10/03 2030 Tabr f parous 42.8 9.75 N 32o 19' 44.9" W 110o 47' 54.4" 3035 03/22/03 2040 Tabr f parous 43.8 12.25 N 32o 18' 56.2" W 110o 48' 37.5" 3038 03/22/03 2150 Tabr f parous 43.3 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 3039 03/22/03 2150 Tabr f null 42.3 8.75 N 32o 18' 56.2" W 110o 48' 37.5" 3061 04/10/03 2300 Tabr f parous 42.5 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 3092 05/20/03 2245 Tabr f parous 42.7 12.25 N 32o 19' 30.8" W 110o 48' 20.9" 3093 05/20/03 2315 Tabr f parous 45.1 15.00 N 32o 19' 30.8" W 110o 48' 20.9" 3094 05/20/03 2330 Tabr f parous 44.5 14.00 N 32o 19' 30.8" W 110o 48' 20.9" 3103 05/25/03 2235 Tabr f parous 42.2 10.75 N 32o 18' 56.2" W 110o 48' 37.5" 3127 06/09/03 2220 Tabr f preg 42.3 12.50 N 32o 19' 01.1" W 110o 48' 38.8" 3130 06/09/03 2325 Tabr f preg 43.8 11.00 N 32o 19' 01.1" W 110o 48' 38.8" 3131 06/09/03 2330 Tabr f preg 43.4 14.50 N 32o 19' 01.1" W 110o 48' 38.8" 3132 06/09/03 2345 Tabr f preg 43.0 12.00 N 32o 19' 01.1" W 110o 48' 38.8" 3135 06/09/03 2410 Tabr f preg 42.7 11.50 N 32o 19' 01.1" W 110o 48' 38.8" 3136 06/09/03 2440 Tabr f preg 40.6 12.50 N 32o 19' 01.1" W 110o 48' 38.8" 3161 06/18/03 2340 Tabr f preg 42.7 14.00 N 32o 19' 24.7" W 110o 48' 27.8" 3163 06/18/03 105 Tabr f preg 42.3 15.75 N 32o 19' 24.7" W 110o 48' 27.8" 3164 06/18/03 145 Tabr f preg 42.3 14.25 N 32o 19' 24.7" W 110o 48' 27.8" 3185 08/04/03 2200 Tabr f subad 45.1 10.75 N 32o 18' 56.2" W 110o 48' 37.5" 119

TABLE 4.1. Continued 3235 09/05/03 2325 Tabr f null 41.3 9.00 N 32o 19' 44.9" W 110o 47' 54.4" 3244 09/12/03 2235 Tabr f parous 42.6 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 3253 10/03/03 1900 Tabr f regress 43.3 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 3307 10/25/03 2040 Tabr f regress 43.3 15.25 N 32o 18' 56.2" W 110o 48' 37.5" 3308 10/25/03 2040 Tabr f parous 42.0 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 3318 11/15/03 1849 Tabr f regress 45.0 16.25 N 32o 18' 56.2" W 110o 48' 37.5" 3321 11/15/03 2155 Tabr f parous 43.0 13.25 N 32o 18' 56.2" W 110o 48' 37.5" 3322 11/15/03 2155 Tabr f parous 43.0 12.75 N 32o 18' 56.2" W 110o 48' 37.5" 3323 11/15/03 2233 Tabr f parous 42.9 15.00 N 32o 18' 56.2" W 110o 48' 37.5" 3326 12/21/03 2045 Tabr f null 43.0 13.75 N 32o 19' 44.9" W 110o 47' 54.4" 4006 01/11/04 1855 Tabr f parous 43.8 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 4008 01/11/04 1915 Tabr f null 41.7 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 4012 01/11/04 2230 Tabr f parous 42.4 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 4022 02/17/04 2135 Tabr f parous 40.9 10.25 N 32o 19' 48.7" W 110o 47' 39.1" 4023 02/17/04 2135 Tabr f parous 45.3 13.25 N 32o 19' 48.7" W 110o 47' 39.1" 4024 02/17/04 2140 Tabr f parous 42.9 12.75 N 32o 19' 48.7" W 110o 47' 39.1" 4025 02/17/04 2235 Tabr f null 41.0 9.75 N 32o 19' 48.7" W 110o 47' 39.1" 4026 02/17/04 2235 Tabr f null 42.2 11.25 N 32o 19' 48.7" W 110o 47' 39.1" 4033 02/17/04 1207 Tabr f parous 42.0 10.50 N 32o 19' 48.7" W 110o 47' 39.1" 4043 04/14/04 2035 Tabr f parous 41.2 10.30 N 32o 19' 48.7" W 110o 47' 39.1" 4059 05/12/04 2130 Tabr f parous 42.6 12.00 N 32o 18' 54.8" W 110o 48' 40.3" 4061 05/12/04 2214 Tabr f parous 44.5 12.75 N 32o 18' 54.8" W 110o 48' 40.3" 4062 05/12/04 2214 Tabr f parous 43.3 13.00 N 32o 18' 54.8" W 110o 48' 40.3" 4063 05/12/04 2214 Tabr f parous 43.5 13.50 N 32o 18' 54.8" W 110o 48' 40.3" 4069 05/12/04 2340 Tabr f parous 44.6 13.50 N 32o 18' 54.8" W 110o 48' 40.3" 120

TABLE 4.1. Continued 4071 05/12/04 2435 Tabr f - - - N 32o 18' 54.8" W 110o 48' 40.3" 4075 05/20/04 2138 Tabr f preg 42.9 12.50 N 32o 20' 24.9" W 110o 47' 06.4" 4079 06/14/04 2200 Tabr f preg 42.8 13.50 N 32o 18' 56.2" W 110o 48' 37.5" 4108 08/03/04 2125 Tabr f subad 42.7 10.50 N 32o 19' 44.9" W 110o 47' 54.4" 4132 08/08/04 2025 Tabr f regress 42.8 11.00 N 32o 18' 53.2" W 110o 48' 40.8" 4135 08/08/04 2025 Tabr f lac 42.0 11.00 N 32o 18' 53.2" W 110o 48' 40.8" 4140 08/08/04 2140 Tabr f ad not 43.0 10.00 N 32o 18' 53.2" W 110o 48' 40.8" 4144 08/08/04 2225 Tabr f - - - N 32o 18' 53.2" W 110o 48' 40.8" 4157 09/05/04 2100 Tabr f parous 43.0 11.25 N 32o 18' 56.2" W 110o 48' 37.5" 4169 09/05/04 2315 Tabr f regress 43.2 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 4179 10/23/04 1840 Tabr f regress 44.6 14.00 N 32o 20' 24.9" W 110o 47' 06.4" 4183 10/23/04 1930 Tabr f null 43.4 8.75 N 32o 20' 24.9" W 110o 47' 06.4" 4186 11/11/04 1820 Tabr f parous 43.7 14.30 N 32o 18' 56.2" W 110o 48' 37.5" 4187 11/11/04 1820 Tabr f parous 44.1 12.25 N 32o 18' 56.2" W 110o 48' 37.5" 4189 11/11/04 1850 Tabr f parous 43.1 14.25 N 32o 18' 56.2" W 110o 48' 37.5" 5001 01/16/05 1830 Tabr f parous 43.2 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 5002 01/16/05 1855 Tabr f parous 44.4 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 5007 01/16/05 2110 Tabr f parous 43.3 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 5009 01/16/05 2219 Tabr f parous 42.1 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 5016 04/05/05 2200 Tabr f parous 43.4 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 5018 04/05/05 2220 Tabr f parous 43.2 12.59 N 32o 18' 56.2" W 110o 48' 37.5" 5021 04/05/05 2300 Tabr f parous 42.6 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 5022 04/05/05 2300 Tabr f parous 44.8 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 5060 09/03/05 200 Tabr f parous 43.6 11.00 N 32o 20' 24.9" W 110o 47' 06.4" 121

TABLE 4.1. Continued 5077 11/04/05 1830 Tabr f used 44.2 12.75 N 32o 20' 00.4" W 110o 47' 24.4" 5078 11/04/05 1842 Tabr f used 41.2 10.00 N 32o 20' 00.4" W 110o 47' 24.4" 5087 11/04/05 2115 Tabr f used 42.6 12.50 N 32o 20' 00.4" W 110o 47' 24.4" 5089 11/04/05 2200 Tabr f used 43.6 16.50 N 32o 20' 00.4" W 110o 47' 24.4" 2003 3/22/02 1921 Tabr m - 43.5 10.00 N 32o 18' 56.2" W 110o 48' 37.5" 2004 3/22/02 1921 Tabr m - 41.9 10.40 N 32o 18' 56.2" W 110o 48' 37.5" 2005 3/22/02 1928 Tabr m - 42.9 9.50 N 32o 18' 56.2" W 110o 48' 37.5" 2006 3/22/02 1930 Tabr m - 43.2 10.00 N 32o 18' 56.2" W 110o 48' 37.5" 2009 3/22/02 2003 Tabr m - 43.8 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 2014 3/22/02 2220 Tabr m - 43.5 11.25 N 32o 18' 56.2" W 110o 48' 37.5" 2027 05/13/02 1955 Tabr m not 44.0 11.00 N 32o 18' 53.7" W 110o 48' 40.4" 2035 05/13/02 2045 Tabr m not 44.2 10.25 N 32o 18' 53.7" W 110o 48' 40.4" 2037 05/13/02 2112 Tabr m not 45.0 10.25 N 32o 18' 53.7" W 110o 48' 40.4" 2039 05/13/02 2150 Tabr m not 42.3 9.50 N 32o 18' 53.7" W 110o 48' 40.4" 2044 05/13/02 2306 Tabr m not 43.8 10.00 N 32o 18' 53.7" W 110o 48' 40.4" 2046 05/13/02 2355 Tabr m not 44.6 11.00 N 32o 18' 53.7" W 110o 48' 40.4" 2052 05/13/02 2430 Tabr m not 43.6 11.50 N 32o 18' 53.7" W 110o 48' 40.4" 2055 05/13/02 2440 Tabr m not 43.4 11.50 N 32o 18' 53.7" W 110o 48' 40.4" 2056 05/13/02 2445 Tabr m not 43.5 11.50 N 32o 18' 53.7" W 110o 48' 40.4" 2057 05/13/02 2445 Tabr m not 43.4 9.50 N 32o 18' 53.7" W 110o 48' 40.4" 2058 05/13/02 110 Tabr m not - - N 32o 18' 53.7" W 110o 48' 40.4" 2059 05/13/02 110 Tabr m not 42.7 11.00 N 32o 18' 53.7" W 110o 48' 40.4" 2060 05/16/02 1930 Tabr m not 43.0 8.50 N 32o 18' 51.8" W 110o 48' 41.2" 2061 05/16/02 1940 Tabr m not 43.5 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 122

TABLE 4.1. Continued 2066 05/16/02 2000 Tabr m not 42.1 9.75 N 32o 18' 51.8" W 110o 48' 41.2" 2067 05/16/02 2000 Tabr m not 43.5 9.50 N 32o 18' 51.8" W 110o 48' 41.2" 2068 05/16/02 2000 Tabr m not 41.9 9.90 N 32o 18' 51.8" W 110o 48' 41.2" 2074 05/16/02 2021 Tabr m not 40.9 9.25 N 32o 18' 51.8" W 110o 48' 41.2" 2076 05/16/02 2037 Tabr m not 42.6 9.50 N 32o 18' 51.8" W 110o 48' 41.2" 2083 05/16/02 2308 Tabr m not 42.5 11.75 N 32o 18' 51.8" W 110o 48' 41.2" 2098 05/16/02 2458 Tabr m not 44.0 10.50 N 32o 18' 51.8" W 110o 48' 41.2" 2101 05/16/02 105 Tabr m not 43.0 11.00 N 32o 18' 51.8" W 110o 48' 41.2" 2112 05/16/02 353 Tabr m not 44.0 10.00 N 32o 18' 51.8" W 110o 48' 41.2" 2139 06/11/02 2045 Tabr m not 47.4 9.90 N 32o 19' 01.1" W 110o 48' 38.8" 2177 06/30/02 2430 Tabr m not 43.9 10.50 N 32o 19' 24.7" W 110o 48' 27.8" 2181 06/30/02 335 Tabr m not 43.4 10.75 N 32o 19' 24.7" W 110o 48' 27.8" 2182 06/30/02 335 Tabr m not 43.5 9.75 N 32o 19' 24.7" W 110o 48' 27.8" 2192 08/06/02 2055 Tabr m not 44.6 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 2193 08/06/02 2105 Tabr m not 42.5 13.00 N 32o 18' 56.2" W 110o 48' 37.5" 2236 09/04/02 1245 Tabr m not 44.2 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 2237 09/04/02 110 Tabr m not 43.5 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 2238 09/04/02 110 Tabr m not 42.6 11.25 N 32o 18' 56.2" W 110o 48' 37.5" 2239 09/04/02 2150 Tabr m not 42.7 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 2258 10/05/02 1958 Tabr m adnr 42.3 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 2286 10/05/02 350 Tabr m adnr 33.3 10.75 N 32o 19' 44.9" W 110o 47' 54.4" 2300 11/05/02 1840 Tabr m not 43.5 12.75 N 32o 20' 24.9" W 110o 47' 06.4" 2301 12/07/02 1800 Tabr m not 42.2 10.25 N 32o 18' 53.7" W 110o 48' 40.4" 123

TABLE 4.1. Continued 2303 12/07/02 1800 Tabr m not 43.2 11.75 N 32o 18' 53.7" W 110o 48' 40.4" 2304 12/07/02 1800 Tabr m not 44.1 13.00 N 32o 18' 53.7" W 110o 48' 40.4" 2306 12/07/02 1810 Tabr m not 42.2 13.50 N 32o 18' 53.7" W 110o 48' 40.4" 2315 12/07/02 2000 Tabr m not 41.2 10.50 N 32o 18' 53.7" W 110o 48' 40.4" 2319 12/15/02 1840 Tabr m not 42.5 10.75 N 32o 20' 24.9" W 110o 47' 06.4" 2322 12/28/02 1812 Tabr m not 42.2 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 2323 12/28/02 1812 Tabr m not 43.0 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 3007 01/05/03 1810 Tabr m not 42.4 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 3023 03/10/03 1915 Tabr m not 43.5 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3028 03/10/03 2135 Tabr m not 43.3 10.75 N 32o 19' 44.9" W 110o 47' 54.4" 3040 03/22/03 2150 Tabr m not 41.4 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 3044 04/10/03 1911 Tabr m not 44.1 10.75 N 32o 19' 44.9" W 110o 47' 54.4" 3045 04/10/03 1915 Tabr m not 43.2 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3046 04/10/03 1915 Tabr m not 41.4 10.00 N 32o 19' 44.9" W 110o 47' 54.4" 3047 04/10/03 1915 Tabr m not 42.9 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3048 04/10/03 1940 Tabr m not 43.8 11.75 N 32o 19' 44.9" W 110o 47' 54.4" 3050 04/10/03 1950 Tabr m not 44.6 10.75 N 32o 19' 44.9" W 110o 47' 54.4" 3051 04/10/03 1955 Tabr m not 41.5 10.75 N 32o 19' 44.9" W 110o 47' 54.4" 3053 04/10/03 2035 Tabr m not 41.7 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 3055 04/10/03 2045 Tabr m not 45.7 11.00 N 32o 19' 44.9" W 110o 47' 54.4" 3057 04/10/03 2145 Tabr m gular 42.7 13.00 N 32o 19' 44.9" W 110o 47' 54.4" 3059 04/10/03 2205 Tabr m not 43.4 12.75 N 32o 19' 44.9" W 110o 47' 54.4" 3060 04/10/03 2205 Tabr m gular 41.5 11.50 N 32o 19' 44.9" W 110o 47' 54.4" 3067 05/01/03 2120 Tabr m not 43.0 9.75 N 32o 19' 44.9" W 110o 47' 54.4" 124

TABLE 4.1. Continued 3068 05/01/03 2210 Tabr m not 42.8 9.75 N 32o 19' 44.9" W 110o 47' 54.4" 3069 05/01/03 2230 Tabr m not 43.0 9.50 N 32o 19' 44.9" W 110o 47' 54.4" 3100 05/25/03 2215 Tabr m not 44.1 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 3105 05/25/03 2235 Tabr m not 43.3 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 3120 06/09/03 2050 Tabr m not 43.1 11.50 N 32o 19' 01.1" W 110o 48' 38.8" 3121 06/09/03 2050 Tabr m not 42.5 10.75 N 32o 19' 01.1" W 110o 48' 38.8" 3129 06/09/03 2255 Tabr m not 42.3 10.00 N 32o 19' 01.1" W 110o 48' 38.8" 3133 06/09/03 2400 Tabr m not 43.0 12.00 N 32o 19' 01.1" W 110o 48' 38.8" 3134 06/09/03 2408 Tabr m not 43.5 10.50 N 32o 19' 01.1" W 110o 48' 38.8" 3139 06/09/03 225 Tabr m not 42.2 10.25 N 32o 19' 01.1" W 110o 48' 38.8" 3157 06/18/03 2300 Tabr m not 44.1 11.75 N 32o 19' 24.7" W 110o 48' 27.8" 3159 06/18/03 2318 Tabr m not 43.8 12.50 N 32o 19' 24.7" W 110o 48' 27.8" 3162 06/18/03 2355 Tabr m not 42.4 11.25 N 32o 19' 24.7" W 110o 48' 27.8" 3182 08/04/03 2055 Tabr m ad,not 41.9 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 3186 08/04/03 2300 Tabr m ad,not 44.0 12.00 N 32o 18' 56.2" W 110o 48' 37.5" 3188 08/04/03 2309 Tabr m ad,not 42.2 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 3189 08/04/03 2309 Tabr m ad,not 43.9 12.50 N 32o 18' 56.2" W 110o 48' 37.5" 3190 08/04/03 2309 Tabr m ad,not 43.4 10.75 N 32o 18' 56.2" W 110o 48' 37.5" 3194 08/04/03 120 Tabr m ad,not 42.8 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 3200 08/06/03 2120 Tabr m ad,not 42.4 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 3202 08/06/03 2140 Tabr m - - - N 32o 18' 56.2" W 110o 48' 37.5" 3205 08/06/03 2340 Tabr m ad,not 41.6 10.25 N 32o 18' 56.2" W 110o 48' 37.5" 3206 08/06/03 2340 Tabr m ad,not 41.0 10.25 N 32o 18' 56.2" W 110o 48' 37.5" 3208 08/06/03 2400 Tabr m ad,not 42.5 11.75 N 32o 18' 56.2" W 110o 48' 37.5" 3209 08/06/03 2420 Tabr m ad,not 42.6 10.25 N 32o 18' 56.2" W 110o 48' 37.5" 125

TABLE 4.1. Continued 3222 09/05/03 2055 Tabr m unk 42.9 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3234 09/05/03 2325 Tabr m ad 43.0 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3245 09/12/03 2405 Tabr m subad 42.6 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3246 09/12/03 2420 Tabr m ad 42.8 10.25 N 32o 19' 44.9" W 110o 47' 54.4" 3276 10/03/03 2230 Tabr m ad not 43.1 9.50 N 32o 18' 56.2" W 110o 48' 37.5" 3279 10/03/03 2320 Tabr m ad not 43.6 11.25 N 32o 18' 56.2" W 110o 48' 37.5" 3314 10/25/03 215 Tabr m ad not 42.4 14.50 N 32o 18' 56.2" W 110o 48' 37.5" 3319 11/15/03 1903 Tabr m not 42.5 14.00 N 32o 18' 56.2" W 110o 48' 37.5" 3325 12/21/03 1945 Tabr m not 41.5 11.25 N 32o 19' 44.9" W 110o 47' 54.4" 4002 01/11/04 1820 Tabr m ad not 41.7 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 4007 01/11/04 1855 Tabr m ad not 44.2 16.50 N 32o 18' 56.2" W 110o 48' 37.5" 4014 01/11/04 2305 Tabr m ad not 42.7 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 4020 02/17/04 2120 Tabr m no gular 43.5 11.75 N 32o 19' 48.7" W 110o 47' 39.1" 4021 02/17/04 2135 Tabr m no gular 44.9 10.75 N 32o 19' 48.7" W 110o 47' 39.1" 4027 02/17/04 2235 Tabr m no gular 41.8 11.25 N 32o 19' 48.7" W 110o 47' 39.1" 4028 02/17/04 2235 Tabr m not 42.9 10.75 N 32o 19' 48.7" W 110o 47' 39.1" 4030 02/17/04 2250 Tabr m no gular 42.6 12.00 N 32o 19' 48.7" W 110o 47' 39.1" 4034 02/17/04 1207 Tabr m no gular 42.4 11.00 N 32o 19' 48.7" W 110o 47' 39.1" 4037 04/14/04 1940 Tabr m not 43.3 11.50 N 32o 19' 48.7" W 110o 47' 39.1" 4040 04/14/04 2005 Tabr m not 44.4 12.00 N 32o 19' 48.7" W 110o 47' 39.1" 4044 04/14/04 2035 Tabr m not 42.5 11.30 N 32o 19' 48.7" W 110o 47' 39.1" 4051 04/19/04 1940 Tabr m not 42.9 10.00 N 32o 19' 01.1" W 110o 48' 38.8" 4053 04/19/04 2220 Tabr m not 43.9 11.50 N 32o 19' 01.1" W 110o 48' 38.8" 4055 04/24/04 1925 Tabr m not 42.6 11.00 N 32o 19' 48.7" W 110o 47' 39.1" 126

TABLE 4.1. Continued 4058 05/12/04 2130 Tabr m not 42.7 10.30 N 32o 18' 54.8" W 110o 48' 40.3" 4060 05/12/04 2130 Tabr m not 42.8 10.75 N 32o 18' 54.8" W 110o 48' 40.3" 4064 05/12/04 2245 Tabr m not 44.0 10.75 N 32o 18' 54.8" W 110o 48' 40.3" 4065 05/12/04 2316 Tabr m not 43.2 11.00 N 32o 18' 54.8" W 110o 48' 40.3" 4066 05/12/04 2316 Tabr m not 43.0 12.25 N 32o 18' 54.8" W 110o 48' 40.3" 4070 05/12/04 2425 Tabr m not 44.3 12.00 N 32o 18' 54.8" W 110o 48' 40.3" 4120 08/08/04 2020 Tabr m ad not 39.4 11.00 N 32o 18' 53.2" W 110o 48' 40.8" 4124 08/08/04 2010 Tabr m ad not 43.4 11.00 N 32o 18' 53.2" W 110o 48' 40.8" 4126 08/08/04 2025 Tabr m ad not 43.4 10.75 N 32o 18' 53.2" W 110o 48' 40.8" 4127 08/08/04 2043 Tabr m ad not 42.6 11.25 N 32o 18' 53.2" W 110o 48' 40.8" 4129 08/08/04 2043 Tabr m ad not 42.0 10.75 N 32o 18' 53.2" W 110o 48' 40.8" 4134 08/08/04 2025 Tabr m ad not 42.3 10.25 N 32o 18' 53.2" W 110o 48' 40.8" 4136 08/08/04 2121 Tabr m subad 44.1 9.50 N 32o 18' 53.2" W 110o 48' 40.8" 4137 08/08/04 2120 Tabr m ad not 44.1 11.00 N 32o 18' 53.2" W 110o 48' 40.8" 4138 08/08/04 2120 Tabr m ad not 43.1 9.50 N 32o 18' 53.2" W 110o 48' 40.8" 4139 08/08/04 2140 Tabr m subad 41.7 8.50 N 32o 18' 53.2" W 110o 48' 40.8" 4141 08/08/04 2145 Tabr m ad not 42.9 10.50 N 32o 18' 53.2" W 110o 48' 40.8" 4142 08/08/04 2215 Tabr m subad 41.6 9.50 N 32o 18' 53.2" W 110o 48' 40.8" 4162 09/05/04 2136 Tabr m not 42.4 11.25 N 32o 18' 56.2" W 110o 48' 37.5" 4166 09/05/04 2205 Tabr m ad not - - N 32o 18' 56.2" W 110o 48' 37.5" 4167 09/05/04 2255 Tabr m ad not 43.1 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 4168 09/05/04 2255 Tabr m ad not 43.4 11.25 N 32o 18' 56.2" W 110o 48' 37.5" 4170 09/05/04 2315 Tabr m ad not 43.1 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 4173 09/05/04 2355 Tabr m - - - N 32o 18' 56.2" W 110o 48' 37.5" 127

TABLE 4.1. Continued 4193 12/13/04 1800 Tabr m not 42.9 11.25 N 32o 19' 48.7" W 110o 47' 39.1" 5005 01/16/05 1908 Tabr m not 42.3 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 5008 01/16/05 2130 Tabr m not 42.7 9.00 N 32o 18' 56.2" W 110o 48' 37.5" 5013 04/05/05 2035 Tabr m - 43.4 11.00 N 32o 18' 56.2" W 110o 48' 37.5" 5017 04/05/05 2200 Tabr m - 42.8 12.09 N 32o 18' 56.2" W 110o 48' 37.5" 5020 04/05/05 2300 Tabr m - 43.7 10.50 N 32o 18' 56.2" W 110o 48' 37.5" 5023 04/05/05 110 Tabr m - 42.5 11.50 N 32o 18' 56.2" W 110o 48' 37.5" 5035 09/03/05 2050 Tabr m ad not 42.3 10.50 N 32o 20' 24.9" W 110o 47' 06.4" 5042 09/03/05 2315 Tabr m ad not 46.6 - N 32o 20' 24.9" W 110o 47' 06.4" 5043 09/03/05 2320 Tabr m ad not 43.5 10.75 N 32o 20' 24.9" W 110o 47' 06.4" 5044 09/03/05 2320 Tabr m not 43.0 11.50 N 32o 20' 24.9" W 110o 47' 06.4" 5045 09/03/05 2330 Tabr m - 42.7 9.50 N 32o 20' 24.9" W 110o 47' 06.4" 5046 09/03/05 2340 Tabr m - 44.6 11.50 N 32o 20' 24.9" W 110o 47' 06.4" 5047 09/03/05 2340 Tabr m - 41.7 9.50 N 32o 20' 24.9" W 110o 47' 06.4" 5049 09/03/05 2345 Tabr m subad 42.7 11.00 N 32o 20' 24.9" W 110o 47' 06.4" 5050 09/03/05 2350 Tabr m ad not 43.1 10.75 N 32o 20' 24.9" W 110o 47' 06.4" 5066 11/04/05 1802 Tabr m not 43.1 9.80 N 32o 20' 00.4" W 110o 47' 24.4" 5074 11/04/05 1820 Tabr m not 44.1 12.00 N 32o 20' 00.4" W 110o 47' 24.4" 5076 11/04/05 1830 Tabr m not 43.3 13.00 N 32o 20' 00.4" W 110o 47' 24.4" 5088 11/04/05 2115 Tabr m not 43.1 12.25 N 32o 20' 00.4" W 110o 47' 24.4" 128

APPENDIX E: COMPARATIVE ANALYSIS

Bats partition resources using difference techniques such as foraging at different elevations (i.e., height above ground) or habitats (Schnitzler and Kalko 1998). This section investigates comparative data, including different elevations in the net that the seven most frequently captured species were caught (Table 5.1), different directions that species were flying upon capture (Table 5.2) and plots of sex ratios of captured bats during 2002-2003 (Fig. 5.1) and 2004-2005 (Fig. 5.2). Recent bat research in Mohave (Williams et al. 2006) and Chihuahuan Deserts (Higgenbotham 1999) reflects lower capture rates (Table 5.3) than our Sonoran Desert study. Although this may be, in part, to different sampling protocol, it also suggests that the Sonoran Desert has more available resources for bats. The Mojave Desert gets most of its precipitation (~95.6 mm/year) during winter months (Turner 1994). The Chihuahuan Desert also has a unimodal rainfall pattern but gets its precipitation (~250 mm/year) during summer (Brown 1994). The Sonoran Desert has a bimodal rainfall pattern and is the most lush of the four Western U.S. deserts (Turner and Brown 1994). The Sonoran rainfall pattern may contribute to greater foraging resources for bats. Winter rains restock water holes and encourage growth in winter forbs and grasses, providing forage for insect prey. Summer storms provide much needed water during the hottest time of the year, replenishing water holes that dried in early summer and providing critical water for plants dependent on summer precipitation. Other researchers looking at bat distributions in the Lower Colorado Subdivision of the Sonoran Desert Biome (Cutler 1996, Schmidt 1999, Kuenzi 2001) also demonstrated the importance of water for bats in xeric landscapes. Although they captured fewer species than our study, they sampled for bats at wildlife water developments not associated with riparian landscapes. It may be that montane canyons with riverine environments provide greater structural complexity and offer more foraging opportunities to bats (Grindal et al. 1999). 129

TABLE 5.1. To capture free-flying bats we used 2-ply 50 denier, 38 mm mesh nylon mist nets of either 6, 9, or 12 m stretched across flowing or pooled water along Sabino Creek (Kunz and Kurta 1988). We placed our nets perpendicular to the creek’s flow and recorded the direction that the bat was flying (upstream vs. downstream) when captured to evaluate any directional preference by bats (Adams and Simmons 2002). Of the seven most frequently captured species, N. femorosaccus was the only species that indicated any preference for flight direction during 2002-2005.

Species N* US DS χ2

Eptesicus fuscus 64 50.0 50.0 1.00

Lasiurus cinereus 28 53.6 46.4 0.71

Myotis yumanensis 23 52.2 47.8 0.83

Nyctinomop femorosaccus 229 44.3 55.7 0.09

Nyctinomop macrotis 19 52.6 47.4 0.82

Pipistrellus hesperus 167 55.1 44.9 0.19

Tadarida brasiliensis 234 48.5 51.5 0.65

* directional data missing for some bats 130

TABLE 5.2. When netting we stacked two nets vertically with a single shelf overlap between upper and lower nets (covering a ~3.6 m vertical flight window) at each site and recorded the height a bat was flying upon capture. Of the seven most frequently captured species, only P. hesperus and M. yumanensis showed strong evidence for foraging just above the water’s surface along Sabino Creek during 2002-2005.

Species N* High Net Low Net χ2

Eptesicus fuscus 64 51.5 48.5 0.81

Lasiurus cinereus 27 48.1 51.9 0.85

Myotis yumanensis 24 16.7 83.3 0.01

Nyctinomop femorosaccus 221 54.3 45.7 0.20

Nyctinomop macrotis 23 65.2 34.8 0.14

Pipistrellus hesperus 149 32.2 67.8 <0.001

Tadarida brasiliensis 237 52.3 47.7 0.47

* height data missing for some bats 131

FIGURE 5.1. Sex ratios of bats captured along Sabino Creek during 2002-2003, prior to post-Aspen Fire sedimentation of pools. males females 140

120

100

Number of individuals 40

0 Nyfe Tabr Pihe Epfu Nyma Myyu Laci Eupe Myve Chme Myca Laxa Labl Coto Lano Myth Myau Species 132

FIGURE 5.2. Sex ratios of bats captured along Sabino Creek during 2004-2005, after post-Aspen Fire sedimentation of pools.

Males Females

30 Number of individuals 20

0 Nyfe Tabr Pihe Epfu Nyma Myyu Laci Eupe Chme Myca Labl Coto Myau Species 133

TABLE 5.3. Results of comparative bat research across three Southwestern deserts, with hourly capture rates. These data suggest resource levels for bats are variable across different biotic communities and elevations

Hourly Year of Number Total Capture Study Location Report of Species Captured Hours Netting Rate Elevation

Sonoran Desert (Lower Colorado Subdivision)1 1996 7 164 ~72 2.28 ~300 m Sonoran Desert (Arizona Upland Subdivision)2 1999 9 1,153 795 1.45* ~270 - 390 m Sonoran Desert (Lower Colorado & AZ Upland)3 2001 6 718 1,552 0.46* ~395 - 910 m Chihuahuan Desert 4 1999 18 1,978 1,334 1.48 ~550 - 1500 m Mojave Desert 5 2006 13 394 312 1.26 ~520m Sonoran Desert (Arizona Upland Subdivision)6 2007 17 961 385 2.50 750 - 960 m

1 Cutler, 2 Schmidt, 3 Kuenzi, 4 Higgenbotham, 5 Williams et al., 6 Buecher * Capture rate calculated using net lengths x hours and may not be comparable to other studies

Note: Some studies also trapped bats at roosts but analysis of hourly capture rate includes results from netting along riparian corridors or at water sources only 134

APPENDIX F: SPECIES ACCOUNTS, SEASONAL DISTRIBUTIONS AND COMPARATIVE MEASUREMENTS OF THE SEVEN MOST FREQUENTLY CAPTURED SPECIES ALONG SABINO CREEK, 2002-2005

Eptesicus fuscus pallidus (Big brown bat):

Big brown bats are one of the most cosmopolitan temperate bat species, ranging from Central America north to Canada. They are found throughout Arizona, primarily in woodlands (Hoffmeister 1986), but also found in desertscrub and grassland (Sidner

1997). They forage throughout the night with greatest activity during the second hour after sunset (Kurta and Baker 1990). Big brown bats were our fourth most commonly netted species (n = 70) along Sabino Creek, making up 7.3% of total captures. Although big brown bats are documented year-round at lower elevations in southern Arizona

(Hoffmeister 1986), I only captured them in spring, summer and early fall (Fig. 6.1) at nine sites along Sabino Canyon, suggesting that they seasonally migrate from the area (or up in elevation). Females form maternity colonies during summer but males are generally solitary. I did not capture lactating females, so little evidence exists for a maternity colony within lower Sabino Canyon. Sexual dimorphism occurs in big brown bats (Sidner 1997), with female forearms longer than males (Fig. 6.2), which I observed in big browns captured along Sabino Creek (n male = 35, n female = 35, t = 2.85, df = 68, p =

0.006: FAmale = 46.39mm, 95% CI 45.80-46.98; FA female = 47.53mm, 95% CI 46.99-

48.08). I caught 7 subadults and their body mass was not significantly lower than that of adults (t = 1.35, df = 68, p = 0.18, Mass subadults = 12.96, 95% CI 11.34-14.59, Mass adults =

14.49, 95% CI 13.76 –15.22). 135

FIGURE 6.1. Monthly distribution of sexes for big brown bats along Sabino Creek 2002- 2005, with limited netting during July.

20 Females Males 15

10 Number of Individuals 5

0 J FMAMJ J ASOND Month

FIGURE 6.2. Differential forearm lengths between male and female big brown bats – note sexual dimorphism with females having longer forearms (n = 70).

45 Average Forearm Lengths

44 Female Male 136

Nyctinomops macrotis (Big free-tailed bat): Big free-tailed bats are the largest member of genus Nyctinomops and range from southern Bolivia, through Central America and southwestern United States (Milner et al.

1990). They prefer rugged canyons and besides early records from Sabino Canyon

(Hoffmeister 1986), they are documented from Big Bend, Texas (Higgenbotham 1999),

Chiricahua Mountains, Arizona (Barbour and Davis 1969), northern New Mexico, southern Colorado and Utah (Adams 2003). Although this species is found throughout

Arizona, it occurs in low numbers (Hoffmeister 1986) and due to its large size, has the ability to fly long distances each night (~32 km - Corbett 2005). Big free-tailed bats have low frequency sonar calls (i.e., within human hearing) and can be heard flying high above

Sabino Canyon. I captured big free-tails at six pools, typically netting them in spring or fall (Fig. 6.3). This species was never captured in large numbers except in October 2003, when I caught 9 one evening at Anderson Dam. Because I captured that many big free- tailed bats at one time, I suspect they were migrating through the area.

In April 2004 I caught an adult female big free-tailed bat and attached a Holohil

Systems (Ontario, Canada) BD-2 (0.72 g) radio transmitter (< 5% of body mass), tracking her to a crevice high on a cliff near Thimble Peak. I was only able to follow her for two nights and suspect she either migrated from the area or successfully groomed the transmitter off beyond range of my radio receiver. I later rappelled down to a ledge below the crevice where I had tracked her and found a large pile of guano, suggesting that this roost is either used on a regular basis or has been used sporadically for many years. 137

As with all free-tailed bats, big free-tails have long narrow wings producing a high-aspect ratio (i.e., stall at higher flight speeds) so prefer to drink from larger pools

(Barbour and Davis 1969). Requisite pool conditions are limited along Sabino Canyon during dry months of the year. However, animals can fly long distances and may day- roost in the Santa Catalina Mountains but drink from large ponds at nearby golf courses.

There is weak evidence of reverse sexual dimorphism (Fig. 6.4), with the forearms of the males longer than those of the females (n female = 15, n male = 9, t = -1.88, df = 22, p = 0.07, FA male = 62.14mm (95% CI 61.09-63.20); FA female = 60.92mm (95%

CI 60.10-61.74). Pregnant big free-tailed bats may have reduced maneuverability with long narrow wings, especially if the late stage fetus is ~16% of the female’s mass (Milner et al. 1990) and natural selection may favor slightly shorter, but broader wings in females. During future work along Sabino Creek, I plan look at overall wing shape (i.e., measuring wing surface area) of big free-tailed bats to determine if this hypothesis is correct. 138

FIGURE 6.3. Monthly distribution of sexes for big free-tailed bats along Sabino Creek, Arizona, 2002-2005.

15 Females 12 Males

Numbers of Individuals 3

0 JFMAMJJASOND Months

FIGURE 6.4. Differential forearm lengths between male and female big-free-tailed bats - note reverse sexual dimorphism than usually seen between sexes in bats (n = 24).

63.0

62.0

61.0 Average Forearm Length

60.0 Female Male 139

Lasiurus cinereus cinereus (Hoary bat):

Hoary bats are the most widespread North American bat species, ranging from

Mexico to central Canada (Shump and Shump 1982). Because this species does not use caves or similar protected roost-sites, it is heavily furred including the interfemoral membrane, providing additional thermoregulatory insulation. Unfortunately, wind turbines along eastern United States and western Canada are negatively impacting this species. There are recent records of hundreds of deaths of migrating lasiurine bats associated with large wind farms (Cryan and Brown 2007).

I caught hoary bats every month except June-August (Fig. 6.5), when they may forage at higher elevations. Female lasiurine bats have two pair of anterior mammary glands and nipples (Racey 1988) and hoary bats often bear twins but can have up to four young/yr (Barbour and Davis 1969). There is sexual dimorphism in hoary bats (Fig. 6.6), with females larger than males. Although I had a small sample size for hoary bats (n female = 10, n male = 19), I documented differences in forearm lengths (t = 3.87, df = 1, p =

0.0006, FA female = 54.94mm, 95% CI 54.29 - 55.59; FA male = 52.78mm, 95% CI 52.07 -

53.50) and differential body mass (t = 6.12, df = 25, p = 0.0001; mass female = 28.65, 95%

CI 26-97 – 30.33, mass male = 22.69, 95% CI 21.54 – 23.84) between sexes.

April 2006 I radio-tracked a male hoary caught near Sabino Canyon Dam. For 5 days he day-roosted in large willow trees (Salix gooddingii) and an ash tree (Fraxinus velutina) in the same area as his capture. For three nights he consistently foraged along the dense riparian vegetation just above Sabino Canyon Dam before he groomed off the transmitter on Day 6. 140

FIGURE 6.5. Monthly distribution of sexes for hoary bats along Sabino Creek, 2002- 2005 - with limited netting during July.

7 Females Males 6

2 Number of Individuals 1

0 JFMAMJJASOND Month

FIGURE 6.6. Differential forearm lengths between male and female hoary bats (n = 29).

52 Average Forearm Lengths 51

Female Male 141

Tadarida brasiliensis mexicana (Mexican free-tailed bat):

The only member of the genus Tadarida in North America, Mexican free-tailed bats congregate in the largest aggregations of mammals known worldwide. This species is widespread throughout southwestern U. S. and one of the largest known maternity sites

(prior to a collapse in population) was at Eagle Creek Cave, Graham County (Barbour and Davis 1969). Mexican free-tailed bats range from southern South America through central North America (Wilkins 1989). Males have a gular gland on their upper chest that becomes enlarged when they are reproductive and both sexes have a distinctive

‘musky’ odor, helpful when identifying abandoned roosts. As with all free-tailed bats, the wings of Mexican free-tails are long and narrow and these animals have high wing loading, resulting in rapid flight but reduced maneuverability (Wilkins 1989).

Mexican free-tailed bats day-roost in crevices under urban concrete bridges in the

Tucson Basin (Buecher unpubl. data, Wolf 2003) and crevices in the University of

Arizona football stadium (R. Sidner pers. comm.). Although there are no known

Mexican free-tailed bat day-roosts in Sabino Canyon, these animals can easily travel 50 km nightly from their roosts (Davis et al. 1962) and could commute from urban structures to forage and obtain water along Sabino Creek. Mexican free-tailed bats are year-round residents in Tucson, although they occur in reduced numbers in the winter. During 2002-

2005, I captured Mexican free-tailed bats throughout the year at ten sites along Sabino

Creek (Fig. 6.7).

There is no difference in forearm lengths between male and female Mexican free- tailed bats (Fig. 6.8). 142

FIGURE 6.7. Monthly distribution of sexes for Mexican free-tailed bats along Sabino Creek, Arizona, 2002-2005 - with limited netting during July. 70

60 Females 50 Males

Number of Individuals 20

0 JFMAMJJASOND Month

FIGURE 6.8. No difference in forearm lengths between male and female Mexican free-tailed bats (n = 281)

43.2

43.1

43.0

42.9 Average Forearm Lengths 42.8 Female Male 143

Nyctinomops femorosaccus (Pocketed free-tailed bat):

Nyctinomops femorosaccus is generally tropical in distribution, with its U.S. range extending along southern Texas, New Mexico, Arizona and California. Pocketed free-tailed bats were the most frequently captured species in Sabino Canyon during 2002-

2003 (32% of overall captures) however, after post-Aspen Fire sedimentation of pools their numbers dropped (14% of overall captures). I captured a number of pregnant, lactating and post lactating females and in late summer I caught subadults (Fig. 6.9). In

August 2003 I radio-tracked two adult, post-lactating females for 2+ weeks until I located their roost in upper Bear Canyon. This roost was ~50m above the floor of the canyon, allowing animals a substantial drop into flight. This maternity roost is a significant discovery because little is known regarding this species’ natural history (Kumirai and

Jones 1990, Adams 2003). In June 2003 the Aspen Fire burned the east side of Bear

Canyon directly across from the day-roost. Fortunately the site is on a vertical cliff and fire is not a potential threat, although the smell of smoke likely disturbed females during parturition.

During 2002-2005 I caught 253 pocketed free-tailed bats at eleven sites along

Sabino Creek. Body mass of subadult pocketed free-tailed bats was significantly lower than adults (t = 10.17, df = 245, p = 0.0001, body mass adults = 14.52g, 95% CI 14.30 –

14.75, body mass subadults = 12.81g, 95% CI 12.59 – 13.02) during early autumn. As with big free-tailed bats (N. macrotis) captured in Sabino Canyon, there is strong evidence of sexual dimorphism in forearm length (Fig. 6.10). The males’ forearms are significantly longer than those of females (n female = 145, n male = 106, t = -4.43, df = 249, p = 0.0001), 144

FA male = 46.97mm (95% CI 46.79 – 47.15); FA female = 46.44mm (95% CI 46.29 –

46.59). Long narrow wings, especially for pregnant females, affect power (Norberg

1995) and may contribute to reduced maneuverability, which could be a disadvantage when foraging. As a result, females may be selected for slightly shorter, but broader wings in this genus. Future work will investigate overall wing shape to determine differences in aspect ratio (wingspan2/wing area) between sexes. This will involve measuring the length of the 5th finger and recording the ratio of chiropatagium to plagiopatagium (Altringham 1996). As with Mexican free-tailed bats and western pipistrelles, pocketed free-tailed bats were active year-round, although in reduced numbers during winter. 145

FIGURE 6.9. Monthly distribution of sexes for pocketed free-tailed bats (Nyctinomops femorosaccus) along Sabino Creek, Arizona, 2002-2005.

80 70 Females l 60 Males 50 40 30 20 Number of Individua 10 0 JFMAMJJASOND Month

FIGURE 6.10. Differential forearm lengths between male and female pocketed free- tailed bats – note that the females have shorter forearms than males (n = 251)

47.5

47.0

46.5

46.0

Average Forearm Length 45.5

45.0 Female Male 146

Pipistrellus hesperus hesperus (Western pipistrelle bat):

Western pipistrelles are the smallest bats in North America, the earliest to begin foraging and often emerge to forage again just at sunrise (Adams 2003, Cockrum and

Cross 1964, Cross 1965). Pipistrelles are distinguished by slow erratic flight and forage in the vicinity of day-roosts (Cross 1965). Although most female pipistrelles have a body mass <5g, they commonly bear twins. Pipistrelles are found from hot arid lower Sonoran

Desert (Kuenzi and Morrison 2003, Rabe and Rosenstock 2005), through desertscrub

(Cross 1965) and into semiarid grassland and adjacent woodlands (Hoffmeister 1986).

Another common name for western pipistrelles is ‘canyon bat’ which is descriptive of the habitat that these animals prefer. Western pipistrelles range from Mexico through western canyons as far north as Idaho but never ranging too far from appropriate day roosts (Adams 2003, Wilson and Ruff 1999). Western pipistrelles use small cliff crevices along canyons (Cross 1965), living in small colonies (typically 5-12 individuals including young) and are not particularly loyal to one roost. Much of our current knowledge of pipistrelle day-roosts was documented along Sabino Canyon (Cross 1965) and I often caught pregnant, lactating and post-lactating females, confirming that Sabino Canyon still maintains maternity colonies (Fig. 6.11). Perhaps because of their small size and multiple offspring (2 pups/year), female pipistrelles have longer forearms (t = 8.95, df =

185, p = 0.0001, FA male = 29.42, 95% CI 29.20 – 29.64, FA female = 30.69, 95% CI 30.52

–30.87 – Fig. 6.12) and greater body mass than males (t = 8.93, df = 125, p < 0.0001,

Mass female = 4.09, 95% CI 3.95 – 4.22, Mass male = 3.31, 95% CI 3.20 – 3.41). Western pipistrelles were the third most frequently captured species along Sabino Creek and 147

documented from 12 sites. Body mass differed between adults and subadults in the fall (t

= 3.63, df = 154, p = 0.0004, body mass adults = 3.71, 95% CI 3.60 – 3.82, body mass subadults = 3.27, 95% CI 3.11 – 3.43). This supports evidence that, at least for some bat species, young-of-the-year are underweight in fall, a condition that may contribute of low survivorship during winter (Sidner 1997).

Western pipistrelles were one of three bat species captured year-round and often emerged to obtain drinking water and/or forage for small insects on warm winter evenings. Work in Nevada (O’Farrell and Bradley 1970, O’Farrell et al. 1967) also found P. hesperus active during winter months. O”Farrell and Bradley (1977) recorded rectal body temperatures (Tb ) of P. hesperus captured in winter and found that they had lower Tb and were active at lower ambient temperatures (Ta) than other seasons. It is hypothesized that bats flying in winter are not foraging on insects but searching for drinking water because light-trapping has shown a paucity of available insect prey and examination of stomach contents indicated that animals were not eating insects

(O”Farrell and Bradley 1977, Whitaker and Rissler 1993). Additionally, Nevada bats showed adequate fat reserves when captured in winter. It may be that P. hesperus caught on warm winter evenings along Sabino Creek are day-roosting in sites inappropriate for hibernation and lose water reserves that must be periodically replenished. 148

FIGURE 6.11. Monthly distribution of sexes for western pipistrelles, in 2004 the only available water during June was in Bear Canyon

70 Females 60 Males

20 Number of Individuals

0 J FMAMJ J ASOND Month

FIGURE 6.12. Differential forearm lengths between male and female western pipistrelles - female pipistrelles carry twins and may have a longer forearm to compensate (n = 192)

28 Average Forearm Length 26 Female Male 149

Myotis yumanensis (Yuma myotis): Yuma myotis are broadly distributed throughout western United States and western Canada. This species is a small little brown bat with large feet, usually associated with water (Duff and Morrell 2007) where it forages on emergent insects or insects trapped on the water’s surface tension (Findley 1993, Adams 2003). I caught 26

Yuma myotis at six pools (Fig. 6.13), of which 21 were captured in 2002-2003, prior to extensive post-Aspen fire sedimentation of pools along Sabino Creek. In the hand, Yuma myotis can be confused with little brown bats (M. lucifugus), although little brown bats are not commonly found in Sonoran desertscrub. During acoustic sampling, Yuma myotis can be confused with California myotis (M. californicus) because both species are

50 kHz myotis with similar shaped ultrasonic calls. Female and male Yuma myotis along

Sabino Creek did not have different forearm lengths (t = 0.48, df = 23, p = 0.64, FA male

= 33.65, 95% CI 32.99 – 34.32, FA female = 33.88, 95% CI 33.15 – 34.62 – Fig. 6.14)).

Yuma myotis is one species that might well benefit from maintenance of pools throughout the year along Sabino Creek. 150

FIGURE 6.13. Monthly distribution of sexes for Yuma myotis – capture rates dropped dramatically after reduced water availability

6 Females Males 5

2 Number of Individuals

0 J FMAMJ J ASOND Month

FIGURE 6.14. There is no difference in forearm lengths between male and female Yuma myotis ( n = 25).

33 Average Forearm Length 32 Female Male 151

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