Seasonal and Nightly Activity of Mexican Long-Nosed Bats

SEASONAL AND NIGHTLY ACTIVITY OF MEXICAN LONG-NOSED BATS

(LEPTONYCTERIS NIVALIS) IN BIG BEND NATIONAL PARK, TEXAS

A Thesis

Presented to the

Faculty of the College of Graduate Studies

Angelo State University

In Partial Fulfillment of the

Requirements for the Degree

MASTER OF SCIENCE

ERIN RANELLE ADAMS

December 2015

Major: Biology

SEASONAL AND NIGHTLY ACTIVITY OF MEXICAN LONG-NOSED BATS

(LEPTONYCTERIS NIVALIS) IN BIG BEND NATIONAL PARK, TEXAS

ERIN RANELLE ADAMS

APPROVED:

Dr. Loren K. Ammerman

Dr. Nicholas J. Negovetich

Dr. Amaris R. Guardiola

Dr. Laurence F. Jones

November 6, 2015

APPROVED:

Dr. Susan E. Keith Dean, College of Graduate Studies

DEDICATION

I dedicate this thesis to Bryan Adams, “Everything I do, I do it for you”;

and to the bats.

iii

ACKNOWLEDGMENTS

I owe many thanks Dr. Loren K. Ammerman. She took a chance on me as a non- traditional graduate student with an over-abundant sense of gumption. I have valued her

instruction, direction, work-ethic, confidence, and wisdom over these past two years. Her expertise in bat biology and her courage in research have showed me what it takes to be an effective, well-rounded biologist. I am grateful she gave me many opportunities to develop and focus my research skills without dampening my excitement for wildlife biology. I could not have asked for a greater mentor or friend in my graduate career.

In addition I would like to thank the following people for their logistical help,

brainstorming, thoughts, prayers, and participation to make this project possible.

Thank you to my thesis committee for your guidance; Dr. Nicholas Negovetich, Dr.

Robert Dowler, Dr. Casey Jones, and Dr. Amaris Guardiola. I would also like to thank the

ASU Faculty and Staff who provided me assistance and advice; Dr. Ben Skipper, Dr.

Michael Dixon, Dr. Terry Maxwell, Dr. Laurel Fohn, Jody Casares, and Katie Plum.

I am grateful for the input, support, and encouragement from other biologists and

friends during this project; Raymond Skiles, Cyndee Watson, Holly Niederriter, Kevin

Oxenrider, Sarah Bouboulis, Lauren Johnson, Matthew Fisher, Christina Kocer, Becky

Rosamond, Jeff Gore, Steven Thomas, Carl Herzog, Winifred Frick, Rodrigo Medellín and

his laboratory at UNAM, and Laramie Lindsey.

Thank you to those I have leaned on for support; my family John, Ranelle, Shannon,

John, Meghan, and Jess, plus Gene and Kathy Morris and Bryan Lopez.

A special thanks goes out to those who joined me for field work and endured the car

rides, the trail miles, the heat and hard work in Big Bend National Park. Your help and hard

work was essential to the project; Caitlin Bruns, Antonio Cantu, Natalie Craig, Sarah

Gonzales-Perez, Riccay Harrison, Thomas Horsley, Korry Huddleston, Malorri Hughes, Ana

Ibarra, P. Citlally Jimenez, Mary Jones, Katie Kuzdak, Lajitas Stables (Janelle, Tiffany and

Rosie), Clint Morgan, Austin Osmanski, Doug Raybuck, Kaitlin Thogmartin, T. Marie Tipps,

Leonora Torres-Knoop, and Megan Wallrichs. Thank you to the staff in Big Bend National

Park including Diana Edwards and Bunny Larson.

Most of all to my husband and best friend, Bryan Adams, without you this all would

have never come to pass. Thanks for your love, support, prayers, encouragement, prodding,

pack-muling, hiking, laughter, comforting, and all-around sacrifice in order to make this project happen from start to finish. We did it!

I am grateful to the following organizations for funding through donations, grants and scholarships to support this project; Angelo State University (ASU), ASU Center for

Innovative Teaching and Research, ASU Center for International Studies, Head of the River

Ranch, Southwestern Association of Naturalists - Howard McCarley Student Research

Award, Batteries Plus – San Angelo, and Bat Conservation International.

This work was performed under Texas Parks and Wildlife permit SPR-0994-703,

Resource Activity permits from the National Park Service (#BIBE-2012-SCI-0031 and

#BIBE-2014-SCI-0029), and an Endangered Species permit from the U. S. Fish and Wildlife

Service (TE127287-1).

All animal handling protocols were consistent with guidelines published by the

American Society of Mammalogists (Sikes et al. 2011), and were approved by the IACUC committees of Angelo State University (#1310) and the National Park Service

(#IMR_BIBE_Ammerman_Mexican long-nosed Bat_2014.A2).

ABSTRACT

The seasonality and activity of endangered Mexican long-nosed bats (Leptonycteris nivalis) was studied by Passive Integrated Technology (PIT) tagging in Big Bend National

Park, Texas. Activity of 79 bats (out of 104 total tagged bats) at Mount Emory Cave was monitored via a serpentine antenna from 26 April – 1 September 2014 and 16 June – 19

August 2015. First year return rates included 42% adult females, 50% juvenile females, and

8% juvenile males. Cave use varied by demographic; on average juvenile males were active over longer duration each night (p<0.01) and greater range of dates (p<0.02) than lactating females. The most detections of bats occurred in the morning hours (p<0.005). Individuals were present for an average of 13.9 nights (±10.3, range: 1-39). Since some tagged

individuals return to the cave annually, apparent survival and redetection probabilities can be

generated for this colony over time and improve our understanding of this species.

TABLE OF CONTENTS

Page

DEDICATION ...... iii

ACKNOWLEDGMENTS ...... iv

ABSTRACT ...... vi

TABLE OF CONTENTS ...... vii

LIST OF TABLES ...... ix

LIST OF FIGURES ...... x

INTRODUCTION ...... 1 Conservation status ...... 1 Distribution ...... 2 Monitoring with technology ...... 4 Study objectives ...... 6 METHODS ...... 8

Study site ...... 8 Capture and marking techniques ...... 9 Remote PIT tag detection system ...... 10 Seasonality ...... 14 PIT tag record analysis...... 15 Nightly activity ...... 17 RESULTS ...... 18

Seasonality ...... 21 PIT tag record analysis...... 21 Preliminary apparent survival estimates ...... 27 Nightly activity ...... 27 DISCUSSION ...... 33

Seasonality ...... 33 vii

PIT tag detection ...... 34 Apparent survival and probability of redetection estimates ...... 35 Nightly activity ...... 37 Future directions ...... 39 LITERATURE CITED ...... 43

APPENDIX 1 ...... 48

APPENDIX 2 ...... 50

viii

LIST OF TABLES

TABLE 1. Demographics of Mexican long-nosed bats (Leptonycteris nivalis) captured and implanted with PIT tags at Mount Emory Cave in 2014 and 2015 ...... 20

TABLE 2. Results from program MARK for modeling the apparent survival and capture probabilities of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) at Mount Emory Cave during the 2014-2015 study period over two week intervals across the study ...... 28

TABLE 3. Top two model estimate results for apparent survival and probability of redetection for PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) at Mount Emory Cave during the 2014-2015 study period ...... 29

TABLE 4. The average duration of activity of Mexican long-nosed bats (Leptonycteris nivalis) per age and reproductive group based on the time between maximum emergence and return triggers in the same night, generated from nights in 2014 and 2015 ...... 31

LIST OF FIGURES

FIGURE 1. A sketch of the serpentine antenna configuration used during this study to detect PIT tags of Mexican long-nosed bats tagged at Mount Emory Cave ...... 13

FIGURE 2. Yearly totals of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) at Mount Emory Cave including 12 redetections. Eight individuals were detected in both years and 4 were tagged in 2014 but only detected in 2015...... 22

FIGURE 3. All days of detection of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) in 2014 and 2015 at Mount Emory Cave ...... 24

FIGURE 4. Percentage of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) detected at Mount Emory Cave in 2014 (n= 38) and 2015 (n=104) ...... 25

FIGURE 5. Total nights each PIT tagged Mexican long-nosed bat (Leptonycteris nivalis) was detected at Mount Emory Cave in 2014 and 2015 ...... 26

FIGURE 6. Total detections of Mexican long-nosed bat (Leptonycteris nivalis) activity at Mount Emory Cave in July 2014 and 2015 ...... 32

INTRODUCTION

Bats in the genus Leptonycteris (Phyllostomidae: Glossophaginae) are migratory nectar feeders and are a group of conservation concern in the United States and Mexico.

Mexican long-nosed bats, Leptonycteris nivalis, are considered rare but widespread and are primarily found in Mexico and a few summer roost sites in the southwestern United States

(Ammerman et al. 2012). Throughout this document I refer to the Mexican long nosed bat

(Leptonycteris nivalis) and the Lesser long-nosed bat (L. yerbabuenae) following the taxonomy of Simmons (2005).

Surveys in Mexico in the mid-1900s documented caves containing Mexican long- nosed bats in the thousands. Surveys repeated in the 1970s-80s indicated that these roosts no longer housed the large colonies once reported (USFWS 1994). The cause of the declines is unknown (Arroyo-Cabrales et al. 2008) but likely due to a combination of natural and human-mediated factors. These include human disturbances, colony movement between roosts, reduction of food sources as Agave was harvested for tequila production, taxonomic misassignment between Leptonycteris species at surveyed caves, an unknown disease, or survey timing and methods that missed the bulk of the seasonal population. These declines prompted action to provide the bats with protected status.

Conservation status

Mexican long-nosed bats were listed as federally endangered by the U.S. Fish and

Wildlife Service in the U.S. in 1988 and also by Mexico in 1991 (USFWS 1994). The recovery plan for this species anticipated that downlisting criteria would be met by 2014

(USFWS 1994). Unfortunately this goal has not been met, in part due to the lack of

______Journal of Mammalogy 1

information about the ecology and behavior of Mexican long-nosed bats to inform a comprehensive conservation plan. This further increases the need to understand this species.

Little was known about their movements and habits during early surveys (USFWS

1994). Subsequently, there have been few studies since they were assigned protected status that have increased our understanding of the natural history, biology, and migration of

Mexican long-nosed bats (Moreno-Valdez. et al. 2004; Sánchez and Medellín 2007;

Ammerman et al. 2009; England 2012). This is in part due to a low number of stable colonies to study, their seasonal nature, and unknown migratory routes.

Distribution

Mexican long-nosed bats are thought to be comprised of locally-migrating female

populations from southern Mexico and Guatemala north to southeastern Arizona, southern

New Mexico, and western Texas, United States, though there are no records from Arizona to

date (Simmons 2005). Some researchers suggest that Mexican long-nosed bats are capable of

a 1200km or more migration across their range via nectar corridors (Moreno-Valdez et al.

2000). In a recent study there was genetic evidence for a single Mexican long-nosed bat

population (Brown 2008). Males and likely some females stay in central Mexico year-round

(Segurajáuregui et al. 2006; Ammerman et al. 2012), and in their very northwestern and

southern range Mexican long-nosed bats are found in sympatry with Lesser long-nosed bats

(Hoyt et al. 1994).

In southern Mexico, Cueva del Diablo, Morelos, is the only documented site where

male and female Mexican long-nosed bats are known to gather and breed in winter (Sánchez

and Medellín 2007). Additional caves in Mexico are used by Mexican long-nosed bats, but

this species is considered scarce in its range in Mexico (Arita 1993; Arita and Santos-del-

Prado 1999).

In the United States they are found at few locations in arid regions where Agave

species bloom. They roost in colonies in higher-elevation caves and occasionally in buildings

(USFWS 1994). They only have been confirmed at a few localities in the United States;

Mount Emory Cave in Big Bend National Park in western Texas and the Animas Mountains in Hidalgo County, New Mexico (Hoyt et al. 1994).

Davis and Schmidt first reported Mexican long-nosed bats in Mount Emory Cave in the Chisos Mountains of Texas in 1937 (Borell and Bryant 1942) where they collected adult and juvenile females, juvenile males, and a ‘baby’. Gaps remain in our understanding of

Mexican long-nosed bat migration and how these bats use the caves that they are known to inhabit (Moreno-Valdez et al. 2000). According to Easterla (1972) historical survey estimates of the colony at Mount Emory Cave in Texas fluctuated widely from 0-10,000 bats from late

May-August. Other surveys also suggested fluctuating colony estimates based on multiple survey methods; visual estimates of ceiling counts of roosting Mexican long-nosed bats, guano splatter on the floor of the cave, or from surveys poorly timed with the bats’ migration

(USFWS 1994). Female Mexican long-nosed bats are thought to give birth to a single pup in spring (Davis 1974). It has been assumed that females and their volant young only migrate to and frequent Mount Emory Cave as a spill-over site when conditions permit (USFWS 1994).

Mount Emory Cave is likely used annually. A thermal imaging census was conducted to count the colony size upon emergence one night in July during 2005 and again annually from 2008 to 2015. Colony numbers from 2008 to present have ranged from 294 to 3238 with a mean of 2111 (Ammerman et al. 2009; Ammerman pers. comm.), indicating more

3 regular usage of this site than originally suggested. In addition to a mid-summer colony count, monitoring the arrival and departure of the annual Mexican long-nosed bat colony at

Mount Emory Cave would provide insight into their movements and use of the cave. The earliest northern migration records for Lesser long-nosed bats at a transient roost in the coastal town of Bahia de Kino, Mexico is 26 March (Fleming 2004), located at a similar latitude to Big Bend National Park. There are records of pregnant females in Big Bend

National Park in late April which is generally a few weeks before their Agave food source blooms locally (Higginbotham and Ammerman 2002; Brown 2008; this study). Therefore, this location might be used by Mexican long-nosed bats earlier in the season than originally thought.

Monitoring with technology

Bats are challenging to study. They are small, nocturnal, inconspicuous, and evasive.

Any useful monitoring technology must be small enough to be carried by a bat, and the data must be retrieved through signal detection or data download. Bat activity studies have utilized radio transmitters to monitor short-term (<21 day) home range locations (England

2012, Neubaum et al. 2005). Others have used marking methods to produce a recapture dataset at roosts to examine colony dynamics, survival, and recapture probabilities over multiple years (Lettink and Armstrong 2003; Pryde 2003; O’Shea et al. 2010).

Technological improvements in wildlife monitoring equipment have allowed researchers to gain access to the biology of species like never before, which is especially useful for studying threatened or endangered species. Passive Integrated Transponder (PIT) tagging systems were initially developed and applied in fisheries biology as a permanent, individual marking system for monitoring fish passage and has expanded to be used in many

taxa (reviewed in Gibbons and Andrews 2004). PIT tags are small, lightweight inert glass

capsules that contain an individual identification code that is recorded when a tagged animal

passes within the detection range of a PIT tag receiver and antenna. The capsule does not

contain a battery and is minimally invasive to insert under the animal’s skin, making it a safe

method to mark and “reencounter” an animal without the need to recapture and handle

individuals to read external identification tags.

This technology has been widely applied to other taxa than fish, and bat biologists

have used these tags as a biologically-appropriate tool to identify and monitor bats at roosts with the added benefit of avoiding band injuries or tag loss. The study species have been most often vespertilionids at roosts (Neubaum et al. 2005; Wimsatt et al. 2005; Britzke et al.

2014), hibernacula (Johnson et al. 2012; Ingalls 2014) or over water (Adams and Hayes

2008), but at least one study targeted phyllostomids (Lang et al. 2005) with success. PIT tagging studies have demonstrated little-to-no harm to bats, and seem to be most useful for gathering long-term data sets at roosts to study bat emergence order, survivorship, and body condition of tagged individuals recaptured over time (Neubaum et al. 2005; O’Shea et al.

2010; Gillam et al. 2011). A pilot study of Lesser long-nosed bats explored this technology to overcome band and tag injuries while monitoring movement away from maternity roosts

(USFWS 2005), but beyond that no data involving PIT tags in Leptonycteris species have been published.

The limited detection range of PIT tag antennas determines where this technology can be applied to passively scan for them in the environment. Typically antennas have been installed where bats exit their roost from a small hole or where they drink over shallow pools

(Adams and Hayes 2008; Johnson et al. 2012). A close size match between antenna style,

size and roost exit diameters have been a limiting factor to the application of this technology

at bat colonies, in order to successfully prevent bats from navigating around the antenna upon

exit and avoiding the detection range. Many bat colonies live in roosts like Mount Emory

Cave with larger entrances than can be adequately monitored by fly-through hoop-style

commercial PIT tag antenna systems.

Study objectives

Long-term studies are necessary to understand population trends in bats but short-

term datasets can aid in gathering information on the sampling feasibility, potential for

gathering useable survivorship parameters, and maintaining long-term monitoring efforts

(O’Donnell 2009). The purpose of this study was to use PIT tag technology to 1) monitor

Mexican long-nosed bats at Mount Emory Cave to detect seasonality as they transitioned

between Mexico and the United States, 2) to estimate apparent survival of marked bats and 3)

examine nightly activity of marked bats. Two of the recovery tasks within the USFWS

Leptonycteris nivalis recovery plan (USFWS 1994) that guided this study were:

1. Monitor known occupied and unoccupied roost sites (1.21, 1.22)

2. Obtain demographic data and determine and monitor migration

times, routes and habitats (4.1, 4.2).

Mount Emory Cave is likely used more extensively during the year than suggested by

historical, summer observations during June, July and August (Easterla 1972; USFWS 1994).

Part of the reason cave use has been difficult to verify is the remote location of the site and

lack of access to all the roosting areas within the cave for a comprehensive survey

(Ammerman et al. 2009). Mexican long-nosed bats migrate to Big Bend National Park from

Mexico, but our understanding of the population structure, migratory stop-overs, and lack of

marking studies have thus-far prevented researchers from tracing the extent of migration and

seasonality throughout the range of Mexican long-nosed bats.

I hypothesized that PIT tag data could be used to monitor the colony at Mount Emory

Cave and generate estimates of apparent survival of bats within the colony over time.In addition to estimating apparent survival, I hypothesized that Mexican long-nosed bat activity

(as measured via PIT tag records) within a season would vary by sex and age. Some bat species can move between resource areas with young and are not dependent on a roost site

while others, like Leptonycteris species, use a central roost and commute to foraging areas

making them ‘central place foragers’ (Ober et al. 2005; Rainho and Palmeirim 2011).

Leptonycteris species have been shown to commute approximately 20-30 km to foraging

areas (Ober et al. 2005; England 2012). Historically the understanding of this colony’s

dynamics are that the adults arrive at the cave mid-summer with volant juveniles (Easterla

1972; USFWS 1994) and if this is the case, Mexican long-nosed bats would not be expected

to return frequently to the cave throughout the night to feed non-volant juveniles. It is

possible that few pregnant bats arrive in spring and give birth in the cave (Brown 2008), which would then require repeated nightly visits until the young were volant. If some adult

females are supporting non-volant juveniles within Mount Emory Cave, then I expected PIT

tag detections to indicate activity differences that are specific to their reproductive condition.

METHODS Study site

Big Bend National Park is located in the Chihuahuan Desert ecoregion and the cave is located at a high elevation (above 2130m) surrounded by pine-oak woodlands (Easterla

1973a, b; Poulos and Camp 2010). Mount Emory Cave is composed of three rooms with crevices, hallways and chambers, most of which are inaccessible without technical climbing equipment. Mexican long-nosed bats roost with small numbers of Myotis thysanodes and

Corynorhinus townsendii in the cave (Brown 2008), and the site is used by occasional

Antrozous pallidus, Eptesicus fuscus, Myotis ciliolabrum, and Myotis volans (pers. obs).

Temperature recordings from summer 2005 in the main room of this cold-air cave ranged from 15.2º - 18.3º C (Ammerman et al. 2009) and a hop-hornbeam tree (Ostrya chisosensis)

obstructs much of the main cave exit. A cool breeze consistently moves through the cave indicating other openings; one is a known skylight into the cave, though bats are not known

to consistently use any other exit from the cave. The flyway between the cave entrance and

the inner rooms is made of an irregular quadrilateral 1.1 m wide at the bottom, 2.1 m wide at

the top and 2.6 m high.

In addition to the work at Mount Emory Cave I was involved with a PIT tagging

effort at Cueva Del Diablo, Tepoztlán, Morelos, Mexico in November 2013. It is a warm, deep cave with multiple exits where Leptonycteris nivalis, Desmodus rotundus, Pteronotus parnellii, and other species roost. Cueva del Diablo is approximately 1200 km from Mount

Emory Cave, and this site is further described by Segurajáuregui et al. (2006). It is unknown if individuals make a migration that extends between these two sites, or if other mating caves occur in Mexico.

Capture and marking techniques

To detect the activity of Mexican long-nosed bats, individuals were captured via harp

trap or mist net near the cave entrance and then marked and released for this study. Bats were

captured in 2013 at Cueva del Diablo with collaborators from the University Nacional

Autónoma de México. In 2014 I targeted marking 250 Mexican long-nosed bats at Mount

Emory Cave over 13 nights between May-August, and two nights in June 2015 when adult females were lactating and juveniles were volant. Standard measurements were collected from each bat: age, sex, reproductive condition, weight (g), and forearm (mm) (Kunz et al.

2009). Juveniles were limited to young-of-the-year based on the epiphyseal gap when transilluminated and gray coloration, instead of the gray/brown/yellow pelage of adults

(Ammerman et al. 2012). Without knowing their age upon initial marking, non-reproductive, non-young-of-the-year females were classified as adults in this study.

Captured bats were hand-scanned (Biomark 601, Biomark, Boise, Idaho) for an existing PIT tag identification code from Cueva del Diablo, Mexico or previous tagging efforts at Mount Emory Cave. When no PIT tag identification code was detected, a new PIT tag (Biomark, 12 mm) was implanted subcutaneously in the lower lumbar region of each bat.

I used standard PIT tag implantation methods and applied surgical glue to close the implant site and minimize the risk of tag expulsion (Wimsatt et al. 2005). This size tag has been used successfully in species half this size with only an approximate 10% tag loss rate (Johnson et al. 2012; Rigby et al. 2012). After processing a maximum of 50 individuals per night as per the permit conditions, and prior to release, bats were offered a dilute sugar-water solution (1 sugar: 4 water) with a disposable pipette. White-nose syndrome has not yet been found in

Texas but equipment decontamination actions were performed as a precautionary measure

(USFWS 2012). All animal handling protocols were consistent with guidelines published by

the American Society of Mammalogists (Sikes et al. 2011), and were approved by the

IACUC committees of Angelo State University (#1310) and the National Park Service

(#IMR_BIBE_Ammerman_Mexican long-nosed Bat_2014.A2).

Remote PIT tag detection system

To detect PIT tags I used a passive antenna and transponder system, the Biomark

IS1001, with a 15.2m flexible PIT tag antenna cable capable of sensing a PIT tag at any point along the cable length. This system was originally designed to be used in an aquatic environment such as a stream bed, and its flexible design could be manipulated to provide customized coverage for the remote sampling area. Successful remote monitoring depended on the presence of tagged bats and the distance between tagged bats and the antenna, making the flexible antenna option desirable for use at Mount Emory Cave.

I selected the portable enclosure option to weatherproof and house the Biomark

IS1001. I transported equipment to the cave with the help of volunteers and a pack mule.

There I installed the cable antenna at a doorway between the cave entrance area and the first room of Mount Emory Cave. I hung the flexible cable antenna from scaffolding made of polyvinyl chloride pipe (PVC) using a combination of nylon parachute cord and outdoor zip ties because metal was known to interfere with antenna performance per the manufacturer’s

suggestion. The horizontal crossbars of the PVC scaffolding were covered with flexible

fiberglass screen (2 mm mesh) to smooth its appearance and prevent bats from exiting around

portions of the PVC scaffolding that produced a gap near the cave ceiling. The system was powered by a 120 watt portable solar array (ZS-120-P; ZAMP Solar, Bend, Oregon) with built-in charge regulator, and two 12 volt batteries connected in parallel that were housed in a

lockable water resistant storage container hidden outside the cave entrance. Both ends of the antenna cable attached to the Biomark IS1001 enclosure to properly operate and emit the 134 kHz signal used to detect PIT tags. The antenna produced a detection range of approximately

15 cm. The detection range could be increased or decreased according to the orientation and proximity of the cable to itself. When cable sections are close together the detection ranges

can be configured to cover the combined area between them. The antenna was auto-tuned and tested with a Biomark test-tag to determine maximum possible detection range and best flyway coverage.

From 26 April to 14 June 2014 I set the flexible antenna in a spiral as recommended by the manufacturer and similar to an ongoing project with the same system in Vermont

(Ingalls 2014). This created approximately two, 1 m diameter loops in the upper half of the

flyway where bats are known to exit the cave. Per the manufacturer’s suggestion I lashed the

first 30 cm lengths of both ends of the antenna 15 cm apart starting at their connection with

the Biomark IS1001 to promote proper antenna tuning. To attempt a more powerful detection

coverage area, I doubled the antenna once (2 x 7.62 m sections) and wrapped it in a single

ring without spiraling it as it hung from the PVC scaffolding. This configuration had strong

detection range but left a small coverage gap over the center of the flyway where most bats

tend to exit. The antenna was removed from 14 June to 3 July 2014 to minimize obstruction

while Mexican long-nosed bats returned for the season and while I developed a final

configuration and settings appropriate for the site (Adams and Ammerman, in press;

Appendix 1).

An annual census of the Mexican long-nosed bat colony at Mount Emory Cave was conducted on 1 July 2014 with thermal imaging video equipment (FLIR SC660; FLIR

Systems, Boston, MA), and after confirming typical colony activity and numbers at the cave with the scaffolding in place (Ammerman, pers. comm.), I reinstalled the PIT tag antenna on

3 July 2014. I used a serpentine pattern to create a detection-range plane over the doorway

(Figure 1). I modified the fly-through configuration and hung two serpentine sections in succession 25-30 cm apart to cover the width and draped 2.1 m in length to cover the entire cave doorway in PIT tag detection coverage. Excess cable antenna was draped on the cave floor, but if the entryway had been wider or longer the excess could have covered an additional serpentine section of the flyway. The serpentine antenna used at Mount Emory

Cave produced a curtain of PIT tag detection coverage that encompassed almost the entire doorway and substantially increased the zone of detection compared to the previous looped configurations. I covered all the equipment with 3 mil contractor plastic based on comments by Ingalls (2014, pers. comm.) to reduce low levels of ultrasonic noise produced by the equipment and solar panel electronics, but also to deter gnawing from the occasional curious animal in the cave.

After reinstalling the antenna and using the serpentine antenna configuration on 3

July, at emergence I verified Mexican long-nosed bat activity and ability to fly through the antenna by collecting a brief recording of the emergence with the thermal imaging video equipment. I observed another emergence on 23 July using an infrared night-vision scope to verify continued emergence activity through the serpentine antenna. The antenna recorded bat emergences successfully throughout the season and continued to monitor for activity until the battery strength fell below the threshold embedded in the IS1001 software and the system went on standby in mid-October.

FIG. 1.— A sketch of the serpentine antenna configuration used during this study to detect PIT tags of Mexican long-nosed bats tagged at Mount Emory Cave. Antenna cable were within 2.5-5 cm from the surrounding rock walls. Dimensions in text. (Artist: Terry Maxwell).

I suspect this power-down could have been avoided or delayed if a rock had not fallen on and broken one half of the surface of the solar panel. Once the system was on standby, the remaining solar panel section was able to recharge the battery completely until the following season. The antenna was moved to the side of the entrance over the winter to accommodate a cave-mapping group and reinstalled in March 2015 for the season. At this time a second flexible 120 watt solar panel and RA-9 charge controller (PowerFilm Inc., Ames, IA), were added to the power system to compensate for damages to the original solar panel. The system registered feedback from the charge controller in the IS1001 system during the times of day when it experienced the most direct sunlight, but the feedback did not impact the performance of the Biomark system in use at night.

Seasonality

All analyses of PIT tag detections were limited to the bats tagged at the Mount Emory

Cave study site. Any tags detected from Cueva del Diablo were primarily intended to provide seasonal and migratory information. Using the entire set of detection records from May 2014

– August 2015 I calculated the number of days bats were detected by the PIT tagging system or captured at the cave to examine the seasonal activity at the site and report the demographics of bats redetected after initial tag insertion.

I examined the average number of nights bats were detected during the study using an analysis of variance test (ANOVA with Tukey’s test) and verified statistical significance with a randomization test. I generated a null distribution from the data since it likely violated normality assumptions, and used the null distribution from the data to detect differences between the average number of nights detected by age, sex, and further examined by group

(adult female, juvenile male, juvenile female).

I calculated the average maximum duration of activity between emergence and return

for individuals that triggered the PIT tag system at emergence prior to 23:30, or after

03:30am within the same night. Three bats that were tagged as juveniles in 2014 triggered the

system the second year, their 2015 records were excluded from the analysis because their

reproductive condition was unknown in 2015. I used a Linear Mixed Effects Model in the

statistics program R (R Core Team 2014) with the package lme4 (Bates et al. 2015) to

perform a regression analysis of all the means of the nights across groups, accounting for the

repeated measures of PIT tagged individuals across multiple nights. I generated a full and

alternate model, created a likelihood ratio, and compared the results to a null distribution of

the likelihood ratio from a randomized dataset, and additionally I tested subsets of the reproductive groups for significant differences in mean activity durations.

PIT tag record analysis

In order to test if survival parameters could be estimated for Mexican long-nosed bats in this colony in this study (and therefore over time), I used a Cormack-Jolly-Seber (CJS) live recapture analysis in the program MARK (White and Burnham 1999) to calculate

‘apparent survival’ (ф) and capture (reencounter) probability (p) estimates (Lettink and

Armstrong 2003; Pryde 2003). Apparent survival is the probability a bat will survive from one time interval to the next and remain in the study population. It often underestimates true survival (O’Donnell 2009) because emigration and death are indistinguishable. Note that the results of this analysis are not estimating true survival, since the analysis cannot distinguish between bats exiting the roost without detection, migration from the site, use of a different roost, or death (Neubaum et al. 2005; O’Shea et al. 2010). The recapture probability is how likely a bat in the study will be redetected per sampling period.

Raw redetection data were converted into an encounter history matrix to be analyzed in program MARK. In this encounter history matrix, each individual was recorded as present or absent (1 or 0). Realistic encounter history intervals are tailored to individual research objectives. I used a 2-week interval to estimate the ‘apparent survival’ of adults and juveniles in this study, but other studies often use monthly or annual intervals. My first interval began at the first tagging event in May 2014, and continued for 13 intervals through August 2015, with interval 8 spanning the final 2 week period in 2014 and also the winter migration period.

Continuous PIT tag detection data violates one of the CJS assumptions of instantaneous sample collection (Barbour et al. 2012) where both marked and unmarked individuals are collected during capture events. However, the small sample size and the short duration of this study (2 seasons) prevented the use of more sensitive analyses for continuous PIT tagging data, such as the Barker (Barbour et al. 2013) or Multi-state models which account for tagged juveniles aging in the population (Ellison et al. 2007).

I pooled the age and demographic information for individuals to create group variables for adult female, juvenile female and juvenile male only. Because the single adult male tagged in this study was not re-detected, adult males were excluded from the analysis. I held the capture probability (p) constant over time and group (adult female, juvenile female, juvenile male) as I did not expect any biological reason to assume an age or sex difference as the bats flew through the antenna and were detected. Apparent survival (ф) was held constant and also varied by time, group, and age to test four different apparent survival models as follows: ф(.)p(.), ф(groups by age and sex)p(.), ф (age)p(.) and ф(sex)p(.). To select the best model from those tested, I used Akaike’s Information Criterion (AIC) values and to correct

for small sample sizes I considered the model with the highest AICc weight as the best model

(Burnham and Anderson 2002).

Nightly activity

I examined hourly bat activity in July of both years. I binned the PIT tag records into

15 minute increments per hour after sunset and tested if detections per hour were significant.

I tested the activity of individuals by age, sex and hour after sunset using mixed effects

models (Bates et al. 2015). The data likely violated normality assumptions so I used a

randomized likelihood ratio test to simplify my models and identify significant groups. I

generated null distributions from the data in order to run a mixed effects regression. I used a

generalized mixed effects logistic model (glmer, binomial) to examine if individual activity was significant by hour, and a linear mixed effects model (lmer) to test if age, sex, and hour had an effect on total activity. Where appropriate, significance levels (p values) are expressed as <0.05 or lower, and where randomization tests generated a null distribution from the data, no degrees of freedom were reported.

RESULTS

At Cueva del Diablo in 2013, collaborators and I PIT tagged 100 female Mexican long-nosed bats. Over two seasons at Mount Emory Cave (2014-2015) I captured and marked an additional 104 individual Mexican long-nosed bats comprised of 47% adults and 53% juveniles (Table 1, Appendix 2). The PIT tagging system in the serpentine configuration was successful at detecting bats at Mount Emory Cave, however none of the Mexican long-nosed bats tagged at Cueva del Diablo were detected at Mount Emory Cave. Once the antenna was hung from the PVC scaffolding in the serpentine style on 3 July 2014, I noted that the colony emerged as anticipated through the serpentine configuration, instead of avoiding the non- serpentine clutter as observed at emergence earlier in summer 2014. While recording the 3

July 2014 emergence video I could hear what sounded like a few bats experience wing- strikes against the plastic sleeve on the antenna in addition to the typical wing-strikes at a hop-hornbeam tree blocking much of the cave entrance. I noted from approximately 20 minutes of video footage collected during the peak of emergence that some bats landed briefly near the antenna as if to inspect it. A subsequent observation of the emergence on 23

July 2014 showed an emergence pattern that more closely resembled census recordings prior to the antenna installation in both number of bats using the exit and behaviors (Ammerman et al. 2009). I did not note any bats displaying the investigative landing behaviors while watching the second emergence. The wing-strike sound was mostly eliminated from the antenna area, though it could still be heard near the tree as bats brushed the leaves as they flew by. The vespertilionids in the cave did not demonstrate behaviors indicating that they were avoiding the antenna (Britzke et al., 2014) nor did they abandon the roost when the

18 antenna was in any configuration during this study. They were regularly captured in the harp trap throughout all trapping efforts.

TABLE 1.— Demographics of Mexican long-nosed bats (Leptonycteris nivalis) captured and implanted with PIT tags at Mount Emory Cave in 2014 and 2015. Number of reproductive females indicated in parenthesis. Age Sex 2014 2015 Total

Adult Female 18 (15) 30 (20) 48 Male 1 0 1

Juvenile Female 8 16 24 Male 11 20 31

Total 38 66 104

Seasonality

Mexican long-nosed bats were present at Mount Emory Cave in 2014 from at least 26

April – 1 September. Noteworthy captures include pregnant Mexican long-nosed bats on 26

April and 19 May (n=4), an adult male on 29 May 2014, and lactating females between 29

May and 3 July 2014. PIT tag records documented bats present from 8 May – 1 September

2014. However, bats were only detected from 16 June – 19 August the subsequent year based

on PIT tag records.

PIT tag record analysis

I detected 79 individuals (out of a total of 104 marked) based on PIT tag records

during this study. Twenty-five PIT tagged bats were not detected during this study after

initial tagging; 29% of bats tagged in 2014, and 21% of the 2015 bats. Twelve of the 38 bat

tagged in 2014 were redetected (Figure 2) for a first-year return rate of 32%, however when

examined by age and sex 42% of adult females, 50% of juvenile females, and 8% of juvenile

males returned to Mount Emory Cave the year after they were PIT tagged. Some of these

first-year redetections included individuals not detected after tagging in 2014. None of the

bats captured and PIT tagged throughout this study were subsequently recaptured, only

redetected via the PIT tag system.

Fig. 2.— Yearly totals of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) at Mount Emory Cave including 12 redetections. Eight individuals were detected in both years and 4 were tagged in 2014 but only detected in 2015.

Not all bats were detected on all nights (Figure 3). The antenna was able to detect and monitor the passing of individual Mexican long-nosed bats through the cave entrance for 99 total nights over both seasons. Examining a subset of dates where activity was detected by the serpentine antenna (3 July to 18 August of both years) the highest percentage of PIT tagged bats, 31% and 41%, were detected on 13 July (Figure 4), and detections dropped gradually through the summer.

More individuals were detected longer in the season in 2014 than 2015. Some late season residents of the cave were tagged in 2014 but not in 2015, which may have influenced the late season detection of those bats. The longest number of total nights any Mexican long- nosed bat detected was a juvenile male over 39 nights (Figure 5). Of the top 10 most detected bats, 9 were 2015 juveniles and one adult originally tagged in 2014, and the top four most detected juvenile were males. Bats were detected for an average of 13.9 ±10.3 days (mean ±

SD; range 1-39), but adults (9.8 ±9.2 days) were detected on average for fewer nights than juveniles (16.6 ±10.2 days). Mexican long-nosed bats and were detected more frequently during morning return to the cave than at emergence (Figure 6). Juvenile males were detected for significantly more nights than adult females (p<0.02).

FIG. 3.— All days of detection of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) in 2014 and 2015 at Mount Emory Cave. Light gray rows represent individuals PIT tagged as adults, while black rows represent juveniles. The upper horizontal group were records in 2014, and the two lower horizontal blocks were redetections and records in 2015. Not all individuals are detected on all nights, some were only detected once.

FIG. 4.— Percentage of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) detected at Mount Emory Cave in 2014 (n= 38) and 2015 (n=104).

FIG. 5.— Total nights each PIT tagged Mexican long-nosed bat (Leptonycteris nivalis) was detected at Mount Emory Cave in 2014 and 2015. Results grouped by age: Juveniles and Adults.

Preliminary apparent survival estimates

The PIT tagging records analyzed in MARK generated two models that best represented the apparent survival and probability of redetection for the study overall, and for different age groups. The highest-ranking model (AICc weight) for apparent survival in this study was ф(age)p(.), representing a variable age structure with time and probability of redetection both constrained to constant (Table 2). Juvenile apparent survival 91.86% (95%

Confidence Interval: 86.15-93.35) was slightly higher than adult apparent survival 89.07%

(95% CI: 76.74-90.76) in that model (Table 3), but these results are similar because of the overlapping confidence intervals. No adult males were included in these calculations because only one adult male was tagged in 2014 and he was never redetected. For constant parameters of time, age, and probability of redetection, the apparent survival of bats in this study was 89.31% (95% CI 84.23-92.89) and probability of redetection was 38.9% (95% CI:

31.70-46.74).

Nightly activity

On nights when bats were detected, some triggered both emergence and return records.

However, many did not, greatly reducing the full nights available for activity analysis or determining the frequency of entries and exits throughout the night. Of the 79 individuals detected, 66% (n=52) triggered the system to generate a maximum nightly activity duration from emergence to return one or more times, while 34% of bats (n=27) only triggered emergence or return detections to verify presence, but not an activity record. Mexican long- nosed bats were active an average of 7 h 42 min per night (range: 4 h 05 min – 9 h 09 min) based on the first and last time-stamped activity records over 321 activity-nights for the total

TABLE 2.— Results from program MARK for modeling the apparent survival and capture probabilities of PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) at Mount Emory Cave during the 2014-2015 study period over two week intervals across the study.

Model Parameters AICc ∆AICc AICc weight

ф(age)p(.) 3 532.69 0.00 0.46 ф(.)p(.) 2 533.85 1.16 0.26 ф(group)p(.) 4 534.59 1.89 0.18 ф(sex)p(.) 3 535.50 2.81 0.11

TABLE 3.— Top two model estimate results for apparent survival and probability of redetection for PIT tagged Mexican long-nosed bats (Leptonycteris nivalis) at Mount Emory Cave during the 2014-2015 study period

Model Group Estimate 95% CI

ф(age)p(.) Adult ф 85.07 76.74-90.76 Juvenile ф 91.86 86.15-93.35 p(.) 39.81 32.51-47.60

ф(.)p(.) ф(.) 89.31 84.23-92.89 p(.) 38.96 31.70-46.74

29 study (Table 4). When examined by age and reproductive group, differences were detected in the average activity per night. The linear mixed effects model regression suggested that lactating adult females were active for shorter durations on average than non-reproductive adults and juveniles (p<0.01), and there was no significant difference between adult and juvenile non-reproductive bats.

More morning detections occurred than night (emergence) detections. A generalized linear mixed effects model suggested that hour was a significant predictor of a bat’s presence

(p<0.005), and a linear mixed effects model demonstrated that individuals were detected more frequently during the morning (return) hours (p<0.005; Figure 6).

TABLE 4.— The average duration of activity of Mexican long-nosed bats (Leptonycteris nivalis) per age and reproductive group based on the time between maximum emergence and return triggers in the same night, generated from nights in 2014 and 2015.

Reproductive Number Range of nights Total nights Average time active Condition of bats detected detected (minutes)

Adult Female lactating 7 1-4 28 410.0 post-lactating 1 5 5 371.0 non reproductive 4 2-20 29 467.3

Juvenile female 19 1-23 120 461.5 Juvenile male 20 1-21 132 476.9

FIG. 6.— Detections of Mexican long-nosed bat (Leptonycteris nivalis) activity per hour after sunset at Mount Emory Cave in July 2014 and 2015.

DISCUSSION

I hypothesized that PIT tag data could be used to monitor the colony at Mount Emory

Cave and generate estimates of apparent survival of bats within the colony over time. I

successfully applied PIT tagging technology to monitor tagged Mexican long-nosed bats in

2014 and 2015 at Mount Emory Cave. The mark-recapture analyses showed promise for a continued study to monitor Mexican long-nosed bats at Mount Emory Cave. Within the season I found that individuals were detectable for up to 39 nights, and across seasons I found that some bats were detectable from year to year. Others were not detected after initial tagging, which was not surprising considering the migratory nature of this species. It is possible that these tagged bats will be redetected in future seasons. Adult females appeared to depart the cave earlier in the season than juveniles.

Seasonality

This study, based on combined results from capture efforts and PIT tag detections, documented an extension of the dates bats use Mount Emory Cave, and included pregnant females which had only been captured in April at another site in Big Bend National Park,

Glenn Springs (Higginbotham and Ammerman 2002). The records of pregnant Mexican long-nosed bats roosting in the cave on 26 April 2014 confirmed the earliest reproductive females roosting at this site as suggested by Brown (2008). In May 2014 an additional pregnant female was captured, and lactating females were captured in June and July of both years of this study. Easterla (1973) did not capture any pregnant females at the cave nor in the park during his surveys in April, June, July or August 1967-1971.

This study confirmed the presence of individuals before and after the local blooming

period for Agave havardiana in the Chisos Mountains. These plants typically bloom in the park from mid-May through mid-August, though occasionally an early or late bloomer has been seen in the lowlands in April and the highlands in October (Wauer and Flemming

2002). The latest detection was in the early morning of 1 September 2014, eight days after the last PIT tag detection of the last previous bat, and approximately two weeks after the

Agave bloom had ended for the season in the Chisos Mountains (pers. obs.). It is possible that

Agave or other nectar sources were blooming within commuting distance of Mount Emory

Cave, or that individual bat record is atypical for this site. The winter diet of Mexican long- nosed bats near Cueva del Diablo consists of nectar and pollen from at least 21 plants from

10 families, such as Bombax, Ceiba, Ipomoea, Calliandra, Pinus, and grasses (Sánchez and

Medellín 2007, Arroyo-Cabrales et al. 2008). From April to October while Mexican long-

nosed bats are in their northern range, their diet has been tightly correlated to availability of

Agave (USFWS 1994; Moreno-Valdez et al. 2004). In Big Bend National Park, Agave

havardiana is the only known nectar source for Mexican long-nosed bats (Kuban et al. 1983;

Kuban 1989; USFWS 1994; England 2012) with speculation that they supplement their diet on the later-blooming A. chisoensis (Easterla 1972), insects or cactus fruits. No comprehensive diet study has verified if other diet items are part of the diet for this northern- most colony. If dependent on Agave for nectar and pollen as their sole food source in the region, Mexican long-nosed bats would have been expected to migrate west to feed on later-

blooming Agave in New Mexico (England 2012) or south to Mexico by mid-August.

PIT tag detection

Redetections of PIT tagged Mexican long-nosed bats were low across the extent of this study and some possible reasons for it included bat behavior, tag loss, or problems with the detection method. Some bats that were marked and never detected may have left the study area, as they are highly migratory and likely have other local roosts like other researchers have suggested and looked for but have not found yet (USFWS 1994, England

2012). It is possible that PIT tags were ejected through the injection site and lost prior to detection with the PIT tagging system. Rigby et al. (2012) reported up to a 10% tag loss in their PIT tagging study. Johnson et al. (2012) also reported tag loss and were able to confirm lost tags by scanning the floor of the roost for ejected tags. During this study, cave structure

and logistics prevented me from detecting any shed PIT tags to exclude from analysis or

generate a tag loss estimate (Johnson et al. 2012) because I could not access the parts of the

cave below where the bats roosted to scan for ejected PIT tags.

Apparent survival and probability of redetection estimates

The apparent survival over this study was higher than reported in other studies which report on annual apparent survival estimates from banding and PIT tagging studies

(Neubaum et al. 2005, O’Shea et al. 2010). The likelihood that bats were detected throughout this study after initial marking was a probability of redetection approximately 40%, though approximately 76% of individuals were redetected at least once during the study, mostly

within the season they were also initially marked. A third season of detections would have

helped refine the estimates (Neubaum et al. 2005) and enhance the apparent survival model

results (O’Donnell 2009). This study was not long enough in duration to provide the data

needed to generate a multi-state age model to address recruitment and survival over time.

Barker model analyses are better able to handle the continuous detection data, but estimates

are generated through evaluating a large number of parameters and the models are best when

applied to datasets gathered over multiple years (Barbour et al 2013).

I expected juvenile survival to be lower than adult survival (O’Shea et al. 2010), but

in this study that was not the case, though the 95% confidence intervals for both adult and

juvenile survival estimates largely overlapped. The sex and age structure of this colony may

present challenges with analyses until a larger sample size is achieved, especially considering

that adults seem to trigger the system for fewer days than juveniles do. This could be a

function of the adults knowing of alternate exits or roosts in the area compared to juveniles,

evading detection more often than juveniles, or that adults and volant juveniles migrate independently at the end of the season. The sample size and encounter history of this study

was too small with only two seasons to employ those analyses using age structure parameters. A study of at least 5-10 years would provide more useful survival estimates for this colony (O’Donnell 2009). The lack of adult males made it hard to support a detectable difference between adult and juvenile apparent survival at this time, though over multiple seasons these estimates have been generated for maternity colonies where bachelor males are not regularly found (O’Shea et al. 2010).

One previous study in the 1960s-70s marked 568 Mexican long-nosed bats at three sites around Big Bend National Park with tags with only a single tag recovery (Easterla

1973a, b). This PIT tagging study represents the first time a coordinated effort to mark bats in the United States and Mexico to potentially detect migratory movement between a known

mating site Cueva del Diablo and the northern maternity site, Mount Emory Cave. There are no other studies yet published generating apparent survival or detection probabilities for this genus with which to compare my results. I anticipated that only females would be redetected from year to year, as males are not known to make the same long-distance migration that females and volant juveniles do (Ammerman et al. 2012). The majority of PIT tagged bats from 2014 that returned in 2015 were female, but the return of one juvenile male could mean that males use the cave more regularly than expected, or that sub-adult Mexican long-nosed bats use the cave in addition to reproductive adult females and their young of the year.

Nightly activity

The major limitation to addressing the activity pattern hypotheses of this study were the low sample size of PIT tagged Mexican long-nosed bats in 2014, and the imperfect detection in both seasons. Initially the lack of detector read-range, flyway clutter, and few numbers of bats PIT tagged meant that data for early in 2014 were sparse. Despite the limitations I learned that the activity and survival of Mexican long-nosed bats was more dynamic than hypothesized by early researchers (Easterla 1972a, b, USFWS).

The records in this study indicate that some bats do return to the cave throughout the night, and the patterns of activity at the cave vary by individual. Without increasing the number of PIT tagged bats at nightly detections at Mount Emory Cave, PIT tagging may not

be the preferred method for studying activity at this site in future studies due to the imperfect

detection coverage. Imperfect detection during emergence and return could have been a

result of bats using alternate exits, flight pattern changes in response to the clutter of the

original antenna configurations in early 2014, or bats roosting in nearby unknown roosts on

some nights.

Another possibility for why individuals triggered the system during only some times

of the night may be related to behavioral differences in the flight pattern at emergence or return per night. More detections occurred during the morning, and some individuals were

only ever detected from morning records. As bats exit the cave and pass the antenna, they are

flying down a slope in the cave and continue out of the cave, and often they briefly circle outside the cave entrance with social chatter (pers. obs.). There may be an urgency to disperse and feed for the night that is not present during the morning returns after bats have

fed for the night. In addition, bats have to fly up-slope into the cave entrance which may require a slower approach and promote better detection by the antenna. Without further

research the exact cause for higher detectability in the morning compared to the evening

cannot be determined.

A concern of this study was that the fly-through antenna system could be a deterrent to Mexican long-nosed bats or other species in the cave and trigger roost abandonment as

observed from some cave gating projects at other sites (USFWS 2005). Some individuals

appeared to be undetectable or leave the study area after PIT tagging. Since the antenna was

arranged to hang through the flyway the bats may have viewed the antenna as an obstruction

similar to a cave gate (Spanjer and Fenton 2005). Little is known about how Leptonycteris react to cave gating. There are a few examples of Lesser long-nosed bat colonies in cave- gating situations, and they seem to have variable responses (USFWS 2005). In one case an experimental PVC gate had no effect while its steel replacement caused the colony to

abandon the roost. Other Lesser long-nosed bat colonies changed their response to the gate in relation to moon phases and predation risk, and others abandoned their roosts once caves

were gated (USFWS 2005).

In this study there was little lasting indication that the colony was impacted by the

antenna support system or the antenna itself. I based this conclusion on emergence video

monitoring and more importantly, the PIT tagging record data. The video and tag records in

2014 and 2015 suggest that activity was similar to annual census activity from previous years

(Ammerman, unpublished data).

Future directions

England (2012) studied the summer foraging of Mexican long-nosed bats in Big Bend and found that the foraging home ranges of juveniles were larger than that of adults, and noted that adult females foraged near Mount Emory Cave, while the adult males or juvenile males and females radio tracked for that study made excursions to areas where there was little or no Agave available for foraging. England’s (2012) results corroborate with the shorter average duration records for reproductive adult females compared to longer nights of activity on average for juvenile males and females seen in this study. The adult females are likely focusing their foraging on Agave rich areas and minimizing energy spent on commuting. The daily temporal patterns of some fruit eating Phyllostomus hastatus, suggest they spend at least three hours away from their cave roost per night (Kunz et al. 1998). A radio tracking study of leaf-roosting Rhinophylla pumilio showed that lactating females spent significantly more time in flight and were more active throughout the night than non- reproductive females (Henry and Kalko 2007). As detection methods at Mount Emory Cave

39 are refined, and more Mexican long-nosed bats are tagged with their initial reproductive status noted, differences in PIT tag activity records may be sensitive enough to indicate if and when females are non-reproductive, lactating, or post-lactating at this site.

I speculate that the probability of redetection for Mexican long-nosed bats with PIT tags could be related to their proximity the antenna, the sensitivity of the detection system to register a passing tag, and the flight speed of some individuals. Flight speed of bats could not be calculated by the number of PIT tag records per pass due to filter settings in the Biomark

IS1001 software. Assuming bats are not typically using alternate local roosts, this study suggests that speed or behavior at emergence reduces the detectability of bats as they emerge from the roost but return behaviors increase detection. However, if the detections recorded are an accurate representation of the nights that bats use the cave, it was surprising to see how much sooner the adults move from the site than juveniles did in both seasons, since they are assumed to migrate to the site together (USFWS 1994). In addition the bats may be using a local night roost while foraging, as suggested by England (2012) and others, but that site might also represent a day roost the bats use on the days they were unrecorded by the PIT tag antenna at Mount Emory Cave.

Biomark technology is ever-improving to address needs in fisheries research. For example, they now make a fast-flow PIT tag and antenna system designed to use with fish in swift river systems. I would recommend researchers in future studies consider the improvements and limitations available with this type of equipment for PIT tag monitoring efforts with this group of bats. Additionally, PIT tag detection system manufacturers do not anticipate that their equipment can detect the tag of other brands (Gibbons and Andrews

2004). If other biologists plan to scan for PIT tags and implant new tags in Leptonycteris of

any species, I would recommend using the same systems across all sites to increase the

chance of detecting bats tagged from other locations. I would also recommend that

researchers invest in a PIT tag hand-scanner and make scanning a part of their capture

protocol.

PIT tagging technology at Mount Emory Cave shows promise to detect the population

dynamics and survival of Mexican long-nosed bats at this northern maternity roost over time, but shows less promise at this site to detect activity patterns. Larger sample sizes of all available demographic groups using the cave (adult females, juvenile males and females) could increase the power of this analysis, and over time the detection probabilities could increase to levels useful for generating more refined population estimates.

The data from this study and the thermal imaging censuses indicate that the spill-over hypothesis for Mexican long-nosed bat presence in Big Bend National Park is ready to be refined. Mount Emory Cave is an important site populated by individuals who express a level of annual site fidelity, and their young. If Mexican long-nosed bats are PIT tagged at other locations in their range, Mount Emory Cave could prove to be a site from which migratory movements can be detected from arrivals of the bats tagged at other sites.

The rare and widespread distribution of Mexican long-nosed bats, along with their

endangered status means that continued monitoring at this site and at the few other studied

sites in their range is important to detect any long-term population trends. With the

increasing interest in the study of Leptonycteris and a developing network of researchers

using PIT tagging technology, there is a possibility for collaboration to study movement

41 between monitored roosts over time. Decreases in the population are likely to occur faster than increases based on impacts this species faces compared to slow reproductive rates.

Monitoring increases or decreases in population numbers of Mexican long-nosed bats over time will be an integral part of conservation efforts for this species in the United States and

Mexico.

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APPENDIX 1

Example of Full Status Report (RFS command) for Biomark IS1001 with data logger board systems and settings used at Mount Emory Cave in 2014-2015.

INF: Start of Full Status Report Reader: Reader ID: 01 Reader SN: 1347.0023 Reader Date: 06/21/2015 Reader Time: 22:45:18 Application Version: IS1001-12V 2.4.5 Reader Network Mode: IS1001 Standalone Operation Mode: Scan Sync. Mode: Standalone Reader Beeper: Disabled Tag Display Format: DEC Reader Initiation Delay: Disabled Alarms: Antenna Current Low Alarm: 1.0 A Noise High Alarm: 80% Tuning Capacit. High Alarm: 970 Tuning Capacit. Low Alarm: 50 Alarms Unique Delay: 60 sec Antenna/Tuning: Tuning Capacitors: 157 Dynamic Tuning: Enabled Target Phase: 391 Phase Deviation Threshold: 6 Measurements: Antenna Current Gain: 120 Antenna Current Offset: 110 Communication: Local Port Speed: 115200 Tags To Local Port: Enabled Alarms To Local Port: Enabled Messages To Local Port: Enabled Remote Port Protocol: ASCII Detection: HDX Tag Detection: Disabled Detection Counter Enabled: Yes Detection Counter: 85 Unique Mode: Delay Unique Delay: 20 sec

FDXB Detection Scan Time: 120 ms VTT Level: 128 Auto VTT Delay: 360 min Memory: Tags Memory Size: 100015 Tags In Memory: 145 (0%) Status Reports Memory Size: 1023 Status Reports In Memory: 177 (17%) Save Tags To Memory: Enabled Save VTT To Memory: Enabled Save Stat. Reports To Memory: Enabled Reports: Auto Noise Report Delay: 240 min Auto Status Report Delay: 240 min Sensors: Input Voltage: 12.4 V Exciter Voltage: 10.0 V Antenna Current: 3.6 A FDXB Signal Level: 84 mV (9%) Antenna Phase: 395 Relative Phase: -4 Sync. Input: N/A Temperature: 22.2 C Active Alarms: No Active Alarms

APPENDIX 2

Mexican long-nosed bats (Leptonycteris nivalis) captured via harp trap and subcutaneously marked with 12mm Biomark PIT tags for identification for remote detection at a fly-through serpentine antenna (Biomark IS1001) located at Mount Emory Cave. Reproductive status Preg=pregnant, NR=non-reproductive, Lact=lactating, Post lact=post lactating.

Date Time Weight Reproductive (m/dd/yyy) (12h) (g) Age Sex Status Forearm (mm) Tag Number 4/26/2014 9:30 pm 36 Adult Female Preg 57.5 released no tag 4/26/2014 9:30 pm 35 Adult Female Preg 55 released no tag 4/26/2014 9:30 pm 34 Adult Female Preg 56 released no tag 5/19/2014 5:53 am 34 Adult Female Unknown 57.5 PIT tag 989.001003147006 5/19/2014 6:13 am 36 Adult Female Preg 59 PIT tag 989.001003147033 5/19/2014 4:55 am 27 Adult Female Unknown 57.6 PIT tag 989.001003147040

50 5/29/2014 4:15 am 37 Adult Male NR 57.1 PIT tag 989.001003147013 5/29/2014 4:55 am 35 Adult Female Lact 56 PIT tag 989.001003147062 5/29/2014 5:35 am 36 Adult Female Lact 57.3 PIT tag 989.001003147082 6/08/2014 9:29 pm 30 Adult Female Lact 59.7 PIT tag 989.001003147051 6/08/2014 9:37 pm 27 Adult Female NR 56.2 PIT tag 989.001003147063 7/01/2014 9:30 pm 33 Adult Female Lact 56.7 PIT tag 989.001003147007 7/02/2014 NA NA Juvenile Male NR NA PIT tag 989.001003147042 7/03/2014 9:50 pm 29 Adult Female Lact 60 PIT tag 989.001003147011 7/03/2014 10:10 pm 30 Juvenile Female NR 57.1 PIT tag 989.001003147019 7/03/2014 9:45 pm 34 Adult Female Lact 57.5 PIT tag 989.001003147023 7/03/2014 9:45 pm 30 Juvenile Male NR 59.7 PIT tag 989.001003147060 7/03/2014 9:45 pm 29 Juvenile Male NR 58.5 PIT tag 989.001003147067 7/03/2014 9:45 pm 32 Adult Female NR 59 PIT tag 989.001003147078

Date Time Weight Reproductive (m/dd/yyy) (12h) (g) Age Sex Status Forearm (mm) Tag Number 7/03/2014 10:10 pm 29 Adult Female Lact 57.2 PIT tag 989.001003147086 7/13/2014 NA 28 Adult Female Post lact 57.3 PIT tag 989.001003147022 7/13/2014 NA 29 Juvenile Male NR 61.1 PIT tag 989.001003147029 7/13/2014 NA 32 Juvenile Female NR 57.5 PIT tag 989.001003147037 7/13/2014 NA 24 Juvenile Male NR 57.7 PIT tag 989.001003147043 7/13/2014 NA 31 Juvenile Male NR 55.7 PIT tag 989.001003147045 7/13/2014 NA 29 Adult Female NR 58.4 PIT tag 989.001003147046 7/13/2014 NA NA Juvenile Female NR 56.4 PIT tag 989.001003147054 7/13/2014 NA NA Adult Female Post lact 58 PIT tag 989.001003147059 7/13/2014 NA 31 Juvenile Male NR 59.5 PIT tag 989.001003147061 7/13/2014 NA NA Juvenile Female NR 59.1 PIT tag 989.001003147069

51 7/13/2014 NA 29 Juvenile Female NR 60.1 PIT tag 989.001003147072

7/13/2014 NA 32 Adult Female Post lact 58.4 PIT tag 989.001003147076 7/13/2014 NA 30 Juvenile Female NR 57.7 PIT tag 989.001003147090 7/13/2014 9:18 pm 34 Adult Female Post lact 55 PIT tag 989.001003147092 7/13/2014 Na 33 Juvenile Female NR 57.3 PIT tag 989.001003147093 7/13/2014 NA NA Juvenile Male NR 56.8 released no tag 8/08/2014 NA 27 Juvenile Male NR 57 PIT tag 989.001003147071 8/08/2014 NA 29 Juvenile Male NR 58 PIT tag 989.001003147083 8/08/2014 NA 31 Juvenile Male NR 56 PIT tag 989.001003147085 8/08/2014 NA 25 Juvenile Female NR 55 PIT tag 989.001003147094 8/08/2014 NA 27 Adult Female NR 57 PIT tag 989.001003147097 8/09/2014 NA 31 Juvenile Male NR 58 PIT tag 989.001003147102 6/19/2015 10:07 pm 29 Adult Female Lact 56 PIT tag 989.001003147005

Date Time Weight Reproductive (m/dd/yyy) (12h) (g) Age Sex Status Forearm (mm) Tag Number 6/19/2015 10:14 pm 26 Juvenile Female NR 59 PIT tag 989.001003147008 6/19/2015 10:14 pm 27 Juvenile Male NR 58 PIT tag 989.001003147010 6/19/2015 9:53 pm 30 Adult Female Lact 57 PIT tag 989.001003147016 6/19/2015 10:43 pm 29 Juvenile Male NR 56 PIT tag 989.001003147017 6/19/2015 9:48 pm 30 Adult Female Lact 56 PIT tag 989.001003147031 6/19/2015 10:54 pm 30 Juvenile Male NR 59 PIT tag 989.001003147032 6/19/2015 11:44 pm 33 Adult Female NR 58 PIT tag 989.001003147036 6/19/2015 11:50 pm 29 Juvenile Male NR 57 PIT tag 989.001003147038 6/19/2015 11:48 pm 26 Juvenile Male NR 55 PIT tag 989.001003147048 6/19/2015 10:35 pm 26 Juvenile Female NR 57 PIT tag 989.001003147053 6/19/2015 9:51 pm 30 Adult Female NR 56 PIT tag 989.001003147056

52 6/19/2015 10:12 pm 31 Juvenile Male NR 59 PIT tag 989.001003147065

6/19/2015 10:50 pm 31 Juvenile Male NR 57 PIT tag 989.001003147080 6/19/2015 11:44 pm 24 Juvenile Female NR 58 PIT tag 989.001003147089 6/19/2015 10:38 pm 27 Juvenile Male NR 56 PIT tag 989.001003147099 6/21/2015 3:18 am 34 Adult Female NR 59 PIT tag 989.001003383157 6/21/2015 4:38 am 32 Adult Female NR 58 PIT tag 989.001003383173 6/21/2015 2:46 am 32 Adult Female Lact 55 PIT tag 989.001003383177 6/21/2015 2:52 am 36 Adult Female Lact 59 PIT tag 989.001003383204 6/21/2015 10:48 pm 29 Juvenile Male NR 58 PIT tag 989.001003147009 6/21/2015 9:50 pm 29 Adult Female Lact 56 PIT tag 989.001003147012 6/21/2015 9:39 pm 32 Adult Female Lact 60 PIT tag 989.001003147014 6/21/2015 10:07 pm 19 Adult Female Lact 55 PIT tag 989.001003147015 6/21/2015 10:23 pm 26 Juvenile Female NR 55 PIT tag 989.001003147018

Date Time Weight Reproductive (m/dd/yyy) (12h) (g) Age Sex Status Forearm (mm) Tag Number 6/21/2015 10:30 pm 32 Adult Female Lact 56 PIT tag 989.001003147020 6/21/2015 1:24 am 30 Juvenile Female NR 59 PIT tag 989.001003147021 6/21/2015 12:56 am 26 Juvenile Female NR 57 PIT tag 989.001003147024 6/21/2015 1:20 am 30 Adult Female Lact 59 PIT tag 989.001003147025 6/21/2015 1:03 am 34 Adult Female NR 57 PIT tag 989.001003147026 6/21/2015 10:36 pm 27 Juvenile Female NR 56 PIT tag 989.001003147027 6/21/2015 10:57 pm 30 Juvenile Male NR 55 PIT tag 989.001003147028 6/21/2015 1:10 am 27 Juvenile Female NR 56 PIT tag 989.001003147030 6/21/2015 12:43 am 41 Adult Female Lact 59 PIT tag 989.001003147034 6/21/2015 12:44 am 30 Juvenile Male NR 56 PIT tag 989.001003147035 6/21/2015 10:30 pm 29 Juvenile Male NR 58 PIT tag 989.001003147039

53 6/21/2015 10:20 pm 34 Juvenile Male NR 58 PIT tag 989.001003147041

6/21/2015 2:39 am 29 Adult Female NR 56 PIT tag 989.001003147044 6/21/2015 9:53 pm 33 Adult Female Lact 57 PIT tag 989.001003147047 6/21/2015 9:52 pm 24 Adult Female NR 58 PIT tag 989.001003147049 6/21/2015 11:37 pm 35 Adult Female Lact 60 PIT tag 989.001003147050 6/21/2015 11:54 pm 27 Juvenile Male NR 56 PIT tag 989.001003147052 6/21/2015 1:18 am 28 Juvenile Male NR 58 PIT tag 989.001003147055 6/21/2015 2:30 am 34 Adult Female Lact 60 PIT tag 989.001003147057 6/21/2015 1:40 am 27 Juvenile Female NR 57 PIT tag 989.001003147058 6/21/2015 9:45 pm NA Juvenile Female NR 57 PIT tag 989.001003147064 6/21/2015 9:39 pm 30 Adult Female NR 57 PIT tag 989.001003147066 6/21/2015 11:40 pm 29 Juvenile Female NR 56 PIT tag 989.001003147068 6/21/2015 2:26 am 32 Adult Female Lact 54 PIT tag 989.001003147070

001003147096 001003147098 001003147100 001003147101 001003147103 001003147104 001003147073 001003147074 001003147075 001003147077 001003147079 001003147081 001003147084 001003147087 001003147088 001003147091 001003147095 Tag NumberTag PIT tag 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT PIT tag 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT 989. tag PIT

58 57 53 61 58 60 58 55 59 55 58 59 61 58 57 55 60 Forearm (mm)

ductive Status NR NR Lact NR NR Lact NR Lact Lact NR NR NR Lact NR NR NR NR Repro

Sex Male Male Female Male Female Female Female Female Female Female Female Male Female Female Female Female Male

Age Juvenile Juvenile Adult Juvenile Adult Adult Juvenile Adult Adult Juvenile Juvenile Juvenile Adult Juvenile Juvenile Adult Juvenile

29 32 34 30 30 31 26 27 33 32 30 28 32 32 30 38 30 (g) Weight Weight

am am am am am

pm pm pm pm pm pm pm pm pm pm pm pm

1:20 2:20 1:50 1:10 9:53 9:56 9:54 9:49 Time (12h) 12:56 11:53 11:23 10:00 11:33 10:37 11:33 10:32 11:11

Date 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 6/21/2015 (m/dd/yyy) 54