ranid frog conservation and management - native fish lab of marsh

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RANID FROG CONSERVATION AND MANAGEMENT Michael J. Sredl, Editor Nongame Branch, Wildlife Management Division Arizona Game and Fish Department itclmical Report 121 Nongame and Endangered Wildlife Program Program Chief; lerry B. Johnson Arizona Game and Fish Department 2221 West Gteenway Road Phoenix, Arizona 85023-4399 June 1997

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Page 1: ranid frog conservation and management - Native Fish Lab of Marsh

RANID FROG CONSERVATION AND MANAGEMENT

Michael J. Sredl, Editor Nongame Branch, Wildlife Management Division

Arizona Game and Fish Department

itclmical Report 121 Nongame and Endangered Wildlife Program

Program Chief; lerry B. Johnson Arizona Game and Fish Department

2221 West Gteenway Road Phoenix, Arizona 85023-4399

June 1997

Page 2: ranid frog conservation and management - Native Fish Lab of Marsh

RECOMMENDEO CITATION

Sredl, M.J., editor. 1997. Ranid frog conservation and management. Nongame and Endangered Wildlife Program Technical Report 121. Arizona Qame and Fish Department, Phoenix, Arizona.

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TABLE OF CONTENTS

Mark-Recapture Studies of Arizona Leopard Frogs ................................................................... 1 Introduction .................................................................................................................... 3 Methods ........................................................................................................................... 3

Site Descriptions ................................................................................................ 4 Field Methods ..................................................................................................... 7 Statistical Methods .............................................................................................. 7

Results .............................................................................................................................. 8 Population Estimates and Survivorships .............................................................. 8 Body Size, Growth, and Sex Ratios ................................................................ 12

Discussion .................................................................................................................... 14 Demographics of Arizona Leopard Frog Populations ..................................... 14 Ecological Factors Affecting Arizona Leopard Frog Populations .................... 17

Literature Cited ............................................................................................................ 19

Validation of Visual Encounter Surveys ................................................................................. 21 Introduction ................................................................................................................. 23 Methods ......................................................................................................................... 24

Site Description and History ............................................................................ 24 VES Validation ................................................................................................ 25

Results ........................................................................................................................... 27 Phenology of Earthen Cattle Tanks .................................................................. 27 Count Results ................................................................................................... 28 Anecdotal Observations .................................................................................... 29

Discussion .................................................................................................................... 29 Phenology of Stock Tanks ............................................................................... 29 Counts ............................................................................................................... 30 Anecdotal Observations .................................................................................... 32

Literature Cited ............................................................................................................. 34

Status and Distribution of Arizona's Native Ranid Frogs ...................................................... 37 Introduction .................................................................................................................. 39 Methods ......................................................................................................................... 40

Survey Methodology ......................................................................................... 40 Target Ranid Frogs ........................................................................................... 41 Database Design .............................................................................................. 43 Data Analysis ................................................................................................... 43

Results ........................................................................................................................... 44 Database Overview ........................................................................................... 44 Distribution and Status of Arizona Leopard Frogs .......................................... 44

Discussion .................................................................................................................... 52 Generalizations from Statewide Surveys ......................................................... 52

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Factors Leading to Population Declines ........................................................... 53 Spatial Distribution of Extant Populations ...................................................... 53 Management Strategies - Conservation and Management Zones .................... 54 Information Needs ........................................................................................... 57

Literature Cited ............................................................................................................. 58 Appendix A. Riparian Herp Survey Form ................................................................... 63 Appendix B. Locality distribution by county and native ranid observation status 79 Appendix C. Locality distribution by drainage and native ranid observation status . 80 Appendix D. Locality distribution by management category and native ranid observation

status .................................................................................................................. 81 Appendix E. Elevational ranges within drainages by native ranid species ................. 82 Appendix E Number of ranid frog localities per county by species ........................... 83 Appendix G. Native ranid species status within management category ...................... 84 Appendix H. Native ranid species status within drainage .......................................... 87

MARK-RECAPTURE TABLES

Table 1. JOLLYAGE estimates of capture probabilities for Big Spring and Tule Creek .......... 8 Table 2. JOLLYAGE estimates for Big Spring .......................................................................... 9 Table 3. Lincoln-Peterson estimates for all sites ..................................................................... 11 Table 4. JOLLYAGE estimates for Tule Creek ........................................................................ 11 Table 5. e test of the null hypothesis: sex ratios are not different from 1:1 (sites x year) . 13

MARK-RECAPTURE FIGURES

Figure 1. JOLLYAGE population parameter estimates for Big Spring ..................................... 9 Figure 2. Total monthly rainfall for Big Spring ........................................................................ 10 Figure 3. JOLLYAGE population parameter estimates for Tule Creek ................................... 12 Figure 4. Total monthly rainfall for Tule Creek ..................................................................... 14 Figure 5. Box and whisker plot of frog SVL for Big Spring .................................................. 14 Figure 6. Box and whisker plot of frog SVL for Tule Creek .................................................. 15 Figure 7. Histograms of SVL for Big Spring .......................................................................... 16 Figure 8. Von Bertalanffy growth curves .................................................................................. 17

VISUAL ENCOUNTER SURVEY VALIDATION TABLES

Table 1. VES study site names, numbers, legal descriptions and elevations ........................... 26

VISUAL ENCOUNTER SURVEY VALIDATION FIGURES

Figure 1. Map of VES study sites modified from Gisela (1972) and Mazatzal Peak (1972) . 25 Figure 2. Tank surface area and rainfall .................................................................................. 27 Figure 3. Maximum counts of Rana yavapaiensis per trip ...................................................... 28

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STATEWIDE SURVEY TABLES

Table 1. Elevational ranges of native ranid localities ................................................................... 45 Table 2. Locality status designations for native ranid species ...................................................... 46

STATEWIDE SURVEY FIGURES

Figure 1. Stacked bar chart of aquatic habitat types grouped by species ................................... 46 Figure 2. Stacked bar chart of status designations grouped by species ...................................... 47

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MARK-RECAPTURE STUDIES OF ARIZONA LEOPARD FROGS

Michael J. Sredl, E. Patricia Collins, and Jeffrey M. Howland Nongame Branch, Wildlife Management Division

Arizona Game and Fish Department

Technical Report 121 Nongame and Endangered Wildlife Program

Program Chief: Terry B. Johnson Arizona Game and Fish Department

2221 West Greenway Road Phoenix, Arizona 85023-4399

June 1997

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RECOMMENDED CITATION

Sredl, M.J., E.P. Collins, and J.M. Howland. 1997. Mark-recapture of Arizona leopard frogs. Pages 1-20 in M.J. Sredl, editor. Ranid frog conservation and management. Nongame and Endangered Wildlife Program Technical Report 121. Arizona Game and Fish Department, Phoenix, Arizona.

ACKNOWLEDGMENTS

For assistance with field work, we thank: Linda Allison, Maggy Bathory, Rob Brauman, Ed Burns, Bruce Christman, Brian Dickey, Tracy Ertz-Berger, Matt Goode, Morgan Heath, Jacques Hill, Ron Hill, Wendy Hodges, Angie Kiselyk, Chris Klug, Charlie Painter, Bruce Pavlick, Ben Robles, Loralei Saylor, Kelly Schwartz, Sharon Seim, Farraday Sredl, Eric Wallace, Dana Waters, Dave Weedman, and John Windes.

Linda Allison assisted with many aspects of the analysis; Ken Burnham and Jim Hines provided insight into analysis of mark-recapture data, and Jim Collins provided background information on Tule Creek and amphibian life history which greatly improved the study. We are grateful to Phil Fernandez as a cooperator in the skeletochronolgy analysis. We also thank Roy Murray for help with growth data analysis, Dana Waters for writing data conversion programs, and Sharon Seim for analyses early in the study. We recognize Donald Pinkava and Barry Spicer for assistance with plant identification. Finally, we thank Randy Babb for donating the cover illustration.

PROJECT FUNDING

Funding for this project was provided through: the Arizona Game and Fish Department's Heritage Fund; voluntary contributions to Arizona's Nongame Wildlife Checkoff; U.S. Fish and Wildlife Service Project E5, Jobs 12 and 26, under Title VI of the Endangered Species Act; U.S. Fish and Wildlife Service Partnerships For Wildlife project, Job 02, administered by the National Fish and Wildlife Foundation.

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RANID FROG MARK-RECAPTURE

Michael J. Sredl, E. Patricia Collins, and Jeffrey M. Howland

INTRODUCTION

Taxonomy of the Rana pipiens complex, particularly populations of those frogs inhabiting the western United States, Mexico, and Central America, has been problematic (Hillis 1988). The focus of most studies of this group has therefore been taxonomic (Mecham 1968; Moore 1944; Platz and Platz 1973; Platz 1976; Post and Pettus 1966). Three new species of leopard frogs from Arizona have been described since 1979, and the Chiricahua leopard frog will soon be split (Platz 1993; Platz and Frost 1984; Platz and Mecham 1979; J.E. Platz, pers. comm.).

Reports of recent population declines in native ranids of the western United States (Jennings and Hayes 1994), including members of the "pipiens complex" within Arizona (Clarkson and Rorabaugh 1989), have made baseline demographic data a high priority. To understand the specific mechanisms which shape population dynamics, an important first step is distinguishing natural fluctuations from fluctuations brought about by human impacts (Pechmann et al. 1991).

The population dynamics of species within this complex in the arid Southwest have received little attention. To gather baseline population data, we conducted mark-recapture studies on the lowland leopard frog (R. yavapaiensis) and Chiricahua leopard frog (R. chiricahuensis). R. yavapaiensis is a desert ranid found in rivers, streams, cienegas (=wetland), stock tanks, and other permanent waters. In Arizona, which includes most of the geographic distribution of this species, habitats range in elevation from 146-1817 m (480-5960 ft) (for additional status and distribution data, see below). The Chiricahua leopard frog ranges from the southern edge of the Colorado Plateau in Arizona and New Mexico into SE Arizona, SW New Mexico, and the Sierra Madre Occidental in Mexico (Platz and Mecham 1979). Within Arizona, it has been found from 1160-2963 m (3480-8890 ft). We studied population dynamics of these two species at seven sites. We chose six sites across much of the historical range of R. yavapaiensis and representative of various habitats (e.g. stock tanks, intermittent streams with perennial pools, and permanent streams). We chose a single study site, consisting of two stock tanks, for mark-recapture study of R. chiricahuensis. Over the six year study period, we accumulated the largest baseline data set on southwestern leopard frog population dynamics that has yet been produced.

METHODS

From 1991 through 1996, we carried out mark-recapture studies of Rana yavapaiensis at one or more of six sites. We conducted a mark-recapture study of R. chiricahuensis at a single site in 1994. Initial criteria for site selection were: high degree of geographic closure, relatively natural or minimally modified habitat, and sufficient number of frogs to yield adequate captures and

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recaptures. Sites initially chosen for study of R. yavapaiensis population dynamics were: Alamo Canyon, Big Spring, Reed Spring, Thicket Spring, and Tule Creek. When it became apparent, through our statewide surveys (see chapter 3), that earthen cattle tanks were important as leopard frog habitat in Arizona, we chose two additional sites, Horsefall Canyon for R. chiricahuensis and Barnhardt Mesa for R. yavapaiensis, to study the dynamics of cattle tank populations.

Site Descriptions: Alamo Canyon, Pima Co. (T12S, R14E, Sec. 9, NE1/4 of NE 1/4), drains W off the Santa Catalina Mountains near Tucson, Arizona. There is no perennial flow in this drainage, and the only "permanent" water is found in tinajas (scoured bedrock pools filled by surface runoff). The Alamo Canyon study site (AC) consists of three such pools varying in size from 1 m x 2 m (3 ft x 6 ft) to 3 m x 4 m (9 ft x 13 ft) and .5 to 1 m (1.5-3 ft) in depth depending on the time of year. Lack of permanent flow allows little development of riparian vegetation, but a few scattered Fremont cottonwood (Populus fremonti) and willows (Salix spp.) were present. The largest, deepest pools had small stands of cat-tail (Typha angustifolia). Surrounding vegetation was Arizona Upland upper division (Brown 1994) dominated by mesquite (Prosopis spp.), saguaro (Cereus giganteus), and ocotillo (Fouquieria splendens). Turpentine bush (Ericameria laricifolia) and Arizona grape (Vitis arizonica) grew on the gentle slopes of the canyon. Study site elevation is 960 m (2880 ft). We sampled this site twice a year, in August and October, between 1991 and 1992. We used rainfall data from the University of Arizona, 20.1 km (12.5 mi) away, at an elevation of 745 m (2444 ft), as representative for the AC site. Annual rainfall for 1894 to 1996 averaged 286 mm (11.28 in).

Barnhardt Mesa, Gila Co., Arizona, is in the eastern foothills of the Mazatzal Mountains. The Barnhardt Mesa study site (BM) consisted of a series of three earthen cattle tanks filled by surface runoff. Because of seasonal variation in rainfall, water level of these tanks is subject to dramatic fluctuation, with peak water levels coincident with winter and summer rainy seasons. The tanks are W of State Highway 87 (T9N, R9E, Sec. 24, SW 'At of SW %), approximately 130 km (81 mi) NE of Phoenix. At 1134 m (3720 ft) elevation, Barnhardt Mesa Tank is the uppermost tank in this system. Springtime diameter of 25 m (82 ft) and maximum depth of 1 m (3 ft) are typical. The tank is ephemeral, usually drying by mid summer. There is little vegetation at the tank. Mesa Tank, the largest of the three tanks with typical spring diameter of 44 m (145 ft) and maximum depth of 1.5 m (5 ft), is 300 m (984 ft) downstream from Barnhardt Mesa Tank. Vegetation included spike rush (Eleocharis), and pondweed (Potomageton). One side of the tank had several willows (Salbc spp.) and a sugar sumac (Rhus ovata). At 1085 m (3560 ft) elevation, an unnamed tank 900 m (2970 ft) from Mesa Tank was the lowest and deepest of the three study tanks. It reached a maximum depth of 2.7 m (9 ft) with springtime diameter of 36 m (119 ft). This pond also contained spike rush and pondweed, and was surrounded by introduced bermuda grass (Cynodon dactylon), mesquite (Prosopis spp.), and scrub oak (Quercus turbinella). The surrounding rolling hills were formerly desert grasslands, but heavy grazing and fire suppression have caused reduction in bunch grass cover with replacement by prickly pear (Opuntia spp.), mesquite, yucca (Yucca spp.), one-seed juniper (Juniperus monosperma), and wait-a-minute bush

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(Mimosa biuncifera) (Brown 1982). BM tanks were studied April, June, August and October of 1994 and 1995. Surveys in June and August of 1995, however, did not yield any frogs. An apparent red-leg epidemic (opportunistic bacterial infection caused by Aeromonas hydrophila) in early spring of 1995 decimated the BM frog population, leading us to stop work there in October, 1995. Rainfall data from Gisela, 16.1 km (10 mi) E, elevation 884 m (2900 ft), show average annual rainfall of 429 mm (16.9 in) from 1941 to 1970.

Big Spring, Graham Co., Arizona, 13 km (8.1 mi) W of Safford and 3.2 km (2 mi) N of the Gila River, is a small spring producing approximately 7.6 1(2 gal) of water per minute. Surface flow, except when unusually high, does not reach the Gila River. Big Spring study site (BS) (T6S, R25E, Sec. 5, NE% of SE%) consists of a 183 m (600 ft) section of Big Spring Wash containing perennial water. When the study began, frog habitat consisted of at least four pools connected by shallow riffles. Several high flow events gradually silted in all but one pool. This pool, 5 m x 3 m (16 ft x 10 ft) and 1 m deep (3 ft), was maintained by scouring action of water flowing over a small concrete dam approximately 54 m (177 ft) downstream from the spring head. Uplands surrounding BS were characterized as Sonoran desertscrub, Arizona Upland subdivision (Brown 1994), dominated by creosote bush (Larrea tridentata). The riparian corridor was dominated by introduced salt cedar (Tamarix chinensis), but some native trees and shrubs were present, including Goodding willow (Salbc gooddingii), seepwillow (Baccharis glutinosa), and Fremont cottonwood. Alkali sacaton (Sporobolus airoides), common reed (Phragmites australis), yerba-mansa (Anemopsis califomica), and plantain (Plantago spp.) formed terrestrial ground cover, with cat-tail the predominant aquatic cover. The study area was closed to cattle grazing in 1980. A pipe at the spring head carried water to a drinker outside the riparian area. We sampled BS twice a year (August and October) beginning in 1991, and four times a year (April, June, August, and October) from 1993 through 1996. The elevation of the BS site is 914 m (3000 ft). Rainfall at Safford Agricultural Center, 18.2 km (11.3 mi) SE at 900 m (2954 ft) elevation, averaged 233 mm (9.18 in) from 1948 to 1996.

Horsefall Canyon, Cochise Co., Arizona, is a small, mostly dry canyon draining into North Fork Pinery Creek. Pinery Creek drains the NW side of the Chiricahua Mountains, just S of Chiricahua National Monument. Horsefall Canyon study site (HC) consists of two earthen cattle tanks. Headquarters Windmill Tank (HW) (T17S, R30E, Sec. 7, SW 'A of NW %) is at the confluence of Horsefall Canyon and North Fork Pinery Creek, at 1768 m (5800 ft) elevation. It is fed by spillover from a large, metal drinker supplied by a windmill. Frogs occupied both the drinker and the tank. Monkey Tank (MT) (T17S, R30E, Sec. 17, need 1/4), at 1853 m (6080 ft) elevation, is 1.6 km (1 mi) up Horsefall Canyon from HW. Surrounding upland vegetation is Madrean Evergreen Woodland (Brown 1994), dominated by Chihuahua pine (Pinus leiophylla var. chihuahuana), Apache pine (Pinus engelmannii), southwestern white pine (Pinus strobifonnis), ponderosa pine (Pinus ponderosa), border pinyon (Pinus discolor), Emory oak (Quercus emoryi), Arizona white oak (Quercus arizonica), and Mexican blue oak (Quercus oblongifolia). We began sampling HC in May, 1994. A massive die-off at HW in June, 1994, possibly stimulated by

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hydrogen sulfide poisoning, brought our brief study of R. chiricahuensis demographics to a close. Rainfall at Chiricahua National Monument, 5.8 km (3.6 mi) N, elevation 1634 m (5360 ft), averaged 500 mm (19.63 in) from 1909 to 1996.

Reed Spring, Gila Co., Arizona, is a perennial spring (T8N, R10E, Sec. 34, SW 1/4

of SW 1/4) in Reed Gulch, 1.6 mi (2.6 km) West of Tonto Creek, 90 km (56 mi) NE of Phoenix. Reed Gulch is a tributary of Tonto Creek, but they are connected by surface flow only after heavy rainfall events. Reed Spring study site (RS) consisted of the entire 100 m (328 ft) stretch of perennial water: seven to nine small pools connected by shallow surface flow. The largest plunge pool was approximately 3 m (9.8 ft) across and 1 m (3.3 ft) deep. Some pools contained dense aquatic vegetation while others were bare. The drainage was 2-10 m (6-33 ft) below surrounding terrain. Vegetation in the drainage included seepwillow, Fremont cottonwood, willows, netleaf hackberry (Celtis reticulata), Arizona grape, and tamarisk, with one-seed juniper, velvet mesquite (Prosopis velutina), catclaw acacia (Acacia greggii), barberry (Berberis sp.), desert hackberry (Celtis pallida), squawbush (Rhus trilobata) and sugar sumac above the banks. The site lies in a transition zone between Sonoran desert scrub/Arizona upland subdivision and semi-desert grassland (Brown 1994) and dominant vegetation of the uplands above Reed Gulch includes saguaro, prickly pear (Opuntia phaeacantha), jojoba (Simmondsia chinensis), catclaw acacia, and one-seed juniper. RS elevation is 865 m (2840 ft). We sampled RS in August and October of 1991 and 1992. Rainfall at Gisela, 9.6 km (6 mi) E, is reported under BM.

Thicket Spring is in Bloody Basin, 80 km (50 mi) N of Phoenix, Yavapai Co., Arizona, in a small, dry wash near Red Creek, an intermittent tributary of the Verde River. Thicket Spring study site (TS) (T1ON, R10E, Sec. 34, SE% of SW 1/4) is a tear drop shaped pond formed by excavation in and around the spring itself. The long axis of the pond varied from 23-25 m (75-82 ft) and depth exceeded 1 m (3 ft) at the deepest point. Vegetation around the pond included introduced rabbitfoot grass (Polypogon monspeliensis), junipers (Juniperus sp.), and mesquites. Two large netleaf hackberries provided approximately 20% canopy cover. The pond surface was entirely covered by duckweed (Lemna sp.) and other aquatic macrophytes, including filamentous algae. Surrounding terrain is hilly and in a transition zone between semi-desert grassland and Sonoran upland desertscrub (Brown 1994). Dominant vegetation includes prickly pear, ocotillo, mesquite, cheesebush (Hymenoclea salsola), catclaw acacia, and one-seed juniper. The site elevation is 902 m (2960 ft). We sampled TS in August and October of 1991; surveys in 1992 yielded few or no frogs. Rainfall at Childs, 19.6 km (12.2 mi) NE, at 807 m (2650 ft) elevation, averaged 480 mm (19 in) from 1961 to 1990.

Tule Creek, Yavapai Co., Arizona, is an intermittent tributary of the Agua Fria River and is the type locality for R. yavapaiensis. Tule Creek (TC) study site (T8N, R1E, Sec. 28, NW 'A of SW 'A) is an approximately 1.6 km (1 mi) perennial stretch of this drainage, located 53 km (33 mi) N of Phoenix. Surrounding terrain is steep and rugged Sonoran desertscrub, Arizona Upland subdivision, dominated by the paloverde (Cercidium spp.)-saguaro community. Riparian overstory

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consisted of Fremont cottonwood, Goodding willow, catclaw acacia, velvet mesquite, and seepwillow. Canyon ragweed (Ambrosia ambrosioides), arrow-weed (Tessaria sericea), three-square (Scirpus americanus), yerba-mansa, and introduced Bermuda grass were the predominant ground cover. Cat-tail was present in larger pools. One year into the study, the site was designated a riparian conservation area and excluded from cattle and feral burro grazing. Once fenced, vegetation quickly became too dense to allow thorough sampling of frogs, so the site was dropped from further study after 1993. Elevation of TS is 658 m (2160 ft). TC was surveyed twice a year beginning in 1991 (August and October) and in April, June, August and October of 1993. Rainfall at Castle Hot Springs, 9.8 km (6.1 mi) W, 640 m (2100 ft) elevation, averaged 404 mm (16 in) from 1959 to 1996.

Rainfall at all sites is biseasonal, with peaks in late summer and winter.

Field Methods: Beginning after sunset, we sampled leopard frogs by repeatedly searching a 50 m (164 ft) section of a lotic system or quarter section of a lentic system and capturing as many frogs as possible, by hand or dipnet. Frogs were placed in plastic zipper bags or moist mesh cloth bags. We searched areas until repeated passes yielded no further captures. Animals captured for the first time were marked using a modified Martof (1953) scheme (thumbs were not removed). For each animal marked, we recorded mass (±0.5 g), snout-vent length (SVL)( ±1 mm), sex, section of capture, and microhabitat (marsh, small pool, large pool, upland, debris, riffle) where the frog was first sighted.

Except for the first two years of the study, we visited sites four times a year between April and October, the times of highest frog activity. Because of low numbers of captures and recaptures, studies at Reed Spring, Thicket Spring, and Alamo Canyon were terminated after 1992.

Statistical Methods: We used program JOLLYAGE to compute estimates for a two age class Jolly-Seber open population model. The Jolly-Seber model assumes the population of interest is open, allowing additions and deletions to the population (Pollock et al. 1990). When it became apparent that, for most sites, our recapture rates were not sufficient to compute the necessary parameters to test the model assumptions and fit of Jolly-Seber models, we used the Chapman bias adjusted Lincoln-Peterson equation to estimate population size for all sites. The general Lincoln-Peterson model assumes demographic closure (i.e. no additions or deletions to the population)(White et al. 1982). We computed population estimates for individuals >55.0 mm SVL, because individuals below this size are likely to represent recently metamorphosed individuals and, therefore, additions to the population. In years with four sampling efforts, we combined the first two trips into a single pre-monsoon sampling period, and the last two trips into a single post-monsoon sampling period.

We used the Von Bertalanffy equation to fit growth curves to data collected on male and female frogs captured at BS between 1991-1996. We fit our curve using SVL at first and last capture and

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time between captures for each individual included. Juveniles whose sex was unambiguously known at second capture were included in the curve of that sex. Juveniles whose sex was indeterminable at second capture were included in the curves of both males and females.

For all descriptive statistics, analyses of size and growth, and plots, we used SPSS (Version 7.5) with a =0.05.

RESULTS

Population Estimates and Survivorships: Because of low numbers of recaptures at most study sites, we were able to compute Jolly-Seber estimates for TC and BS only. Capture probabilities for BS ranged from 0.13-0.58 (Table 1).

Table 1. JOLLYAGE estimates of capture probabilities for Big Spring and Tule Creek (site abbreviations as in text; ne=not estimable).

Site Month 1991 1992 1993 1994 1995 1996

BS Apr 0.44 t 0.18 0.58 ± 0.35 0.39 ± 0.21 0.23 ± 0.28

Jun 0.26 ± 0.14 0.25 ± 0.25 0.32 ± 0.19 0.25 ± 0.31

Aug ne 0.13 ± 0.06 0.34 ± 0.16 0.40 ± 0.19 0.36 ± 0.16 0.18 ± 0.33

Oct 0.16 ± 0.07 0.37 ± 0.15 0.32 ± 0.30 0.25 ± 0.17 0.40 ± 0.23

TO Apr 0.58 ± 0.50

Jun 0.06 ± ne

Aug ne 0.10 ± 0.17 0.09 ± 0.07

Oct 0.04 ± 0.08 0.13 ± 0.27

These low capture probabilities resulted in large confidence intervals for many of the Jolly-Seber estimates for both sites (Table 2). Jolly-Seber population estimates from BS (Fig. la) varied from a high of 632 animals in August of 1992, followed by a large decline to 80 individuals between October 1992 and April 1993. After April 1993, the BS population had a few "peaks" in numbers of frogs, which tended to take place late in the season (August through October). These peaks occurred in 1993, 1994, and 1995, the last full year of our study. In spite of these late season peaks in frog numbers, population size at BS generally decreased throughout our study (Fig. la; Table 2).

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Table 2. JOLLYAGE estimates for Big Spring: population size and adult and juvenile survivorships (±95% Cl).

Date Population Recruitment Adult

Survivorship Juvenile

Survivorship

Aug 1991 1.18 ± 0.38 0.51 ±0.19

Oct 1991 532 ± 200 279 ± 2506 0.66 ± 0.25 0.27 ± 0.18

Aug 1992 633 ± 253 -127 ± 6259 0.95 ± 0.42 0.15 ± 0.17

Oct 1992 474 ± 191 11 ± 168 0.15 ± 0.07 0.03 ± 0.05

Apr 1993 80 ± 30 101 ±128 0.85 ± 0.36 1.69 ± 0.47

Jun 1993 169 ± 85 11 ± 666 0.79 ± 0.37 0.85 ± 0.43

Aug 1993 145± 61 52 ± 236 0.81 ± 0.83 0.71 ± 0.75

Oct 1993 169 ± 164 9 ± ne 0.06 ± 0.07 0.0 ± 0.0

Apr 1994 19 ± 11 48 ± ne 0.72 ± 0.23

Jun 1994 62 ± 55 58 ± 666 1.72 ± 0.69 0.7 ± 0.33

Aug 1994 164 ± 71 182 ± 205 0.56 ± 0.43 1.83 ± 1.16

Oct 1994 274 ± 180 -8 ± ne 0.12 ± 0.08 0.0 ± 0.0

Apr 1995 26 ± 12 49 ± ne 1.03 ± 0.25

Jun 1995 76 ± 40 0.36 ± 387 1.24 ± 0.41 1.08 ± 0.53

Aug 1995 94 ± 37 -4 ± ne 0.78 ± 0.47 0.0 ± 0.0

Oct 1995 70 ± 38 3 ± ne 0.56 ± 0.69

Apr 1996 42 ± 48 6 ± ne 0.32 ± 0.48

Jun 1996 19 ± 21 20 ± 46 1.10 ± 2.17 0.31 ± 0.65

Aug 1996 41 ± 74

Chapman-bias adjusted Lincoln-Peterson estimates for BS are shown in Table 3. Low capture probabilities similarly affected the confidence intervals of our Lincoln-Peterson estimates. Like JOLLYAGE, the Lincoln-Peterson model calculated the highest population estimate in 1992. This estimator also indicated that a sudden decrease in adult population size took place between 1992-1993, followed by a gradual decrease in population size for the duration of the study (Table 4).

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Arizona Game and Fish Department NGTR 121: Ranid Frog Conservation and Management

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20

The estimates of adult and juvenile a) survivorships for BS also vary and, in general, are higher for adults than juveniles, with one notable exception taking place in April 1993 (Fig. lb; Table 2). Adult survivorship estimates ranged from 0.06 to > 1.00 throughout the study period, while that of juveniles ranged from 0.03 to > 1.00. The lowest survivorship estimates for both adults and juveniles were calculated for the same interval, October 1993 to April 1994 (winter). Besides winter 1993-94, juvenile survivorship was "0" for winter 1994-95 and August-October 1995. "Recruitment" estimates the number of individuals caught at time "I" that survived to time "1+1". These estimates for BS ranged from a maximum of 279 juveniles in October 1991 to -127 in August 1992 (Table 2). In every year for which we have more than one value including an October estimate, the recruitment estimate for October was always the lowest. These estimates did not exhibit any seasonal pattern, but in the last two years of the study, decreased to near zero in many intervals, indicating little recruitment at BS.

Estimates of both adult and juvenile survivorship appear to follow a seasonal pattern. Within a given year, survivorships were always lowest in the winter. In 3 of 4 years for which we have estimates for all four intervals, wintertime survivorship was by far the lowest, for both age classes.

Peak rainfall occurred in November through February and August or September of each year (Fig. 2), with the exception of August, 1995 which was a very dry year at BS.

date

Figure 1. JOLLYAGE population parameter estimates for Big Spring. a) Population estimates b) Survivorship plots for juveniles (triangles, dashed lines), and adults (squares, solid line).

11 Aug 91 Aug 92 Aug 93 Aug 94 Aug 95 Aug 96

Feb 92 Feb 93 Feb 94 Feb 95 Feb 96

date (mmm yy)

Figure 2. Total monthly rainfall for Big Spring.

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JOLLYAGE estimates of capture probabilities for TC ranged from 0.04-0.58 (Table 1). The 0.04 capture probability estimated for the October 1991 interval was the lowest estimate calculated for either site, while the April 1993 estimate of 0.58 tied the high estimate.

Table 3. Lincoln-Peterson estimates for all sites. Estimates (±95% Cl) were computed for individuals >55.0 mm SVL (site abbreviations as in text; ne=not estimable).

1991 1992 1993 1994 1995 1996

AC 41 t 46 41 t 49

BM 863 t 303

BS 313±111 443±111 156 t 37 134 t 85 92 t 25 70 t 47

HW 59 t 39

MT 24 tne

RS ne 19 t 21

TS 73 ± 36

IC 704 t 471 887 t 803 1806 t 1874

Table 4. JOLLYAGE estimates for Tule Creek: population size and adult and juvenile survivorships (±95% Cl; ne=not estimable).

Date Population Recruitment Adult

Survivorship Juvenile

Survivorship

Aug 1991 3.59 ± 8.00 2.09 ±4.77

Oct 1991 3284 ± 7417 111 ± 5172 0.04 ± 0.09 0.08 ± 0.16

Aug 1992 231 ± 412 904 ± 2746 0.48 ± 1.16 0.26 ± 0.73

Oct 1992 1015 ± 2294 -11 ± ne 0.02 ± 0.03 ne

Apr 1993 14 ± 15 226 ± ne 3.92 ± ne ne

Jun 1993 282 ± ne ne ne 0.36 ± 0.36

Aug 1993 1647 ± ne ne ne ne

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b) 4.0

3.0

2.0

1.0

0.0 Aug 91

Oct 91 Feb 92 Jun 92 Oct 92 Feb 93 Jun 93 Apr 92 Aug 92 Dec 92 Apr 93 Aug 93

Arizona Game and Fish Department NGTR 121: Ranid Frog Conservation and Management

a)

June 1997 Page 12

Jolly-Seber population estimates for TC (Fig. 3a) fluctuate from a high of 3283 frogs in October of 1991 to a low of 14 frogs in April of 1993. Late season population estimates, like those at BS, tended to be among the highest calculated for a given year (Table 4). Survivorship estimates for TC ranged from 0.03 for adults in October of 1991 to > 1.00 for several sampling intervals (Fig. 3b). Wintertime survivorship of TC adults and juveniles exhibited a pattern similar to BS. We were only able to compute recruitment estimates for four intervals. In the one year for which we had more than one recruitment estimate including an October estimate, the October estimate was the lowest, as for BS.

Rainfall at TC generally remained below 150 mm per month throughout the study, except for January 1993, when 291 mm of rain was recorded (Fig. 4).

Lincoln-Peterson estimates for all other sites appear in Table 4.

Body Size, Growth, and Sex Ratios: A seasonal fluctuation in body size is evident at TC and BS (Figs. 5 and 6). At both sites median SVL is highest in frogs measured in April, October.

date

Figure 3. JOLLYAGE population parameter estimates for Tule Creek. a) Population estimates b) Survivorship plots for juveniles and adults (symbols as in Fig. 2).

lowest in June, and gradually increases through

Additional seasonal changes in size structure are evident in histograms of SVL measured between October 1992-1994 from BS (Fig. 7). SVL distribution was bimodal in October 1992, with the first mode comprised of individuals which metamorphosed from eggs oviposited in spring of 1992. In contrast to this pattern of recruitment of "same year" animals, the vast majority of animals recruited to BS in 1993 and 1994 appear to have been individuals which had overwintered as tadpoles that metamorphosed in June of each year. One fmal pattern occurred in 1994. At the beginning of that year, the BS population was extremely small. Modal size in subsequent samples appeared to track the cohort of frogs that metamorphosed in June as they survived and grew through October.

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Table 5. x2 test of the null hypothesis: sex ratios are not different from 1:1 (sites x year). "*" designates significance at a=0.05; site abbreviations as in text; ne=not estimable.

Site Year Males Females X2 values

AC AC 1991 10 5 1.67 0.1967

1992 1 10 7.36 0.0067*

BM 1994 137 180 5.83 0.0157*

1995 4 8 1.33 0.2482

BS 1991 75 68 0.34 0.5583

1992 97 143 8.82 0.0030*

1993 61 73 1.07 0.2999

1994 16 54 20.63 0.0000*

1995 15 77 41.78 0.0000*

1996 - 11 24 4.83 0.0280*

HW 1994 14 20 1.06 0.3035

MT 1994 16 10 1.38 0.2393

RS 1991 ne ne

1992 3 4 0.14 0.7055

TS 1991 21 23 0.09 0.7630

TC 1991 76 98 2.78 0.0954

1992 80 97 1.63 0.2013

1993 142 150 0.22 0.6397

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Results of nonlinear regression give us estimates of mean asymptotic SVL of 63.1 mm for males and 76.4 mm for females (male ms residuals =30.51; female ms residuals =23.74). The characteristic growth rates are 0.21 for males and 0.16 for females, indicating that males grow faster than females (Fig. 8), at least at small sizes.

We tested the hypothesis that sex ratios differ from 1:1 at all sites by year using a X2

analysis. At five out of eight sites, we found no difference in sex ratios (Table 5). At two sites, AC and BM, we found a female bias in the years 1992 and 1994, respectively. At BS, a female bias was present during four out of the six years of the study.

DISCUSSION

Demographics of Arizona Leopard Frog Populations: Estimates of all population parameters fluctuated dramatically. In general, population estimates calculated by Jolly-Seber and Lincoln-Peterson agreed closely, and increased as capture probabilities became >0.25. But in order to make an effective comparison, we would have to collapse our JOLLYAGE intervals from four to two. A straight estimate by estimate comparison is not possible. The most notable "discrepancy" between JOLLYAGE and Lincoln-Peterson estimates occurs in October 1991, which was also the interval which had the lowest Jolly-Seber capture probability of our study.

Aug 91 Dec 91 Apr 92 Aug 92 Dec 92 Apr 93 Aug 93 Oct 91 Feb 92 Jun 92 Oct 92 Feb 93 Jun 93

date (mmm yy)

Figure 4. Total monthly rainfall for Tule Creek.

snou

t-ven

t len

gth (

mm

)

90

80

70

60

50

40

TRIP

30

20 91 92 93 94 95 96

Year

Figure 5. Box and whisker plot of frog SVL for Big Spring. Dark line represents median, box includes 50% of individuals, and whiskers represent largest and smallest individuals that are not outliers (trip 1=April, trip 2=June, trip 3=August, trip 4=October).

300

250

200

150

100

50

Frog SVL varied considerably both within and between sites. The largest variation in size at BS occurred early in the study when frogs were most numerous. As the population decreased, median SVL seemed to reflect growth of a single cohort of newly metamorphosed frogs. Thus, for years where four data points are available, a conspicuous pattern emerges, where frogs seen in April are

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comparatively large. The pulse of metamorphosing frogs in early summer brings the median size of frogs down to its lowest annual level, and increasing median SVLs for August and October reflect the rapid growth of newly recruited individuals. Median SVL changed little at TC from 1991 to 1992, with the age spread of individuals being larger in August than in October for both years. In 1993, where four data points are available, we see a similar pattern as at BS where median frog SVL is elevated in April, at its lowest in June, and reflects the growth of juvenile frogs in August and October.

100

90

80

70

60

50

snou

t-ven

t len

gth

(mm

)

TRIP

40

0

0 2

30

3

20 IM 4

Figure 6. Box and whisker plot of frog SVL In 1991, the numbers of frogs captured at for Tule Creek (graphing conventions and RS and AC was comparably low (n=34 and abbreviations as in Fig. 5).

n=29, respectively), yet the summary plots of SVL show them to be demographically very different. In 1991, the range of SVL of AC frogs was representative of the known range for the species, suggesting a heterogeneous mixture of newly metamorphosed individuals and reproductive adults. In August of 1992, the spread was even wider, though the number of frogs captured decreased by roughly one third (n=21). The median SVL was quite low, however (approximately 30 mm) and increased to over 60 mm by October, reflecting once again the growth of young of the year. RS on the other hand appears to have been on the downside of a decline when we sampled there in 1991. All of the 34 frogs captured there measured under 30 min SVL, suggesting few reproductive adults were present. Median SVL in October of 1991 reflects the growth of these young as does the considerably elevated median of August 1992. Sample sizes for 1992 were too low (n=9) to draw any strong conclusions about the SVL of individuals there, but they do suggest the population was reduced to a single cohort.

We found female biased sex ratios at AC and BM in one out of the two years studied at each site. Sample sizes at AC are too small to be conclusive however, as is the case for BM in 1995. At BS we found a female bias in four out of the six years of the study. The female bias first appeared in 1992, when the population was at its densest. After flooding and the die-off of December 1992, numbers of males and females were again comparable. In the three subsequent years, however, the female bias was pronounced. We are unable to pinpoint a cause for this phenomenon and we cannot discount incorrect sexing as one possible source of the bias. Sexual dimorphism in terms of capture probability is another possible explanation.

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0■00. 000111

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Arizona Game and Fish Department NGTR 121: Ranid Frog Conservation and Management

June 1997 Page 16

Oct 1992

no mo no 410 40 WO MO WO 00 MO no mo ma ma mo

MO WO MO 400 40 WO MO WO 00 MO no mo mo mo mo

Jun 1993 Jun 1994

Po• 12 0 1491 • 00 7 N•33703

00

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100 92 Dee • 07 Nean•63• N•3703

Apr 1993 Apr 1994 mo

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Oct 1993 so

10 • 03•1153 /Men. 511 5•6700

MO VO MO Q0 40 WO WO WO WO MO no ma 00 WO 00

Oct 1984

S O 056. 372 1,1901•5115 u•noa

Figure 7. Histograms of SVL for Big Spring Rana yavapaiensis.

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90

80

70

60

50

V;

40

30

20

10

as se ass 5 5 5

10 20 30

Arizona Game and Fish Department June 1997 NGTR 121: Ranid Frog Conservation and Management Page 17

The mean asymptotic length of females at BS was more than 10 mm larger than that of males. The growth curve generated by our data indicate that males approached asymptotic length sooner than females, consistent with growth rate of males. Depending on how tightly sexual maturity is tied to body size, this could mean females attain sexual maturity later than males. Mean maximum length is reached between 22 and 24 months for both sexes. Our growth curve will be useful in comparison in future studies on leopard frog growth, age at sexual maturity, and longevity.

time post-metamorphosis (months)

Figure 8. Von Bertalanffy growth curves for male (+) and female (m) Rana yavapaiensis from Big Spring.

Ecological Factors Affecting Arizona Leopard Frog Populations: In addition to providing insights into population dynamics, mark-recapture has given insights into the ecological factors that create those dynamics. At the beginning of our study at BS, total surface area of pools measured approximately 200 m2 . In the summer of 1992, one or more high flow events reduced pool surface area to 35 m2. Even though these flow events were strong enough to fill the main breeding pool with gravel, survivorship of adults and juveniles did not appear to be affected (Fig. 1). Like many Sonoran Desert fish populations (Meffe and Minckley 1987), lowland leopard frog populations may persist following all but the most severe flash flood events. Comparison of densities of frogs between BS and TC, our next most densely populated site (AGFD unpubl. data), showed densities at BS to be as high as 10 times that at TC. Red-leg, a bacterial infection caused by Aeromonas hydrophila and other opportunistic bacteria, can be brought on by stress (Carey 1993). Within freshwater aquatic systems, these bacteria are ubiquitous (Reed and Toner 1942) and appear to be psychrophilic (=cold loving). They are also common in the intestines of amphibians at low temperature (Carr et al. 1976). Extremely low temperatures in December 1992 (National Climatic Data Center 1997), further stressing the dense BS population, was the likely cause of a major red-leg epidemic which killed a large portion of the BS population. This die-off in a stressed population following low temperatures is consistent with the hypothesized mechanism of declines in boreal toads (Carey 1993). Since this epidemic, the BS population has not recovered to its pre-epidemic population size. This lack of recovery is most likely due to the siltation and reduction of pool habitat within the drainage.

Desiccation is likely to be another stressor affecting arid land leopard frog populations, especially those populations which occupy more ephemeral aquatic habitats. For small isolated systems such

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as earthen cattle tanks, the importance of this factor is likely to be especially important. Our BM study consisted of three such sites. During the summer of 1994, Mesa Tank (BM) dried completely. As the surface water shrank, 154 frogs moved about 250 m (825 ft) upstream to Barnhardt Mesa Tank and 4 moved 900 m (2970 ft) downstream to Unnamed Tank. The following spring, Mesa Tank refilled, but the effects of this drying event likely stressed this population, and in April, 1995, a red-leg epidemic reduced this population to just a few individuals.

We have witnessed other large die-offs which we have attributed to factors than disease. Prior to our second mark-recapture visit to Pinery Canyon in June 1994, frogs began dying at HW. We sampled dead and moribund frogs and water, and had both assayed for bacteria which cause red-leg. These results indicated that disease was an unlikely cause of death (Arizona Veterinary Diagnostic Lab unpubl. data). However, levels of hydrogen sulfide in the water were high enough to be toxic to aquatic wildlife (Herman and Meyer 1990). We suspect that a high detritus load in the pond, coupled with lowering of water level, high water temperature, and low concentrations of dissolved oxygen, combined to form an anoxic environment suitable for proliferation of sulphur producing bacteria. This event reduced a population of 60-80 adult frogs to fewer than 10.

Disturbances such major floods also affect Arizona leopard frog populations. Following the installation of a livestock exclosure at TC, the open water habitats, the preferred habitats of leopard frogs, became chocked with vegetation. In January 1993, a major scouring flood impacted the TC study site. This flood removed a great deal of sediment and increased the open water habitats, and may be partly responsible for the increase in numbers of frogs we saw at the end of our study.

Considering that nearly half of our study populations went extinct while they were under study, the probability of extinction of Arizona leopard frog populations appears high. These populations are vulnerable to large-scale mortality on a frequent basis by the causative factors outlined above and others. Mortality may be density-independent (e.g. floods or sulphur toxicity) or density-dependent (e.g. red-leg). However, the different mechanisms share a common result: potentially disastrous mortality with possible extinction of local populations and destabilization of metapopulations.

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LITERATURE CITED

Brown, D.E. 1994. Warm temperate grasslands. Pages 123-131 in D.E. Brown, editor. Biotic communities: southwestern United States and northwestern Mexico. University of Utah Press, Salt Lake City.

Carey, C. 1993. Hypothesis concerning the causes of the disappearance of boreal toads from the mountains of Colorado. Conservation Biology 7:355-362.

Carr, A.H., R.L. Amborski, D.D. Culley, Jr., and G.F. Amborski. 1976. Aerobic bacteria in the intestinal tracts of bullfrogs (Rana catesbeiana) maintained at low temperatures. Herpetologica 32:239-244.

Clarkson, R.W. and J.C. Rorabaugh. 1989. Status of leopard frogs (Rana pipiens complex: Ranidae) in Arizona and southeastern California. Southwestern Naturalist 34(4):531-538.

Herman R.L. and F.P. Meyer. 1990. Fish kills due to natural causes. Pages 41-44 in F.P. Meyer and L.A. Barclay, editors. Field manual for the investigation of fish kills. US Fish and Wildlife Service Resource Publication 177, Washington, DC.

Hillis, D.M. 1988. Systematics of the Rana pipiens complex: puzzle and paradigm. Annual Review of Ecology and Systematics 19:39-63.

Jennings, M.R. and M.P. Hayes. 1994. Decline of native ranid frogs in the desert Southwest. Pages 183-211 in P.R. Brown and J.W. Wright, editors. Herpetology of the North American Deserts: proceedings of a symposium. Southwestern Herpetologists Society, Van Nuys, CA.

Martof, B.S. 1953. Territoriality in the green frog, Rana clamitans. Ecology 34:165-174.

Mecham, J. S. 1968. Evidence of reproductive isolation between two populations of the frog, Rana pipiens, in Arizona. Southwestern Naturalist 13:35-44.

Meffe, G.K. and W.L. Minckley. 1987. Persistence and stability of fish and invertebrate assemblages in a repeatedly disturbed Sonoran Desert system. American Midland Naturalist 117:177-191.

Moore, J.A. 1944. Geographic variation in Rana pipiens Schreber of eastern North America. Bulletin of the American Museum of Natural History 82:345-370.

National Climatic Data Center. 1997. Monthly precipitation data for U.S. cooperative & NWS sites. [http://www.ncdc.noaa.gov].

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Pechmann, J.H.K., D.E. Scott, R.D. Semlitsch, J.P. Caldwell, L.J. Vitt, and J.W. Gibbons. 1991. Declining amphibian populations: the problem of separating human impacts from natural fluctuations. Science 253:892-895.

Platz, J.E. 1976. Biochemical and morphological variation of leopard frogs in Arizona. Copeia 1976:660-672.

Platz, J.E. 1993. Rana subaquavocalis, a remarkable new species of leopard frog (Rana pipiens complex) from southeastern Arizona that calls under water. Journal of Herpetology 27:154-162.

Platz, J.E. and J.S. Frost. 1984. Rana yavapaiensis, a new species of leopard frog (Rana pipiens complex). Copeia 1984:940-948.

Platz, J.E. and J.S. Mecham. 1979. Rana chiricahuensis, a new species of leopard frog (Rana pipiens complex) from Arizona. Copeia 1979:383-390.

Platz, J.E. and A.L. Platz. 1973. Rana pipiens complex: hemoglobin phenotypes of sympatric and allopatric populations in Arizona. Science 179:1334-1336.

Pollock, K.H., J.D. Nichols, C. Brownie, and J.E. Hines. 1990. Statistical inference for capture-recapture experiments. Wildlife Monographs No.107:1-97.

Post, D.D. and D. Pettus. 1966. Variation in Rana pipiens (Anura: Ranidae) of eastern Colorado. Southwestern Naturalist 11:476-482.

Reed, G.B. and G.C. Toner. 1942. Proteus hydrophilus infections of pike, trout, and frogs. Canadian Journal of Research 20 D:161-166.

White, G.C., D.R. Anderson, K.P. Burnham, and D.L. Otis. 1982. Capture-recapture and removal methods for sampling closed populations. Los Alamos National Laboratory, Los Alamos, New Mexico. 1-235 pp.

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VALIDATION OF VISUAL ENCOUNTER SURVEYS

Jeffrey M. Howland, Michael J. Sredl, and J. Eric Wallace Nongame Branch, Wildlife Management Division

Arizona Game and Fish Department

Technical Report 121 Nongame and Endangered Wildlife Program

Program Chief: Terry B. Johnson Arizona Game and Fish Department

2221 West Greenway Road Phoenix, Arizona 85023-4399

June 1997

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RECOMMENDED CITATION

Howland, J. M., M. J. Sredl, and J. E. Wallace. 1997. Validation of visual encounter surveys. Pages 21-36 in M.J. Sredl, editor. Ranid frog conservation and management. Nongame and Endangered Wildlife Program Technical Report 121. Arizona Game and Fish Department, Phoenix, Arizona.

ACKNOWLEDGMENTS

We would like to thank the many individuals who contributed to the development and implementation of this project. AGFD personnel involved: Carl Lutch provided site specific leopard frog observations and other information of the area which was essential to the project. Sharon Seim provided input with original study design, Loralei Saylor aided with database design, and John Windes assisted with all the field work, data entry, and ideas along the way. Bruce Christman helped with field work. Rhonda O'byrne of Tonto National Forest provided information regarding the USFS acquisition of water rights for 8 of the 9 tanks. Austin Haught, a long time local resident and rancher provided insight into the ranching history of the area and absolute cooperation with access. Anna Mae Peace provided rainfall data. Brenda Healy and Verna Miera made comments which substantially improved this report. Finally, we thank Randy Babb for donating the cover illustration.

PROJECT FUNDING

Funding for this project was provided through: the Arizona Game and Fish Department's Heritage Fund; voluntary contributions to Arizona's Nongame Wildlife Checkoff; U.S. Fish and Wildlife Service Project E5, Jobs 12 and 26, under Title VI of the Endangered Species Act; U.S. Fish and Wildlife Service Partnerships For Wildlife project, Job 02, administered by the National Fish and Wildlife Foundation.

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VALIDATION OF VISUAL ENCOUNTER SURVEYS

Jeffrey M. Howland, Michael J. Sredl, and J. Eric Wallace

INTRODUCTION

The visual encounter survey (VES), as described by Crump and Scott (1994), is one of the most commonly employed techniques used for determination of amphibian abundance and population status. This is due mainly to its simplicity: 1) a VES involves very little equipment, 2) can be carried out by one person, and 3) involves little initial training (depending on ease of positive field identification of species of interest). In this technique, field personnel move systematically through a designated habitat and record visual observations of target species.

Accurate and precise determination of leopard frog population size requires use of mark-recapture methods (Donnelly and Guyer 1994). These studies are time and resource intensive and provide information that is often specific to the study site. Such studies provide estimates that allow determination of population status through tracking of its size through time. VES may provide an index of abundance, though influenced by a variety of factors ranging from diel and seasonal activity patterns, to weather, to observer bias. Because we are responsible for conservation and management of populations statewide, we need survey methods that will allow determination of status of large numbers of populations on a geographically extensive basis. This need led us to attempt to evaluate precision and bias of our leopard frog VES through comparison of VES abundance indices to population estimates derived at the same sites through mark-recapture studies.

Further incentive to evaluate and refine the YES technique for use on Arizona leopard frogs came in 1994, when the North American Amphibian Monitoring Program was developed to address concerns of North American amphibian declines. The primary goal of this endeavor was to: "Develop a statistically defensible program to monitor the distributions and abundance of amphibians in North America, with applicability at the state, provincial, ecoregional, and continental scales." (North American Amphibian Monitoring Program 1996).

Ideally, we would like to determine the relationship between abundance indices derived from VES (conducted at many sites statewide) and true population size (as determined at a small number of sites through intensive mark-recapture population studies; see Davis and Winstead (1980) for discussion of population indices and estimates). It is important not only to determine accuracy (e.g. sources and direction of bias), but also to determine the precision (e.g. confidence limits on population estimates) of the VES method by attempting to discern and then control variables that significantly affect counts.

Building on preliminary work (Wallace and Sredl 1995) we attempted to evaluate and refine our VES technique, by carrying carried out repeated VES at ten aquatic sites within a relatively small

23

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geographic area. The selected VES sites included three sites where a mark-recapture study on lowland leopard frogs (Rana yavapaiensis) was in progress. Our intent was to make comparisons of the results of the two approaches to determine the nature of any relationship between them and the extent to which we might rely on extensive use of VES to monitor population status of native ranid frogs on a statewide basis. A variety of biological and environmental factors combined to make these determinations impossible (see Results), but in the course of our efforts we made a number of important observations on natural history and ecology of leopard frogs and phenology of their stock tank habitats.

METHODS

Site Description and History: Our study area lies in Tonto Basin, on the eastern slopes of the Mazatzal Mountain foothills, between 963-1231 m (3178-4062 ft) elevation. This region is characterized as semi-desert grasslands (Brown 1994). It is bordered by Great Basin Conifer Woodland and Interior Chaparral above and Arizona Upland Subdivision of the Sonoran Desert below. Historically, vegetative cover was dominated by native perennial bunch grasses (e.g. grama grasses, Bouteloua spp.), but with the onset of intensive grazing in the 1880s and the subsequent reduced incidence of fire there has been a dramatic shift to a shrub/scrub dominated landscape (= disclimax grassland) (Brown 1994; Croxen 1926).

Climatically this region is characterized by seasonal drought and drying winds with extended drought periods not uncommon. Rainfall is distributed fairly evenly between winter precipitation (December-March) and warm-season rainfall (July-September) with May-June being the driest months (Brown 1994; National Climatic Data Center 1997).

The study area is somewhat centrally located within the known range of the lowland leopard frog, Rana yavapaiensis (Stebbins 1985). Specific aquatic sites sampled were nine man-made earthen tanks and one 400 meter intermittent stretch of Rye Creek (into which all sites drain) (see Table 1 for site names, numbers and locations). Perennial water is available elsewhere in Rye Creek and in Tonto Creek (of which Rye Creek is a tributary). Study area selection was based on recent observations of robust populations of lowland leopard frogs at several of the sites (C. Lutch pers. comm.; AGFD unpubl. data) and the fact that a mark-recapture study was already in progress at VES07-09 (Fig. 1; see mark-recapture chapter for results).

All tanks lie within natural drainages and are impounded by earthen berms. Estimated surface area of tanks at capacity ranges from -220 - 1500 m2 (2367 - 16,137 ft2) with depth ranging from "0.4 - 2.7 m (1.3 - 8.9 ft). These variables, combined with position in watershed, result in different tanks retaining water for differing periods of time after capture of runoff. In most years, at least 5 of the 9 tanks hold some water throughout the year (A. Haught pers. comm.). All tanks were built prior to 1955 (R. O'byrne pers. comm.).

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VES Validation: We designed a standard VES, based on our statewide surveys, that was simple, repeatable, and generated information from which inferences about abundance could be made.

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Figure 1. Map of VES study sites modified from Gisela (1972) and Mazatzal Peak (1972) U.S.G.S. 7.5" topographic maps. Numbers are as in Table 1.

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Upon approaching a tank, we glassed the site with binoculars from "20 m, searching for and recording frogs floating in water away from the bank. We then proceeded around the entire perimeter of tanks, or along 400 meters of the Rye Creek site, recording any additional frogs seen or heard. While walking along banks, we used long handled dipnets to sweep vegetation to flush frogs that did not respond to our approach. After the initial perimeter survey, we searched mud cracks and divots formed by cattle hooves. Counts included visual observations and audible "plops" of frogs escaping into water.

In addition to counts of frogs, we recorded: time and date, air and water temperature, water pH and conductivity, and relative humidity. We also measured habitat characteristics, including: water clarity (scale of 1-5), vegetation types present (submerged, emergent, perimeter, floating, canopy), and soils (clay, sand, gravel, cobble). Local weather conditions (e.g. cloud cover, precipitation, wind velocity) and sign of potential vertebrate and invertebrate predators were also noted. We dipnetted intensively about every ten meters to determine presence of amphibian larvae and/or invertebrate predators. Once each trip, we measured two preset axes of each tank to estimate current surface area of available water. Water depth was recorded from graduated poles set in the deepest part of each tank. Photographs were taken at a preset point to document physical changes in the habitat throughout the year.

Table 1. VES study site names, numbers, legal descriptions and elevations.

Site Number Legal Description

Elevation

m ft

Bear Creek Tank VES01 T8N, R9E, Sec. 1, NW% of SW% 1036 3419

Shake Ridge Tank VES02 T8N, R9E, Sec. 3, NW% of NE% 1231 4062

Table Top Tank VES03 T8N, R9E, Sec. 2, NE% of NW% 1097 3620

Suicide Tank #1 VES04 T9N, R9E, Sec. 36, NVV% of SW% 1018 3359

Suicide Tank #2 VES05 T9N, R9E, Sec. 36, NW% of SW% 1018 3359

Lower Barnhardt Tank VES06 T9N, R10E, Sec. 30, SW% of NE% 1000 3300

Unnamed Tank VES07 T9N, R9E, Sec. 24, NW% of SE% 1082 3571

Mesa Tank VES08 T9N, R9E, Sec. 24, SVV1/4 of SVV% 1127 3719

Barnhardt Mesa Tank VES09 T9N, R9E, Sec. 24, SW% of SW% 1143 3772

Rye Creek VES10 T9N, R10E, Sec. 18, S% of SE% 963 3178

On each sampling day, VES surveys were conducted at all 10 sites by two surveyors. The ten sites were numbered (Table 1) and surveyed in ascending or descending order (direction chosen by coin

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a)

1.1 Apr 95 J.696 Aug 95 0065 0.6 66 FGO 96 *95 96 A65;96 609 96 ace 06566 F6067

b) M

axim

um sur

face

are

a (m

2)

1200

900

1003

GOO

400

so

50

40

A00

20

Data (mmm yy)

Figure 2. Tank surface area and rainfall. a) Surface area per trip. b) Total monthly rainfall for Barnhardt Mesa. Symbols as follows: Bear Creek Tank (0), Shake Ridge Tank (N), Table Top Tank (o), Suicide Tank #1 (.), Suicide Tank #2 (+) Unnamed Tank (0).

Arizona Game and Fish Department June 1997 NGTR 121: Ranid Frog Conservation and Management Page 27

toss each sampling day), with site 1 following 10 in ascending order (vice versa in descending order) with the initial site in the sequence being selected randomly. The first surveyor began sampling between 0730-0900 MST, with the second surveyor beginning at the same initial site approximately two hours later and sampling sites in the same order.

RESULTS

Near the beginning of this project, leopard frogs in the three tank mark-recapture study on Barnhardt Mesa (VES07-09) suffered a massive die-off (see Chapter 1). Frog numbers at most of the other YES sites were lower than prior surveys had indicated as well (C. Lutch pers. comm.; AGFD unpubl. data). At Table Top Tank, for example, 70 adult frogs were observed in May 1993, while the highest count during the study was 18. These reductions in frog numbers made it impossible to thoroughly evaluate precision and bias of our YES counts. However, we present a subset of our data from sites VES01-05,07, and 10 to illustrate some of the important observations that were made during the study (VES06, 08-09 had few or no frogs throughout the study, although populations at VES08-09 were formerly large enough that they were selected as part of our Barnhardt Mesa mark-recapture site: see mark-recapture chapter).

Phenology of Earthen Cattle Tanks: Surface area and depth of available water fluctuated dramatically at all sites. Rainfall during 1995 was 415 mm (16.4 in), slightly lower than the 30-year average of 478 mm (18.8 in) for data collected from 1966-1996. Rainfall in 1996 was 208 mm (8.2 in.) (National Climatic Data Center 1997; A.M. Peace pers. comm.), less than half of normal. During our study, Shake Ridge Tank (Fig. 2a) dried and refilled three times, Suicide Tank #1 and Bear Creek Tank dried and refilled twice, and Unnamed Tank, Suicide Tank #2 and Table Top Tank dried completely only once, in summer 1996. When the study began in spring 1995, all tanks were near capacity, but none returned to these levels until spring 1997. At Rye Creek, surface water was initially present along the entire 400 m reach included in our surveys, but by

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October 1995, surface water was available only as scattered, small pools. The system eventually dried, and remained so until spring 1997, when flow was restored by late winter and spring rains (Fig 2b).

Count Results: Our counts of frogs fluctuated throughout the study, yet a few general trends were noted (Fig. 3). All counts presented in Figure 3 for each tank are the highest of all counts (usually two to six counts) for the given sampling period (usually a month). Counts at tanks decreased during August 1995, the driest period of that summer. After summer thunderstorms refilled tanks during the latter part of August, at all tanks dropped to 1. Counts remained Suicide Tank #1 did counts subsequently excee at any tank (Fig. 3).

30

S 20

10

0 Apr 95 Jun 95 009 95 Oehr5 Ow 96 Fob 96 Apr 96 J3r7915 A496 NC 95 Feb 97

Date (mmrn yy)

Figure 3. Maximum counts of Rana yavapaiensis per trip from April 1995-March 1997 at six sites (VES01- 05, 07). Vertical dashed line represents trip in which Rye Creek (VES10) high count equaled 80 newly metamorphosed frogs (Symbols as in Figure 2).

counts increased at all tanks. By December, counts low to zero for the remainder of the study. Only at d two, and after July 1996, no frogs were observed

In spite of universally low counts in 1995 (preliminary counts at the sites prior to this project were considerably higher than any counts recorded during the study), there were clear and consistent differences in counts among some of the sites. With the exception of August 1995, when several sites dried, counts for Suicide Tank #1 and Table Top Tank were consistently higher than counts at Bear Creek Tank and Shake Ridge Tank, while counts at Suicide Tank #2 began high and never recovered after the August 1995 drying. Conversely, 1995 counts at Rye Creek and Unnamed Tank increased from zero or one frog in July to large numbers of metamorphs in August. Counts then dropped through late summer and fall and no frogs were observed at either site from December 1995 through completion of the project. Follow-up visits in February and March 1997 yielded no frog observations at any of the study sites.

Count values of 0 were uncommon at sites that were known or suspected to have frogs, except in winter. For April through October of 1995, when frog populations were small to moderate in size at Bear Creek Tank, Shake Ridge Tank, Table Top Tank, and the two Suicide tanks, we recorded a cumulative total of only 16 counts of 0 from a total of 124 surveys (12.9%). Seven of the 16 were from Shake Ridge Tank, whose frog population was very small (maximum count of 4 frogs for the entire study). Nine of the 16 zeros were recorded in August, when four of the six tanks went completely dry, and one was recorded on October 26, at the very end of the activity season. For this same set of tanks, frogs persisted through the winter and were observed again in the 1996 active season, yet for the period of November 1995 through March 1996, we recorded 83 counts of 0 from a total of 110 surveys (75.5%).

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We tested for observer bias in counts by comparing paired counts by the two observers at each site on each sampling day to determine whether they differed consistently. In order to avoid large numbers of zero counts, we used only counts from April through October (for both 1995 and 1996) at sites VES01-05,07, and 10. Of 73 paired counts, the two observers had the same count 23 times, observer #1 had the higher count 34 times, and observer #2 had the higher count 16 times. A Wilcoxon signed-ranks test showed that this inter-observer variation was significant (Z = -2.528; P — (2-tailed) = 0.011). At the beginning of the study, observer #1 (higher counts) had about 1.5 field seasons of experience with our VES technique (or similar, earlier versions). Observer #2 had a full field season of leopard frog survey experience in Arizona, but not with our survey protocol.

Anecdotal Observations: Although our original design was somewhat disrupted by drought and frog population declines, repeated visits to the same sites over two years provided some interesting and important incidental observations on the behavior and ecology of lowland leopard frogs. In August 1995, Shake Ridge Tank was dry, yet we observed frogs in contact with wet soil in mud cracks and cattle hoof prints. Use of these types of microhabitats was also observed at other tanks that were not completely dry.

Twice we observed frogs at locations remote from study sites, near the Suicide tanks. One frog was observed at Clover Well (Fig. 1), a pump-fed metal cattle drinker -1.0 km (3300 ft) down drainage from the Suicide tanks. The other was observed in a rain pool in a normally dry wash -0.2 km (660 ft) overland from any tank.

DISCUSSION

With the onset of ranching in AZ in the late 1800s and early 1900s, construction of artificial water catchments for livestock, generally in small upland drainages, became common. Many of these waters provided permanent or semi-permanent aquatic habitat that could be colonized by leopard frogs from naturally permanent riparian areas lower in the drainages. As most natural systems were still relatively intact at this time, the supplementary aquatic habitat provided by stock tanks may actually have allowed native frog populations to increase for a time. Eventually, as natural aquatic habitats were altered, degraded, or lost due to surface water diversion, ground water pumping, livestock grazing practices, introduction of non-native predators and competitors, and other causes, cattle tanks became essential refugia for leopard frogs, and we contend that they should be managed as such.

Phenology of Stock Tanks: Earthen stock tanks are generally filled by surface runoff from rainfall or snowmelt. Permanence of water in a given cattle tank is dependent upon factors such as size of the tank (maximum depth and surface area), permeability of substrate, and size of watershed from which it receives runoff. Other factors affect evaporative water loss. Aspect of the tank and

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surrounding topography may affect exposure to wind and solar radiation. Species composition and density of bank and emergent vegetation affect exposure of water surface to sun and wind and also influence evapotranspirational water loss.

Frequent drying and refilling of a tank such as Shake Ridge Tank is due to its size and location within the watershed. It had the smallest surface area of all the study tanks and was furthest up in a drainage (thus fed by a small watershed). This tank dried after any substantial period of low rainfall and quickly refilled after rain. Suicide Tank #1 filled quickly with rain, but was relatively quick to dry due to its shallowness. In contrast, Table Top Tank is large and deep and holds water throughout the year, at least during periods of normal rainfall (A. Haught pers. comm). These examples illustrate the dynamic and sometimes unstable nature of these habitats, which are now, from an applied conservation perspective, perhaps the best and most important leopard frog habitats available throughout much of Arizona. The more permanent tanks seem to host more stable leopard frog populations. Management efforts should consider this as .an important factor in priority ranking sites for protection or as potential sites for reintroduction or other management actions. We should also consider modification of tanks to enhance permanence as a management tool for leopard frogs.

Counts: The August 1995 reduction in counts is easily explainable. As available surface water and aquatic habitat became scarce along with increasing heat and aridity, frogs reduced activity while seeking refuge, thus becoming less detectable by our survey techniques. Increased counts in September 1995 were due to influx of water from summer rains. Decreased counts in November 1995 through March 1996 were presumably due to reduced activity with low winter temperatures. When spring rains came and temperatures rose by April 1996, the failure of counts to return to the previous year's level was unexpected. Only Suicide Tank #1 ever had a count >2, and its counts, along with all other sites, dropped to zero by September. We also failed to find frogs in spring 1997. We strongly suspect that our counts reflect a local extinction event throughout the study area, caused, at least in part, by drying of nearly all available aquatic habitat in an unusually dry year. Based on Corn and Fogelman (1984) we would need to support this contention with return visits which yielded zero counts over one full reproductive cycle ("2-3 yrs).

We suspect that observed counts are a true reflection of leopard frog abundance, at least for populations in lightly vegetated stock tanks. Unfortunately, the die-off that occurred at our mark-recapture sites just prior to beginning this study makes it impossible to evaluate the accuracy and precision of VES counts as an indicator of actual population size. Nevertheless, we think the consistent differences in counts among some of the sites (consistently higher counts at Table Top Tank and Suicide tanks #1 and #2; consistently low counts at Bear Creek Tank and Shake Ridge Tank) reflect real differences in population size among the sites. However, counts alone may be misleading. At Unnamed Tank and Rye Creek, high counts in July and August (Fig. 3; AGFD unpubl. data) were due solely to pulses of metamorphosis (see further discussion below).

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For several years, the meaning, importance, and reliability of surveys resulting in frog counts of zero has been uncertain. An answer to the question of how to interpret these zero counts has particular relevance to the overall results of our statewide surveys. Because of limited resources, we are forced to priority rank site visits to emphasize sites that are known present or former habitat for leopard frogs or that have high potential to host leopard frog populations. Given the large number of potential new sites that have never been visited and the number of known sites that need to be monitored, we have chosen not to revisit most sites where one or two initial visits have failed to find frogs. We have questioned whether failure to find frogs in one or two attempts really means that frogs are not present at the site. The results of our intensive, repeated VES in this study provide an excellent data set for evaluating the meaning of zero counts.

With the exception of periods of extreme drying (e.g. August 1995), zero counts were rare during April through October, the primary activity season. Even at the five study sites analyzed, where frog populations were relatively small, we detected their presence in 93 of 100 (93%) VES conducted during the daytime in April through October 1995, excepting August 8-10 when most aquatic habitat dried. Our interpretation of this result is that zero counts in our VES really are significant if surveys are conducted during the activity season when habitats are not dry or nearly dry. For surveys conducted under such conditions, we have a fairly high level of certainty that if frogs are present at a site, even in low numbers, we will detect them. Conversely, if we fail to detect frogs, they are probably not present, at least not in substantial numbers. This high level of detectability could probably be increased by restricting surveys to periods favorable for frog activity as measured by such simple factors as air and water temperature, cloud cover, and wind speed.

This result greatly increases our confidence in the reliability of negative results recorded during our statewide surveys. However, we must be careful not to over-extrapolate from the results of the present study, which was conducted at simple stock tank habitats lacking heavy vegetation (i.e. frog visibility was high). This is, however, a common habitat type for native ranids throughout much of Arizona. In addition to limiting extrapolation to other habitats, the activity season should be adjusted appropriately. For example, the activity season for frogs at higher elevations, such as the White Mountains and Mogollon Rim, would be somewhat shorter than the April through October season we use in the present study.

As an interesting side note on count results, prior to the mass metamorphosis of tadpoles that occurred at Unnamed Tank and Rye Creek in July and August, we recorded six counts of zero at Unnamed Tank (out of six surveys) and two counts of one at Rye Creek (out of two surveys). These sites apparently had very small populations of adult frogs. Eight tadpoles were observed in late spring surveys of Unnamed Tank and large numbers (estimated at over 300) were seen in Rye Creek during July 1995 surveys. The pulse of metamorphs witnessed in August (high count of 18 frogs at Unnamed Tank and 80 at Rye Creek), coincident with low counts at all other sites, probably resulted from stimulation of metamorphosis by drying of habitats (Werner 1986). The

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subsequent decrease in counts was probably due to mass emigration or in situ mortality when habitat dried (completely at Rye Creek, partially at Unnamed Tank). Metamorphosis events were not observed at any of the other tanks. Because metamorphs often disperse and have poor survivorship, and because large numbers of metamorphs can result from only one or a very few egg masses (data on size of R. yavapaiensis egg masses are lacking, but egg mass size in other leopard frogs ranges from a few hundred to several thousand eggs, Stebbins 1985), high counts of metamorphs should not, by themselves, be interpreted as evidence of a healthy population. In spite of large numbers of metamorphs, neither Unnamed Tank nor Rye Creek ever had frogs after early November 1995, making them apparently the first two of the seven primary study tanks to die out (the other five sites all had frogs persisting into spring 1996). Successful recruitment is certainly important, but a stable and healthy population must also include reasonable numbers of mature animals.

Our comparison of paired counts by two observers demonstrates inter-observer bias as an important determinant of frog counts. This occurred in spite of the fact that both observers were well-trained and experienced in leopard frog surveys in Arizona. We can only speculate on causes of the observed bias. As noted above, observer #1 had more experience with our particular VES protocol than observer #2, and this may have been a factor in the higher counts recorded by observer #1, but a sample size of one comparison is inadequate to make any conclusions. Individual abilities such as hearing acuity, ability to orient toward auditory stimuli, visual acuity, peripheral vision, and ability to concentrate or focus may all contribute to individual differences in aptitude. Except by making protocols as simple and repeatable as possible (which we already do), these types of factors are difficult to control through study design, so perhaps we should focus on training and experience. Our results indicate that it may be important to use well trained and experienced personnel to conduct surveys. Unfortunately, we are aware that many survey projects of this type are funded and administered in such a way that personnel are hired on a seasonal basis, and annual turnover in personnel may approach 100%. Under this type of project structure, it is often difficult to maintain a highly trained and experienced group of field biologists. We have been fortunate in having a diversity of funding sources that have allowed us to employ individual biologists for years at a time, and we think this has enhanced our ability to sustain high quality field projects.

Anecdotal Observations: The anecdotal observations we made shed some light on leopard frog behavior in stock tank habitats. The present importance of these sites to leopard frog conservation makes it important for us to understand how the habitats are utilized, so that we may manage them more effectively. Historically, lowland leopard frogs occupied perennial wetland habitats such as springs, seeps, cienegas, streams, and rivers. Harsh surrounding uplands restricted the frogs to these permanent aquatic sites. Unlike many other anurans of the desert southwest, leopard frogs are obligate wetland species (Stebbins 1985). They lack adaptations that allow others to endure long periods of drought. Spadefoot toads (Scaphiopus spp.), for example, remain underground during extended dry periods, reducing metabolic rate and protected from water loss by a layer of

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keratinized skin (Mayhew 1965). Our observations of frogs in mud cracks demonstrate at least some behavioral adaption to short periods of drying. We suspect they may also use rodent burrows and other microhabitats that provide shelter from heat and desiccation.

Our observations of frogs in temporary waters near known habitats provide evidence of dispersal ability that would allow colonization of somewhat remote sites (see discussion of Barnhardt tanks in mark-recapture chapter). That we found frogs at Shake Ridge Tank attests to their ability to disperse, as this tank is up a steep narrow drainage -1.0 air mile from the nearest known perennial water (Table Top Tank, which is downstream). Similar dispersal ability has been noted for R.

blairi (Frost and Bagnara 1977).

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LITERATURE CITED

Brown, D.E. 1994. Warm Temperate Grasslands. Pages 123-131 in D.E. Brown, ed. Biotic Communities: Southwestern United States and Northwestern Mexico. University of Utah Press, Salt Lake City.

Corn, P.S. and J.C. Fogleman. 1984. Extinction of montane populations of the northern leopard frog (Rana pipiens) in Colorado. Journal of Herpetology 18:147-152.

Croxen, F.W. 1926. History of grazing on Tonto. Tonto Grazing Conference. Phoenix, Arizona, November 4-5, 1926.

Crump, M.L. and N.J. Scott, Jr. 1994. Standard techniques for inventory and monitoring-visual encounter surveys. Pages 84-92 in W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, M.S. Foster, editors. Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, DC.

Donnelly, M.A. and C. Guyer. 1994. Estimating populations size - mark-recapture. Pages 183-200 in W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, M.S. Foster, editors. Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, DC.

Davis, D.E. and R.L. Winstead. 1980. Estimating the numbers of wildlife populations. Pages 221-245 in S.D. Schemnitz editor. Wildlife management techniques manual. The Wildlife Society, Washington, D.C.

Frost, J.S. and J.T. Bagnara. 1977. Sympatry between Rana blairi and the southern form of leopard frog in southeastern Arizona (Anura: Ranidae). Southwestern Naturalist 22(4):443-453.

Mayhew, W.W. 1965. Adaptations of the amphibian Scaphiopus couchii to desert conditions. American Midland Naturalist 74: 95-109.

National Climatic Data Center. 1997. Monthly precipitation data for U.S. cooperative & NWS sites. [http://www.ncdc.noaa.gov].

North American Amphibian Monitoring Program 1996. Protocols and strategies for monitoring North American amphibians. (Unpubl.).

Stebbins, R.C. 1985. A field guide to western reptiles and amphibians. Houghton Mifflin company, Boston Massachusetts.

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Wallace, J.E. and M.J. Sredl. 1995. Preliminary evaluation of visual encounter surveys for southwestern leopard frogs. Annual Meeting of the Southwestern Working Group of the Declining Amphibians Population Task Force. Phoenix, Arizona, January 5-6, 1995.

Werner, E.E. 1986. Amphibian metamorphosis: growth rate, predation risk and the optimal size transformation. American Naturalist 128:319-341.

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STATUS AND DISTRIBUTION OF ARIZONA'S NATIVE RANID FROGS

Michael J. Sredl, Jeffrey M. Howland, J. Eric Wallace, and Loralei S. Saylor Nongame Branch, Wildlife Management Division

Arizona Game and Fish Department

Technical Report 121 Nongame and Endangered Wildlife Program

Program Chief: Terry B. Johnson Arizona Game and Fish Department

2221 West Greenway Road Phoenix, Arizona 85023-4399

June 1997

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RECOMMENDED CITATION

Sredl, J.M. Howland, J.E. Wallace, and L.S. Saylor. 1997. Status and distribution of Arizona's native ranid frogs. Pages 37-89 in M.J. Sredl, editor. Ranid frog conservation and management. Nongame and Endangered Wildlife Program Technical Report 121. Arizona Game and Fish Department, Phoenix, Arizona.

ACKNOWLEDGMENTS

For assistance with field work and logistical support, we thank: Linda Allison, Maggy Bathory, Rob Brauman, Bill Burger, Ed Burns, Mike Burroughs, Dave Carrothers, Cheryl Carrothers, Bruce Christman, Pat Collins, Brian Dickey, Cecilia Dargan, Tracy Ertz-Berger, Dan Groebner, Matt Goode, Greg Goodwin, Morgan Heath, Tom Hildebrandt, Jacques Hill, Ron Hill, Wendy Hodges, Mike Ingraldi, Angie Kiselyk, Chris Klug, Susie MacVein, Keith Menasco, Rick Miller, Charlie Painter, Bruce Pavlick, Don Pollack, Mike Pruss, Natalie Robb, Ben Robles, Mike Ross, Kelly Schwartz, Sharon Seim, Farraday Sredl, Bob Vahle, Dana Waters, Dave Weedman, Mark Whitney, and John Windes. We thank Randy Babb, Kurt Bahti, Joseph Bagnara, Dave Bradford, Tom Britt, Jerry Bradley, Rob Clarkson, Kevin Cobble, Jim Collins, Troy Corman, Bob Csargo, Dave Dorum, Charles Drost, Rick Dreyer, Philip Fernandez, Mary Franks, Glen Frederick, John Frost, Eric Gardner, Stephen Hale, Randy Jennings, Hans Koenig, Thomas Jones, Chris Klug, Tom Liles, Mike Lopez, Carl Lutch, Debbie Lutch, Matthew Magoffin, Vir McCoy, Jack Meyer, Terry Myers, Tom Newman, Charlie Painter, David Parizek, Bruce Palmer, James Platz, James Rorabaugh, Philip Rosen, Cecil Schwalbe, Joan Scott, Sheridan Stone, Brian Sullivan, Luke Thirkill, Barney Tomberlin, Jim Vial, Dana Waters, John Windes, Tom Wood, Dale Ward, and Richard Zweifel for providing records of Arizona leopard frogs. Terry Johnson provided advice, assistance, and support in development of conservation and management planning and implementation. Brenda Healy and Verma Miera made comments that improved the manuscript. Finally, we thank Randy Babb for donating the cover illustration.

PROJECT FUNDING

Funding for this project was provided through: the Arizona Game and Fish Department's Heritage Fund; voluntary contributions to Arizona's Nongame Wildlife Checkoff; U.S. Fish and Wildlife Service Project E5, Jobs 12 and 26, under Title VI of the Endangered Species Act; U.S. Fish and Wildlife Service Partnerships For Wildlife project, Job 02, administered by the National Fish and Wildlife Foundation; several Challenge Cost-Share Agreements with USDA Forest Service (Apache-Sitgreaves, Coconino, Kaibab, and Tonto national forests); Purchase Order 43-8167-5-0291 from Coconino National Forest; and contract #DABT63-95-P-2237 with U.S. Department of Defense, Ft. Huachuca.

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STATUS AND DISTRIBUTION OF ARIZONA'S NATIVE RANID FROGS

Michael J. Sredl, Jeffrey M. Howland, J. Eric Wallace, and Loralei S. Saylor

INTRODUCTION

As currently recognized, Arizona's ranid frog fauna includes the Tarahumara frog (Rana tarahumarae) and six or seven species of leopard frogs. This leopard frog fauna is surprisingly diverse and is amongst the largest in North America (Platz et al. 1990), including a disjunct population of the plains leopard frog (R. blairi), the southwestern end of the distribution of the northern leopard frog (R. pipiens), the northern end of the distribution of the Chiricahua leopard frog (R. chiricahuensis; soon to be split into two distinct species, a Mogollon Rim form and a southern form), the core of the distribution of the lowland leopard frog (R. yavapaiensis), the only known populations of the Ramsey Canyon leopard frog (R. subaquavocalis), possibly a portion of the small range of the relict leopard frog (R. onca), and a healthy and expanding population of the non-native Rio Grande leopard frog (R. berlandieri). In fact, the only leopard frogs native to the United States not known or suspected to occur in Arizona are the southern leopard frog (R. utricularia) and Vegas Valley leopard frog (R. fisheri).

Amphibian population declines have been reported around the world (Barinaga 1990; Blaustein and Wake 1990; Vial and Saylor 1993). Many factors have been implicated in these declines, including: 1) introduction of exotic organisms (Bradford et al. 1993; Hayes and Jennings 1986; Rosen et al. 1995, 1996); 2) higher levels of UV radiation due to thinning of the ozone layer (Blaustein et al. 1994); 3) acid precipitation and toxic substances (Berri11 et al. 1993; Carey and Bryant 1995; Dunson et al. 1992; Hale and Jarchow 1988); 4) disease (Carey 1993; N.J. Scott, Jr. unpubl. data); 5) destruction and fragmentation of habitats (Bradford et al. 1993; Hedges 1993; Sjogren 1991); and 6) weather, including drought and flooding (Corn 1994).

Interest in amphibian declines has spread beyond the realm of academic herpetologists and conservationists. Because of their basic biology, amphibians may be important indicator species (Vitt et al. 1990) of global environmental health. Pertinent characters include: 1) the biphasic life cycle of most amphibians, aquatic larvae that are usually herbivorous and terrestrial adults that are exclusively carnivorous, exposes them to both air and water borne toxicants that may be concentrated in either plant or animal foods; 2) they occur in habitats ranging from deserts and rain forests, to alpine and subpolar tundra, across the globe (only the marine environment lacks a significant amphibian fauna); 3) amphibians are an important constituent of the energy and nutrient cycles of many ecosystems, often comprising a significant fraction of total biomass (Burton and Likens 1975); 4) their moist, permeable skin leaves amphibians vulnerable to a variety of environmental insults from pollutants (both air and water borne) to UV radiation.

39

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In the western United States, populations of anurans within the true toad (Bufonidae) and true frog (Ranidae) families appear to have been affected to the greatest degree (Corn 1994). Within Arizona, some concern has been expressed over the continued viability of the Arizona toad (Bufo microscaphus), however, recent work indicates persistence of healthy breeding populations where riparian corridors have not been severely altered (Sullivan 1993, 1995). Most recent concern in Arizona involves native ranid frogs, many of which have been extirpated or reported to have declined (Clarkson and Rorabaugh 1989; Hale and Jarchow 1988; Sredl 1993a). While it is unclear exactly when declines of these populations began, they were noticed during population studies of the Tarahumara frog in the late 1970s (Hale 1992; Hale and Jarchow 1988; Hale and May 1983). They also suspected concurrent declines of native leopard frogs, but comparative baseline data were unavailable. Clarkson and Rorabaugh (1989) made the first systematic investigation of the status of southwestern leopard frogs when they surveyed for four species of native leopard frogs at 56 historical and seven new localities from 1983-1987. They concluded that all leopard frogs examined were declining. Our survey expands the scope of this seminal work.

We assessed the current status and distribution of all species of native Arizona ranids by conducting statewide visual encounter surveys (sensu Crump and Scott 1994) of historical and high potential habitats. Because of the confused taxonomic history of this group (Hillis 1988) and the high likelihood that specimens of this complex will be misidentified even in museum collections (Jennings 1994), we used localities from selected sources in the published literature, or local museums whose collections reflect the current taxonomy, to help us establish our historical baseline. In this report, we present results of these surveys and discuss implications relative to conservation and management of native Arizona ranids.

METHODS

Survey Methodology: We surveyed for ranid frogs and other riparian amphibians and reptiles by walking steadily along, around, or in aquatic habitats while looking and listening for herpetofauna activity. To maximize the chance of encounter during surveys, we constantly scanned the shoreline, embankments, or other appropriate areas from one to ten meters from our position as we walked. A long-handled dipnet aided in capture of animals, and the handle was used to comb and probe dense bushes and grasses. We also looked under rocks, logs, and other debris for amphibians and reptiles. Whenever possible, animals were captured, positively identified to species, photographed, and released (or retained as voucher specimens if population was sufficiently large or specimen identification was ambiguous). If animals were not captured or positively identified to species, we assigned an uncertain code to that observation.

Most surveys were conducted during the daytime, between dawn and dusk, from April through October. The length of habitat searched depended on the size of the system surveyed. Shores of stock tanks, small lakes, and springs were searched in their entirety by walking in and out of the

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water whenever possible. The shoreline and banks of larger lakes, streams and rivers were partially searched. For lakes or wetlands that were approximately 8.0 hectares (20 acres) or larger, only a portion of the perimeter was surveyed. Streams and rivers were surveyed by walking a segment of at least 400 meters (one-quarter mile). Inaccessibility or absence of habitat caused some surveys to be curtailed at lesser distances.

Numbers of individuals of target species encountered were recorded as exact numbers when possible. It was often impossible, however, to make an exact count, particularly with larvae and large populations of adults or metamorphs. For non-target amphibians and reptiles, at least presence/absence was recorded. If more than one trip was made around the perimeter of an enclosed body of water (most stock tanks and lakes), encounters with herpetofauna were recorded only during the first walk around the perimeter, unless numbers and individuals encountered were clearly different on the second pass. For linear aquatic systems (small lotic and narrow lentic systems), numbers of herps encountered were recorded while walking one direction only, unless individuals observed were unambiguously different.

Upon completion of a survey, we filled out a Riparian Herp Survey Form (Appendix A) for each site visited. The data form is divided into three sections. The top section on the front of the form is for site-specific locality data, the bottom section is for data relating to herpetofauna observations, and the back of the data sheet pertains to habitat conditions. Depending on site history and survey outcome, as outlined below, we completed the back of the form (conditions data). This section includes atmospheric and aquatic conditions, measures of habitat quality, relative abundance of important predators of target species, and comments on incidence of disease and other impacts which may negatively affect target species. We completed this section for historical localities, regardless of the survey outcome, and for non-historical localities where target species were found. For non-historical localities where target species were not found, we completed only the top (locality data) and bottom (herpetofauna observations) sections. For further explanation of survey form data fields refer to Riparian Herp Survey Form Instructions (Appendix A).

Target Ranid Frogs: Plains Leopard Frog (Rana blain). The geographic range of the plains leopard frog is centered on the central and southern Great Plains, where it is an inhabitant of aquatic habitats in prairie and desert grassland ecosystems (Stebbins 1985). It is found in temporary and permanent aquatic habitats, including streams, stock tanks, irrigation ditches, and playa lakes. In Arizona, this species is restricted to the Sulphur Springs Valley, Cochise Co. (Frost and Bagnara 1977), an area disjunct from other populations of the species by approximately 250 km (C.W. Painter pers. comm.). Specimens have also been collected from Ashurst Lake, Coconino Co., SE of Flagstaff (J. E. Platz pers. comm.), but recent surveys have failed to find them there and it is unclear whether their occurrence at the site was natural.

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Chiricahua Leopard Frog (Rana chiricahuensis). The range of the Chiricahua leopard frog includes a disjunct portion along the southern edge of the Colorado Plateau and headwater drainages to the south in Arizona (the White Mountains and Mogollon Rim) and New Mexico. Frogs in this area, hereafter referred to as Rim form R. chiricahuensis, are soon to be described as a distinct species from those to the south (J.E. Platz unpubl. data). Frogs in the southern portion of the range, hereafter referred to as southeastern form R. chiricahuensis, inhabit much of SE Arizona (drainages of the Madrean Archipelago and surrounding desert grasslands (Sredl and Howland 1995), portions of SW New Mexico, and the Sierra Madre Occidental in Mexico (Platz and Mecham 1979).

Relict Leopard Frog (Rana onca). The relict leopard frog has the dubious distinction of being the first North American amphibian thought to have gone extinct (Platz 1984) and be rediscovered (Jennings et al. 1995). Historically, it was known from the Virgin River drainage of southern Utah and Nevada. Presently, there are three known extant populations near the Overton Arm of Lake Mead, and two recently discovered populations S of Hoover Dam in springs adjacent to the Colorado River (D.F. Bradford pers. comm.), all in Nevada. No Arizona records of this species exist. However, clarification of the relationships of lowland leopard frog populations near Littlefield, Arizona and the "Lake Mead frogs" may result in the former being re-classified as conspecific with the latter (Jennings et al. 1995).

Northern Leopard Frog (Rana pipiens). Until the late 1960s, all currently recognized leopard frogs, including those found throughout the Southwest, were classified as Rana pipiens. At that time, the range of "R. pipiens" stretched from NE North American to Central America. During the 1960s, taxonomists realized that "R. pipiens" was really a multispecies complex (Hillis 1988). Since that time, numerous distinct species within this complex have been described (Mecham et al. 1973; Platz 1993; Platz and Frost 1984; Platz and Mecham 1979). As currently recognized, R. pipiens, the northern leopard frog, ranges from Newfoundland and New England across the northern plains and southern boreal forests to the Canadian Rockies and south across the Great Basin and Colorado Plateau, occurring in a wide variety of aquatic habitats at elevations from sea level to 3350 m (11,000 ft). Both local and geographically extensive declines, as well as recoveries, have been reported for this species in many parts of its range (Corn and Fogleman 1984; various authors in Bishop and Pettit 1992).

Ramsey Canyon Leopard Frog (Rana subaquavocalis). The Ramsey Canyon leopard frog is the most recently described Arizona leopard frog (Platz 1993). It is known only from the Huachuca Mountains in SE Arizona.

Tarahumara Frog (R. tarahumarae). The Tarahumara frog inhabits seasonal and permanent rock bound pools and bouldery streams in the Sierra Madre Occidental and its foothills in Mexico and three mountain ranges in extreme south-central Arizona. Coincident with extirpation in Arizona (see Results), some populations in northern Sonora also declined. Periodic surveys since that time

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have failed to find any Tarahumara frogs in Arizona (Hale 1992; Hale and Jarchow 1988; Hale and May 1983). Recent reports indicate that some northern Sonora populations may have rebounded.

Lowland Leopard Frog (Rana yavapaiensis). Distribution of the lowland leopard frog is mainly in Arizona, with populations formerly found in SE California, SW New Mexico, and N Sonora. While status and distribution in Sonora are poorly known, the species is believed to be extirpated from California (Jennings and Hayes 1994) and has not been recently documented in New Mexico (Degenhardt et al. 1996), though healthy populations in Arizona exist within just a few miles of each state. They inhabit aquatic systems in desertscrub to pinyon-juniper habitats (Platz and Frost 1984), and were formerly present in irrigation canals in California.

Database Design: Data from surveys have been entered into a dBASE we database. We chose a relational database structure consisting of four main database files, rather than a single "flat file" structure. Locality, habitat conditions, source records, and herpetofauna observation data each reside in a separate database file (Waters et al. 1994). These files, with the exception of the "sources database", reflect the three sections of our field data form (Appendix A). The locality file contains data describing a site in space (e.g. UTM coordinates, Township-Range information). Data which change from visit to visit such as date, time, surveyor, temperatures, and water chemistry are entered into the conditions database file (sites receive a new conditions record for each visit). Species observations are entered into the herpetofauna observations database, which contains the smallest unit of information. A "herp obs" is an observation of a developmental stage of a single target species on a particular date. Data from all sources reside in the source record data file. In addition to target native ranid frogs, we compiled records for other riparian reptiles and amphibians encountered.

Output from the database has been forwarded for inclusion in the Arizona Game and Fish Department (AGFD) Heritage Data Management System (HDMS). Completed survey forms and photocopies of topographic maps with survey areas delineated are available upon request from the Amphibians and Reptiles Program, Nongame Branch, AGFD.

Data Analysis: Status Determination. We present analysis of our survey data through May 1997 for all species of Arizona ranids except the Tarahumara frog, which has been recently reviewed elsewhere (Hale et al. 1995). Historical sites are those where native ranid frogs were recorded prior to January 1, 1993, roughly coinciding with the beginning of this project. Each site where native ranid frogs have ever been recorded is assigned one of four status codes: present historical (any site where frogs have been recorded both before and after January 1, 1993); absent historical (frogs recorded prior to January 1, 1993, but not observed in more recent surveys); unsurveyed historical (frogs recorded prior to January 1, 1993, but no surveys have been conducted at the site since then); and new (frogs recorded at the site only after January 1, 1993). Because of large temporal differences

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in activity and numbers of many amphibians, determining their presence or absence from a locality is difficult and requires multiple visits (but see Chapter 2). Whenever possible, we determined presence or absence at a given site by conducting multiple site visits during times of peak activity (April-October).

Data Compilation. For all descriptive tables, we imported our dBASE. files into SPSS. (Release 7.5) and compiled these data using the General Tables and Stacked Bar procedures. We calculated status codes by using a dBASE' application program, which evaluated the observations of each species at all sites using the criteria outlined above.

RESULTS

Database Overview: We have 1623 locality records in our database. These records represent information gathered through searches of museum records, published literature, technical reports, and knowledgeable individuals, as well as surveys conducted by AGFD personnel, and cover a time span from 1884 to the present. Of these locality records, 595 have native leopard frog observations associated with them, while the remaining 1028 do not.

Our database has native leopard frog records from every county in Arizona (we have historical R. yavapaiensis records from Yuma Co.; the absence of these records from our database output is an isolated artifact of the database entry process for these sites, not a real pattern). Over 50% of our native leopard frog localities occur in four counties: Cochise, Gila, Santa Cruz, and Yavapai. Over 75% are associated with the greater Gila River watershed. Sixty-six percent of all native leopard frog localities are on lands managed by the state or federal government (over 75% of these are on U.S. Forest Service [USFS] lands). The remaining 34% of native leopard frog localities are on Indian Reservations and private lands.

Between September 1990 and April 1997, AGFD conducted 2089 surveys at 1274 localities. These surveys resulted in a total of 3208 riparian herpetofaunal observations (including mud turtles, garter snakes, and all amphibians observed). Seven hundred and seventy-seven of these observations were of native leopard frogs.

Distribution and Status of Arizona Leopard Frogs: In the following species specific distribution accounts, information on elevational ranges, county and watershed occurrences, and land management classifications is based on all known localities for a given species, whether historical or new, whether we visited them or not. Aquatic habitat types are based on both historical and new localities that we have actually visited and characterized. Descriptions of present status are based on all known localities.

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In Appendix B, we summarize the distribution of locality records in the database (all localities, whether native ranids have ever been recorded or not) among Arizona counties. In Appendix C, we provide a similar summary of locality distribution among drainages. Finally, Appendix D summarizes locality distribution among land management categories.

Plains Leopard Frog (Rana blairt). Elevational range of R. blairi records in our database, from the Sulphur Springs Valley, Cochise Co., is 1237-1798 m (4060-5900 ft). This region consists of two distinct drainage systems. A closed basin system, Pluvial Lake Cochise, is central to the valley. Whitewater Draw drains the southern part of Sulphur Springs Valley into the Rio Yaqui System. In the early 1970s, J.E. Platz (pers. comm.) collected R. blairi at Ashurst Lake, Coconino National Forest, at 2168 m (7114 ft) elevation (elevational distributions are summarized in Table 1 and Appendix E).

Table 1. Elevational ranges of native ranid localities.

Species 1 Meters Feet ' Minimum Maximum 1 Minimum Maximum n

Rana blairi 1237 2168 4060 7114 22 Rana chiricahuensis Rim 1067 2710 3500 8890 57

Southeast 1 1061 2012 3480 6600 145 Rana pipiens I

1 I 952 2789 3122 9150 93 Rana subaquavocalis 1501 1829 4925 6000 9 Rana tarahumarae 1109 1775 3640 5823 8 Rana yavapaiensis 146 1817 480 5960 316

Known plains leopard frog habitats (Fig. 1) in the Sulphur Springs Valley are mostly lentic systems (67% of known localities), primarily muddy earthen stock ponds and irrigation sloughs. Thirty-three percent of known localities are lotic systems, mostly intermittent streams with permanent pools, draining the west side of the Chiricahua Mountains. One locality consists of flowing effluent and ash settling ponds at a coal-burning power plant.

With the exception of the Ashurst Lake record from Coconino Co., all known records of R. blairi are from Cochise Co. (95%) (Appendix F).

We conducted 182 surveys within the historical range of the plains leopard frog in southeastern Arizona. We visited 17 of 18 historical localities, observing frogs at two (Table 2, Fig. 2a). We located four new sites for this species.

Five of six (83%) extant localities are in that portion of the Sulphur Springs Valley within the San Pedro River drainage (including Pluvial Lake Cochise) (Appendix G). The other is in the Rio Yaqui drainage. Four of the known extant populations (67%) are on private land (Appendix H), including active agricultural lands and a power plant. Two are on state lands (one managed by Arizona State Land Department [ASLD] and one by AGFD). All six extant populations are on the

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Figure 1. Stacked bar chart of aquatic habitat types grouped by species for sites surveyed by Arizona Game and Fish.

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Sulphur Springs Valley floor in slightly to highly modified habitats. We conducted three surveys at Ashurst Lake from 1991-1995, and made no observations of this or any other species of leopard frog.

Table 2. Locality status designations for native ranid species.

Species Absent Historical Present Historical Unsurveyed

Historical New Total

Cnt Rw% Col% Cnt Rw% Col% Cnt Rw% Col% Cnt Rw% Col% Cnt Col% Rana blairi 15 68% 7% 2 9% 2% 1 5% 0% 4 18% 3% 22 3% Rana chiricahuensis

rim 21 37% 10% 4 7% 5% 21 37% 10% 11 19% 8% 57 9% SE 67 46% 32% 17 12% 21% 17 12% 8% 44 30% 31% 145 22%

Rana pipiens 30 32% 14% 12 13% 15% 35 38% 16% 16 17% 11% 93 14% Rana subaquavocalis

3 33% 4% 6 67% 4% 9 1%

Rana tarahumarae 6 75% 3% 2 25% 1% 8 1% Rana yavapaiensis 72 23% 34% 43 14% 53% 140 44% 65% 61 19% 43% 316 49% Total 211 32% 100% 81 12% 100% 216 33% 100% 142 22% 100% 650 100%

The status of R. blairi in Arizona is tenuous. We suspect the existence of additional breeding populations on state and private lands in the Sulphur Springs Valley, but the combination of land ownership patterns, habitat alteration (including presence of non-native predators and competitors), and small number of remaining populations makes this species a management challenge. It may be the most endangered leopard frog in Arizona.

Chiricahua Leopard Frog - Rim form (Rana chiricahuensis). Elevational range of Rim form R. chiricahuensis (occurring along the Mogollon Rim and White Mountains, north of the Gila River) records in our database is 1067-2710 m (3500-8890 ft) (Table 1). Most (79%) Rim Chiricahua leopard frog localities are in higher elevation headwaters of the Salt, Verde, and upper Gila rivers, with the remaining 21% in the Little Colorado River drainage (Appendix E).

Fifty percent of known Rim form R.

chiricahuensis sites are natural lotic systems and 50% are lentic, primarily livestock tanks (39%), but also natural lakes and artificial reservoirs (11%) (Fig. 1).

Rim form Chiricahua leopard frogs are known primarily from Apache (32%), Gila (21%), and Coconino (16%) counties, with smaller numbers of localities in Graham, Greenlee, Navajo, and Yavapai counties (Appendix F).

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160

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Eight hundred and seventy-one surveys were a) conducted within what we consider the range of Rim form R. chiricahuensis. We surveyed 25 historical localities, confirming presence of frogs at four (Table 2, Fig. 2b). We did not survey 21 historical sites, of which 14 are on the San Carlos Apache (n=6) and White Mountain Apache (n=8) reservations, two are on private lands, and five are on the Apache-Sitgreaves (n=4) and Coconino (n=1) national forests. We

b) located 11 new sites.

The 15 sites where frogs have been observed since 1993 are distributed across the Salt (n=7), Verde (n=5), Upper Gila (n=2), and Little Colorado (n=1) river drainages (Appendix G). Thirteen sites are managed by USFS: Apache-Sitgreaves (n=2), Coconino (n=4) and Tonto (n=7) national forests. One site is owned by AGFD and one is of unknown management status (Appendix H).

Figure 2. Stacked bar chart of status designations grouped

by species. a) Bar segments represent the number of

localities with a given status designation for each native

ranid species. New localities have observations recorded

since 1993 only. Unsurveyed historical localities have

observations before 1993, but have not been visited after

that date. Present historical localities have had

observations both before and after 1993. Absent historical

localities have observations prior to but not after 1993. b)

Split bar chart of status designations for Rana

chiricahuensis grouped by Rim and southeastern form.

The Rim form Chiricahua leopard frog has declined dramatically, particularly in the White Mountains. We know of only a few localities where more than a few frogs have been observed since 1993, and they are absent from 84% of historical sites surveyed. Because much of the known and potential habitat for this frog is managed by USFS or AGFD, management potential is good, and we have begun active management of populations in two areas, one in the White Mountains and one in the Gentry Creek drainage.

Chiricahua Leopard Frog - Southeastern form (Rana chiricahuensis). Elevational range of southeastern Arizona (south of the Gila River in Cochise, Santa Cruz, Pima, and Graham counties) Chiricahua leopard frog records in our database is 1061-2012 m (3480-6600 ft) (Table 1). Seventy-nine percent of known localities are in the San Simon, San Pedro, and Santa Cruz river drainages, the major watersheds of this region draining northward into the Gila river, and

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21% are in the headwaters of the Rio Concepcion and Rio Yaqui, both of which flow south into Mexico (Appendix E).

Southeastern R. chiricahuensis have been found in natural lotic systems (53% of localities) and in livestock tanks (45% of localities). We know of two occurrences in artificial reservoirs (Fig. 1).

Southeastern form R. chiricahuensis are known primarily from Cochise (49% of localities), Santa Cruz (30%), and Graham (17%) counties, with the remainder from Pima Co. (Appendix F).

We conducted 656 surveys within what we consider the range for southeastern form R. chiricahuensis (south of the Gila River). We found frogs at 17 of 84 historical sites surveyed (Table 2, Fig. 2b). We did not survey 17 historical sites, four on National Wildlife Refuges, six on Coronado National Forest, six on privately owned lands, and one on military land. Forty-four new sites were located.

We know of 61 extant sites for southeastern Chiricahua leopard frogs. Major river drainages with extant sites are the San Pedro (n=36), Santa Cruz (n=10), Rio Concepcion (n=8), Rio Yaqui (n=4), and San Simon (n=3) (Appendix G). Seventy-nine percent of extant sites are on Coronado National Forest (n=48). Remaining sites are on state lands (n=5), National Wildlife refuges (n=1), and private lands (n=7) (Appendix H).

Although southeastern form R. chiricahuensis have declined, the number and distribution of extant populations and land management status combine to make this entity perhaps the second-most stable and manageable leopard frog in the state, behind R. yavapaiensis.

Relict Leopard Frog (Rana onca). There are no confirmed Arizona records of R. onca (however, the identity of leopard frogs at Littlefield, long considered to be R. yavapaiensis, is uncertain; Jennings et al. 1995). The historical distribution of the species in Nevada and Utah is such that it could potentially occur in the Virgin River drainage of Arizona, as well as in the drainages of smaller tributaries to the Colorado River from Grand Wash to Davis Dam, from elevations of perhaps less than 300 m to over 1000 m (roughly 900-3500 ft, but this range is strictly conjectural). We have conducted six surveys within the potential Arizona range of the relict leopard frog, and found frogs only at Littlefield. Recent surveys of springs along the Colorado River below Hoover Dam have found leopard frogs (again of uncertain identity, but likely R.

onca) at two sites in Nevada and failed to find them in Arizona (D. Bradford pers. comm.), but additional sites in both states remain to be surveyed. Land ownership of potential habitat in Arizona is mostly federal (Bureau of Land Management and National Park Service), but some state (ASLD) and private lands may also be involved, especially along the Virgin River and Beaver Dam Wash.

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Northern Leopard Frog (Rana pipiens). Elevational range of R. pipiens records in our database is 952-2789 m (3122-9150 ft) (Table 1). Nearly half (47%) of known localities for northern leopard frogs are within the Little Colorado River drainage, the major drainage of the Colorado Plateau, between 1341-2789 m (4399-9150 ft) elevation. Another 47% are below the Mogollon Rim and southern slope of the White Mountains in headwater tributaries of the Gila, Salt, and Verde rivers. Two lower elevation localities are along the Colorado River. One at Glen Canyon is at 952 m (3123 ft) and the other, Kanab Creek (a tributary), is at 1134 m (3720 ft) elevation. Truxton Wash, a tributary of Red Lake Playa at 1250 m (4100 ft), is a peripheral locality, -100 km W of the nearest known extant population (Appendix E).

Over two-thirds (68%) of known northern leopard frog observations have been in lentic habitats, with stock tanks constituting 46% of all known localities (Fig. 1). Other lentic habitats include natural wetlands and lakes and artificial reservoirs on the Colorado Plateau. The remaining 32% of R. pipiens observations have been in natural lotic systems.

Almost all (94%) known localities for R. pipiens are in Coconino, Navajo, and Apache counties, with a handful of records from Gila, Greenlee, Mohave, and Yavapai counties (Appendix F). These political entities coincide closely with the Colorado Plateau.

We conducted 904 surveys within the historical range of the northern leopard frog. We surveyed 42 historical localities, confirming presence of frogs at 12 sites (Table 2, Fig. 2a). Thirty-five (38% of total) historical localities, 22 (63%) of which are on non-state/federal lands, have never been surveyed by us. We located 16 new localities. We also know of recent survey results from the Navajo Indian Reservation (J. Meyer pers. comm.) that failed to find frogs at 57 sites (including eight historical sites) in the Chuska Mountains.

Of the 28 extant northern leopard frog sites, most are in the Little Colorado River drainage (n=13) and the sub-Mogollon Rim headwaters of the Verde River (n=11). The remaining four are associated with the Grand and Glen canyon sections of the Colorado River (Appendix G). Twenty-three extant sites are managed by USFS on Apache-Sitgreaves (n=13), Kaibab (n=4), and Coconino (n=6) national forests, four are on non-state/federal lands, and one is in Grand Canyon National Park (Appendix H).

Northern leopard frogs have apparently disappeared entirely from the White Mountains and the mainstem Little Colorado River. Substantial declines have also occurred along the Mogollon Rim, where we know of only three extant population centers. We know of a handful of isolated populations scattered elsewhere in the state. Little information is available for the Navajo and Hopi Indian reservations (but see above), which include a large proportion of the potential range of R. pipiens in Arizona. Overall, the status of northern leopard frogs in Arizona is rather poor. However, other than those on Native American lands, most known populations are on federally managed land, so management should be possible.

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Ramsey Canyon Leopard Frog (Rana subaquavocalis). The Ramsey Canyon leopard frog is found exclusively in the San Pedro River Valley on the eastern slopes of the Huachuca Mountains. Elevational range of records in our database is 1501-1829 m (4925-6000 ft) (Table 1, Appendix E).

R. subaquavocalis is known from nine localities, including eight artificial ponds or stock tanks and one natural lotic system (Fig. 1).

All known localities of the Ramsey Canyon leopard frog are in Cochise Co. (Appendix F).

We have conducted 149 surveys in the Huachuca Mountains within the probable range of R. subaquavocalis. We have observed frogs at the three historical sites and located six new sites (Table 2, Fig. 2a).

R. subaquavocalis is known from nine localities in Ramsey, Brown, and Tinker canyons, which drain into the San Pedro River (Appendix G). The nine known sites are on Coronado National Forest (n=1), Fort Huachuca Military Reservation (n=2), and privately owned lands (n=6) (Appendix H).

The future of the Ramsey Canyon leopard frog is uncertain. We have established productive partnerships and initiated cooperative management actions with several agencies, landowners, and other interested parties. Land ownership of known and potential habitat is favorable for further management and a multi-party Conservation Agreement has been completed. Unfortunately, the small number of breeding populations, all consisting of relatively small numbers of individuals, make this species highly vulnerable to extinction by demographic stochasticity, natural disasters, or anthropogenic causes.

Tarahumara Frog (Rana tarahumarae). Elevational range of R. tarahumarae records in our database is 1109-1775 m (3640-5823 ft) (Table 1). The Tarahumara frog was known to occur in the Santa Rita, Tumacacori, and Pajarito mountains in the Santa Cruz River and Rio Concepcion drainages (Appendix E).

R. tarahumarae was known to inhabit natural lotic systems, especially plunge pools in mountain canyons (Hale 1992) (Fig. 1).

All known localities for the Tarahumara frog in Arizona are in Santa Cruz Co. (Appendix F).

Seven of the eight records are in the Santa Cruz River watershed, and one is in a headwater tributary of the Rio Concepcion.

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We conducted surveys in the potential range of the Tarahumara frog. They were absent from all six historical sites we surveyed (Table 2, Fig. 2a).

R. tarahumarae was last observed in Arizona in the early 1980s and has been considered extirpated since 1983 (Hale and Jarchow 1988; Hale and May 1983; Hale et al. 1995).

Lowland Leopard Frog (Rana yavapaiensis). Based on records in our database, R. yavapaiensis occurs between 146-1817 m (480-5960 ft) elevation (Table 1). Documented records exist for the lower Colorado and Gila rivers in Yuma County (Clarkson and Rorabaugh 1989), at elevations approaching sea level, but these are absent from our database. Eighty percent of the known lowland leopard frog localities are within the region drained by the Gila River and its tributaries, with 17% in the Bill Williams River drainage (Appendix E). A few localities (2%) are in the headwaters of the Rio Concepcion and the Rio Yaqui at 1097-1378 m (3600-4520 ft).

Of the known lowland leopard frog localities, 82% are natural lotic systems and 18% are lentic habitats, primarily stock tanks (Fig. 1).

R. yavapaiensis has been found in every county except Apache and Navajo (Appendix F). Over one-half (57%) of all localities in Arizona are from the centrally located counties of Gila, Maricopa, and Yavapai.

We conducted 1104 surveys within the historic range of lowland leopard frogs. We observed lowland leopard frogs at 43 of 115 historical localities surveyed (Table 2, Fig. 2a). We have not been to 140 historical sites since 1993 and we located 61 new sites.

Eighty-five percent of extant lowland leopard frog sites are in the Bill Williams (n=21), Salt (n=22), Upper Gila (n=18), Verde (n=14), and Agua Fria (n=13) watersheds (the latter four representing the Greater Gila watershed) (Appendix G). Extant sites in southeastern Arizona are in the San Pedro (n=9), Santa Cruz (n=5), and Rio Concepcion (n=1) watersheds. The 104 extant localities of R. yavapaiensis are on non-state or federal lands (n=43), federally managed lands (n=56), and state managed lands (n=5) (Appendix H). Federally managed lands with extant lowland leopard frog localities are managed by Bureau of Land Management (n=12), Tonto (n=26), Apache-Sitgreaves (n=8), Coronado (n=4), Prescott (n=3), and Coconino national forests, and Bill Williams River National Wildlife Refuge (n=2).

The lowland leopard frog is extirpated from the lower Gila and Colorado Rivers of Arizona and adjacent California. It may be extirpated from New Mexico and it has declined fairly dramatically in southeastern Arizona (AGFD unpubl. data). Nevertheless, its status in central Arizona seems good. It is certainly the most stable native ranid in Arizona.

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DISCUSSION

Our intent in carrying out statewide surveys for determining status of Arizona's native ranid frogs was to quantify, at least to some extent, anecdotal reports and observations of population declines. These determinations are helpful in placing status of the various native ranids into a statewide and even global context, and they form the basis for planning and prioritization of conservation actions. We now have baseline data for each native ranid that can be used to track future changes in status. We have a fairly clear picture of magnitude and geographic extent of declines of each species, and we have information on current distribution to allow us to determine which populations or metapopulations are most in need of management intervention, what sorts of management actions are necessary, which populations can be expected to benefit most, and which are the most manageable. All of these are important considerations in setting priorities for expenditure of the limited resources available for ranid frog conservation.

Generalizations from Statewide Surveys: The results of our statewide surveys confirm findings of earlier studies indicating that nearly every native ranid frog in Arizona has declined over the past two or three decades (Clarkson and Rorabaugh 1989; Sredl 1993a). We have strong survey data supporting this contention for the Tarahumara frog and at least three of the five native Arizona leopard frog species: the northern, Chiricahua, and lowland leopard frogs. For two of these three species, northern and Chiricahua leopard frogs, the statewide pattern of occupancy of historical localities is similar: native leopard frogs are apparently absent from large numbers of historical localities that supported populations of leopard frogs as recently as the late 1970s to mid 1980s. Though never broadly distributed in Arizona, the Tarahumara frog is now extirpated, having disappeared from all known localities in Arizona during the late 1970s and early 1980s. The relict leopard frog was never documented from Arizona, so it would not be possible to demonstrate a decline. However, its historical distribution makes it hard to imagine that it was not present here, at least in the Virgin River drainage (and as noted elsewhere, a leopard frog population near Littlefield may be conspecific).

Lowland leopard frog populations have not shown a severe, rangewide decline. This species appears stable in central Arizona (Sredl and Howland 1992), but our surveys verified Clarkson and Rorabaugh's (1989) finding that this species is doing poorly in southeast Arizona and the lower Gila and Colorado rivers. This result is geographically consistent with the apparent extirpation from New Mexico (C.W. Painter and R.D. Jennings pers. comm.) and California (Clarkson and Rorabaugh 1989; Jennings and Hayes 1994).

The status of Ramsey Canyon and plains leopard frogs is less clear. Populations of the newly described Ramsey Canyon leopard frog (Platz 1993) are restricted to a few canyons in the Huachuca Mountains. These populations appear stable (Platz pers. comm.), but their small number and size make them vulnerable to a variety of potential threats. Preliminary data for the plains leopard frog indicate a severe decline, but these data are incomplete.

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Factors Leading to Population Declines: Separating human impacts from natural causes of amphibian population fluctuations is difficult (Peclunann et al. 1991), but some of the anthropogenic factors implicated in declines of amphibian populations in Arizona and elsewhere are: 1) Non-native organisms appear to be at least partially responsible for many declines. The bullfrog (R. catesbeiana), introduced throughout much of the western United States, is both a predator and competitor of native ranid frogs (Moyle 1973), including leopard frogs in Arizona (Schwalbe and Rosen 1988; AGFD unpubl. data). Bass and sunfish, both relatively recent additions to Arizona's fauna, eat amphibians (Hayes and Jennings 1986). In general, our data and those of others (Rosen et al. 1995, 1996) show a strong negative association in occurrence of native ranids and non-native fishes. The importance of crayfish is less clear, but negative impacts through predation on various life stages of fishes (e.g. White 1995) and amphibians (Gambradt and Kats 1996) have been demonstrated. The introduced Rio Grande leopard frog (R. berlandieri) may prove to be problematic if its continued range expansion (Platz et al. 1990) brings it into contact with native ranids. It is now nearly parapatric with R. yavapaiensis (AGFD unpubl. data). 2) Acid rain has been implicated as a factor leading to the decline of the tiger salamander in different parts of its range (Harte and Hoffman 1989; Pierce 1985). Closer to home, there is some evidence that elevated pH (coupled with other factors) led to the demise of the Tarahumara frog (Hale and Jarchow 1988). 3) Habitat destruction and habitat degradation have been responsible for numerous declines (National Research Council workshop 1990), and have affected some historical localities in Arizona. 4) Heavy metals leeched by acidic precipitation have been hypothesized as the responsible agent in the decline of the Tarahumara frog in Arizona (Hale and Jarchow 1988) and other amphibians in different parts of the globe.

Other factors of decline have been identified elsewhere but not yet implicated in Arizona: 5) Agricultural pesticides and herbicides, as well as other toxins, have been implicated in some declines (Hall and Henry 1992). 6) Thinning of the ozone has led to increased levels of incident ultraviolet radiation. While not directly implicated in any specific decline we are aware of, it was demonstrated to be potentially harmful to the eggs of some amphibians (Blaustein et al. 1994). 7) Global warming may have caused declines of some amphibian species (Wyman 1990).

Finally, some of these factors may interact synergistically to cause problems where either factor on its own might not be harmful. Such a synergism between UV and pH has been demonstrated to reduce viability of R. pipiens embryos and larvae in a laboratory setting (Long et al. 1995).

Spatial Distribution of Extant Populations: A likely contributing factor to leopard frog declines in the arid Southwest is habitat reduction and fragmentation resulting in small, isolated, unstable local populations (Sredl and Howland 1995). Damming, draining, and diverting of water have fragmented formerly contiguous aquatic habitats. In many areas, fragmentation has been accentuated by introduced predatory fishes, crayfish, and bullfrogs, leaving potential dispersal corridors between available aquatic habitats impassable (Bradford et al. 1993). Leopard frog life history makes their populations very dynamic. Rates of reproduction, recruitment, and other

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population attributes are highly variable and dependant upon rainfall and other environmental influences (see Chapter 1). Leopard frogs are highly aquatic and vulnerable to desiccation. Stimuli such as sudden cold snaps can result in disease outbreaks (Carey 1993), especially at local sites where overcrowding occurs (due to high reproductive success, or habitat contraction from drought or sedimentation of pools). Since local populations of leopard frogs are prone to extinction (see Chapter 1; Sredl 1993b; Sredl and Seim 1993), it is important to maintain dispersal corridors for recolonization.

Our locality data indicate that most extant leopard frog populations occur in small, isolated aquatic sites distributed in clusters, rather than showing random spatial distribution (Sredl and Howland 1995). In most cases, habitable aquatic sites are either: remnants of historically larger perennial aquatic systems (which are now contracted, see Hendrickson and Minckley 1984), which probably hosted large populations of frogs; or they are peripheral to a large aquatic system (large lake, stream, or river) which presumably hosted a large central source population. In either case, we suspect that extant population clusters are remnants of former metapopulations. This hypothesis is supported by some historical records and anecdotal reports of large numbers of frogs in big lakes and rivers that are no longer inhabited.

In summary, our observations of leopard frog populations lead us to believe that patches of aquatic habitat, connected by drainages that can be traveled by dispersing leopard frogs, at least intermittently, form the microgeographic basis for functioning metapopulations (see SjOgren 1991 on the importance of connectivity for metapopulations of aquatic frogs). With the aquatic faunas of remaining large aquatic habitats being dominated by non-native species, habitat for native ranids is reduced to small, isolated pockets that are capable of supporting only small, unstable populations. Large core populations no longer exist. Essential dispersal corridors are impassable due to lack of water or, in the case of perennially flowing corridors, the presence of non-native species. The life history of southwestern leopard frogs predisposes them to high rates of local extinction and recolonization. We suspect that population declines among southwestern leopard frogs can, at least in part, be attributed to disruption of normal metapopulation dynamics by various human disturbances causing increased rates of extinction and decreased rates of recolonization.

Management Strategies - Conservation and Management Zones: We have begun to develop and test management strategies to restore functioning metapopulations to appropriate areas. Potential conservation techniques include: 1) ex situ captive breeding and/or rearing of tadpoles for release (as adults) to the wild; 2) translocating wild eggs, tadpoles, and frogs; 3) removing non-native species; and 4) improving or creating habitat. Our intent is to use a coordinated mix of these and other techniques, tailored to the needs of particular situations, to reconstitute functioning metapopulations in areas where frogs may be destined for extirpation in the absence of active conservation.

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One of our first steps in formalizing our approach to conservation and management of Arizona leopard frogs is development of the concept of Conservation and Management Zones (CMZs). Using our statewide survey results, we will identify areas of critical conservation need. We must first establish criteria for priority ranking importance of populations from a statewide conservation perspective. These criteria may include: 1) overall status of the species, both statewide and global (a highly sensitive species endemic to Arizona receives higher conservation priority than a geographically widespread species whose status in Arizona is less critical); 2) geographical context of a population or cluster of populations (those in a region of severe decline or in a remote area that is unlikely to be naturally recolonized in the event of a local extinction event receive highest priority); 3) evolutionary context of a population or cluster of populations (those that are evolutionarily important due to genetic distinctness or diversity receive highest priority); 4) manageability of the population or area (those populations in areas where threats are most likely to be controllable and land owners or managers are more willing and able to cooperate receive highest priority); and 5) complexity and cost (those populations that can be stabilized or recovered through use of the fewest, simplest, and most cost-effective conservation actions receive highest priority).

Our statewide surveys have provided, and will continue to provide, most of the information we will need to address the first two criteria. With cooperating academic biologists, we are beginning to gather the sorts of genetic information necessary to address the third criterion. We have some information concerning the last two criteria, but they will require additional investigation specific to individual populations, areas, politics, and other factors that may be peculiar to each proposed CMZ. These will largely be addressed through cooperative, site-specific planning and negotiation with appropriate landowners and resource managers, including public, academic, and interagency review of proposed actions.

After setting priorities, we must begin the cooperative effort of CMZ designation, customized conservation planning, generation of funding, preparation of any necessary environmental compliance documents, and implementation of measures that are appropriate to each area. Although AGFD can coordinate this process and make significant contributions in funding and implementation, it is essential to have active participation and funding by cooperators, especially the affected landowners or managers, if we are to have any hope that this approach will be successful on a large landscape level.

We must also recognize that our initial efforts to designate CMZs and implement conservation measures will be test cases. We must evaluate these first efforts and modify our approaches as needed to make them more effective and efficient. We can expect to encounter difficulties and outright failures at the beginning, but by making methodical evaluations and modifications, it should be possible, with adequate commitments from key resource managers and stewards, to successfully orchestrate stabilization and recovery of Arizona's native ranid frogs.

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We have now made initial efforts to implement and test each of the general conservation techniques mentioned above. Cooperating biologists at The Phoenix Zoo and Grand Canyon University have made tremendous strides in development of captive breeding and rearing, of R.

subaquavocalis, R. yavapaiensis, and Rim form R. chiricahuensis. Their findings are likely to be generalizable to all other native leopard frogs, but further work will be needed for captive husbandry of R. tarahumarae. We have released captive reared R. subaquavocalis, and captive bred Rim form R. chiricahuensis from these facilities into the wild, with varying results. We must continue to evaluate appropriateness of release sites, relative success of releases of different life stages, and seasonal timing of releases.

We have conducted or assisted in translocations of wild metamorph and adult frogs in the White Mountains (Rim form R. chiricahuensis), southern Kaibab Plateau (R. pipiens), and San Bernardino Valley (southeastern form R. chiricahuensis). As mentioned above, further evaluation of site selection, appropriate life stages for translocation, and timing of the action are needed. Again, these actions have had varied success, ranging from rather rapid disappearance of translocated frogs (White Mountains) to persistence of new breeding groups for over three years (San Bernardino Valley).

We have conducted or cooperated in several efforts to remove non-native species from leopard frog habitat. We have tested trapping methods for removal of crayfish, but no concerted effort has been made to eliminate them from a particular site. We successfully eradicated bullfrogs from a pond (formerly inhabited by Rim form R. chiricahuensis) in the Gentry Creek drainage of Tonto National Forest. Similar small scale efforts have occurred at other sites, including potential new sites for R. subaquavocalis. We are involved in a long-term, relatively high intensity effort to control bullfrogs at San Bernardino National Wildlife Refuge (SBNWR; formerly inhabited by R.

yavapaiensis and southeastern form R. chiricahuensis), which has successfully reduced numbers and body sizes of resident bullfrogs, but has failed to approach eradication. A new approach to bullfrog control at this site will be implemented in concert with planned habitat renovation/creation on SBNWR (see below). We need further evaluation of removal techniques, ranging from trapping to poisoning to explosives, for all non-native species. We also need to investigate design and effectiveness of barriers to exclude non-natives from areas where they have never invaded or have been eradicated. There is considerable potential to work with proponents of recovery efforts for native fishes because of parallels in nature of threats (in some cases identical) and needed management actions.

We have been involved in habitat renovation/creation projects consisting of actions as simple as adding brush to increase aquatic and bankside habitat heterogeneity (providing oviposition sites, cover from predators, etc.) and as complex as the planned creation of a new habitat. We have worked with Ft. Huachuca to design a new pond that would be slated for establishment of a new breeding population of R. subaquavocalis. We assisted SBNWR in design and management (including translocation of leopard frogs) of four small breeding ponds protected by bullfrog-proof

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fencing, and planning of new bullfrog-free artificial cienega habitats for native fishes and leopard frogs. We coordinated repair of a small check dam at a southeastern R. yavapaiensis site to help maintain an important breeding pool. We are considering enhancement of pool habitat in some stream systems that are inhabited by Rim form R. chiricahuensis. We coordinated release of captive reared R. subaquavocalis into a pond that was dredged after having been filled with sediment. A promising new direction is working with ranchers to manage aquatic habitats for leopard frogs, which are generally compatible with livestock. We have assisted with development of two Stewardship Agreements designed to help ranchers renovate or manage stock tanks to the benefit of both livestock and leopard frogs (southeastern form R. chiricahuensis so far, but the potential exists to pursue these activities for all native ranids). We need to continue to test and evaluate these manipulations, giving special attention to cost-effectiveness because of the tremendous range in cost of different types of habitat renovation or creation.

Information Needs: One of our most basic information needs is clarification of southwestern leopard frog taxonomy. At this point, we are uncertain as to the identity of leopard frogs at Littlefield. Though historically identified as R. yavapaiensis, Jennings et al. (1995) showed that these frogs are morphologically and genetically more closely allied to R. onca populations in Nevada than to R. yavapaiensis in central Arizona. It remains unclear whether R. yavapaiensis is actually distinct from R. onca. The distinctiveness of R. subaquavocalis, southeastern form R.

chiricahuensis, and Rim form R. chiricahuensis are also questionable (J. E. Platz pers. comm.), though limited genetic data suggest at least some level of differentiation (Jennings et al. 1995). The same study presented limited data suggesting differentiation of Sulphur Springs Valley R. blairi from other populations to the east.

From a non-taxonomic perspective, we need information on genetic structure of leopard frog metapopulations (and constituent subpopulations), including knowledge of levels of heterozygosity and genetic exchange within and among metapopulations. This would influence management goals concerning numbers and proximity of breeding populations that need to be established and/or maintained in order for metapopulations to continue or resume functioning. Dispersal capability of leopard frogs is also unknown. This information is important in identifying characteristics of dispersal corridors that will be necessary for establishment or maintenance of functioning metapopulations.

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Vial, J.L. and L.S. Saylor. 1993. The status of amphibian populations: a compilation and analysis. (Unpubl.).

Vitt, Li., J.P. Caldwell, H.M. Wilbur, and D.C. Smith. 1990. Viewpoint: amphibians as harbingers of decay. BioScience 40:418.

Waters, D.L., S.G. Seim, and M.J. Sredl. 1994. Herpetofauna database guide. (Unpubl.).

White, J.N. 1995. Indirect effects of predation by crayfish on Little Colorado spinedace. Northern Arizona University. 46 pp.

Wyman, R.L. 1990. What's happening to the amphibians? Conservation Biology 4:350-352.

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Appendix A. Riparian Herp Survey Form

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ARIZONA GAME AND FISH DEPARTMENT

NGTR 121: RANID FROG CONSERVATION AND MANAGEMENT

JUNE 1997 PAGE 65

NEW: RIPARIAN HERP SURVEY FORM

Locality Data

SEC:

TRS_COMM:

UTM_X:

Date.

no day year

SITE: TWN_RNG R. SITE_AT:

NUM: OLD: 1 1 1 1 1 1 1 1 1 LIII 1 1 1 1

of

1 1 1 1 1 1 1

1 1 1 1

GEN:

1 I I QUAD: MIN:

MGT UNIT:

YR:

COUNTY:

1 1 1 1 1 7.5 15 1 1 1 1

LIII III 1 1 1 1 1 SUBUNIT:

ELEV:

DIRECTIONS.

BASIN.

UTM_Y:

ZONE:

1 1 1 1

11 12

Herpetofauna Observations

SPECIES

CRT STAGE NUM

MESO_HAB

POSIT

COVER OBS

0 1 ELJA

R. A

10

AS AM

COMMENTS:

Rev.: 06/16/1995

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1 1 1

Conditions Data

OBSERVER(S):

START:

EFFORT:

STOP. 1 1 1

1 1 1 HAD PHOTO: 1 1 1

PHO_VOUCH: PP LB

H_CLASS: H20 TYPE: 0 1 1 2 3 4 5 6 7

NOTES.

TP RB

no day year DATE:

COLLECT: BB

Arizona Game and Fish Department June 1997 NGTR 121: Ranid Frog Conservation and Management Page 66

Num.

T_AIR:

T_H20:

PH:

COND:

D02:

RH :

1 I 1 1 1 1 1

1 1 1 1

WIND: 1 2 3 4 5 0 1 2 3 4 5 6

1 CLOUD: 1 1 1 1 1 1 2 3 4 5 rn

PPT: 1 1 1 1 0 1 2 3 4

1 2 3 4 5 6 7 T_STORM:

0 1 2 2 3 4 5 6

CLARITY:

LENTAX_L:

LENTAX_S:

LOT WIDE:

1 1 SUBSTR: PPm 1

PROMINENT SPECIES

FLOAT VEG: SUB VG: EMEiG VEG: PERIM—VEG: CANOP'7_VEG:

PREDATORS:

LEECH CRAY ANSOP BELO BEET WWFSH CWFSH ANTI BFRG TURT GSNK HRON BHWK MAMM

LAND USE: AGRIC DEV GRAZE LOG MINE REC

OTHER ORGS-ORG_OiS:

Rev.: 06/16/1995

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RIPARIAN HERP SURVEY FORM INSTRUCTIONS

> All fields are to be filled out for historical sites and sites with target riparian herpetofauna. > Fields with an asterisk (*) are to be filled out for every survey, regardless of results. The exception to this

is for sites where an NGB survey has been previously done, the only LOCALITY fields you need to fill out are NUM, SITE, SITE_AT, and QUAD.

> Comments enclosed with [ ] are not for database entry. > NT = data Not Taken. > Use a zero value when none are encountered. > Upon return to the office, have each Survey Form checked for completeness, conciseness and clarity by

someone else prior to submitting for entry into the Herp Database.

LOCALITY Fields:

* SITE:

* SITE_AT:

* NUM:

A "site" is any aquatic system (or piece of an aquatic system) that is > 1 mile from any other survey locality, or if less than 1 mile apart, represents a significant change in aquatic habitat types (e.g. riverine vs. lake or cienega). Features with unique names are considered unique sites regardless of how far apart they are. Record the site name as it is marked on the quad. If the site is unnamed on the quad, refer to the corresponding land management map (e.g. Forest Service map, Surface Management Responsibility map). If the site doesn't have a name, write "unnamed" preceding the feature; similarly, if the site is not marked on any map, write "unmarked" preceding the feature (e.g. Unnamed Wash, Unmarked Tank).

This field should always be filled out for unnamed and unmarked sites and for large/long aquatic systems. For other localities, use this field as needed to enhance a site name--to verbally pin-point a site in space. Use such features as the nearest road crossing (e.g. East Verde River at Highway 87) stream confluence (e.g. East Verde River at Webber Creek) or topographic feature (e.g. East Verde River N of Piety Hill).

Put the site number on both the front and back of the survey form. A site number is a unique number that, once assigned to a site, will always be used in conjunction with that site. The site number starts with a 3-letter code that describes the land manager. These 3 letters are followed by a hyphen and then a 4-digit number (e.g. TON-0001, COC-0153). Sites will be numbered in ascending, consecutive order within each management unit.

Arizona Game and Fish -- AGF Bureau of Land Management -- BLM Land Grants — LGR Military -- MIL National Forests — APA, COC, COR, KAI, PRE, TON National Parks/Monuments — NPS National Wildlife Refuges — NVVR Private Lands — PVT State Lands — ARZ

swsinst.wpd, rev: 04/09/97

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Tribal Lands (note: the final "R" = Reservation): Ak Chin -- AKR Fort Apache APR Cocopah OCR Chemehuevi CHR Colorado River CRR Camp Verde CVR Gila Bend -- GBR Gila River GRR Havasupai HAR Hopi HOR Hualapai HUR Kaibab KAR

Fort McDowell -- MCR Fort Mojave MOR Navajo NAR San Carlos SCR Salt River SRR San Xavier — SXR Tonto-Apache — TAR Tohono O'odham — TOR Yavapai-Prescott YPR Fort Yuma -- YUR Zuni — ZUR

OLD_NUM:

QUAD:

When a historical site is visited for the first time, the historical site number must be replaced by a new site number (described above). Put the old, historical site number in this box. Historical site numbers begin with "RAN", "GAR", or "BUM". On rare occasions, a site number has to be changed because the land management unit has changed (e.g. the Forest Service now owns a parcel of land that used to be in private ownership). Put the old site number in this box.

Record the quadrangle name as it appears on the quadrangle except in the situations outlined below.

> Don't use periods > Don't use apostrophes > Change the word "Mountain" to "Mtn" if it appears anywhere in the quad name

other than the first word > Composite polar coordinates (e.g. Southeast, Northwest) should be abbreviated

(e.g. SE, NW) if they appear anywhere in the quad name other than the first word > Never abbreviate the four cardinal directions

* MIN: Circle "7.5" or "15" to note whether the quadrangle series is 7.5 or 15 minutes.

* YR:

Record the year of the quadrangle as it is printed in the lower right corner of the quad. If more than one year appears on the map, record the year of the most recent revision.

* MGT_UNIT:

Use the same 3-letter codes for the management unit as are used in the site number (see "NUM" above).

* SUB_UNIT:

The sub-unit further defines the management unit. Record the full name of the sub-unit as follows:

Arizona Game and Fish Department region (1-6) Bureau of Land Management -- district (Phoenix, Safford, Strip, or Yuma) Land Grants -- land grant name Military -- name of military property National Forest lands -- ranger district (e.g. Beaver Creek, Heber, or Pleasant

Valley)

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National Park/Monument lands -- park or monument name National Wildlife Refuges -- refuge name Private and/or State lands -- nearest city, town, or community Tribal lands -- nearest tribal community or town

* COUNTY:

County is always recorded as the state abbreviation (AZ) followed by a hyphen and then the first 4 letters of the county (e.g. AZ-MARI, AZ-YAVA). The county name can be found in the upper right corner of the quadrangle if the quad covers an area within a single county. For quads that cover areas in two or more counties, the names of the counties will appear somewhere in the topographic region of the quad. National Forest maps and the Arizona Highway road map, and the Arizona Atlas & Gazetteer are also useful in identifying counties.

Please use the following abbreviations for county: APAC, COCH, COCO, GILA, GRAF!, GREE, LAPA, MARI, MOHA, NAVA, PIMA, PINA, SANT, YAVA, YUMA

* ELEV:

* BASIN:

* TWN_RNG:

Record the elevation at which the starting point of the survey occurs. Read the elevation off of the survey quad. Be sure to indicate whether the elevation is in meters (m) or feet (f) by circling the appropriate box. The contour interval and unit (meters or feet) is written in the center of the bottom margin of the quadrangle.

Record the basin number of the survey area as a 8-digit number as it occurs on the 1974 Hydrologic Unit Map of the State of Arizona. Use townships and ranges to help you determine the basin in which the survey site occurs.

Write the township and range within which the starting point of the survey occurs. Write the township as: T#4.#direction (e.g. T17.0N, T02.5S). Similarly, write the range as: R#4#.#direction (e.g. R01.0W, R31.5E).

* SEC:

Write the section within which the starting point of the survey occurs. Write the section as 2 digits (e.g. 02).

* TRS_COMM:

UTM_X:

Write the quarter-quarter and the quarter-section within which the starting point of the survey occurs. Write the comment as one of the 8 cardinal directions followed by 4 or 2. Occasionally a site falls in the absolute center of a quarter-section. When this occurs, use CTR (center). For example: NW4 of NE4 = the northwest quarter of the northeast quarter; S2 of NW4 = the south half of the northwest quarter; CTR of 54 = the center of the south quarter. Be specific: use NW, NE, SW, SE over N, S, E, W, CTR whenever possible.

This field is to be filled out in the office. Record the starting point of the survey as a 6-digit number followed by the letter "E" (E indicates that the number is an easting value). An example of a UTM x-coordinate is 295440E. The first 3 numbers will be found on the top or bottom edge of the quad. These numbers are in 100,000-meter increments. The fourth number describes a point with ± 100-meters accuracy. The fifth number describes a point with ± 10-meters accuracy. The last number will be a zero. Use a coordinate scale to determine the fourth and fifth numbers. Alternatively, use a Global Positioning System (GPS) unit to measure the UTM coordinate. Enter only those GPS values that have been corrected.

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UTM_Y: This field is to be filled out in the office. Record the starting point of the survey as a 7-digit number followed by the letter "N" (N indicates that the number is a northing value). An example of a UTM y-coordinate is 4318410N. The first 4 numbers will be found along the left or right edge of the quad. These numbers are in 1,000,000-meter increments that tell you how far north of the equator you are. The fifth number describes a point with ± 100-meter accuracy. The sixth number describes a point with ± 10-meter accuracy. The last number will be a zero. Use a coordinate scale to determine the fifth and sixth numbers. Alternatively, use a GPS unit to measure the UTM coordinate. Enter only those GPS values that have been corrected.

ZONE: Circle "11" or "12" to note whether the starting point of the survey is in UTM grid zone 11 (west of 114 degrees longitude) or 12 (east of 114 degrees longitude).

GEN: Circle the category that describes how the UTM coordinates were generated:

G — UTMs measured using a GPS unit (corrected values only) M — UTMs mapped on a quadrangle using a coordinate scale C UTMs converted from Township and Range data using ARCINFO

Please don't write uncorrected UTM values from GPS units in the boxes on the data form. If you need to jot down the uncorrected values, write them with a pencil in the margins of the data form, or keep a separate log.

DIRECTIONS/COMMENTS: Write the directions to the site. Keep them short and pertinent (e.g. on FS 105 -4.3 MI N of FS 105/FS 393 jct.). Directions are especially important when there are no roads or when existing roads are not marked on your maps. Use the 8 cardinal directions (N, NE, E, SE, S, SW, W, and NW) instead of "turn right" or "veer left".

!!!!This field can also contain any information you want to convey to other field personnel. For example: "Dry 05/1994"; "Contact landowner for permission to access (602)555-9683"; "Also survey adjacent tank and draw"; etc.

HERPETOFAUNA OBSERVATIONS Fields:

SPECIES: Record all riparian herp species (target or non-target) detected during a survey in this column (put records of non-riparian herpetofauna in the OTHER_ORGS and ORG_OBS fields). Use their unique 4-letter Genus-species code (see "Herpetofauna List - Derived from Stebbins (1985)"). When an organism cannot be identified to species (e.g. "I saw a ranid-like frog", or "I saw an anuran egg mass"), use the 4-letter code corresponding to the taxonomic classification for which you are confident in Your identification. For the examples above, the ranid-like frog would be assigned the code "RANA", and the egg mass would get the code "ANUR". If you are confident you saw a leopard frog, but you are not certain which species you saw, use the code "RAPC".

Do not use historic information to bias your decision on species obs. Record most confident obs. and justify in notes.

CRT: Write a 0 or 1 in the appropriate column to indicate your level of certainty about your identification of each species:

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0 — certain 1 — uncertain

Certainty of identification should be based on species-specific diagnostic characters (i.e. thigh pattern and dorsolateral folds in leopard frogs, scale row of lateral stripes in garter snakes, lack of dorsal stripe and cranial crests in Arizona toads). For information on diagnostic characters of species, see Stebbins (1985), "Characteristics of Arizona Leopard Frogs", and "Garter Snakes of Coconino National Forest".

STAGE: Write an E, L, J or A in the appropriate column to indicate the life stage(s) of each species observed:

E — eggs L — larvae J juvenile (usually <50-55 mm SVL) A — adult and subadult (usually > 50-55mm SVL or obvious sign of breeding

condition (e.g. swollen thumbpads, stretched vocal sacs)

NUM_OBS:

MESO_HAB:

Enter the number of individuals of each species and life stage you encountered. Do not estimate total numbers within the survey area—record only the number that you saw. For egg masses est. # of eggs noting overall size of mass, condition, and stage of embryos in comments (see Gosner 1960).

Meso-habitat is defined on an ordinal scale. Write a 1, 2, or 3 under the meso-habitat type that best indicates the habitat(s) in which the species/life stage was observed most frequently. For example, if 1 is written under "R" and 2 is written under "B" this means that most of the animals were seen in riffle and they were next most abundant in backwater pools. If the species was observed in only one habitat, use only the number 1. Remember, if animals are observed in equal numbers in different habitats, positions or cover the # 1 can be used more than once.

Lotic Habitats:

Lentic Habitats:

Terrestrial Habitats: (areas >1.5 meters from the waters' edge)

R — Riffle A — Active Channel Pool B — Backwater Pool 10 — Inlet/Outlet C — Cove M — Main Body F Flat (without water: mud flats, sand banks,

marshy flats) U — Upland (non-riparian vegetation) T — Terrestrial Riparian Zone

POSIT: Position is defined on an ordinal scale for lotic and lentic systems only (not for terrestrial habitats). Write a 1, 2, or 3 in the appropriate column in order to describe where a species/life stage was observed most frequently relative to the aquatic shoreline:

L — Shoreline (terrestrial area s1.5 meters from the waters' edge; include animals seen half-in/half-out of water)

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S — Shallows (from the shoreline out to the "point" of drop-off, or s0.5 meter deep; include animals that have all 4 feet in water)

D — Deep Water (beyond the "point" of drop-off, or in water 0.5 meter deep)

COVER: Cover is defined on an ordinal scale. Cover is not substrate — it matters not what the animal is sitting on (substrate) but rather what it is sitting in or under (cover). For all habitats (lotic, lentic, and terrestrial), write a 1, 2, or 3 in the column under the code that best describes the most common cover types used by each species/life stage observed:

COMMENTS:

• Trees/Shrubs (do not confuse with canopy cover; only use this code if an individual is found just under, on, or against a tree or shrub)

• Rock/Outcrops • Not Vegetated AS

Aquatic Submerged (submerged aquatic plants like algae, Ceratophyllum, and Potomageton)

AM

Aquatic Macrophytes (cattails, bulrushes, sedges, broad leafed floating veg.) Grasses

• Debris (including dead and down woody debris)

Use this field to elaborate upon species observations. Types of observations to include are as follows: 1) what criteria were used to identify a species; 2) if species identification is uncertain, what was observed (both physical features and behaviors would be of use, e.g. "dorsal spots obs.," "ranid like plop," "no bullfrog peep"); 3) record the collection number (AGFD field tag #) of any voucher specimens taken; 4) note the presence of disease; and, 5) if extra surveys/counts are conducted on same day note pertinent obs. here. Be sure to reference your comments with the species observation to which it relates by using numbers or letters (like a footnote).

CONDITIONS Fields:

* OBSERVER(S): List the names of all people present during the survey. Record the names as: first initial, second initial, and full last name (e.g. M.J. Sredl).

* DATE: Record the date of the survey as eight numbers giving the month first, followed by the day then the year (e.g. 10-27-1993, 06-02-1994).

* START: Record the time the surveyor begins searching for herps using a 24-hour clock.

* STOP: Record the time the surveyor stops searching for herps using a 24-hour clock. Include only the time spent actually searching for herps.

* EFFORT: There are 5 categories of effort:

TP — Total Perimeter PP — Partial Perimeter LB Left Bank RB — Right Bank BB — Both Banks

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Circle the category(s) that apply. For all categories other than TP, record the distance surveyed in meters. The minimum acceptable survey distance for linear systems and large lentic systems (>20 acres) is 400m (0.25 mile). Use category BB for any lotic system in which it is possible for you to access both banks—to meander from shore to shore. Use categories LB and RB for large, deep, and/or swiftly flowing lotic systems in which you are unable to meander shore to shore. LB and RB should always be filled out together even if you didn't survey, or were unable to access, one of the shores (e.g. LB=0000m, RB=0350m; RB=0050m, LB=0200m). Left and right banks are in reference to upstream. During the course of any survey, the surveyor should dipnet, comb through bushes and grasses, turn over rocks, and scan the water and shore for herpetofauna.

To calculate meters walked, use a map wheel to determine the distance in miles. Be sure to use the scale on the map wheel that corresponds to the scale of your map or quad. Multiply the value generated from the map wheel by 5,280 feet/mile. Multiply this new value by 0.3048 meters/foot Round this final result to the nearest 25-meter value.

Alternatively, use the reverse side of the map wheel to determine the distance in kilometers. Multiply your result by 1000 to get meters.

* COLLECT:

Circle "Y" (yes) or "N" (no) as an indication of whether voucher specimens were collected at a site. If "Y" is circled, the collection tag number(s) along with additional relevant data should be written in "COMMENTS" in the "Herpetofauna Observations" section of the survey form.

* HAB_PHOTO:

* PHO_VOUCH:

Note how many habitat photographs were taken at a site. Write the number as 2 digits (e.g. 00, 02). Habitat photos should be taken at any site in which target riparian herps were found, at any historical locality regardless of results, and at any survey site that has suitable habitat even if no target riparian herps were found.

Note how many photo vouchers were taken at a site. Write the number as 2 digits (e.g. 00, 13). Photo vouchers should be close-ups (macro shots) of diagnostic characters (i.e. thigh pattern and dorsolateral folds of leopard frogs, scale row of lateral stripes in garter snakes, dorsal and cranial views of Arizona toads).

* H_CLASS:

Circle the category that best describes the hydrological class of the water body you have surveyed:

0 lentic (still water) 1 — lotic (flowing water)

* H20_TYPE: Circle one category (1-7) that best describes the type of water you have surveyed. Water type categories are based upon lotic/lentic characteristics as well as the size/magnitude of the water body:

1 — canal (concrete diversion of riverine water) 2

plant outflow (sewage and electric plants; any chemical or mechanical processing of water; storm drainages)

3 — riverine (natural flow, from raging rivers to streams to seeps)

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NOTES:

4— wetland (an inland body of water that is primarily emergent vegetation, e.g. cienegas)

5 -- stock tank (an earthen-dammed or dredged basin that catches run-off for livestock or wildlife)

6 — lake (an inland body of water that is primarily open water) 7 — reservoir (a dammed riverine system that is primarily used for recreation

and/or human water supply) 8 — small manmade water holding structures (metal/concrete tanks and drinkers)

Use this field to describe the most outstanding features of a survey site. Don't be redundant with fields already completed. Write short, specific comments that emphasize habitat quality and why you think you did or did not find hems. Be sure to comment on any land use in, around, or in proximity of the survey area that may potentially impact the study site (e.g. large mining operation 0.5 mile upstream of survey site, agricultural spraying 1 mile from survey site). You can also use this field to describe any noteworthy similarities or dissimilarities between the area searched and the total area (e.g. wash devoid of vegetation except in area of survey, survey covered the north end of the lake which was the only area with emergent vegetation).

T_AIR: Take air temperature (degrees Celsius) 1.5 meters above ground and 1.5 meters from the water. Be sure thermometer is shaded and completely dry.

T_H20: Take water temperature (degrees Celsius) 1 centimeter below water's surface and 1 meter from shore. For bodies of water less than 2 meters wide, take temperature from the center. Be sure to shade the thermometer when possible.

PH: Use a pH meter to measure. The water sample should be taken from water column 1 meter from shore. For bodies of water less than 2 meters wide, take the sample from the center. Be sure to: 1) take the cap off the meter before using, 2) keep the level of the water sample below the mark on the meter, 3) turn the meter on before measuring the pH of the sample, and 4) turn the meter off when finished sampling. Meters should be calibrated monthly.

COND: Use a dissolved solids meter to measure. The water sample should be taken 1 centimeter below waters' surface and 1 meter from shore. For bodies of water less than 2 meters wide, take the sample from the center. Multiply LCD reading by ten and record value as 1.4 (micro-Seimens). Be sure to: 1) take the cap off the meter before using, 2) keep the level of the water sample below the mark on the meter, 3) turn the meter on before measuring the conductivity of the sample, and 4) turn the meter off when finished sampling. Meters should be calibrated monthly.

D02: Use a dissolved oxygen test kit to measure. The water sample should be taken 1 centimeter below waters' surface and 1 meter from shore. For bodies of water less than 2 meters wide, take the sample from the center. Record as ppm (parts-per-million). Be sure to: 1) plug in probe, 2) turn meter to the 02 setting, 3) let the meter stabilize in the atmosphere before testing the water sample, 4) let the meter stabilize in the sample before recording the value of the water sample, and 5) turn the meter off when finished sampling. Meters should be calibrated monthly.

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RH: Measure with a psychrometer 1.5 meters above ground and 1.5 meters from water. Record as percent. Be sure to continue to spin the psychrometer until the wet bulb has stabilized.

CLARITY: Circle one value (1-5) that best describes the survey area:

1 — extremely clear 2 somewhat clear 3 — moderately turbid 4 — somewhat heavy turbidity 5 — extremely heavy turbidity

LENTAX_L: For lentic systems, record the longest axis of the system in meters. Measure the entire system (not just the portion surveyed), and use the standing water at the time of the survey as your boundaries (do not use the normal waterline or highwater mark). For larger systems, estimate the length of the long axis using a map (don't rely on a visual guesstimate).

LENTAX_S: For lentic systems, record the shortest axis of the system in meters. This short axis should be the mean perpendicular axis to "LENTAX_L". As with "LENTAX_L", the short axis should reference the entire lentic system, not just the portion surveyed, and should be determined based upon the standing water present at the time of the survey, not the usual waterline or highwater mark. Use a map as a guide for larger systems.

LOT_VVI DE: For lotic systems, select one value (1-7) that best describes the width of water at the time of the survey (not at the normal waterline or at the highwater mark):

1 — 0-2 meters 2 — >2-5 meters 3 —>5-10 meters 4— >10-20 meters 5 — >20-50 meters 6— >50-100 meters 7— >100 meters

SUBSTR: Circle from one to three categories (1-6) as appropriate. All substrate types may be present, but choose only those that best describe the area potentially inhabited by target species. In the box below each category circled, record the percent occurrence of that substrate. Percents should total 100.

1 — mud and sitt(0.001-0.1 mm) 2 — sand (0.1-2 mm) 3 — gravel (2-32 mm) 4 — cobble(32-256 mm) 5 — boulder(>256 mm) 6— bedrock(exposed sheet of rock)

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WIND: Use a wind meter to measure wind speed, then check one category (0-6) as appropriate. Wind should be measured 1.5 meters above the ground and 1.5 meters from the water. Be sure to: 1) hold meter near the top so that you are not blocking any holes, 2) face into the direction of the wind while reading the meter, and 3) use the left scale for wind strengths <10 mph, and use the right scale (by putting your index finger over the red knob on top of the meter) for wind strengths 10 mph. Wind categories are those used in the Beaufort scale:

0 — s1 mph, smoke rises vertically 1 — 1-3 mph, wind direction shown by smoke drift 2 — 4-7 mph, wind felt on face, leaves rustle 3 8-12 mph, leaves and small twigs in constant motion, wind extends light flag 4 — 13-18 mph, raises dust and loose paper, small branches are moved 5 — 19-24 mph, small trees begin to sway, crested wavelets form on inland waters 6 — >24 mph, greater effect than above

CLOUD: Circle one category (1-5) as appropriate. Categories are based on percent cover:

1 —0 - 20% cover 2-21 - 40% cover 3-41 - 60% cover 4 — 61 - 80% cover 5 — 81 - 100% cover (includes fog)

PPT: Circle one category (0-4) as appropriate. Precipitation categories are based upon the type and degree of precipitation:

0 — no precipitation 1 — intermittent rain 2 — steady light rain 3 — steady heavy rain 4 — snow/sleet

T_STORM: Circle one category (0-2) as appropriate. Thunder storm categories are based upon the presence and proximity of thunder and lightening:

0 — no thunder or lightning 1 — thunder and lightning present at a distance (> 10 seconds between lightning

and thunder) 2 — thunder and lightning present nearby (s 10 seconds between lightning and

thunder)

FLOAT_VEG: Record the percent of the area potentially inhabited by target spp. that is covered by floating vegetation(e.g. broad-leafed macrophytes and dense algal mats). Use increments of 5% (1% effectively = 0). Write down the genus name or common name (only if positive id.) of 1-4 of the most prominent species— those species which best describe the surveyed area.

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SUB_VEG:

EMERG_VEG:

Record the percent of the area potentially inhabited by target spp. that is covered by submerged vegetation. Use increments of 5% (1% effectively = 0). Write down the genus name or common name (only if positive id.) of 1-4 of the most prominent species— those species which best describe the surveyed area.

Record the percent of the area potentially inhabited by target spp. that is covered by emergent vegetation (e.g cattails, sedges, rushes) . Use increments of 5% (1% effectively = 0). Write down the genus name or common name (only if positive id.) of 1-4 of the most prominent species-- those species which best describe the surveyed area.

PERIM_VEG: Record the percent of the area potentially inhabited by target spp. that is covered by perimeter vegetation (up to 1 m from waters edge). Use increments of 5% (1% effectively = 0). Write down the genus name or common name (only if positive id.) of 1-4 of the most prominent species— those species which best describe the surveyed area.

CANOPY_VEG: Record the percent of the area potentially inhabited by target spp. that is covered by canopy vegetation. Use increments of 5% (1% effectively = 0). Write down the genus name or common name (only if positive id.) of 1-4 of the most prominent species—those species which best describe the surveyed area.

* PRED: Circle all appropriate boxes so as to indicate the type of predators seen or otherwise detected (i.e. by sign) at a survey site. In the boxes below the predator types, insert a code (1-3) so as to suggest the magnitude of each predators' abundance:

1 — present/detected 2 — moderate numbers present 3 — abundantly present

Most of the predator categories lump together similar organisms and/or organisms with similar effects on riparian herps:

LEECH leeches CRAY crayfish (include claws and carapaces as evidence of presence) ANSOP dragonflies, adults and larvae BELO belostomatids BEET large aquatic beetles: hydrophilids and dytiscids VWVFSH warm water fish: bass, carp, catfish, perch, sunfish, walleye CWFSH cold water fish: trout, pike AMTI

tiger salamander (also write-up in the Herpetofauna Observations table on the front of the survey form)

BFRG

bullfrogs (also write-up in the Herpetofauna Observations table on the front of the survey form)

TURT

mud turtles (also write-up in the Herpetofauna0bservations table on the front of the survey form)

GSNK

garter snakes (also write-up in the Herpetofauna Observations table on the front of the survey form)

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HRON large wading birds: American bittern, black-crowned night heron, egrets, great blue heron, green-backed night heron

BHWK common black-hawk and zone-tailed hawk MAMM medium-sized mammals: skunk, ring-tail, raccoon (include footprints

and scat as evidence of presence)

* LAND_USE:

* OTHER_ORGS:

ORG_OBS:

Circle all appropriate boxes so as to best indicate the type of land use at a survey site. For noteworthy land uses that are not immediately at the survey site but which may potentially impact the study site (e.g. large agricultural fields within 1 mile of survey site, active mining operation 0.5 mile upstream of survey area), fill out the land use field as described here, and also make written comments about the land use in the "NOTES" field. In the boxes below the land-use types, insert a code (1-3) so as to suggest the magnitude of each land-use occurrence:

1 — use detected 2 — moderate usage 3 — heavy usage

The land-use categories are:

AGRIC agriculture (include agriculture fields, diversion canals, etc.) DEV human development (include road construction, dam site, housing

development, etc) GRAZE cattle grazing (include manure, hoofprints, increaser species, and

grass length as evidence of grazing use); note elk/deer grazing in "OTHER_ORGS" and"ORG_OBS", but only if heavy

LOG logging MINE mining (include 50+ year tailings/shafts, currently active mines,

small claims, and large developments) REC recreation (include campsites (developed and primitive), trails,

litter, etc.)

This field is to be used for observations of species other than riparian herpetofauna (riparian herps are to be recorded in the "Herpetofauna Observations" table on the front of the survey form). Use "OTHER_ORGS" to list all non-riparian herps (by 4-letter genus/species code [see "Herpetofauna List -Derived from Stebbins (1985)1), federal or state sensitive species of other organismal groups (by common name), or any other species whose occurrence merits noting (also by common name). No verbiage other than the species name(s) should be listed (e.g. UXOR, SCC, great homed owl, elk). Use the "ORG_OBS" field as needed to expand upon why you listed a species.

This is an optional field. Use this field to write out noteworthy observations about any or all of the species listed in "OTHER_ORGS" (e.g. UXOR observed mating, great homed owl roost site observed, area heavily impacted by elk grazing).

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Appendix B. Locality distribution by county and native ranid observation status.

Observed Not Observed Total

Count Row % Col % Count Row % Col % Count Col % Apache 34 28% 6% 89 72% 9% 123 8% Cochise 98 44% 16% 123 56% 12% 221 14% Coconino 45 15% 8% 252 85% 25% 297 18% Gila 70 36% 12% 126 64% 12% 196 12% Graham 36 51% 6% 34 49% 3% 70 4% Greenlee 25 47% 4% 28 53% 3% 53 3% La Paz 7 88% 1% 1 13% 0% 8 0% Maricopa 43 39% 7% 67 61% 7% 110 7% Mohave 32 50% 5% 32 50% 3% 64 4% Navajo 20 43% 3% 27 57% 3% 47 3% Pima 32 52% 5% 29 48% 3% 61 4%

Pinal 16 42% 3% 22 58% 2% 38 2% Santa Cruz 51 45% 9% 63 55% 6% 114 7%

Yavapai 86 40% 14% 130 60% 13% 216 13%

Yuma 5 100% 0% 5 0%

Total 595 37% 100% 1028 63% 100% 1623 100%

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Appendix C. Locality distribution by drainage and native ranid observation status.

Observed Not Observed Total

Count Row % Col % Count Row % Col % Count Col %

Colorado River - Glen Canyon 1 100% ' 0% 1 0% Colorado - San Juan River 1 100% 0% 1 0% Colorado River - Grand Canyon 6 10% 1% 53 90% 5% 59 4% Little Colorado River 48 24% 8% 151 76% 15% 199 12% Lower Colorado River 5 100% 0% 5 0% Bill Williams River 53 74% 9% 19 26% 2% 72 4% Upper Gila River 54 52% 9% 49 48% 5% 103 6% Middle Gila River 11 55% 2% 9 45% 1% 20 1% San Pedro River 104 45% 17% 126 55% 12% 230 14% Santa Cruz River 61 43% 10% 82 57% 8% 143 9% Salt River 98 33% 16% 200 67% 19% 298 18% Verde River 73 24% 12% 231 76% 22% 304 19%

Agua Fria River 50 41% 8% 73 59% 7% 123 8% Rio Concepcion 11 58% 2% 8 42% 1% 19 1%

Rio Yaqui 24 52% 4% 22 48% 2% 46 3%

Total 595 37% 100% 1028 63% 100% 1623 100%

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Appendix D. Locality distribution by management category and native ranid observation status.

Observed Not Observed Total

Count Row ok

Col % Count Row %

Col % Count Col %

State Arizona Game & Fish 4 31% 1% 9 69% 1% 13 1% State Land Dept 32 55% 5% 26 45% 3% 58 4%

Federal BLM 48 49% 8% 50 51% 5% 98 6% National Park Service 1 11% 0% 8 89% 1% 9 1% National Wildlife Ref 8 80% 1% 2 20% 0% 10 1% Apache-Sitgreaves NF 45 24% 8% 144 76% 14% 189 12% Coconino NF 37 17% 6% 184 83% 18% 221 14% Coronado NF 104 45% 17% 125 55% 12% 229 14% Kaibab NF 5 7% 1% 63 93% 6% 68 4% Prescott NF 13 33% 2% 26 67% 3% 39 2% Tonto NF 93 41% 16% 134 59% 13% 227 14% Military 3 9% 1% 30 91% 3% 33 2%

Other Non State/Federal 202 49% 34% 207 51% 21% 409 26% Total 595 37% 100% 1028 63% 100% 1623 100%

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Appendix E. Elevational ranges within drainages by native ranid species.

Drainage

Meters Feet Minimum Maximum Minimum Maximum

Rana blairi Little Colorado River 2168 2168 7114 7114 San Pedro River 1268 1798 4160 5900 Rio Yaqui 1237 1499 4060 4917

Rana chiricahuensis Little Colorado River 1692 2310 5550 7580 Upper Gila River 1067 2524 3500 8280 San Pedro River 1219 2012 4000 6600 Santa Cruz River 1097 1882 3600 6175 Salt River 1439 2710 4720 8890 Verde River 1399 2057 4590 6750 Rio Concepcion 1061 1317 3480 4320 Rio Yaqui 1134 1975 3720 6480

Rana pipiens Colorado River - Glen Canyon 952 952 3122 3122 Colorado - San Juan River 1689 1689 5540 5540 Colorado River - Grand Canyon 1134 1250 3720 4100 Little Colorado River 1341 2789 4400 9150 Upper Gila River 1463 2524 4800 8280 Salt River 1458 2499 4785 8200 Verde River 1609 2268 5280 7440

Rana subaquavocalis San Pedro River 1501 1829 4925 6000 Rana tarahumarae Santa Cruz River 1109 1775 3640 5823

Rio Concepcion 1213 1213 3980 3980 Rana yavapaiensis Colorado River - Grand Canyon 543 564 1780 1850

Bill Williams River 146 1634 480 5360 Upper Gila River 817 1817 2680 5960 Middle Gila River 579 1341 1900 4400 San Pedro River 616 1524 2020 5000 Santa Cruz River 605 1775 1985 5823 Salt River 320 1585 1050 5200 Verde River 544 1585 1785 5200 Agua Fria River 503 1585 1650 5200 Rio Concepcion 1097 1378 3600 4520 Rio Yaqui 1135 1135 3725 3725

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Appendix F. Number of ranid frog localities per county by species.

Rana blairi

Rana chiricahuensis

Rana pipiens

Rana subaquavocalis

Rana tarahumarae

Rana yavapaiensis rim southeast

Count Count Count Count Count Count Count

Apache 18 28 Cochise 21 71 9 11 Coconino 1 9 40 1 Gila 12 1 58 Graham 4 25 9 Greenlee 6 2 19 La Paz 7 Maricopa 43 Mohave 2 30 Navajo 3 19 Pima 6 29 Pinal 16 Santa Cruz

43 8 13

Yavapai 5 1 80 Yuma . Total 22 57 145 93 9 8 316

* Several historical records of the lowland leopard frog from the lower Gila and Colorado rivers (where it was extirpated prior to 1970) are not included in our database due to an oversight.

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Appendix G. Native ranid species status within management category.

Rana blairi

June 1997 Page 84

Absent Historical Present Historical Unsurveyed Hist New Total ' Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt CI%

State Arizona Game & Fish 1 100% 50% 1 5% State Land Dept 2 67% 13% 1 33% 25% 3 14%

Federal National Wildlife Ref 1 100% 7% 1 5% Coconino NF 1 100% 7% 1 5% Coronado NF 1 100% 7% 1 5%

Other Non State! Federal 10 67% 67% 1 7% 50% 1 7% 100% 3 20% 75% 15 68% Total 15 68% 100% 2 9% 100% 1 5% 100% 4 18% 100% 22 100%

Rana chiricahuensis, Rim form

AbsentHhistorical Present Historical Cnt

Unsurveyed Rw%

Hist CI% Cnt

New Rw% CI% Cnt

Total CI% Cnt Rw% CI% Cnt Rw% CI%

State Arizona Game & Fish 1 50% 5% 1 50% 9% 2 4% Federal Apache-Sitgreaves

NF 8 57% 38% 1 7% 25% 4 29% 19% 1 7% 9% 14 25%

Coconino NF 6 55% 29% 1 9% 5% 4 36% 36% 11 19% Coronado NF 1 100% 5% 1 2% Tonto NF 2 22% 10% 3 33% 75% 4 44% 36% 9 16%

Other Non State / Federal 3 15% 14% 16 80% 76% 1 5% 9% 20 35% Total 21 37% 100% 4 7% 100% 21 37% 100% 11 19% 100% 57 100%

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Rana chiricahuensis, southeastern form

Absent Historical Present Historical Unsurveyed Hist New Total

Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt Rw`)/0 CI% Cnt CI%

State State Land Dept 1 17% 1% 1 17% 6% 4 67% 9% 6 4%

Federal BLM 3 100% 4% 3 2%

National Wildlife Ref 1 20% 6% 4 80% 24% 5 3%

Coronado NF 34 39% 51% 13 15% 76% 6 7% 35% 35 40% 80% 88 61%

Military 1 100% 6% 1 1%

Other Non State / Federal 29 69% 43% 2 5% 12% 6 14% 35% 5 12% 11% 42 29%

Total 67 46% 100% 17 12% 100% 17 12% 100% 44 30% 100% 145 100%

Rana olniens

Absent Historical Present Historical Unsurveyed Hist New Total

Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt CI%

State Arizona Game & Fish 1 50% 3% 1 50% 3% 2 2%

State Land Dept 2 100% 6% 2 2%

Federal National Park Service 1 100% 6% 1 1%

NF Apache-Sitgreaves 11 38% 37% 4 14% 33% 5 17% 14% 9 31% 56% 29 31%

Coconino NF 16 62% 53% 5 19% 42% 4 15% 11% 1 4% 6% 26 28%

Kaibab NF 1 20% 8% 1 20% 3% 3 60% 19% 5 5%

Other Non State / Federal 2 7% 7% 2 7% 17% 22 79% 63% 2 7% 13% 28 30%

Total 30 32% 100% 12 13% 100% 35 38% 100% 16 17% 100% 93 100%

Rana subaquavocalis

Present Historical New Total Cnt Rw% CI% Cnt Rw% CI% Cnt CI%

Federal Coronado NF 1 100% 33% 1 11%

Military 2 100% 33% 2 22%

Other Non State / Federal 2 33% 67% 4 67% 67% 6 67%

Total 3 33% 100% 6 67% 100% 9 100%

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Rana tarahumarae

Cnt Absent Historical Unsunreyed Hist Total

Rw% CI% Cnt Rw% CI% Cnt CI%

Federal Coronado NF 6 75% 100% 2 25% 100% 8 100%

Total 6 75% 100% 2 25% 100% 8 100%

Rana yavapalensis

Absent Historical Present Historical Unsurveyed Hist New Total

Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt Rw% CI% Cnt CI%

State State Land Dept 5 23% 7% 3 14% 7% 12 55% 9% 2 9% 3% 22 7%

Federal BLM 10 21% 14% 8 17% 19% 25 53% 18% 4 9% 7% 47 15%

National Wildlife Ref 1 25% 1% 1 25% 1% 2 50% 3% 4 1%

Apache-Sitgreaves NF 2 17% 5% 4 33% 3% 6 50% 10% 12 4%

Coconino NF 3 75% 4% 1 25% 2% 4 1%

Coronado NF 14 70% 19% 2 10% 5% 2 10% 1% 2 10% 3% 20 6%

Prescott NF 10 77% 7% 3 23% 5% 13 4%

Tonto NF 26 31% 36% 14 17% 33% 32 38% 23% 12 14% 20% 84 27%

Other Non State / Federal 13 12% 18% 13 12% 30% 54 49% 39% 30 27% 49% 110 35%

Total 72 23% 100% 43 14% 100% 140 44% 100% 61 19% 100% 316 100%

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Appendix H. Native ranid species status within drainage.

Rana Wahl

June 1997 Page 87

Absent Historical Present Historical Unsurveyed Historical New Total

Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % CI % Cnt Rw % CI % Cnt CI `)/0

Little Colorado River 1 100% 7% 1 5%

San Pedro River 8 62% 53% 1 8% 50% 4 31% 100% 13 59%

Rio Yaqui 6 75% 40% 1 13% 50% 1 13% 100% 8 36%

Total 15 68% 100% 2 9% 100% 1 5% 100% 4 18% 100% 22 100%

Rana chiricahuensis, Rim form

Cnt Absent Histor'cal Present Historical Unsurveyed Historical New Total

Rw % Cl % Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % Cl % Cnt Cl %

Little Colorado River 8 67% 38% 3 25% 14% 1 8% 9% 12 21%

Upper Gila River 7 47% 33% 6 40% 29% 2 13% 18% 15 26%

Salt River 3 14% 14% 4 19% 100% 11 52% 52% 3 14% 27% 21 37%

Verde River 3 33% 14% 1 11% 5% 5 56% 45% 9 16%

Total 21 37% 100% 4 7% 100% 21 37% 100% 11 19% 100% 57 100%

Rana chiricahuensis, southeastern form

Absent Historical Present Historical Unsurveyed Historical New Total Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % CI % Cnt Cl %

Upper Gila River 10 71% 15% 3 21% 18% 1 7% 6% 14 10%

San Pedro River 24 36% 36% 6 9% 35% 6 9% 35% 30 45% 68% 66 46%

Santa Cruz River 20 59% 30% 5 15% 29% 4 12% 24% 5 15% 11% 34 23%

Rio Concepcion 2 20% 3% 2 20% 12% 6 60% 14% 10 7%

Rio Yaqui 11 52% 16% 1 5% 6% 6 29% 35% 3 14% 7% 21 14%

Total 67 46% 100% 17 12% 100% 17 12% 100% 44 30% 100% 145 100%

Rana pipiens

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Absent Historical Present Historical Unsurveyed Historical New Total

Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % CI % Cnt Cl %

Colorado River - Glen Cny 1 100% 6% 1 1%

Colorado - San Juan River 1 100% 3% 1 1%

Colorado River - Grand Cny 3 100% 19% 3 3%

Little Colorado River 16 36% 53% 4 9% 33% 15 34% 43% 9 20% 56% 44 47%

Upper Gila River 5 71% 17% 2 29% 6% 7 8%

Salt River 1 6% 3% 15 94% 43% 16 17%

Verde River 8 38% 27% 8 38% 67% 2 10% 6% 3 14% 19% 21 23%

Total 30 32% 100% 12 13% 100% 35 38% 100% 16 17% 100% 93 100%

Rana subaquavocalis

Present Historical New Total Cnt Rw % Cl % Cnt Rw % Cl % Cnt Cl %

San Pedro River 3 33% 100% 67% 100% 9 100% Total 3 33% 100% 6 67% 100% 9 100%

Rana tarahumarae

Absent Historical Unsurveyed Historical Total Cnt Rw % Cl % Cnt Rw % Cl % Cnt Cl %

Santa Cruz River 5 71% 83% 2 29% 100% 7 88% Rio Concepcion 1 100% 17% 1 13% Total 6 75% 100% 2 25% 100% 8 100%

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Rana yavapaiensis

Cnt Absent Historical Present Historical Unsurveyed Historical New Total

Rw % Cl % Cnt Rw % Cl % Cnt Rw % Cl % Cnt Rw % Cl % Cnt CI %

Colorado River - Grand Cny 2 67% 3% 1 33% 2% 3 1%

Bill Williams River 7 13% 10% 6 11% 14% 25 47% 18% 15 28% 25% 53 17%

Upper Gila River 1 4% 1% 4 16% 9% 6 24% 4% 14 56% 23% 25 8%

Middle Gila River 11 100% 8% 11 3%

San Pedro River 7 28% 10% 7 28% 16% 9 36% 6% 2 8% 3% 25 8%

Santa Cruz River 15 48% 21% 2 6% 5% 11 35% 8% 3 10% 5% 31 10%

Salt River 19 28% 26% 12 18% 28% 26 39% 19% 10 15% 16% 67 21%

Verde River 14 31% 19% 3 7% 7% 17 38% 12% 11 24% 18% 45 14%

Agua Fria River 3 6% 4% 7 14% 16% 34 68% 24% 6 12% 10% 50 16%

Rio Concepcion 4 80% 6% 1 20% 2% 5 2%

Rio Yaqui 1 100% 1% 1 0%

Total 72 23% 100% 43 14% 100% 140 44% 100% 61 19% 100% 316 100%

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