Sh vol 27 2014 09 reduced

25 Volume 27

January 2012 September 2014

THIS MONTH’S PROGRAM

FUTURE SPEAKERS

Roger A. Repp A Photographic History of the Suizo Mountain Project

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21 Oct 2014: B. K. Sullivan

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12 Nov 2014: Jim Jarchow

F E AT U R E A R T I C L E

7:15 PM; Tuesday, 16 September “Recital Hall” of Pima Community College West Campus, 2202 W. Anklam Road, Tucson, AZ 85709

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B y vocation, Roger is a tool and die maker, currently employed as Instrument Shop Su-

pervisor for National Optical Astronomy Observatory. By avocation, Roger is a rabid field herpetologist, who, since 1989, has averaged 870 hours per year studying the herpetofauna of the Sonoran Desert. His methods involve radio-telemetry, mark and recapture studies, as well as hands off observations.

Roger’s presentation will first touch on the discovery of Iron Mine Hill in August of 1992. Over the next nine years, Iron Mine Hill gradually became his favored herpetological playground. In March of 2001, Roger joined forces with Dr. Gordon Schuett to begin a radio telemetry study of five species of venomous reptiles there. The “Suizo Mountain Project” continues to this day, making it the longest continuous multi-species venomous reptile radio telemetry study in the history of Arizona. Eight peer-reviewed publications have arisen from this study, and several more are in the hopper.

Number 13

ISSN 2333-8075

“Reproduction in the Ocellated Skink, Chalcides ocellatus (Squamata: Scincidae) in Israel, already established in Mesa, AZ”

by Stephen R. Goldberg

100-MILE CIRCLE 61

“Chiricahua Leopard Frog (Lithobates chiricahuensis)” by Rorabaugh and Sredl

CO N S E R VAT I O N 71

Threatened Narrow-headed Gartersnakes Born at Phoenix Zoo

I N T E R N AT I O N A L Gordon Schuett and Roger Repp with Heloderma suspectum #1, “Geronimo.” Photo by David L. Hardy, Sr., 16 March 2001. Valid permits authorized the handling of H. suspectum for scientific purposes.

But the focus of this presentation is not about people or papers. It centers on images of the real stars of the study—the herpetofauna of the Suizo Mountains, and the ground that they live on and under.

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“Biodiversity of Reptiles and Amphibians of India” by Suman Pratihar and Kaushik Deuti

E N D A N G E R E D S P. 74

“Flat-Tailed Horned Lizard Proposed Rule” by Richard Campbell

ANNOUNCEMENT

N AT U R A L H I S TO RY

Desert Snakes: Mystery and Intrigue

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“Mating in the Coachwhip, Masticophis (= Coluber) flagellum”

by Brian K. Sullivan

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“Encounter between a Striped Racer (Coluber lateralis) and a Southern Alligator Lizard (Elgaria multicarinata)” by Howard Clark

Y ucca Valley, CA—William Hayes will introduce participants to a remarkable variety

of venomous and non-venomous snakes that inhabit Southern California’s deserts in this all-day field class. The class will explore the surrounding desert to identify reptile habitat and hopefully see some snakes!

Date/Time: Sun., Sept. 28, 0900–1600 Meet at: Black Rock Visitor Center, 9800 Black Rock Cyn Rd, Yucca Valley, CA Non-Credit Fee: $60 for JTNPA/PINE members, $70 non-members Credit Fee: $85 for JTNPA/PINE members, $95 nonmembers

BOOK REVIEWS— PAG E 80

SONORAN HERPETOLOGIST 27 (3) 2014

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FUTURE SPEAKERS

21 October 2014:

Brian K. Sullivan Professor of Herpetology

Seasonal Variation in Spatial Ecology of Sonoran Desert Tortoises 7:15 PM; “Recital Hall” of Pima Community College West Campus, 2202 W. Anklam Road, Tucson, AZ 85709

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ullivan first searched for amphibians and reptiles in Arizona in 1970; he returned to attend graduate school in 1979, and has been in residence for most years since. He first taught at ASU at the West Campus in 1984, and arrived as an Assistant Professor of Zoology at ASU in 1989. He was promoted to Full Professor of Herpetology in 2002, and served as the Editor of the Journal of Herpetology from 2000 through 2005.

He is interested in the behavior, ecology and conservation of amphibians and reptiles around the world, and especially in desert regions like the southwest. He has authored over 100 articles, book reviews, technical reports, and book chapters, including many coauthored with students and colleagues in the Phoenix area.

His talk will focus on recent work with Sonoran Desert Tortoises (Gopherus morafkai) near Phoenix. In 2010 he and his family initiated a radio-telemetry study of a population of G. morafkai in the Union Hills on the northern edge of Phoenix metropolitan region.

Preliminary results provide insights on consistency in

Brian records the body temperature of a Sonoran Gopher Snake (Pituophis catenifer affinis) at his long-term study plot in north Phoenix. Photo by Keith O. Sullivan, 2011.

refuge use seasonally, and movements associated with foraging behavior in the fall. Virtually all adults exhibited movement from lower elevation areas associated with drainages to relatively higher elevation, northfacing slopes following the monsoon rains, apparently to forage on abundant and diverse plants on those slopes. These results are consistent with the notion that tortoises in this population exhibit a pattern of annual migration for forage not unlike large herbivorous mammals. These results and other aspects of the natural history of these long-lived animals will be discussed.

12 November 2014:

Sullivan first searched for amphibians and reptiles in Arizona in 1970; he returned to attend graduate school in 1979, and has been in residence for most years since.

He first taught at ASU at the West Campus in 1984, and arrived as an Assistant Professor of Zoology at ASU in 1989.

Jim Jarchow, DVM Associate Veterinarian, Orange Grove Hospital Consulting Veterinarian, Arizona-Sonora Desert Museum Consulting Herpetological Veterinarian, Reid Park Zoo

The perils our native herps face sharing their habitat with us humans

Jim preparing a WesternDiamond-backed Rattlesnake (Crotalus atrox) for anesthesia prior to a surgical implantation of a transmitter, 19 July 2003. Photo by Roger A. Repp.

7:15 PM; Tucson City Council Ward 3, 1510 East Grant Road Tucson, AZ 85719

D r. Jim Jarchow began his career as a reptile keeper in 1964 at the Columbus Zoo in Columbus Ohio.

He received his DVM degree from The Ohio State University in 1971, and has been in practice in Tucson since 1972. Since that time period, Jim has used his talents in too many ways to enumerate here to help both the animals that he loves, and the people who study them. It was his steadfast, stalwart, and selfless dedication that earned him the very first THS Jarchow Conservation Award in 1992. This distinguished award remains in his name to this very day.

Jim’s presentation will discuss various cases that fall under his care. Jim is often the recipient of wild herps found injured by the public, wildlife rehabilitators, and government employees. The cases will reveal a variety of hazards these animals face, and treatment techniques that are used to restore health and function.

In the best of all worlds, the care that Jim provides prepares them for release back to their home ranges.

Jim promises lots of images in his presentation, and very little text or analytical data.

SONORAN HERPETOLOGIST 27 (3) 2014

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RESEARCH ARTICLE

Reproduction in the Ocellated Skink, Chalcides ocellatus (Squamata: Scincidae) in Israel, already established in Mesa, AZ Stephen R. Goldberg, Whittier College, Department of Biology, Whittier, CA 90608; sgoldberg1109@gmail.com

T he Ocellated Skink, Chalcides ocellatus (Figure 1), is native to northern

Africa, the Middle East and the Mediterranean (Schleich et al. 1996).

It is viviparous (Mosauer 1934) and has a long history of human assisted introductions and now occurs in numerous sites outside its native range (Kraus 2009, Gunn et al. 2012) including Mesa, Arizona (Gunn et al. 2012). A detailed account of C. ocellatus reproduction in Egypt is in Badir (1959) and Badir and Hussein (1965). Anedoctal information on Figure 1. Ocellated Skinks (Chalcides ocellatus) breeding, Mesa, Arizona. reproduction of C. ocellatus consist Photo by Robert Bowker. mainly of reports of clutch sizes in lumina of the seminiferous tubules are lined by sperm regional field guides and other experimental studies or clusters of metamorphosing spermatids. The period utilizing C. ocellatus include: Mosauer (1934), Daut and Andrews (1993), Corso et al. (1978), Ebaid (2011). The of spermiogenesis included March, April and May. Chalcides ocellatus with testes exhibiting recrudescence purpose of this paper is to present data on reproducoccurred during September to April. The smallest tion on C. ocellatus from Israel, part of its original reproductively active male measured 68 mm SVL range. Bar and Haimovitch (2011) reported C. ocellatus (TAUM 840) and was collected in April. produced 3 to 10 young in July in Israel. Comparisons The mean SVL of the female sample (n = 32) was are made between reproduction of C. ocellatus outside significantly greater than that of the male sample (n its native range. = 28) (unpaired t-test, t = 2.5, df = 58, P = 0.014).

A sample of 66 C. ocellatus from Israel consisting This has also been reported for C. ocellatus from Egypt of 28 adult males (mean SVL = 82.8 mean snout(Badir 1959). Four stages were observed in the ovarian vent length, ± 7.6 SD, range = 68–94 mm), 32 adult cycle (Table 2): (1) quiescent, no yolk deposition. (2) females (mean SVL = 88.0 mm ± 8.2 SD, range = yolk deposition, basophilic yolk granules present in 75–108 mm) and six juvenile females (mean SVL = ooplasm. (3) enlarged follicles > 4 mm. (4) oviductal 68.2 mm ± 2.9 SD, range = 65–73 mm) deposited eggs, no visible embryos. The period of reproducin the herpetology collection of Tel Aviv University tive activity encompassed March to May. The small(TAUM), Tel Aviv, Israel was examined. Skinks were collected 1942 to 1956. Regions where C. ocellatus were est reproductively active female (yolk deposition in progress) measured 75 mm SVL (TAUM 846) and was collected in Israel are in the appendix. collected in March. Six smaller females (mean SVL A small incision was made in the lower part of the abdomen and the left gonad was removed for histolog- = 68.2 mm ± 2.9 SD, range = 65–73 mm) contained quiescent ovaries and were considered to be juveniles. ical examination. Gonads were embedded in paraffin, Mean litter size for four females (follicles > 4 mm or sections were cut at 5 µm and stained with Harris hematoxylin followed by eosin counterstain (Presnell and oviductal eggs) was 3.8 ±0.96 SD, range = 3–5.

My mean clutch size and range 3.8 ±0.96 SD, 3–5 Schreibman 1997). Histology slides were deposited at is within that reported for C. ocellatus from other areas TAUM. Enlarged ovarian follicles or oviductal eggs (Table 3). There was a significantly positive correlation were counted. An unpaired t-test (Instat, vers. 3.0b, between number of embryos and female body size Graphpad Software, San Diego, CA) was used to test for C. ocellatus from Egypt (Badir and Hussein 1965). for differences between male and female mean SVLs. However, my sample of gravid females from Israel Two stages were noted in the testicular cycle (Table 1): (1) recrudescence, the germinal epithelium is was too small (n = 4) to examine this relationship.

The smallest reproductively active female C. ocellatus replenishing cells for the next period of spermiogenfrom Israel (75 mm SVL) was 15 mm smaller than esis. Primary spermatocytes predominate in early recrudescence; secondary spermatocytes and spermatids the smallest female from Egypt (90 mm SVL) (Badir predominate in late recrudescence. (2) spermiogenesis, 1957). The testes of C. ocellatus in both Egypt (Badir

SONORAN HERPETOLOGIST 27 (3) 2014

The Ocellated Skink, Chalcides ocellatus, is native to northern Africa, the Middle East and the Mediterranean.

It is viviparous and has a long history of human assisted introductions and now occurs in numerous sites outside its native range including Mesa, Arizona.

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1959) and Israel (Table 1) began sperm production in March. The length of the period of sperm production in C. ocellatus from Israel is not known as no males from June, July and August were examined. However, because spermiogenesis in Egypt ends in June (Badir 1959) it seems likely that sperm production also terminates in Israel in June. In both Egypt (Badir 1957) and Israel (Table 1) recrudescence (= renewal of testicular germinal epithelium) proceeds during autumn in preparation fo spermiogenesis in the spring. Clutch (litter) sizes are variable in different parts of the C. ocellatus range (Table 3).

It has been shown that there are significant geographic differences in timing of events of the C. ocellatus reproductive cycle (Badir and Hussein 1965).

Chalcides ocellatus from Egypt commenced reproduction five weeks earlier than C. ocellatus from Europe (Badir and Hussein 1965). Reproduction in Egypt terminated later (mid-September) in contrast to Europe where it terminated in August to early September (Badir and Hussein 1965).

The success of C. ocellatus colonizing different areas is partly due to its toleration of diverse habitats and environmental conditions (Schleich et al. 1996, Schlüter 2006 ). While monthly data on reproduction of C. ocellatus from Mesa, AZ, are not yet available, one can surmise from previous works (Badir and Hussein 1965, this paper) that breeding will occur in spring and one or possibly two litters may be produced (Badir and Hussein 1965). In view of its ability to colonize in different environments and the possibility of escape from pet owners, steps should be taken to limit its importation into the United States.

Acknowledgments—I thank Shai Meiri (TAUM) for permission to examine C. ocellatus, Erez Maza (TAUM) for facilitating the loan and the National Collections of Natural History at Tel Aviv University for providing samples of C. ocellatus for study.

Table 1. Monthly stages in the testicular cycle of 28 adult male Chalcides ocellatus from Israel.

Month

n

Recrudescence

Spermiogenesis

January

1

1

0

February

3

3

0

March

13

7

6

April

5

2

3

May

1

0

1

September

1

1

0

November

2

2

0

December

2

2

0

Table 2. Monthly stages in the ovarian cycle of 32 adult female Chalcides ocellatus from Israel.

Month

n

Quiescent

Early yolk deposition

Follicles > 4 mm

Oviductal eggs

Jan

2

2

0

0

0

Feb

3

3

0

0

0

March

9

6

2

1

0

April

7

5

0

0

2

May

1

0

0

1

0

Sept

3

3

0

0

0

Oct

1

1

0

0

0

Nov

3

3

0

0

0

Dec

3

3

0

0

0

Table 3. Clutch (litter) sizes for Chalcides ocellatus from different geographic locations; UAE = United Arab Emirates.

Location Italy

Clutch (litter) size 6-8

North Africa

2-20, 4-8 usually

Source Corso et al. 1978 Schleich et al. 1996

Turkey

2-10

Baran and Atatur 1998

UAE

3-10

Jongbloed 2000

Literature Cited

Jordan

4-10

Disi et al. 2001

Badir, N. 1959. Seasonal variation of the male urogenital organs of Scincus scincus L. and Chalcides ocellatus Forsk. Zeitschriftn für wissenschaftliche Zoologie 160:290-351.

Badir, N., and M.F. Hussein. 1965. Effect of some ecological conditions on the reproduction of Chalcides ocellatus (Forsk.) Bulletin of the Faculty of Sciences, Cairo University 39:159-168.

Baier, F., D.J. Sparrow, and H. Wiedl. 2013. The Amphibians and Reptiles of Cyprus. Edition Chimaira, Frankfurt am Main, Germany.

Bar, A., and G. Haimovitch 2011. A Field Guide to Reptiles and Amphibians of Israel. Pazbar Ltd., Haerzilya, Israel.

Baran, I., and M.K. Atatur. 1998. Turkish Herpetofauna (Amphibians and Reptiles). Ankara, Turkey.

Corso, G., Lissia Frau, A.M., and Pala, M. 1978. Struttura dell’ovario adulto di Chalcides ocellatus tiligulu

Eastern Palearctic

4-5

Szczerbak 2003

Sri Lanka

4-10

Somaweera and Somaweera 2009

Europe

5-15

Kwet 2010

Ethiopia and Eritrea

4-8

Largen and Spawls 2010

Israel

3-10

Bar and Haimovitch 2011

Egypt

3-11

Ebaid 2011

Cyprus

3-10

Baier et al. 2013

UAE, Oman

4-6

Gardner 2013

The success of C. ocellatus colonizing different areas is partly due to its toleration of diverse habitats and environmental conditions. While monthly data on reproduction of C. ocellatus from Mesa, AZ is not yet available, one can surmise from previous works that breeding will occur in spring and one or possibly two litters may be produced.

(Gmelin) durante il corso dell’ anno. Archivo italiano di Anatomia e di Embriologia 83:207-223.

Daut, E.F., and R. M. Andrews. 1993. The effect of pregnancy on thermoregulatory behavior of the viviparous lizard Chalcides ocellatus. Journal of Herpetology 27:6-13.

Disi, A. M., D. Modry, P. Necas and L. Rifai. 2001.

SONORAN HERPETOLOGIST 27 (3) 2014

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Amphibians and Reptiles of the Hashemite Kingdom of Jordan. An Atlas and Field Guide. Edition Chimaira. Frankfurt am Main, Germany.

Ebaid, H.M. 2011. Associations between reproductive cycle and occurrence of intestinal helminths infecting the Egyptian lizard, Chalcides ocellatus. Egyptian Academy Journal of Biological Sciences 3:19-26.

Gardner, A. S. 2013. The Amphibians and Reptiles of Oman and the UAE. Edition Chimaira, Frankfurt am Main, Germany.

Gunn, J., R.W. Bowker, K.O. Sullivan, and B.K. Sullivan. 2012. An Old World skink, Chalcides ocellatus, with along history of anthropogenically assisted dispersal, now established in Mesa, Arizona, Arizona, USA. Herpetological Review 43:551-553.

Jongbloed, M. 2000. Wild about Reptiles. Field Guide to the Reptiles and Amphibians of the UAE. Atlas Printing Press, Dubai, UAE. Kraus, F. 2009. Alien Reptiles and Amphibiens: a Scientific Compendium and Analysis/ Springer, Dordrecht, Netherlands.

Kwet, A. 2010. Reptilien und Amphibien Europas, 190 Arten mit Verbreitungskarten. Franckh-Kosmos

Verlags-GmbH & Co. Stuttgart, Germany Largen, M., and S. Spawls. 2010. The Amphibians and Reptiles of Ethiopia and Eritrea. Edition Chimaira, Frankfurt am Main, Germany.

Mosauer, W. 1934. The reptiles and amphibians of Tunisia. University of California at Los Angeles, Publications in Biological Sciences 1:49-64.

Presnell, J.K., and M.P. Schreibman. 1997. Humason’s Animal Tissue Techniques, 5th edition, The Johns Hopkins University Press, Baltimore, MD. Schleich, H.H., W. Kastle and K. Kabish. 1996. Amphibians and Reptiles of North Africa. Biology.

Systematics, Field Guide. Koeltz Scientific Books, Koenigstein, Germany.

Schlüter, U. 2006. Der Gefleckte Walzenskink Chalcides ocellatus. Natur und Tier – Verlag Gambh, Munster, Germany.

Somaweera, R., and N. Somaweera. 2009. Lizards of Sri Lanka, A Colour Guide with Field Keys. Edition Chimaira, Frankfurt am Main, Germany.

Szcerbak, N.N. 2003. Guide to the Reptiles of the Eastern Palearctic. Krieger Publishing Company, Malabar, Florida.

Appendix: Chalcides ocellatus examined from the herpetology collection of Tel Aviv University (TAUM ) by Israel region. Central Negev TAUM 821, 822, 835, 840, 851, 936, 2160-2169, 3905, 7206, 7208, 7440; Dead Sea Area TAUM 828, 845, 846, 5170; HaSharon TAUM 5759, 12809; Jordan Valley TAUM 814, 830, 12816; Karmel Ridge TAUM 817, 5152, 5153; Northern Coastal Plain TAUM 12871, 12873; Northern Negev TAUM 812, 819, 839, 852, 961, 1748, 1951, 1952, 1976, 2107, 2108, 2199, 2912, 2913, 3011, 3907, 4249, 4433, 4874, 5989, 13174; Shomeron TAUM 2097; South Negev TAUM 848; Upper Galil TAUM 826; Yehudah Desert TAUM 850, 890, 910, 911, 3357; Yehudah Mountains TAUM 3351; Yizreel Valley TAUM 813, 2235.

H E R P E TO FAU N A O F T H E 100- M I L E C I R C L E

Chiricahua Leopard Frog (Lithobates chiricahuensis) James C. Rorabaugh, P. O. Box 31, Saint David, AZ 85630; jrorabaugh@hotmail.com Michael J. Sredl, Arizona Game and Fish Department, 5000 W. Carefree Highway, Phoenix, AZ 85086

T he relationships of anurans commonly called “leopard frogs” or “meadow

frogs” have baffled herpetologists for over 100 years (Moore 1975, Hillis 1988).

From the mid-1920s through the early1960s, most authorities listed only one to three species of leopard frog in the southwestern United States (Moore 1944, Wright and Wright 1949, Stebbins 1951), including the broad-ranging Rana pipiens, as well as R. fisheri and R. onca of southwestern Nevada and adjacent portions of Utah and Arizona (although some considered the latter two to be subspecies of pipiens, or to be synonyms – see Pace 1974). Wright and Wright (1949) recognized considerable variation within R. pipiens in North America, even singling out questionable frogs in a section of their book called “Arizona Puzzles.” In addition, they were among the first authorities to refer to the leopard frogs as the R. pipiens complex. Interestingly,

Figure 1. Chiricahua Leopard Frog (Lithobates chiricahuensis), Coconino County, AZ. Photo by James C. Rorabaugh.

The relationships of anurans commonly called “leopard frogs” or “meadow frogs” have baffled herpetologists for over 100 years. From the mid-1920s through the early-1960s, most authorities listed only one to three species of leopard frog in the southwestern United States, including the broad-ranging Rana pipiens, as well as R. fisheri and R. onca of southwestern Nevada and adjacent portions of Utah and Arizona.

they noted large leopard frogs from Carr Canyon, Huachuca Mountains, Arizona that they keyed out to R. montezumae of Mexico, and this was among the first recognition of a frog that would later be named Lithobates chiricahuensis. Mecham (1968) quantified the

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Figure 2. Chiricahua Leopard Frog (Lithobates chiricahuensis), Huachuca Mountains, AZ. Photo by James C. Rorabaugh.

morphological differences between the “southern form” (= L. chiricahuensis) and the Northern Leopard Frog (L. pipiens) in the White Mountains of Arizona.

He stated that this southern form resembled frogs from southwestern New Mexico and southeastern Arizona. Employing electrophoresis of frog hemoglobin, Platz and Platz (1973) presented evidence for three species of leopard frogs in Arizona, including the northern form (= L. pipiens), Mecham’s southern form, and the lowland form. Platz and Mecham (1979) subsequently described the southern form as the Chiricahua Leopard Frog (Rana [= Lithobates] chiricahuensis) and Platz and Frost (1984) described the lowland form as the Lowland Leopard Frog (Rana [= Lithobates] yavapaiensis). The type locality for the Chiricahua Leopard Frog is Herb Martyr Reservoir in Cave Creek of the Chiricahua Mountains, Arizona. Frost et al. (2006) proposed the genus name Lithobates for all leopard frogs and other related ranid frogs, which was subsequently adopted by the Society for the Study of Reptiles and Amphibians (Crother 2008, 2012), but remains controversial (see Pauly et al. 2009).

Based primarily on its propensity to call underwater, large, knob-like toe-tips, and unique body proportions, Platz (1993) described the Ramsey Canyon Leopard Frog (Rana [= Lithobates] subaquavocalis) as distinct from the Chiricahua Leopard Frog. However, subsequent genetic studies did not support this designation (Goldberg et al. 2004, Hillis and Wilcox 2005), and subaquavocalis is now considered synonymous with L. chiricahuensis (Crother 2012). Using a 50-loci starch gel survey, morphometrics, and analyses of nuclear DNA, Platz and Grudzien (1999) presented evidence that northern populations of the Chiricahua Leopard Frog (north of the Gila River) represent a species distinct from the southern populations. Goldberg et al.

(2004) demonstrated that frogs from these two regions showed a 2.4% average divergence in mitochondrial DNA sequences. Hillis and Wilcox (2005) suggested the northern populations could be the Vegas Valley Leopard Frog (Rana [=Lithobates] fisheri), a species not seen since 1942 and thought to be extinct (Jennings

2005). In a recent study, Hekkala et al. (2011) confirmed that mitochondrial and nuclear DNA differed between most northern and southern populations, and that populations in the western part of the Mogollon Rim were concordant with Rana (= Lithobates) fisheri. The Gila River is a rough boundary between the two forms in Arizona, but the southern populations include frogs from Dix Creek and adjacent areas north of the Gila River, and in west-central New Mexico, the boundary between these taxa may be an east-west, rather than north-south divide. However, further work in the lab of Dr. Melanie Culver at the University of Arizona using samples collected throughout the range of the species should clarify these relationships. Until this study is complete, we consider the northern and southern populations to be L. chiricahuensis.

Description and Similar Species The Chiricahua Leopard Frog is a large (< 125 mm SVL) and stocky frog with a relatively rugose or tuberculate dorsum and dorsolateral folds that are broken posteriorly and inset medially (Platz and Mecham 1979, Platz et al. 1997, Figs. 1 and 2). The eyes are protuberant, positioned relatively high on the head, and have been called “upturned” (Stebbins 2003). The snout is rounded. Males possess prominent vocal sacs, one on each side just below and behind the tympanum. Webbing is lacking between the digits of the hands, but usually extends to the knuckle between the second to the last and distal phalanges on all but the third (and longest) digit of the feet. Webbing extends from about half to most of the distance along that third digit. Males from the southern populations have rudimentary vestigial oviducts, whereas northern males generally lack any evidence of oviducts (Platz and Mecham 1979, Degenhardt et al. 1996, Rorabaugh pers. obs.).

The dorsal background color varies from light brown to dark brownish-gray or green. In some populations, such as in the Huachuca Mountains, most frogs are a bright green over most of the dorsum. In other areas, green may be present only atop the head and on the upper back, or it may be absent entirely.

Frogs from northern populations are much less likely

The Chiricahua Leopard Frog is a large (< 125 mm SVL) and stocky frog with a relatively rugose or tuberculate dorsum and dorsolateral folds that are broken posteriorly and inset medially.

The eyes are protuberant, positioned relatively high on the head, and have been called “upturned”.

The snout is rounded.

Figure 3. Chiricahua Leopard Frog thigh pattern, Santa Cruz County, AZ. Photo by James C. Rorabaugh.

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Figure 4. Late stage Chiricahua Leopard Frog tadpole, Huachuca Mountains, AZ. Photo by James C. Rorabaugh.

to be green. The back, sides, and head have numerous small dark brown to black spots. One or more spots are usually present on the head anterior to the eyes.

Most individuals lack a light spot on the tympanum.

The hind limbs have dark crossbars, and the front limbs are similarly barred or have dark spots. Dorsal spots and crossbars often fade with age and are absent in some very large frogs. The rear of the thigh in adults from southern populations typically is a salt and pepper pattern consisting of a very dark background overlain with numerous white-tipped pustules that are most prominent near the urostyle (Fig. 3). Juveniles may exhibit a dark reticulation on the rear of the thigh or have the salt and pepper pattern. The salt and pepper pattern is also present in many northern frogs, but some juveniles and adults exhibit a dark and light reticulate pattern. There is no discernible light upper lip stripe anterior to the eyes, although that area may be a green that is somewhat lighter in color than surrounding pigment. The venter is cream to yellowish cream, often with grayish mottling on the chin and throat (Platz and Mecham 1979, Degenhardt et al.

1996, Dodd 2013, Rorabaugh pers. obs.). Individual Chiricahua Leopard Frogs can darken their skin color in response to low water temperature and reduced reflectance off the water’s surface (Fernandez and Bagnara 1991, 1993).

Lemos-Espinal et al. (2013) noted some differences in Chiricahua Leopard Frogs from southwestern Chihuahua as compared to those in Sonora and the United States. The maximum SVL is 58 mm, and they are dark gray-brown and have fewer spots on the dorsum. The authors suggest these differences may reflect taxonomic distinctiveness.

Chiricahua Leopard Frog tadpoles are relatively stocky for a leopard frog and grow to > 80 mm total length prior to metamorphosis. Small tadpoles are a dark velvety olive dorsally and on the sides, and white to pale gray with bronze patches ventrally. Large tadpoles are generally dark gray or olive dorsally with pale venters and heavy mottling on the tail fin (Scott and Jennings 1985, Fig. 4). Mouth parts are illustrated in Scott and Jennings (1985). Lateral lines of southwestern US leopard frog tadpoles may be diagnostic (Fritts et al. 1984), but these differences need to be explored.

None of the other five leopard frogs in Arizona, or other three leopard frogs in the 100-Mile Circle, ever exhibit a salt and pepper pattern on the rear of the thigh. The Chiricahua Leopard Frog is also the only leopard frog species in our area likely to have a dorsum that is entirely green (although many Chiricahua Leopard Frogs lack green in the dorsal pattern).

The upturned eyes are also diagnostic. In all other leopard frogs in our area the eyes are positioned lower down, so the angle of view is more or less lateral to the head, rather than upturned. These characters combined with a preponderance of small, dark spots on the dorsum and relatively rough skin are usually enough to distinguish this species from other southwestern leopard frogs. The advertisement call of the Chiricahua Leopard Frog is also diagnostic (see Activity and Reproduction). American Bullfrogs (Lithobates catesbeianus) sometimes have irregular dark spots on the dorsum, but they grow to a much larger size (< 203 mm SVL) and lack dorsolateral folds. The Chiricahua Leopard Frog is known to hybridize with the Northern Leopard Frog and Lowland Leopard Frog. In wild populations, rates of hybridization are low (0-7%, Platz and Mecham 1979), although Green and Delisle (1985) noted two of four frogs were hybrids at a site with Northern and Chiricahua Leopard Frogs in Coconino County, AZ. Frost and Bagnara (1977) found no evidence for hybridization between the Chiricahua and Plains (L. blairi) Leopard Frogs.

The distribution of the species as recognized herein is divided into northern and southern disjunct regions, separated by lower elevation deserts and grassland.

These areas do not match precisely Hekkala et al.’s northern and southern genetic units.

Distribution and Habitat Use The distribution of the species as recognized herein is divided into northern and southern disjunct regions, separated by lower elevation deserts and grassland (Platz and Mecham 1984, USFWS 2007, Fig. 5). These areas do not match precisely Hekkala et al.’s northern and southern genetic units discussed above.

The northern distributional unit lies north of the Gila River in Arizona, and north of Interstate 10 in New Mexico. Populations in this unit occur or occurred at scattered localities from the upper Fossil Creek drainage in extreme eastern Yavapai County, AZ east through the mountains (above and below the Mogollon Rim) to the eastern bajada of the Black Range in the Río Grande drainage of Sierra and Socorro coun

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Figure 5. Distribution of the Chiricahua Leopard Frog. The distribution south of Chihuahua is not well known. The type locality for the Vegas Valley Leopard Frog is not included (it is off the map to the northwest).

ties, NM. In Arizona, the species occurred historically as far north as Chevelon Creek and other sites in the Little Colorado River drainage and as far south as Ash Creek on the San Carlos Apache Reservation and sites in the Eagle, Dix, and Coal creek drainages of the Apache-Sitgreaves National Forest in Greenlee County. In New Mexico, northern populations occurred from the Tularosa River in Catron County south to the Mimbres River and vicinity in Grant County. The extirpated Las Vegas Valley, Nevada frogs (Hekkala et al. 2011) occurred about 370 km northwest of the nearest traditionally recognized Chiricahua Leopard Frog population.

To the south, populations occur or occurred from the Baboquivari Mountains in Pima County, AZ, eastward to the Animas Mountains and Playas Valley, Hidalgo County, NM, and south well into Mexico. The northern-most records south of the Gila River in Arizona are two specimens from the base of the Pinaleño Mountains, Graham County. The species is also known from the nearby Galiuro Mountains. In Mexico, the Chiricahua Leopard Frog occurs or occurred in eastern Sonora, western Chihuahua, and adjacent Durango, Mexico (Rorabaugh 2008, Lemos-Espinal and Smith 2007, Streicher et al. 2012). The southern distributional limit is unclear. Diaz and Diaz (1997) reported the species from Aguascalientes; however, chiricahuensis-like frogs of Mexico, including Lithobates montezumae and L. lemosespinali could be confused with the Chiricahua

Leopard Frog, and the relationships of these similar frogs to each other and L. chiricahuensis need further work, particularly in regard to L. lemosespinali.

The Chiricahua Leopard Frog is a species of high valleys and mountains, from about 1,034 m elevation in the Altar Valley, Pima County, to 2,709 m in the White Mountains, AZ. Vegetation communities in these areas include Chihuahuan desertscrub, semidesert grassland, plains grassland, oak woodland, pinyon-juniper woodland, pine-oak woodland, and mixed conifer forest. Within these communities, the Chiricahua Leopard Frog is typically found at or near ciénegas, pools, livestock tanks, mine adits, wells, lakes, reservoirs, streams, and rivers with permanent or nearly permanent surface water. This species is rare or absent in high gradient, shallow flowing water, and more likely to be found in deep, still pools.

Some other ranid frogs (e.g., Northern Leopard Frog) use different habitats seasonally, and move among those habitats as the seasons change. No marked seasonal use has been noted in the Chiricahua Leopard Frog, except that when the summer rains come, the frogs often spread out along drainages into ephemeral reaches and likely use the uplands to a greater degree than during the dry foresummer. Based on our anecdotal observations, juveniles may use shallow water and marshy areas more than adults, as well. During drought, when water is scarce, Chiricahua Leopard Frogs concentrate into the last remaining pools. At dry cattle tanks, they have been found in cracks in the mud where conditions remain moist.

Individuals are known to travel overland or along arroyos for several kilometers, hence occasional individuals may be found in the uplands or along ephemeral drainages far from permanent water. In 1974, Frost and Bagnara (1977) noted passive or active movement of Chiricahua and Plains Leopard Frogs for 8 km or more along West Turkey Creek in the Chiricahua Mountains. In August 1996, Rosen and Schwalbe (1998) found up to 25 young adult and subadult Chiricahua Leopard Frogs at a roadside puddle in the San Bernardino Valley, AZ. They believed the only possible origin of these frogs was a stock tank 5.4 km away. In September 2009, 15-20 Chiricahua Leopard Frogs were found at Peña Blanca Lake west of Nogales, AZ. The nearest likely source population was Summit Tank, a straight line distance of 5 km overland and approximately 6.6 km along intermittent drainages.

Five of eight frogs tracked with telemetry moved 2202,450 m along a drainage in New Mexico during about six weeks in September and October. The other three moved minimally (Christman et al. 2003).

These frogs often occur in metapopulations that may include one or more sites where water is dependable and breeding occurs in most years, and other satellite occurrences in less suitable habitat where breeding only occurs in wet years, breeding does not result in recruitment, or frogs occur only as dispersing

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Males usually call from the water’s edge, from shallows, or while floating in mats of aquatic vegetation. Most calling occurs by frogs that are partially submerged, but some call underwater.

Calls recorded underwater sound quite different from those recorded out of water.

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individuals that do not breed. These wetland sites are usually no more than 8 km apart, and may be connected via permanent or ephemeral streams (USFWS 2007).

Activity and Reproduction Activity depends primarily on water temperature.

As water temperature drops to 12-13°C or below, frogs are rarely seen. At water temperatures of 14°C and above usually some frogs are found active. Active frogs have been found year-round in warm springs in New Mexico, but where water temperature varies with ambient air temperature, frogs are most active from April through September. Chiricahua Leopard Frogs are easiest to approach and identify at night with a flashlight or headlamp, but they are active day and night.

Juveniles are perhaps more active by day, and adults more active at night (Jennings 1988). Winter retreats have not been identified; however, Northern Leopard Frogs are known to overwinter at the bottom of welloxygenated ponds or lakes, they may bury themselves in mud, and they have been found during the winter in caves (Rand 1950, Emery et al. 1972, Nussbaum et al.

1983, Cunjak 1986, Harding 1997).

Breeding has been observed every month of the year (Frost and Platz 1983, Scott and Jennings 1985, Sredl and Jennings 2005), although most breeding likely occurs from February into September. Breeding in the winter months may be limited to warm springs (Scott and Jennings 1985). Frost and Platz (1983) noted that populations occurring above 1,800 m tend to have a shorter breeding season, primarily from June through August; although Scott and Jennings (1985) noted no elevational differences in breeding activity in New Mexico. Reduced oviposition has been noted in the hot, dry period of May and June (Scott and Jennings 1985). The male advertisement is a snore with a pulse rate of 16.8-38.7/second and lasting 1-2 seconds that may be followed by grunts, mew, or croak calls (Platz and Mecham 1979, Davidson 1996). Frost

and Platz (1983) provide a sonogram. The call can be heard by following this link: http://amphibiaweb.org/sounds/Rana_chiricahuensis65.mp3

Males usually call from the water’s edge, from shallows, or while floating in mats of aquatic vegetation. Most calling occurs by frogs that are partially submerged, but some call underwater. Calls recorded underwater sound quite different from those recorded out of water (compare L. chiricahuensis and L. “subaquavocalis” recordings on the Davidson 1996 CD). But to the ear, heard out of the water, both sound similar (the underwater calls are muffled and not as loud). While calling, males have been observed to engage in fisticuffs with other males, presumably to defend prime calling locations (Sredl and Jennings 2005). No other territorial behavior is known.

Males from Socorro and Catron counties, NM exhibited adult characteristics at > 53.5 and 56.2 mm SVL, respectively (Sredl and Jennings 2005). Size at maturity for females is unknown. In ideal conditions, maturity may be reached in the first year of life; however, Platz (1997) suggested frogs may not mature until age six in Ramsey Canyon. Amplexus is axillary, takes place in water, and may last as long as 20 hours (Platz 1997, Sredl and Jennings 2005). Eggs are laid in spherical masses that are usually attached to submerged vegetation, or may be set atop mats of algae or other submerged aquatic plants. They are usually placed within 5 cm of the water’s surface and contain 300-2,040 eggs (Jennings and Scott 1991, Platz 1997, Fig. 6). Size of the egg mass is typically correlated with female body size. Time to hatching depends at least in part on water temperature and varies from about 8-14 days (Zweifel 1968, Platz 1997, Sredl and Jennings 2005). Tadpoles take from 3-9 months to metamorphose (Jennings 1988, 1990), and they often overwinter. Because of this, and the extended breeding season, often multiple size classes of tadpoles are found at breeding sites.

Skeletochronology of frogs from the east side of the Huachuca Mountains, Cochise County, AZ, revealed that 47 percent of sampled adults were age six or older. The oldest frogs were estimated at 10 years post-metamorphosis (Platz et al. 1997). However, this area may be anomalous, as Durkin (1995) found no evidence of Chiricahua Leopard Frogs living longer than six years. Platz et al.’s work in the Huachuca Mountains was before the onset of periodic diseaserelated die offs. Longevity at that site is probably much less now.

A wide variety of vertebrates and invertebrates likely prey upon Chiricahua Leopard Frogs and their tadpoles and eggs, including aquatic insects, fishes, snakes, birds, American Bullfrogs and possibly other anurans, and a variety of mammals.

Diet

Figure 3. Chiricahua Leopard Frog egg mass, Huachuca Mountains, AZ. Photo by James C. Rorabaugh.

Christman and Cummer (2006) examined stomach contents of 56 Chiricahua Leopard Frogs from New Mexico. The most common prey items were insects (Coleoptera, Hemiptera, and Diptera – 75.2% by frequency). Vertebrates were uncommon in the diet,

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and included one cyprinid fish and remnants of a frog or late stage tadpole. Both terrestrial and aquatic prey were taken in similar proportions (44.6 and 37.3%, respectively). Incidental material in the stomachs included plant material, wood, and stones. The authors concluded that the Chiricahua Leopard Frog is a prey generalist, taking what is available. However, they suggested Bombardier beetles (Brachinus sp.) are avoided.

Those beetles were common at frog collection sites but were not found in stomachs.

Field et al. (2003) noted a Rufous or Allenâ&#x20AC;&#x2122;s Hummingbird taken by a Chiricahua Leopard Frog in Miller Canyon of the Huachuca Mountains, AZ. Captives readily take crickets, mealworm larvae and adults, silkworm larvae, other invertebrates, and small fish (Demlong 1997, USFWS 2007). Cannibalism of larger frogs on smaller frogs occurs in captive colonies and likely occurs in the wild (Jennings 1988). The tadpoles are probably mostly herbivorous. Marti and Fisher (1998) listed the following likely tadpole food items in Ramsey Canyon, Huachuca Mountains: bacteria, diatoms, phytoplankton, filamentous algae, water milfoil, duckweed, and detritus. We have observed wild Chiricahua Leopard Frog tadpoles feeding upon floating algal mats. Large tadpoles have been observed consuming the gelatinous envelopes of conspecific egg masses (Platz 1997). In captivity, tadpoles have been fed boiled spinach, romaine lettuce, cucumber slices, frozen trout, duckweed, spirulina fish food, and rabbit pellets suspended in agar-gelatin blocks (Frost 1982, Demlong 1997). In captivity, tadpoles have been fed rabbit pellets suspended in agar-gelatin blocks, boiled spinach, romaine lettuce, cucumber slices, frozen trout, duckweed, and spirulina fish food (Frost 1982, Demlong 1997).

Predators, Parasites, and Disease A wide variety of vertebrates and invertebrates likely prey upon Chiricahua Leopard Frogs and their tadpoles and eggs, including aquatic insects, fishes, snakes, birds, American Bullfrogs and possibly other anurans, and a variety of mammals (Sredl and Jennings 2005). However, predation in the wild has rarely been reported. Enderson and Bezy (2002) published an image of a Black-necked Gartersnake (Thamnophis cyrtopsis) eating a metamorph Chiricahua Leopard Frog.

Rorabaugh and Jones (in press) observed a Sonoran Whipsnake (Coluber bilineatus) with an adult Chiricahua Leopard Frog in its mouth in the Dragoon Mountains, AZ. Tom Deecken (pers. comm., 2014) watched a Mexican Gartersnake (T. eques) feeding on young Chiricahua Leopard Frogs in the Huachuca Mountains, AZ. In the laboratory, crayfish (Orconectes virilis) can kill and consume tadpole, metamorph, and adult Chiricahua Leopards Frogs, and their presence has been linked to the disappearance of Chiricahua Leopard Frog populations (Fernandez and Rosen 1996, 1999).

Sredl and Howland (1995) found that Chiricahua Leopard Frogs were almost always absent from sites with American Bullfrogs and non-native predatory fishes, such as Bluegill and other sunfishes, Largemouth Bass, and Carp (including Koi). Tiger Salamanders (Ambystoma mavortium) likely prey upon Chiricahua Leopard Frog tadpoles and juvenile frogs. Chiricahua Leopard Frogs coexist with a number of native fishes, and in some sites persist with Mosquitofish (Gambusia affinis, USFWS 2007). The likelihood of coexisting with predators, especially non-native species, is likely correlated with habitat complexity and the presence of escape cover. Chiricahua Leopard Frogs typically jump into water when threatened, but do not emit an alarm call (Sredl and Jennings 2005).

Goldberg et al. (1998) documented six species of trematodes and one species of nematode in Chiricahua Leopard Frogs from Arizona. A generalized discussion of amphibian parasites and their effects is found in USFWS (2007).

Disease was recognized as an important factor in global amphibian decline in the 1990s, although enigmatic, unexplained population decline and disappearance in seemingly pristine areas had been noted well before that, particularly in the 1970s and 1980s (Carey et al. 2003). Berger et al. (1998) were the first to describe the amphibian skin disease (Chytridiomycosis) caused by the fungal pathogen Batrachochytium dendrobatidis (Bd). The disease was found to be the cause of wavelike global mortality of metamorphosed amphibians, rapid population decline, with many mortalities in high elevation sites that were often protected from habitat degradation (Carey 2000). This apparently introduced disease (James et al. 2009) was first detected in Arizona in a Lowland Leopard Frog collected in 1972 from Sycamore Canyon, Pajarito Mountains, AZ (T. Jones, pers. comm., also see Sredl 2000 and Bradley et al. 2002 for additional background). The factors governing the likelihood of Bd causing population decline are likely extraordinarily complex; however, the disease appears to cause the greatest impact to populations of Chiricahua Leopard Frogs north of Interstate 10 in New Mexico, although it is present in many populations, and sometimes is associated with die-offs in southern Arizona. Chiricahua Leopard Frogs at warm springs or otherwise warmer sites appear to survive and persist better and longer than frogs at cooler locales. Ranavirus, another emerging infectious disease causing amphibian, reptile, and fish declines in the northern hemisphere, was found in a Chiricahua Leopard Frog in New Mexico (R. Jennings, pers. comm.), but neither its prevalence in Chiricahua Leopard Frog populations, nor its effects on the species are understood.

The Chiricahua Leopard Frog was proposed for federal listing as a threatened species in a Federal Register notice published on 14 June 2000 (65 FR 37343).

The final rule was published 13 June 2002 (67 FR 40794) and included a special rule promulgated under section 4d of the ESA that excluded livestock operations and maintenance at cattle tanks on non-federal lands from the ESAâ&#x20AC;&#x2122;s section 9 take prohibitions.

Conservation Clarkson et al. (1986) were the first to report declining populations of Chiricahua and other leopard

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frog species in Arizona. Elaborating on that earlier report, Clarkson and Rorabaugh (1989) found Chiricahua Leopard Frogs at only two of 36 sites in Arizona that supported the species in the 1960s and 1970s.

They suggested that predation by introduced American Bullfrogs and fishes, as well as habitat alteration were likely contributing factors in observed leopard frog declines in Arizona, but that disappearances of Chiricahua Leopard Frogs were often in places that lacked introduced predators and significant habitat disturbance. In a status assessment of Mexican and Narrow-headed Gartersnakes (Thamnophis eques and T. rufipunctatus, respectively), Rosen and Schwalbe (1988) noted that American Bullfrogs prey upon and had replaced leopard frogs at some sites in southern Arizona.

They suggested that non-native predatory fish may have the same effect.

In New Mexico, Randy Jennings did not consider the Chiricahua Leopard Frog in jeopardy based on surveys from 1983-87 (pers. comm. in Clarkson and Rorabaugh 1989), but in a subsequent report, he found the species at only 6 of 33 sites that had supported the species during the previous 11 years (Jennings 1995).

The Chiricahua Leopard Frog was placed on the list of candidate species for federal listing under the Endangered Species Act (ESA) in November 1991.

Subsequent studies reinforced earlier findings that the species was declining in Arizona and New Mexico (Hale 1992, Fernandez and Bagnara 1995, Sredl and Howland 1995, Zweifel 1995, Rosen et al. 1996, Sredl et al. 1997, Painter 2000, Rosen and Schwalbe 2002, Boykin and McDaniel 2008, Witte et al. 2008, Rorabaugh 2010, USFWS 2010, and others). Causes of decline are not always clear, but contributing factors mentioned in these reports include predation by introduced species, habitat loss and degradation from a variety of causes, die-offs associated with Bd, loss of metapopulation function, drought, and climate change.

Livestock grazing can degrade Chiricahua Leopard Frog habitat, but small impoundments built as water sources for cattle are often important frog habitats.

As of 2009, the Chiricahua Leopard Frog occupied at least 131 sites in Arizona and New Mexico, but robust breeding populations were very few in number (only 11 in Arizona). USFWS (2010) concluded that from 2002 to 2009 the status of the Chiricahua Leopard Frog was at least stable and probably improving in Arizona, declining in New Mexico, and unknown in Mexico. More recent status summaries are not available; however, we believe these general conclusions are still valid in 2014.

The Chiricahua Leopard Frog was proposed for federal listing as a threatened species in a Federal Register notice published on 14 June 2000 (65 FR 37343).

The final rule was published 13 June 2002 (67 FR 40794) and included a special rule promulgated under section 4d of the ESA that excluded livestock operations and maintenance at cattle tanks on non-federal

lands from the ESAâ&#x20AC;&#x2122;s section 9 take prohibitions.

Collection of the species is prohibited by Arizona Game and Fish Commission Order 41, except with special permits. Collection is similarly prohibited in Mexico where the Chiricahua Leopard Frog is listed as a threatened species by the Mexican government. Chiricahua Leopard Frogs are not protected by state regulations in New Mexico. In a Federal Register notice dated 20 March 2012, 10,346 acres (4,187 hectares) in 39 units in Arizona and New Mexico were designated as critical habitat for the Chiricahua Leopard Frog (77 FR 16347).

The Chiricahua Leopard Frog Recovery Plan (USFWS 2007) is the primary document guiding conservation activities for this species. It includes recommended recovery actions to reduce or eliminate threats, an implementation schedule for those actions, measurable criteria to gauge recovery success, and much other information needed to conserve the species. Recovery criteria need to be met in each of eight recovery units.

The primary recovery actions include 1) protecting remaining populations, 2) identifying, restoring or creating, and protecting unoccupied sites as necessary to support viable Chiricahua Leopard Frog populations and metapopulations, 3) establishing new or reestablishing former populations, 4) augmenting populations as needed to increase persistence, 5) monitoring populations and recovery, 6) conducting research that promotes recovery, 7) developing support for the recovery program, 8) developing cooperative projects with non-federal landowners, 9) amending land use plans as needed, 10) working with Tribal partners, 11) working with Mexican partners, and 12) practicing adaptive management. The recovery program is guided by a recovery team, three regional steering committees, and numerous local recovery groups, most of which meet on a regular basis. The recovery plan and other documents pertaining to the species can be downloaded from the following site: http://www.fws.gov/southwest/es/arizona/CLF.htm In a 5-year review of the frog and the recovery program, USFWS (2010) found that although the requisite number of metapopulations and isolated robust populations needed for recovery had only been met in one recovery unit, through aggressive rearing and introduction, particularly in Arizona (the Phoenix Zoo alone has reared more than 20,000 Chiricahua Leopard Frogs), progress was being made in several other units towards meeting those population criteria. However, in most cases threats had not been adequately addressed, and many management issues still remained to be resolved. Although significant progress has been made, particularly in regard to establishing or reestablishing populations, if on-going recovery actions are interrupted, drought worsens, or other threats intensify, the status of the species could easily deteriorate (USFWS 2010). Chytridiomycosis has been an

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In a 5-year review of the frog and the recovery program, USFWS (2010) found that although the requisite number of metapopulations and isolated robust populations needed for recovery had only been met in one recovery unit, through aggressive rearing and introduction, particularly in Arizona, progress was being made in several other units towards meeting those population criteria.

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especially difficult problem in New Mexico where recovery efforts have often focused on conserving existing populations through establishment of secure refugial sites. Dramatically successful American Bullfrog eradication efforts have been conducted in some areas (Peña Blanca Lake area, Altar Valley, Scotia Canyon and vicinity, Las Cienegas and vicinity) with subsequent recovery of Chiricahua Leopard Frog populations; however, continued vigilance will be necessary to ensure bullfrogs are not reintroduced or reinvade from adjacent areas. Other threat abatement and locking in favorable management practices are also needed to complement establishment of frog populations. Successes (Aguilar et al.

2008, Rorabaugh et al. 2008) need to be expanded upon and the momentum accelerated if this frog is to be recovered to the point that it no longer needs the protection of the ESA. Literature Cited Aguilar, R., J.M. Servoss, J.C. Rorabaugh, M. J. Sredl, V.L. Boyarski, P. Swanson, and T. Sprankle. 2008. Partnerships foster conservation of native riparian herpetofauna.

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Berger L., R. Speare, P. Daszak, D.E. Green, A.A. Cunningham, C.L. Goggins, R. Slocombe, M.A. Ragan, A.D. Hyatt, K.R. McDonald, H.B. Hines, K.R. Lips, G. Marantelli, and H. Parkes. 1998. Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America. Proceedings of the National Academy of Science, USA 95:9031-9036.

Boykin, K.G., and K.C. McDaniel. 2008. Simulated potential effects of ecological factors on a hypothetical population of Chiricahua Leopard Frog (Rana chiricahuensis). Ecological Modelling 218 (2008):175-181.

Bradley, G.A., P.C. Rosen, M.J. Sredl, T.R. Jones, and J.E. Longcore. 2002. Chytridomycosis in native Arizona frogs.

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Carey, C. 2000. Infectious disease and worldwide declines of amphibian populations, with comments on emerging diseases in coral reef organisms and in humans. Environmental Health Perspectives 108:143-150.

Carey, C., A.P. Pessier, and A.D. Peace. 2003. Pathogens, infectious disease, and immune defense. Pages 127-136 in R.D. Semlitsch (ed), Amphibian Conservation. Smithsonian Books, Washington, D.C. Christman, B.L., and M.R. Cummer. 2006. Stomach contents analysis of the Chiricahua Leopard Frog (Rana chiricahuensis) and Plains Leopard Frog (Rana blairi) in New Mexico.

Final report to the New Mexico Department of Game and Fish. Share with Wildlife Contract 05-516.0000.051.

Christman, B.L., C.G. Kruse, and S. Debrott. 2003. Survey and monitoring of Chiricahua Leopard Frog (Rana chiricahuensis) populations on the Ladder Ranch and adjacent Gila National Forest lands: Sierra County, New Mexico.

Turner Endangered Species Fund, Ladder Ranch, NM. Clarkson, R.W., and J.C. Rorabaugh. 1989. Status of leopard frogs (Rana pipiens Complex) in Arizona and southeastern California. Southwestern Naturalist 34(4):531-538.

Clarkson, R.W., J.C. Rorabaugh, and J.E. Platz. 1986. Status and future of ranid frogs in Arizona. Proceedings of the Arizona-Nevada Academy of Sciences 21:21.

Crother, B.I. (ed.). 2008. Scientific and Common Names for Amphibians and Reptiles of North America North of México. Society for the Study of Amphibians and Reptiles, Herpetological Circular No. 37:1-84.

Crother, B.I. 2012. Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding, seventh edition. Society for the Study of Amphibians and Reptiles, Herpetological Circular (39):1-92.

Cunjak, R.A. 1986. Winter habitat of northern leopard frogs, Rana pipiens, in a southern Ontario stream. Canadian Journal of Zoology 64:255 257.

Davidson, C. 1996. Frog and toad calls of the Rocky Mountains. Library of Natural Sounds, Cornell Laboratory of Ornithology, Ithaca, NY. Degenhardt, W.G., C.W. Painter, and A.H. Price. 1996.

Amphibians and reptiles of New Mexico. University of New Mexico Press, Albuquerque.

Demlong, M.J. 1997. Head-starting Rana subaquavocalis in captivity. Reptiles 5:24-33.

Diaz, J.V., and G.E.Q. Diaz. 1997. Anfibios y reptiles de Aguascalientes. Grupo Impressor Mexico, Aguascalientes, Aguascalientes, MX. Dodd, C.K., Jr. 2013. Frogs of the United States and Canada (2 volumes). Johns Hopkins University Press, Baltimore, MD. Durkin, B. 1995. Aging frogs by counting the rings in their bones (yeah, just like with trees!) Unpublished report, Grand Canyon University, Phoenix, AZ. Emery, A.R., A.H. Berst; K. Kodaira. 1972. Under-ice observations of wintering sites of leopard frogs. Copeia 1972:123-126.

Enderson, E.F., and R.L. Bezy. 2002. Field observations of anuran predation by the Black-necked Garternake (Thamnophis cyrtopsis) in southern Arizona. Sonoran Herpetologist 15:114-115.

Fernandez, P.J., and J.T. Bagnara. 1991. Effect of background color and low temperature on skin color and circulating α-MSH in two species of leopard frog.

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Fernandez, P.J., and J.T. Bagnara. 1993. Observations on the development of unusual melanization of leopard frog ventral skin. Journal of Morphology 216:9-15.

Fernandez, P.J., and J.T. Bagnara. 1995. Recent changes in leopard frog distribution in the White Mountains of east central Arizona. Page 4 in abstracts of the First Annual Meeting of the Southwestern Working Group of the Declining Amphibian Populations Task Force, Phoenix, AZ. Fernandez, P.J., and P.C. Rosen. 1996. Effects of the introduced crayfish Oronectes virilis on the native aquatic herpetofauna in Arizona. Report to the Arizona Game and Fish Department, Heritage Program, IIPAM Project No. I94054.

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Field, K.J., T.L. Beatty Sr., and T.L. Beatty Jr. 2003. Rana subaquavocalis (Ramsey Canyon leopard frog); Diet. Herpetological Review 34(3):235.

Fritts, T.H., R.D. Jennings, and N.J. Scott, Jr. 1984. A review of the leopard frogs of New Mexico. Report to the New Mexico Department of Game and Fish, Santa Fe, NM. Frost, D.R., et al. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297:1-370.

Frost, J.S. 1982. A time-efficient, low cost method for the laboratory rearing of frogs. Herpetological Review 13(3): 73-77.

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sessment/conservation strategy. Department of Biology, Creighton University, Omaha, NE. Platz, J.E. 1997. Status survey of the Ramsey Canyon leopard frog, Rana subaquavocalis. Report to the Arizona Game and Fish Department, Phoenix.

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of the Chiricahua Leopard Frog in Arizona mediated by introduced species. Report to Heritage program, Arizona Game and Fish Department, Phoenix. IIPAM Project No. I92052.

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2008. Epidemiological analysis of factors associated with local disappearances of native ranid frogs in Arizona.

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Zweifel, R.G. 1995. Rana revisited: Some R. chiricahuensis sites in the Chiricahua Mountains then and now. Page 12 in Program and Abstracts of the First Annual Meeting of the Southwestern Working Group of the Declining Amphibian Populations Task Force, Phoenix, AZ.

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LO C A L CO N S E R VAT I O N

Threatened Narrow-headed Gartersnakes Born at Phoenix Zoo Article by Phoenix Zoo staff; reprinted with permission

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n 2007, the Phoenix Zoo received a small research population of wild-caught Narrow-headed Gartersnakes (Thamnophis rufipunctatus) in hopes of developing a propagation and release program. On the morning of 2 July 2014, a four-year-old gave birth to 18 neonates in the Zoo’s specially designed outdoor Suzan L. Biehler Herpetarium. All 18 offspring are healthy and were observed capturing live fish within 48 hours. This reproductive event is the culmination of years of husbandry work and scientific research by the Zoo’s conservation staff. This significant birth comes at a critical time since on 7 July 2014 the species was listed as “threatened” under the Endangered Species Act. The Zoo has also developed a husbandry manual for this species that is currently in use by members of the Gartersnake Conservation Working Group (GCWG).

“The birth of the Narrow-headed Gartersnakes here is a fulfillment of Phoenix Zoo’s commitment to supporting native species conservation and recovery”, says Stuart Wells, Director of Conservation and Science at the Phoenix Zoo. “Our dedicated staff has worked tirelessly for many years to achieve this goal.

We are proud of this accomplishment and pleased to contribute to the recovery of this species.” The Narrow-headed Gartersnake is a unique, highly aquatic species. Its numbers have been declining throughout its range in Arizona and New Mexico for over a

decade. Many factors are contributing to the decline including drought, non-native invasive species, wildfire and agricultural/urban encroachment. Beginning in 2006, the GCWG, a multi-partner, collaborative effort, was formed by U.S. Fish and Wildlife Service to help conserve and recover the northern Mexican gartersnake and the Narrow-headed Gartersnake.

“The Narrow-headed Gartersnake is a mid- to high-elevation, stream-dwelling species that is very sensitive to environmental and physical stress”, explains Jeff Servoss, U.S. Fish and Wildlife Service Chair of the GCWG. “These traits make this species a unique challenge for those trying to not only keep them alive in captivity, but also trying to produce offspring. After many years of trying, by many different institutions, the Phoenix Zoo has finally produced viable Narrow-headed Gartersnakes. This achievement is very noteworthy and a testament to the Zoo’s relentless effort to identify the variables that have prevented breeding in the past. This is a significant achievement and a giant step forward for gartersnake conservation.” The offspring are being head-started for a period of six to nine months before the majority are released to the wild. The remaining few will be retained for the breeding program. The Zoo is proud of this accomplishment and appreciates the opportunity to support the conservation of wildlife in Arizona and throughout the world.

The Narrowheaded Gartersnake is a mid- to high-elevation, stream-dwelling species that is very sensitive to environmental and physical stress. These traits make this species a unique challenge for those trying to not only keep them alive in captivity, but also trying to produce offspring.

The Phoenix Zoo’s Arthur L. and Elaine V. Johnson Foundation Native Species Conservation Center has announced the first-ever propagation of the threatened Narrow-headed Gartersnake (Thamnophis rufipunctatus).

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S H O R T CO N S E R VAT I O N A R T I C L E

Biodiversity of Reptiles and Amphibians of India—Conservation awareness campaign in West Bengal, India with 140 school students Suman Pratihar1 and Kaushik Deuti2 1 Department of Zoology, Vidyasagar University, Midnapore, 721102; pratihar_vu@rediffmail.com 2 Zoological Survey of India, Herpetology Division, Kolkata, 700016; kaushikdeuti@gmail.com

A n awareness campaign was arranged to encourage am-

phibian conservation among school students in the junglebased rural area in Midnapore West Bengal, India. One hundred and forty school students from thirty seven schools were selected for this awareness campaign. It was a four day residential science camp in which amphibian conservation was highlighted. Dr. Bidhan Ch Patra, Dean of Science at Vidyasagar University, was the coordinator of this conservation program, and expressed: “Reptiles and amphibians are eye-catching creatures in our living world. This group make our world colorful and gorgeous. Conservation initiatives are needed to ensure the presence of this much threatened group. And we start it with our school students.”

The science camp was held at the Vidyasagar University campus. Dr. Kaushik Deuti, Zoological Survey of India, head of the Herpetology Division at Kolkata, delivered his diversity and conservation speech on 26 June 2014. Dr. Suman Pratihar arranged discussion session with students and many students interacted with general queries on herpetofauna.

India is among the 20 most biodiversity-rich countries in the world. This is because of the country’s biogeographic location at the junction of Oriental, Indo-Chinese, and Indo-Malayan realms and its unique topography and climatic variations. Of the 34 biodiversity hotspots throughout the globe, two occur in India – the Western Ghats and northeastern India.

The rich flora and fauna of India can be judged by the sheer number of species of each group found in the country. Similarly, the herpetofauna is diverse and consists of 526 species of reptiles and 342 species of amphibians currently known.

The reptiles consist of 33 species of chelonians (5 species of marine turtles, 8 species of soft-shelled turtles, 16 species of hard-shelled turtles or terrapins and 4 species of land tortoises), 3 species of crocodilians, 208 species of lizards (77 species of geckos, 49 species of agamids or chameleons, 60 species of skinks, and 4

Dr. Suman Pratihar interacting with participants.

species of monitor lizards), and 282 species of snakes (5 species of boas and pythons, 160 species of colubrids or typical snakes, 37 species of elapids or cobras and kraits, and 23 species of vipers and pit vipers).

The chelonians are covered by a shell (either hard or soft) into which they can retract their head, neck, limbs, and tail except in case of the marine turtles.

They do not have teeth but only serrations on a horny beak. The soft-shell turtles have tubular nostrils on a fleshy pig-like proboscis. The hard-shelled turtles lack this snout but have enlarged heads and strong jaw muscles. The land tortoises have columnar legs covered with heavy scales and a round, heavy shell with horny shields.

The crocodiles have their bodies covered with scutes and large recurved teeth. The lizards have a prominent tympanum, four limbs with claws, and good eyesight. The geckos lack eyelids and have fingers and toes with lamellae and a detachable tail. The agamids have spines on the back, and the chamaleons have a prehensile tail and a long sticky tongue to catch insects.

The skinks have scales arranged on their body just like tiles on a roof. Monitor lizards have well-developed limbs and a long tail.

Snakes do not have either limbs or tympanum and so cannot hear. The poisonous species have poison glands and fangs, with grooves on the front of the fangs (elapids; cobras and kraits), behind the fang (colubrid snakes), or through the fang (vipers).

SONORAN HERPETOLOGIST 27 (3) 2014

An awareness campaign was arranged to encourage amphibian conservation among school students in the junglebased rural area in Midnapore West Bengal, India. One hundred and forty school students from thirty seven schools were selected for this awareness campaign.

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The amphibians consist of 27 species of toads or bufonids, 60 species of discoglossids, 18 species of hill-stream frogs or megophryids, 11 species of dancing frogs or micrixalids, 23 species of burrowing frogs or microhylids, 27 species of night frogs or nyctibatrachids, 31 species of true frogs or ranids, and 107 species of tree frogs or rhacophorids. In addition, there is one species of urodelan or Himalayan salamander, and 36 species of caecilians or limbless amphibians.

The science camp was a tremendous success and more camps are planned for the future. Reaching out to the Indian youth is one way to ensure the conservation of India’s herpetofauna for future generations.

Poster on the university campus.

The science camp was a tremendous success and more camps are planned for the future. Reaching out to the Indian youth is one way to ensure the conservation of India’s herpetofauna for future generations.

Map of India showing locations of schools from where students participated.

Participants of the science camp (left) and Dr. Kaushik Deuti’s awareness lecture (right).

Dr. Bidhan Chandra Patra inaugurated this camp with his inspiring new thoughts.

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F E D E R A L L I S T I N G U P DAT E

The Latest Piece in the Endangered Species Act “Puzzle”: Did the U.S. Fish and Wildlife Adequately Take into Consideration the “Lost Historical Range” of the Flat-Tailed Horned Lizard in its 2011 Withdrawal of the Proposed Rule to List the Lizard as Threatened?

By Richard Campbell, Adjunct Professor of Law, Golden Gate University School of Law

E ditor’s Note: This article is reprinted with permission from the Arizona Riparian Council newsletter. It describes a series

of legal cases where the THS played a major role in changing federal policy about threatened and endangered species.

Richard Campbell is an Adjunct Professor of Law, Golden Gate University School of Law, San Francisco and Attorney for the U.S. Environmental Protection Agency, Region 9. The opinion expressed in this article are the author’s only and do not represent those of the U.S. Environmental Protection Agency or the Golden Gate University School of Law. The latest chapter in the long-running dispute about the listing of the Flat-Tailed Horned Lizard (Phrynosoma mcallii) as a threatened species under the Endangered Species Act (ESA) occurred on 15 March 2011, when the U.S. Fish and Wildlife Service (Service) withdrew its original Clinton-era proposal to list the lizard as a threatened species under the Act. See 58 Federal Register 62624 (29 November 1993); 76 Fed. Reg. 14257 (15 March 2011). The decision by the Service to withdraw its proposed listing was made in response to the Ninth Circuit’s decision in Tucson Herpetological Society v. Salazar, 566 F.3d 870 (9th Cir. 2009).

In that case, the Court agreed with the Society that when the Service determines whether a species is endangered or threatened throughout a significant portion of its range, the Service must take into consideration whether the “lost historical range” of the species (as opposed to its current range) constitutes a significant portion of the range of that species. The Service’s consideration of the lizard’s lost historical range in its 2011 decision not to list it as threatened under the Act is discussed below.

BACKGROUND Flat-Tailed Horned Lizard The Flat-Tailed Horned Lizard is on average three inches long and has managed to adapt to the Sonoran Desert in Arizona (including the Gila and Tinajas Atlas Mountains in Yuma County), the Coachella Valley of California, and in the northernmost Sonoran Desert of Mexico (62 Fed. Reg. 37852, July 15, 1997). Fortytwo percent of lizard habitat occurs on private land (58 Fed. Reg. 62625). The balance of lizard habitat on public land is managed by the Bureau of Land Management (BLM) (58 Fed. Reg. 62628).

Flat-Tailed Horned Lizard (Phrynosoma mcallii) in hand. Photo by Dale Turner.

Proposed Listing Decision

On 29 November 1993, the Service proposed to list the lizard as a threatened species (58 Fed. Reg. 62624). Habitat loss caused by urban development, conversion of desert lands for agriculture, offhighway vehicle usage, and military activities (e.g., Goldwater Bombing Range), coupled with inadequate regulatory mechanisms to stem this habitat loss on at least public lands managed by BLM, were cited as reasons for the proposed listing (58 Fed. Reg. 62626). The Service noted that fragmentation creates isolated subpopulations that, because of their reduced size, have an increased probability of extinction (58 Fed. Reg. 62626-27 (29 November 1993).

In 1996, the Service estimated that man-made factors were responsible for the destruction of 1,103,201 acres of the lizard’s estimated 4,875,624-acre historic range (71 Fed. Reg. 36745, 36749-51; 28 June 2006). In September 1996, a Service biologist maintained listing was the appropriate action to take: Nothing has really changed on the ground; and in some ways, the status of the lizard has continued to deteriorate. If forced to publish a final rule at this time, I do not believe we could make a case that threats have been alleviated to the point that listing is no longer warranted. (As quoted in the Society’s Initial Brief to the Ninth Circuit on 3 July 2000; 2000 U.S. 9th Cir. Briefs LEXIS 40 *8).

A few months later, however, on July 15, 1997, the Service decided it would not place the lizard on the Endangered Species list (62 Fed. Reg. 37852). Three reasons were provided: 1. BLM and other federal and state agencies (including Arizona Game and Fish) had entered into a

SONORAN HERPETOLOGIST 27 (3) 2014

The latest chapter in the long-running dispute about the listing of the Flat-Tailed Horned Lizard (Phrynosoma mcallii) as a threatened species under the Endangered Species Act (ESA) occurred on 15 March 2011, when the U.S. Fish and Wildlife Service (Service) withdrew its original Clinton-era proposal to list the Lizard as a threatened species under the Act.

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Conservation Agreement and agreed to implement a Management Strategy to protect lizard habitat; 2. A significant portion of lizard habitat was no longer threatened by geothermal and oil and gas development and pesticide spraying as it had been in 1993; and 3. Lizard survey methodology was too uncertain to conclusively demonstrate a downward trend in populations (62 Fed. Reg. 37859).

In essence, the Service found that the lizard’s current range on public land was sufficient to prevent listing even though it was extirpated from a large percentage of its historical range and faced continuing threats on private land. Defenders of Wildlife (DOW) challenged the 1997 withdrawal in federal district court (in southern California), but the district court upheld the Service’s decision.

2001 Ninth Circuit Decision DOW appealed the district court decision to the Ninth Circuit (Defenders of Wildlife v. Norton, 258 F.3d 1136 [9th Cir. 2001]). In that case, the Ninth Circuit first found that due to the ambiguity of the phrase “significant portion of its range,” the Service was entitled to deference in its interpretation of the term, so long as the Service articulated a reasoned basis for its decision and articulated a rational connection between the facts and the decision it made (Defenders of Wildlife v. Norton, 258 F.3d 1141 [9th Cir. 2001]). The Ninth Circuit also found that Congress added the “significant portion of its range” language to the ESA, at 16 U.S.C. § 1532(6), to allow the Service to take a flexible approach to wildlife management, i.e., one that would allow the Service to list a species that is threatened in a “significant portion” of its range even if that same species is thriving in other geographic areas (Defenders of Wildlife v. Norton, 258 F.3d 1141 [9th Cir. 2001]).

The Ninth Circuit then noted that the Service’s 1997 withdrawal of its listing decision presented the court with an opportunity to “puzzl[e] out the meaning” of what Congress meant when it told the Service to take into consideration “a significant portion” of a species’ range when making listing decision (Defenders of Wildlife v. Norton, 258 F.3d 1141 [9th Cir. 2001]).

In puzzling out the meaning of “significant portion of its range,” the Ninth Circuit first rejected the Service’s argument that it could rely on only an examination of the lizard’s current range on public land (Defenders of Wildlife v. Norton, 258 F.3d 1138, 1140).

The Ninth Circuit found that the Service’s distinction between public and private land explained much of the dispute between the Service and DOW, and was responsible, in large part, for the shift between the Service’s initial findings that accompanied the proposed rule and its subsequent decision to withdraw the rule.

(Defenders of Wildlife v. Norton, 258 F.3d 1140-1141).

The Ninth Circuit concluded that the Service needed to take a more “flexible” approach and look at both

private and public lands when considering historical range (Defenders of Wildlife v. Norton, 258 F.3d 1145).

The Court next rejected DOW’s assertion that a species should be listed merely because it no longer inhabits a high percentage of its historical range. The Court explained, [I]t simply does not make sense to assume that the loss of a predetermined percentage of habitat or range would necessarily qualify a species for listing. A species with an exceptionally large historical range may continue to enjoy healthy population levels despite the loss of a substantial amount of suitable habitat.

Similarly, a species with an exceptionally small historical range may quickly become endangered after the loss of even a very small percentage of suitable habitat (Defenders of Wildlife v.

Norton, 258 F.3d 1143).

The Ninth Circuit then concluded that “a significant portion of its range” should be interpreted as follows: [A] species can be extinct “throughout ... a significant portion of its range” if there are major geographical areas in which it is no longer viable but once was. Those areas need not coincide with national or state political boundaries, although they can. The Secretary necessarily has a wide degree of discretion in delineating “a significant portion of its range, since the term is not defined in the statute (Defenders of Wildlife v. Norton, 258 F.3d 1143).

The Court granted, however, that if a species has lost a large portion of its historical range, the agency “must at least explain [the] conclusion that the area in which the species can no longer live is not a ‘significant portion of its range’” (Defenders of Wildlife v.

Norton, 258 F.3d 1145). The Ninth Circuit directed the Service to take this into consideration in its next decision on whether to list the lizard under the Act (Defenders of Wildlife v. Norton, 258 F.3d 1145).

2003 Listing Decision In the course of making its next listing decision, the Service solicited the opinion of four lizard experts: Of the four, two recommended listing the species as threatened, one did not express a firm opinion, and one concluded that listing was not warranted. …

Kevin Young, the biologist that did not favor listing, stated that a ‘significant portion of the [lizard’s] range’ has indeed been lost, but concluded that listing would likely direct resources away from efforts to protect the species on public lands, and toward unproductive efforts to protect lizard habitat on private lands (566 F.3d at 875, n. 7, citing 68 Fed. Reg. at 340-41).

Based on this and other evidence, the Service again decided against listing the lizard in 2003. This decision was subsequently challenged in federal district court (this time in Arizona) by the Tucson Herpetological Society (and others, including DOW, Sierra Club, and the Center for Biological Diversity) arguing that the withdrawal did not comply with the Ninth Circuit’s 2001 decision in Defenders.

The district court agreed with the Tucson Herpe

SONORAN HERPETOLOGIST 27 (3) 2014

In essence, the Service found that the lizard’s current range on public land was sufficient to prevent listing even though it was extirpated from a large percentage of its historical range and faced continuing threats on private land.

Defenders of Wildlife (DOW) challenged the 1997 withdrawal in federal district court (in southern California), but the district court upheld the Service’s decision.

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tological Society in a 2005 decision where it found that the Service “assumed without explanation that large swaths of lost habitat were of no significance at all” and ordered the Service to try again (though the district court found the Service’s assessment of threats to the lizard’s current range was adequate). The Service withdrew its 2003 decision and restored the lizard to proposed listing status while it reconsidered its decision (70 Fed. Reg. 72776; Dec. 7, 2005).

2006 Delisting Decision After another public notice and comment period, the Service again decided to withdraw the proposed listing in 2006 (71 Fed. Reg. 36,745; June 28, 2006).

The Service noted that the “sole purpose” of the 2006 decision was to address the lost historical habitat issue that was the subject of the district court’s 2005 decision (71 Fed. Reg. 36749). Again, the Society challenged this decision in district court (Tucson Herpetological Soc’y v. Kempthorne, 2006 U.S. Dist. LEXIS 70736; N.D. Ariz.

In 2007, after hearing the challenge, the district court upheld the Service’s 2006 lost habitat analysis and listing decision: After setting a temporal baseline and defining the subject area, the Secretary proceeded to evaluate the significance of the lost historical habitat. He concluded that the Coachella Valley area [in California], including its lost associated habitat, was insignificant because of its small size relative to the overall range of the species, the high level of fragmentation due to human development, the lack of genetic, behavioral, or ecological differentiation, and the small size and importance of the population in general.… The remaining parcels of lost historical habitat areas near Mexicali and Yuma were also deemed insignificant.

Not only has the species persisted for nearly a century in the face of the steady habitat destruction, but the size of existing lizard populations has not declined and is not likely to decline in the foreseeable future because of the loss of 1,103,201 acres of historic range, the Secretary found.… After surveying the ‘available data concerning population abundance, trends, and threats,’ the Secretary concluded that yesterday’s conversion of suitable habitat to agriculture in the Mexicali and Yuma areas is not significant to the survival of today’s lizards (Tucson Herpetological Soc’y v. Kempthorne, 2007 U.S. Dist. LEXIS 50740 *27-28; N.D. Ariz. 2007).

The Society appealed this decision to the Ninth Circuit, first arguing the Service’s reasoning was inconsistent with the Ninth Circuit’s 2001 decision in Defenders because it merely relied on pointing to some areas where lizard populations persisted to support a finding that threats to the species elsewhere were not significant. The Society argued the ESA requires a more thorough explanation.

Ninth Circuit 2009 Decision In its 2009 Tucson Herpetological Society decision, the Ninth Circuit found, in part, that the Service had relied

on limited and inconclusive studies in its determination that the lizard was persisting in its current range (particularly in Mexico for which there were no studies provided), and that this reliance had adversely impacted the Service’s lost range analysis: The absence of conclusive evidence of persistence, standing alone, without persuasive evidence of widespread decline, may not be enough to establish that the [Service] must list the lizard as threatened or endangered … But this is a different case. The [Service] affirmatively relies on ambiguous studies as evidence of persistence (i.e., stable and viable populations), and in turn argues that this ‘evidence’of persistence satisfies Defenders’ mandate and proves that the lizard’s lost range is insignificant for purposes of the ESA. This conclusion is unreasonable. The studies do not lead to the conclusion that the lizard persists in a substantial portion of its range, and therefore cannot support the [Service]’s conclusion (Tucson Herpetological Soc’y v.

Kempthorne, 2007 U.S. Dist. LEXIS 50740 *879; N.D. Ariz. 2007).

The Ninth Circuit again remanded the decision whether to list the lizard back to the Service for further reconsideration based on better studies. In a dissent, Ninth Circuit Judge Noonan made the following observation that likely captured the Service’s frustration at this point: How many Flat-tailed Horned Lizards are there?

No one knows the answer to that question. Nor does anyone know how many lizards disappeared when portions of their range disappeared. It is supposed that a diminution in range correlates with a diminution in lizards. This hypothesis is plausible.

It has not been shown to be probable. Yet the case turns on what measures are necessary to keep this unknown population in existence. The court concludes that the [Service] erred in finding that the lizard has not lost a significant portions of its range. The old method of counting lizards is out. A new method has not been tried very much. It’s anybody’s guess whether the lizards are multiplying or declining. In a guessing contest one might defer to the government umpire. The court, however, finds the [Service’s] conclusion impacted by over-reliance on fragmenting evidence of the lizard’s persistence; so the court decides to give the [Service] another crack at the problem.

If the [Service] does not know what the lizard population was to begin with, or what it was in 1993, or what it is now in May 2009, how will [it] know if it is increasing, staying the same, or declining?

A style of judging, familiar to readers of the old English reports, characterizes the judge as dubitante. That is probably the most accurate term for me, which leads me to concur in the majority opinion insofar as it rejects the contentions of the Tucson Herpetological Society and to dissent from the remand whose command to the Secretary of the Interior is, guess again (Tucson Herpetological Soc’y v. Kempthorne, 2007 U.S. Dist. LEXIS 50740 *882-883; N.D. Ariz. 2007).

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If the [Service] does not know what the lizard population was to begin with, or what it was in 1993, or what it is now in May 2009, how will [it] know if it is increasing, staying the same, or declining?

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March 15, 2011 Withdrawal of Proposed Rule To List Lizard as Threatened In its March 15, 2011 decision, the Service, as directed by the Ninth Court, addressed the lost historical range issue again. The Service determined the Lizard’s lost historical range did not represent a significant portion of the Lizard’s range for four reasons: 1. Historically lost habitat was lost decades ago and, despite the amount of time that has since transpired, the species has not experienced a continuing range contraction due to the past loss of habitat.

2. Historically lost habitat “did not provide any special or unique features or meet any life history needs of the [L]izards that made those areas any more significant than any other habitat.”

3. Historically lost range was not continuous and contained natural barriers that separated relevant Lizard population segments.

4. The Lizard populations most in jeopardy do not separately contribute substantially to the resiliency, redundancy, or representation of the entire species (76 Fed. Reg. 14258).

The Service then found that threats to the Lizard’s current range (including that in Mexico) “have been reduced, managed, or eliminated, or found to be less substantial than originally thought.” The Service also found that implementation of the Interagency Conservation Agreement and associated Rangewide Management Strategy was reducing threats in the United States and was benefitting the species throughout its current range.

Therefore, we conclude that none of the existing or potential threats are likely to cause the [lizard] as an entire species …to be in danger of extinction or likely to become so within the foreseeable future throughout all or a significant portion of its range (76 Fed. Reg. 14267-8).

Whether the Service’s analysis is legally adequate remains to be seen, and may be subject to further legal challenge. What is clear is that as urbanization continues apace in Arizona, and in northern Mexico’s Baja region, the requirement that the Service take into adequate consideration the lost historical range of species that reside in Arizona and which are proposed for listing under the Act will take on greater significance.

It is also noteworthy that the Ninth Circuit’s 2009 Tucson Herpetological Society decision was very recently followed by the Ninth Circuit in its November 22, 2011, decision to remand back to the Service its decision to delist the Grizzly Bear in the Yellowstone region of the United States. In that case, the Ninth Circuit found the Service did not adequately consider evidence that brought into question the Service’s conclusion that the distinct population of grizzlies in the Yellowstone region was stable: The Yellowstone grizzly has been the focus of a laudable, decades-long cooperative research effort-one

that we hope continues. It may be that scientists will compile data demonstrating grizzly population stability in the face of whitebark pine declines. Such information, however, simply is not in the record before us.

The lack of any data showing a population decline due to whitebark pine loss is not enough (Greater Yellowstone Coalition v. Servheen, 665 F.3d 1015, 1030 (9th Cir. 2011), quoting Tucson Herpetological Soc’y, 566 F.3d at 879 (“If the science on population ... trends is undeveloped and unclear, the Secretary cannot reasonably infer that the absence of evidence of population decline equates to evidence of persistence.”).

The Ninth Circuit’s 2011 decision is notable because its effect is, in general, to delay the removal of ESA protections for the Grizzly in the lower 48 states.

In conclusion, neither the courts nor the Service have been able to provide a definitive answer as to what amount of “lost historical range” is “significant” enough to warrant a listing under the ESA. But the Ninth Court decisions regarding the Lizard in 2009 and the Grizzly Bear in 2011 make clear that whatever decision is made must be backed by firm data. As the following quote from the Tucson Herpetological Society case makes clear certain assumptions no longer apply: There seems to be a tacit assumption that if grizzlies survive in Canada and Alaska, that is good enough. It is not good enough for me …. Relegating grizzlies to Alaska is about like relegating happiness to heaven; one may never get there. – Aldo Leopold, A Sand County Almanac (1966:277) (Quoted by the Ninth Circuit Court of Appeals in Defenders of Wildlife v. Norton, 258 F.3d 1136, [9th Cir. 2001]).

2012 UPDATE In part, as a result of the Flat-tailed Horned Lizard listing decision and resulting litigation, the US Fish and Wildlife Service and NOAA’s National Marine Fisheries Service (Services), the two federal agencies responsible for administering the Endangered Species Act (ESA), proposed a new federal policy that will help clarify which species or populations of species are eligible for protection under the ESA and will provide for earlier and more effective opportunities to conserve declining species. See 76 Fed. Reg. 76987 (9 December 2011).

The proposed policy will define the key phrase “significant portion of its range” in the ESA and provide consistency for how it should be applied, aiding the agencies in making decisions on whether to add or remove species from the federal list of threatened and endangered wildlife and plants. The phrase is not defined in the ESA, but appears in the statutory definitions of “endangered species” and “threatened species” in the ESA. Until the policy is final, the Services have an obligation to meet statutory timeframes and make determinations in response to petitions to list, reclassify, and

SONORAN HERPETOLOGIST 27 (3) 2014

In conclusion, neither the courts nor the Service have been able to provide a definitive answer as to what amount of “lost historical range” is “significant” enough to warrant a listing under the ESA. But the Ninth Court decisions regarding the Lizard in 2009 and the Grizzly Bear in 2011 make clear that whatever decision is made must be backed by firm data.

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delist species. During this interim period, The Services will consider the interpretations and principles in this proposed policy as nonbinding guidance in making individual listing determinations. As nonbinding guidance, the Services will apply these interpretations and principles only as the circumstances warrant, and the agencies will independently explain and justify any decision made in this interim period in light of the circumstances of the species under consideration. The draft policy can be viewed at

Docket No. [FWS-R9-ES-2011-0031].

2014 UPDATE This rule change was finalized in 2014. http://federalregister.gov/r/0648-BA78

http://www.regulations.gov N AT U R A L H I S TO RY N OT E

Mating in the Coachwhip, Masticophis (= Coluber) flagellum Brian K. Sullivan, Arizona State University, PO Box 37100, Phoenix, AZ 85069-7100

O n 15 May 2014, immediately east of Cave Buttes dam on Cave Creek, on the northern edge of the

Phoenix Metropolitan region (33.729°N, 112.039°W), I encountered a mating pair of Coachwhips (Masticophis [= Coluber] flagellum) stretched across the edge of a dirt road at 0925 hrs. The air temperature was 86.9°F (30.5°C), and the ground temperature of the road was ~101°F (~38.5°C; briefly shaded while the temperature was recorded); the ground temperature in the shade of nearby mesquite trees was 85°F (29.5°C).

The snakes were immobile, but the female was apparently moving while dragging the male behind, when I approached in a vehicle. I stopped and exited my vehicle, remaining at a distance of 49 ft (15 m) to avoid

further disturbance, and watched the snakes. Their relatively bright color, and rate of tongue-flicking, suggested they were heating rapidly, and after 12 min, the female reversed direction, and began moving off the road to the apparent shade of mesquite trees on the east side of the roadway. The male was stretched behind the female, with an obvious bulge within the posterior of the female anterior to the vent, presumably due to intromission by the male, but they remained coupled as she moved 102 ft (31 m) and then disappeared into a burrow at the base of a mesquite tree. I subsequently measured the distance moved, and the approximate lengths of the snakes (both were ~59 inches [1500 mm] total length).

Figure 1. Mating pair of Coachwhips (Masticophis [= Coluber] flagellum) when first encountered; female to left, male to right. Photo

by Brian Sullivan.

On 15 May 2014, immediately east of Cave Buttes dam on Cave Creek, on the northern edge of the Phoenix Metropolitan region (33.729°N, 112.039°W), I encountered a mating pair of Coachwhips (Masticophis [= Coluber] flagellum) stretched across the edge of a dirt road at 0925 hrs.

Figure 2 (left). Mating pair of Coachwhips as female begins moving to shade; note bulge anterior to the vent of the female (upper, red colored snake). Figure 3 (right). Mating Coachwhips as female drags male to shade of mesquite trees; note the male is facing in the opposite direction, tongue flicking as he is dragged backwards over 98 ft (30 m) by the female (visible in the shade to the right).

Photos by Brian Sullivan.

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N AT U R A L H I S TO RY N OT E

Encounter between a Striped Racer (Coluber lateralis) and a Southern Alligator Lizard (Elgaria multicarinata) Howard O. Clark, Jr., Senior Wildlife Ecologist, Garcia and Associates, Clovis, CA; hclark@garciaandassociates.com

O

n 27 August 2014 at around 1300, while conducting wildlife surveys in the Angeles National Forest in Los Angeles County, California, I heard rustling in some leaves up slope from where I was parked. Something fell down the road cut onto the dirt road (an appoximate 25 ft [7.6 m] drop). I recognized it as a snake and a few seconds later saw something else thrashing about. Upon closer inspection I discovered that the snake was a Striped Racer (Coluber lateralis) and the other animal was a Southern Alligator Lizard (Elgaria multicarinata). The lizard was missing its tail (I found it in the leaves later) and the snake was periscoping its head searching for its prey item. The tail was likely lost during the skirmish above. The snake lunged forward and grabbed the lizard, but at the same time the lizard opened its mouth and latched onto the snake’s bottom jaw. The snake, with lizard attached, then crossed the dirt road and rested on a soil berm. Both reptiles remained locked together with little further struggle and remained that way until 1500, when I had to leave. I

checked the area the following morning and no sign of either species could be found, however the lizard tail piece was still present.

UTM of encounter: 11S 0405946 3789288; elevation 3545 ft (1080 m). Approximate length of snake: 37 inches (~94 cm); Approximate length of lizard— snout to tail termination: 5 inches (~12 cm). Detached tail length: 4 inches (~10 cm). Temperature in broken shade: 82°F (28°C).

Although Striped Racers have been previously documented predating on Southern Alligator Lizards (Cunningham 1959), photo-documentation of the attempted event is noteworthy.

Acknowledgments—I thank Robert Hansen and Jackson Shedd for their commments and ideas about the encounter.

Literature Cited Cunningham, J.D. 1959. Reproduction and food of some California snakes. Herpetologica 15:17–19.

Although Striped Racers have been previously documented predating on Southern Alligator Lizards (Cunningham 1959), photodocumentation of the attempted event is noteworthy.

Photo sequence of the encounter between a Striped Racer (Coluber lateralis) and Southern Alligator Lizard (Elgaria multicarinata).

All photos by Howard Clark.

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BOOK REVIEW

Biology and Conservation of North American Tortoises

Review by Margaret H. Fusari, UC Santa Cruz—retired, Board member, Desert Tortoise Council, Tucson Herpetological Society David C. Rostall, Earl D. McCoy, Henry R. Mushinsky; editors Johns Hopkins University Press Hardcover, x + 190pp.

ISBN-13: 978-1-4214-1377-8 $63.66 (listed); June 2014

E dited by three tortoise researchers and authored by 33 others (including the editors) this book will be

a valued reference volume for anyone who wants or needs to review various topics in North American tortoise systematics, taxonomy, phylogeny, genetics, morphology, paleontology, physiology, behavior, reproduction, diet, growth, health, ecology, biology, distribution, and perhaps most important—conservation status and issues.

In the words of the frontpiece: “Providing the first comprehensive treatment of North America’s tortoises Biology and Conservation of North American Tortoises brings together leading experts”...who... “carefully combine their own expertise and observations with results from studies conducted by hundreds of other researchers.”

The (current) five species; Gopherus agassizii, G. morafkai, G. berlandieri, G. flavomarginatus, and G. polyphemus (apologies to those who favor use of the genus Xerobates for the first two species) are all in trouble at one level or another (see Chapter 18 for a summary) and are in need of protection. That can only be based on sound science and its interpretation. This volume will help tortoise advocates, professional or amateur, understand their animal(s) better. It will also help a lot of researchers locate references because of the extensive (although not complete) bibliography.

While reading the Preface, one is struck by what the book could not do; come to an agreement as to what to call the tortoises. Gopherus or Xerobates, Agassiz’s or Mojave, Morafkai’s or Sonoran, Mojave or Mohave? The authors of the paper Murphy et al. (2011) were right on the money with their title (The dazed and confused identity of Agassiz’s land tortoise, Gopherus agassizii). But rather than be disappointed that the editors did not get agreement on the names, formal or common, I am impressed that we begin by realizing that we do not know it all yet; that more work does need to be done. This is apparent in many of the chapters as the authors inform us as what they feel would be the next things we need to know.

Of all the names mentioned my favorite terms (Chapter 17) are Cracker chicken, which changed to Hoover chicken signifying a change is why people needed to eat wild tortoises. Names have many differ-

Cover of Biology and Conservation of North American Tortoises edited by D.C. Rostall, E.D. McCoy, and H.R. Mushinsky; 2014; Johns Hopkins University Press.

ent meanings, contexts, and histories. The chapter was making the point that many people used to eat the tortoises and probably moved them around a lot from the earliest times that people interacted with tortoises, a fact we need to remember.

The word “variable” appears frequently and in many chapters. Each group of chapter authors has gathered the references that exist and presented a synopsis of what is known. Because of variable methods used by the various authors and by the differences in species, geography, recent climate and seasons the knowledge we have shows high variability. Some of the references are clearly pseudoreplication in space or time and hence general conclusions will have to wait for more replication.

Beyond the variability of the references I am impressed with that of the tortoises themselves. Being most associated with the Desert Tortoises I am impressed with the variability of foraging and diet, habitat use, reproductive strategy, social interactions, and the overall opportunism that a desert species must show to survive. It would seem from the chapters that the tortoise species from more mesic habitats also show some of that variability in their life history strategies.

Rather than try to summarize everything, even though every chapter has considerable merit, I am going to call out a few notable items that caught my attention.

In the first three chapters we learn that there were two major lineages leading to further speciation events in each. The first chapter presents the anatomical evidence, the third the genetic evidence and the second

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The (current) five species; Gopherus agassizii, G. morafkai, G. berlandieri, G. flavomarginatus, and G. polyphemus (apologies to those who favor use of the genus Xerobates for the first two species) are all in trouble at one level or another (see Chapter 18 for a summary) and are in need of protection.

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an overview of the fossil record. In a context of disagreement about times and names I see a general consensus but a need for more research to identify important details especially on local genetic structure.

Several chapters address food and water in addition to the primary chapter (10), in which the theme of variability is well documented. Variation in forage species composition due to geography, recent climate/ weather, fire, grazing, and weed invasion are clearly of concern in trying to discover “preferred” forage species and identify long term habitat and/or restoration needs. Chapter 9 is clear on the problems with fire and grazing affecting the availability of good forage as well as presenting a valuable summary on the overall habitat needs of tortoises.

Chapter 4 on thermoregulation and energetics gives a fine overview of tortoises as ectotherms that need a microenvironment awareness superior to that of us endotherms. It also makes the potential impacts of climate change all too clear because the local thermal (as well as nutritional and reproductive) environments are going to change out of the adaptive range of the local tortoises.

There is a set of chapters covering the many aspects of tortoise reproduction including the reproductive physiology, the embryology and the temperature dependent sex determination (important to add to any discussion of climate change impacts), and growth patterns (it does depend on resource availability and quality and hence is highly variable). Two other chapters address the nesting and reproductive behavior and the social behavior. Tortoises are social! They live in groups, they know each other, both male-male and female-female interactions are important, and they can reestablish their social structure as changes occur given appropriate opportunities. (In other words moving tortoises around has to be done carefully to provide both a safe and opportune situation.)

It is praiseworthy that the chapter detailing health assessment (Chaper 8) is not only a comprehensive introduction to the multiple diseases the tortoises face but also has a glossary to help readers understand the complex concepts. The warning that stress can exacerbate disease and the reference to the frequency of epizootic episodes is worth a lot of attention as hu-

man behaviors change the way diseases play out in the field and the mortality populations suffer. I wish the book had added a chapter on the epidemiology of the disease(s) because diseases, acting alone and in synergy, are confirmed as a major threat in several chapters and they will almost surely spread. A take home message, again, is that we must be careful moving tortoises around.

Chapter 18, on threats and conservation is a must read to be aware of the complexity of the issues that tortoises face. The lack of protection on private land is important and might be addressed in cooperative ways of the organizations mentioned got together and worked on this and other issues. The need to better understand the issues behind mitigation and augmentation methods in terms of sound science is another important point.

Several chapters include overlapping points with other chapters about the tortoise-environment interactions that obviously have many dimensions and that are important to understand in all their complexity if we are to work to assure the survival of our North American tortoises in the face of climate change and other anthropogenic impacts.

There are a few things that could be called out for better attention. There are references missing (HINT: learn to use Citation Index to locate them). There is no common voice to the chapters, they read very differently and some contain highly technical terms or archaic words that make them hard to understand if one is not already experienced in the field. There is no warning that to handle tortoises almost always required permits and often training, a strange omission in a book with conservation in the title.

Is this book worth it to a person interested in or working with tortoises in North America? Yes it is!

Literature Cited Murphy, R.W., K.H. Berry, T. Edwards, A.E. Leviton, A. Lathrop, and J.D. Riedle. 2011. The dazed and confused identity of Agassiz’s land tortoise, Gopherus agassizii (Testudines, Testudinidae) with the description of a new species, and its consequences for conservation. ZooKeys 113:39–71.

It is praiseworthy that the chapter detailing health assessment (Chaper 8) is not only a comprehensive introduction to the multiple diseases the tortoises face but also has a glossary to help readers understand the complex concepts.

BOOK REVIEW

The Texas Tortoise: A Natural History Review by Taylor Edwards, Tucson Herpetological Society

I always think of the Texas Tortoise, Gopherus berlandieri, as the least studied of the Gopherus species, but this new book by researchers Drs. Francis Rose and Frank Judd compiles an extensive portrait of the smallest of the Gopherus species. This volume is well cited and dense with information. Although a couple other books provide nice summaries of current research on

the species, such as Turtles of the United States and Canada by Carl Ernst and Jeffrey Lovich (Johns Hopkins University Press, 2nd ed. 2009) and The Turtles of Mexico: Land and Freshwater Forms by John Legler and Richard Vogt (University of California Press, 2013), The Texas Tortoise’s focus on this single species allows for greater presentation of anecdotal accounts and speculation. The release of this book also coincides with the

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new volume Biology and Conservation of North American Tortoises, edited by David C. Rostal, Earl D. McCoy and Henry R. Mushinsky (Johns Hopkins University Press, 2014), however the two are very different in style and presentation. Of these three examples, The Texas Tortoise is perhaps the most accessible to a broad audience. The book’s greatest attribute, but also sometimes its greatest liability, is the experience and opinions of its two authors. Although they address many topics, most attention is in relation to their own research and it is thus like a thesis complied from their lifetimes of work. With that said, they present an informative and interesting picture of the species and its natural history.

The book is divided into 12 Chapters as follows: 1. General description and phylogenetic relationships 2. Geographic Range and habitat 3. Animal associates and relationships 4. Morphology and Anatomy 5. Size, Growth, and Sexual Dimorphism 6. Sensory Modalities 7. Temperature regulation, water use and retention, and photoperiod response 8. Behavior 9. Population ecology 10. Environmental resistance to tortoise survival 11. Conservation, Monitoring, and Maintenance 12. The Future: Species significance and bearing on Ecological problems As this book was prepared by a pair of authors as opposed to an edited volume (as with the other examples above), the book primarily focuses on what the authors know best, which may be best summarized as population and physiological ecology. Having single authors provides a consistent voice but also results in a lack of detail where the authors are less knowledge.

Subjects like population genetics are treated more peripherally despite there being a body of knowledge for Gopherus spp. to expand upon. Other sections appear inconclusive, such as a discussion on taxonomy and systematics (pp. 12-13) where the authors briefly introduce the basic principles before addressing this subject in the context of G. berlandieri.

They have a lot to say about some subjects (e.g., home range—8 pages) and little to say regarding others (e.g., habitat selection—1 paragraph), despite supporting studies on the subject that could be elucidated further. Surprisingly, diet is only qualitatively addressed in reference to prickly pear cactus (Opuntia engelmannii). The authors frequently use comparative information for other congeners (Gopher Tortoise, Bolson Tortoise, and Desert Tortoise), but unfortunately do not always choose the most related taxa when there is information available. Since Desert Tortoises are the most closely related group to the Texas Tortoise, they

The Texas Tortoise: A Natural History

by Francis L. Rose & Frank W. Judd University of Oklahoma Press, 2014 Series: Animal Natural History Series (Book 13) Hardcover: 210 pages; Publisher: University of Oklahoma Press (June 5, 2014); Language: English; ISBN10: 0806144513 should be the most appropriate comparison, however, in a discussion of the potential for multiple paternity they reference only the Gopher Tortoise study by Moon et al. (2006) instead of Davy et al. (2011) who studied this in Desert Tortoises.

Where the authors have more expertise, those sections have the most detail; for example discussions on mating systems begin with a detailed presentation of Emlen and Oring’s (1977) classification system (p. 89).

In the section on home range, it is obvious that the author’s personal experience influences their opinion because they conducted some of the few studies (in 1975 and 1983) that estimate home range for G. berlandieri.

The authors give an extensive critique of methods for estimating home range, but curiously leave out widely used kernel estimator techniques or more advanced modern methods of calculation, such as Core Area Estimation (see Wilson et al., Journal of Wildlife Management 2010).

One of the benefits of having single authors with vast amounts personal experience is that they share unpublished and anecdotal information and this may be the greatest benefit of this work to the collected literature. For example, they describe how a male tortoise whose concave plastron is filled with plaster is unable to copulate because it cannot keep its balance on the female (p. 87). One of my favorite anecdotal observations is where the authors describe tortoise urine after the animals have consumed prickly

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Where the authors have more expertise, those sections have the most detail; for example discussions on mating systems begin with a detailed presentation of Emlen and Oring’s (1977) classification system (p. 89).

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pear; “…the odor is not unpleasant, and we can safely vouch that it is bland to the taste, having had ample experience” (p. 109).

One of the most fascinating discussions is their recurring mention of finding eggs laid on the surface, presumably above a nest. They suggest that the egg is laid after the nest is buried. Texas Tortoises may lay more than one nest on different days but the frequency which they observed this odd behavior seems high and not adaptive. The authors can only speculate about this phenomena; reiterating that this behavior is not documented across studies/sites and maybe only occurs in hard packed soils. This highlights that there are a limited number of study sites where research has been conducted on G. berlandieri and a global perspective of the species is severely lacking.

Most often, where there is debate, the prose is often strongly opinionated. I particularly applaud the authors for stating their uncompromising views on issues of conservation. In reference to drivers who intentionally run over tortoises, it is not often enough that academicians stand up to say things like “one might wish those drivers to be reincarnated as a flea on a Hell Hound’s back and be chased by a pair of red-hot fangs for 10,000 years” (p. 153). Reminiscent of our own Roger Repp, in reference to the construction of the border wall between the U.S. and Mexico, they make their opinion clear by stating that it “… will stand mostly as another monument of human stupidity.”

Unfortunately, there are only 15 pages in the chapter on conservation and although highly forthright in its message, it contains limited data.

In expressing their opinion about other researchers, the authors at times take an almost reprimanding tone in describing research that lacks details or statistics in their publications. For example they highlight a contradiction in a paper on mating systems by Johnson et al. (2009) where they find a discrepancy between the paper’s title and its conclusions (p. 89). At times, some statements appear contradictory. In the section on Life Expectancy and Replacement, they make the claim that “Certainly Cameron County females were larger, which suggest that they were older than those from Dimmit and La Salle Counties” (p. 145). Then in the next section on the same page, Regulation of Numbers, they hypothesize that where food is more abundant, tortoises should be bigger and they show supporting evidence for this being true in females. Such contradictions can be distraction to the reader.

Another area that seems a bit misinformed is in the captive care section where the authors recommend using processed Mazuri® tortoise chow over a natural diet and they recommend supplementing with forage like ‘greens and fruit’, without mention of using native foraging plants. While this may suffice for tortoises in captivity, extensive studies of Desert Tortoises suggest that maintaining a diverse, natural diet is optimal for tortoise health. Oddly, in this section on captive hus-

bandry they spend twice as much space detailing instructions for administering CPR in case of drowning (p. 161). Although interesting (and a bit amusing), this may not necessarily be the most important thing that people need to know in the context of properly caring for a tortoise at home.

There are currently only a few other collections that focus on a single species of Gopherus, and this volume takes an important place on the shelf among those; for Gopher Tortoises, there is The Natural History and Management of the Gopher Tortoise Gopherus polyphemus, by Ray and Patricia Ashton (Krieger Publishing Company, 2007) and for our native Morafka’s Desert Tortoise there is The Sonoran Desert Tortoise: Natural History, Biology, and Conservation edited by Thomas R. Van Devender (University of Arizona Press, 2006). Surprisingly, there is not a definitive work yet specifically on G. agassizii in the Mojave Desert despite it being one of the most studied turtle species in the world (although Woodbury and Hardy’s 1948 Ecological Monograph is still an impressively thorough collection of information). Van Devender’s The Sonoran Desert Tortoise (University of Arizona Press, 2002) book has greater focus on population ecology, distribution, husbandry and health, habitat use and conservation. Ashton and Ashton’s Gopher Tortoise book (Natural History and Management of the Gopher Tortoise Gopherus polyphemus [Daudin]; Krieger Publishing Company, 2007) is written more of an instruction manual for monitoring, care and management. Because of the physiological focus of The Texas Tortoise, it includes many points of interest that are not included in these other single species accounts, for example detailed figures and descriptions of reproductive organs and carpal elements in a tortoise’s foot (pp.

57 and 73-74). An example of other details not found in other collections is Chapter 6 which focuses on Sensory Modalities. Although this chapter is only 5 pages, I have yet to see this fascinating subject summarized elsewhere. As such, this volume may be considered a compliment to any tortoise enthusiast’s collection.

While The Texas Tortoise contains a plethora of information, unfortunately what we do know about Texas Tortoises has mostly come from only a handful of sites and information is lacking entirely from Mexico (see summary in Legler and Vogt 2013, The Turtles of Mexico: Land and Freshwater Forms; University of California Press). The authors of The Texas Tortoise are two of the major contributors to what we do know, and their data come mainly from 2 study sites.

This epitomizes where our knowledge falls short for this species because site-to-site variation is frequently observed and there are inconsistencies among studies.

As a result, many sections of the book end with statements like; “This aspect… is a fertile area for further research”, or “…this is certainly an area in need of further research”. Fortunately, much of what is known about Texas Tortoises can now be found in this single volume and hopefully this will inspire a new genera

SONORAN HERPETOLOGIST 27 (3) 2014

Most often, where there is debate, the prose is often strongly opinionated.

I particularly applaud the authors for stating their uncompromising views on issues of conservation.

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tion of biologists and naturalists to find out more about what we don’t know.

Literature Cited (excluding books) Davy, C.M., T. Edwards, A. Lathrop, M. Bratton, M. Hagan, B. Henen, K.A. Nagy, J. Stone, L.S. Hillard, and R.W. Murphy. 2011. Polyandry and multiple paternities in the threatened Agassiz’s Desert Tortoise, Gopherus agassizii. Conservation Genetics 12:1313–1322.

Emlen, S.T., and L.W. Oring. 1977. Ecology, sexual selection, and the evolution of mating systems. Science 197:215–223.

Johnson, V.M., C. Guyer, S.M. Hermann, J. Eubanks, and W.K. Michener. 2009. Patterns of dispersion and burrow use support scramble competition

polygyny in Gopherus polyphemus. Herpetologica 65:214–218.

Moon, J.C., E.D. McCoy, H.R. Mushinsky, and S.A. Karl. 2006. Multiple paternity and breeding system in the Gopher Tortoise, Gopherus polyphemus. Journal of Heredity 97:150–157.

Wilson, R.R., M.B. Hooten, B.N. Strobel, and J.A. Shivik. 2010. Accounting for individuals, uncertainty, and multiscale clustering in core area estimation.

Journal of Wildlife Management 74:1343–1352.

Woodbury, A.M., and R. Hardy 1948. Studies of the Desert Tortoise, Gopherus Agassizii. Ecological Monographs 18:145–200.

MEETING MINUTES BOD minutes can be found here: http://bit.ly/1qcYyGg

Including the THS in your will is an excellent way to support the value of this organization and the conservation of the herpetofauna of the Sonoran Desert.

We would like to recognize and thank anyone who has included the THS in their will. Please contact us so we can express our appreciation. For information about designating the THS in your will, please contact Kent Jacobs, Treasurer, Tucson Herpetological Society, at jacobskent@gmail.com.

MEMBERSHIP

Membership Information Individual Family Student

$20 $25 $14

Sustaining Contributing Life

$30 $50 $500

Time to Renew Your THS membership?

The Tucson Herpetological Society would like to thank existing members and new members for renewing their membership. We appreciate your support and are always looking for members to actively participate in THS activities and volunteer opportunities. It is a great way to be involved with the conservation of amphibians and reptiles in the Sonoran Desert.

Thank you for your membership in the Tucson Herpetological Society. Renewal reminders for upcoming membership expiration will be emailed at the beginning of the month that your membership expires. If you have any questions about your membership or would like to be in touch with a THS member you do not know how to reach, please contact our Membership Coordinator, Robert Villa, by email: cascabel1985@gmail.com.

Tucson Herpetological Society P.O. Box 709, Tucson, Arizona 85702-0709 MEMBERSHIP RENEWAL FORM NAME: ________________________________________________ Date ______________ Address or Personal Information Changes_______________________________________ _________________________________________________________________________ _________________________________________________________________________ MEMBERSHIP DUES [ ] $20 Individual [ ] $25 Family [ ] $14 Student [ ] $30 Sustaining [ ] $50 Contributing [ ] $500 Life $ _______ Jarchow Conservation Award $ _______ Flat-tailed horned lizard Fund

$ _______ Speakers Bureau $ _______ C.H. Lowe Herp Research Fund

$ _______ Total (MAKE CHECK PAYABLE TO: TUCSON HERPETOLOGICAL SOCIETY) The THS newsletter, the Sonoran Herpetologist, is delivered online only. Please indicate the email address you would like to receive the newsletter if you are not currently receiving the newsletter at your preferred address. If you are unable to receive the newsletter online, please contact Robert Villa at cascabel1985@gmail.com. If not already done, please indicate if you want your email added to the THS directory and/or the Monthly meeting announcement (circle one or both). Please return this form with your check to the address above.

Email address ___________________________________________________________

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Sonoran Herpetologist (ISSN 2333-8075) is the newsletter-journal of the Tucson Herpetological Society, and is Copyright © 1988-2014. The contents of Sonoran Herpetologist may be reproduced for inclusion in the newsletters of other herpetological societies provided the material is reproduced without change and with appropriate credit, and a copy of the publication is sent to the Tucson Herpetological Society. Occasional exceptions to this policy will be noted. Contents are indexed in Zoological Record. A complete set of back issues are available in the Special Collections area of the University of Arizona library. They are accompanied by a copy of The Collected Papers of the Tucson Herpetological Society, 1988-1991.

Editor-in-Chief Howard Clark, Jr., editor.sonoran.herp@gmail.com Associate Editors Robert Bezy, robertbezy@gmail.com Dennis Caldwell, dennis@caldwell-design.com Suman Pratihar, pratihar_vu@rediffmail.com Roger Repp, repp@noao.edu Don Swann, donswann@dakotacom.net Art Editor Dennis Caldwell, dennis@caldwell-design.com Book Review Editor Philip Brown, prbrownnaturalist@gmail.com

Information for Contributors Authors should submit original articles, notes, book reviews to the Editor, either via email using an attached word processed manuscript or by mail to the Society’s address. The manuscript style should follow that of Journal of Herpetology and other publications of the Society for the Study of Amphibians and Reptiles. For further information, please contact the editor, at editor.sonoran.herp@gmail.com.

The Tucson Herpetological Society is dedicated to conservation, education, and research concerning the amphibians and reptiles of Arizona and Mexico.

Tucson Herpetological Society is a registered non-profit organization.

President Roger Repp, repp@noao.edu Vice President Krista Schmidt, turtlerad@hotmail.com Secretary René Clark, serpentprincess@comcast.net Treasurer Kent Jacobs, jacobskent@gmail.com Directors: Dennis Caldwell, dennis@caldwell-design.com Margaret Fusari, maggiefusari@gmail.com Rob Nixon, sapo1047@msn.com Don Moll, donandjanice@q.com Ed Moll, e.o.moll@gmail.com Jim Rorabaugh, jrorabaugh@hotmail.com Membership Robert Villa, cascabel1985@gmail.com Past President Robert Villa, cascabel1985@gmail.com Editor Howard O. Clark, Jr., editor.sonoran.herp@gmail.com

Society Activities

Monthly Members Meeting Roger Repp, Program Chair 3rd Tuesday, 7:15 PM Board of Directors Meeting Last Tuesday of each month (except December), 7:00 PM University of Arizona, BIO5/Keating Building 1657 East Helen Street Speakers Bureau (scheduled presentations) Robert Villa & Ed Moll Conservation Committee Dennis Caldwell Herpetological Information Hotline Bob Brandner, (520) 760-0574 Jarchow Conservation Award Roger Repp Publications: Sonoran Herpetologist, Backyard Ponds brochure, Living with Venomous Reptiles brochure, THS Herp Coloring Book, THS Collected Papers, 1988-1991

LCCN permalink: http://lccn.loc.gov/2013273781

Deadline for Sonoran Herpetologist: 15th of Feb, May, Aug, and Nov (based on the quarterly schedule)

Officers

THS Internet World Wide Webpage http://tucsonherpsociety.org Marty Tuegel, Webmaster, mtuegel@cox.net

For more information about the THS and the reptiles and amphibians of the Tucson area visit

tucsonherpsociety.org

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