Sh vol 27 2014 12

25 Volume 27

January 2012 December 2014

Number 14

ISSN 2333-8075

THIS MONTH’S PROGRAM

FUTURE SPEAKERS

Tom Jones

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14 Jan 2015: Jeff Servoss

Amphibians and Reptiles Program Manager for AZ Game and Fish

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11 Feb 2015: Lawrence Jones

Invasive bullfrog removal in the American Southwest: A case study from the Pajarito/Atascosa mountains, Arizona 7:15 PM; Wednesday, 10 December

RESEARCH ARTICLES 88

“Reproduction in the Southern Prairie Lizard, Sceloporus consobrinus (Squamata: Phrynosomatidae), from Oklahoma” by Stephen R. Goldberg

91

“Effects of Buffelgrass Invasion on Density and Condition of Sonoran Desert Tortoises (Gopherus morafkai)” by Katherine M. Gray and Robert Steidl

95

“Herpetofauna of the Canelo Arc”

by Robert L. Bezy and Charles J. Cole

Tucson City Council Ward 3, 1510 East Grant Road, Tucson, AZ 85719

T om Jones is the Amphibians and Reptiles Program Manager for the Arizona Game and

Fish Department, where he oversees conservation and management of amphibians and reptiles in Arizona. He received a B.S. and M.S. at Auburn University, and a Ph.D. at Arizona State University. After postdocs at the University of Michigan he moved to East Carolina University for a year as a Visiting Assistant Professor. From there he moved back to Arizona, where for 11 years he was on the Grand Canyon University faculty. While at GCU he also taught Neotropical herpetology at the Bocas del Toro Biological Station, in Bocas del Toro, Panamá. In 2005, after GCU had become a for-profit institution he joined the Arizona Game and Fish Department, where he has been for a little more than 9 years. His research interests include ecology, population and conservation biology of amphibians and reptiles. He has done field work in much of the continental United States, and in México, Panamá, Costa Rica, and Vietnam.

Most of his recent efforts, when not sitting at his desk being an administrator, have involved efforts to rid bullfrogs from high priority conservation areas.

When it comes to native aquatic wildlife conservation, invasive exotics are among the greatest impediments to management and/ or recovery. Whether the discussion is about fishes, frogs or gartersnakes, invasive species play an enormous role in reducing native populations and preventing us from securing

BOOK REVIEW 102 “The Biology of Chameleons” Review by Russell Ligonn TAXO N O M Y 79 Tom Jones with a Green Ratsnake (Senticolis triaspis) in southern Arizona. Photo by Tom Jones.

“New tree frog named for Ozzy Osbourne” Submitted by Suman Pratihar

or reestablishing native species. The Amphibians and Reptiles Program in the Arizona Game and Fish Department’s Nongame Wildlife Branch has engaged with partners in a number of efforts to eliminate bullfrogs or create buffers to bullfrog dispersal. Tom’s presentation will provide an overview of Tucson Herpetological Society one of the more successful projects cenmeetings are open to the public and tered around Peña Blanca Lake, highlighting are held on the second Wednesday of the successes that can be expected when each month starting at 7:15 PM resources are committed to invasive species eradication.

SONORAN HERPETOLOGIST 27 (4) 2014 86

FUTURE SPEAKER

14 January 2015:

Jeff Servoss

The Decline of Northern Mexican and NarrowHeaded Gartersnakes: Can they be recovered?

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

JerneffIdaho, grew up in eastern Washington State and northchasing around such regional prizes as

Great Basin Gophersnakes (Pituophis catenifer deserticola), Northern Pacific Rattlesnakes (Crotalus oreganus oreganus), and if lucky, Rubber Boas (Charina bottae) (hey … it wasn’t Arizona). Jeff began his studies at Washington State University in the wildlife program, but became increasingly dissuaded by the depressing job market in natural resource management that plagued the early 1990s. Therefore, he switched degrees and graduated with a B.S. in Environmental Science and subsequently worked on soil and groundwater contamination and remediation, as well as surface water quality assessments. Jeff ultimately found his way back to wildlife in 2001 when he started as a Wildlife Biologist with the U.S. Fish and Wildlife Service and currently resides in Tucson, Arizona. He works predominantly on species status evaluations and conservation and recovery planning and project implementation for reptiles and amphibians in the Southwestern United States, in addition to delivering programs to the local community on the venomous reptiles of Arizona. His focus most recently has included the Sonoran Desert Tortoise (Gopherus morakai), and federally threatened species including the Northern Mexican Gartersnake (Thamnophis eques megalops), Narrow-Headed Gartersnake (Thamnophis rufipunctatus), and Chiricahua Leopard Frog (Lithobates chiricahuensis). In his free time, Jeff enjoys recreational field herping, photography, running with his dogs, and is an avid tennis player.

About the talk: There are many threats to our riparian and aquatic ecosystems in the arid Southwest, both hydrologically (diversions, groundwater pumping, dams) and ecologically (nonnative species). Over the last 40-50 years, native riparian and aquatic vertebrate communities have undergone a slow transformation; a transitional shift from native, to increasingly nonnative in complexion. Our native species are largely predator-naïve.

With the exception of large chubs and pikeminnow, our native species did not evolve in the presence of aggressive predators like those from the Mississippi

Jeff releasing a Mexican Gartersnake (Thamnophis eques) reared at the ASDM to Las Cienegas. Photo by Tom Jones.

Basin which are now largely ubiquitous across Arizona (i.e. warm-water sportfish, bullfrogs, crayfish). The result of this shift has been more than deleterious to our riparian and aquatic communities. In Arizona, 19 of 31 (61 percent) of all native fish species are listed under the Endangered Species Act; another two are candidates for listing and may be proposed for listing within a year. A shocking 86 percent of our native fish species have declining trends, ranking Arizona the highest of all 50 States in this statistic. In addition to the threatened Chiricahua Leopard Frog, there is some concern for the future of all leopard frog species in Arizona. Over the past several decades, we have witnessed a slow-speed, ecological, ripple effect by the harm inflicted mostly by these nonnative species, as evidenced by the documented declines in two of our five native gartersnake species, Northern Mexican and Narrow-headed Gartersnakes (the two most-associated with water). Some nonnative species, in their young or larval age class, provide important prey for extant gartersnake populations. However, the science has shown that over time, nonnative species out-compete these gartersnakes for prey which can result in starvation of populations. Nonnatives also prey on neonatal and juvenile gartersnakes themselves, which limits recruitment of new snakes into the population. Is all lost? Not necessarily. Come listen to Jeff discuss these complex ecological relationships and what can be done to save Arizona’s unique natural heritage.

SONORAN HERPETOLOGIST 27 (4) 2014

Over the past several decades, we have witnessed a slowspeed, ecological, ripple effect by the harm inflicted mostly by these nonnative species, as evidenced by the documented declines in two of our five native gartersnake species, Northern Mexican and Narrow-headed Gartersnakes.

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FUTURE SPEAKER

11 February 2015:

Lawrence L. C. Jones Southwest Reptile Research Collective Tucson, Arizona

Lizard activity and the monsoon: is there a connection?

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

L

awrence L. C. Jones hails from southern California.

He received his B.S. and M.S. in biology and zoology from California State University at Long Beach. Larry worked on habitat relationships of mammals (especially American Marten) and stream-dwelling amphibians in the Pacific Northwest for about twenty years, before he had a midlife crisis and moved to southeastern Arizona. He worked for both research and management branches of federal agencies for thirty years and then retired to become a real biologist again. Larry resides in Tucson, where he studies lizard ecology and invertebrates, under the moniker, Southwest Reptile Research Collective. Larry has co-authored three books on reptiles and amphibians, two about amphibians of the Pacific Northwest and one about lizards in the American Southwest. He has also authored about 90 publications on herps and other topics of biology.

He is currently working on a new book, “Venomous Animals of the American Southwest,” which covers terrestrial and marine invertebrates and vertebrates, from California to Texas.

If you ask any herpetologist about when to look for snakes in southeastern Arizona, they will be quick to respond that you should do so during the monsoon.

If you ask the same question about lizards, you will get mixed answers. There is surprisingly little published

Larry Jones with a Sonoran Spiny-tailed Iguana (Ctenosaura hemilopha macrolopha). Phoyo by Mike Sredl.

information about surface activity of lizards during the year, and less about the influence of the monsoon.

Larry’s research near Safford, Arizona, will try to answer some questions about the seasonal patterns of surface activity of lizards in an area where the North American monsoon occurs. His study near Marijilda Canyon has amongst the highest diversity of lizards in the United States. With help from his colleagues, the study revealed interesting patterns of surface activity among lizards. The upshot is that some species are influenced by this seasonal change in weather patterns, while others not so much. Then there is the whole climate change thing. While this slideshow focuses on results from his study, he discusses the bigger picture across the range of lizards in the American Southwest.

His presentation is lean on technical information and heavy on cool herp pictures.

RESEARCH ARTICLE

Reproduction in the Southern Prairie Lizard, Sceloporus consobrinus (Squamata: Phrynosomatidae), from Oklahoma

Larry’s research near Safford, Arizona, will try to answer some questions about the seasonal patterns of surface activity of lizards in an area where the North American Monsoon occurs. His study near Marijilda Canyon has amongst the highest diversity of lizards in the United States.

Stephen R. Goldberg, Whittier College, Department of Biology, Whittier, CA 90608; sgoldberg1109@gmail.com

S2002celoporus consobrinus, as defined by Leaché and Reeder (Fig. 1), is found throughout the central United States; its northern range extends into South Dakota and eastward to southwest Illinois, while the southern range continues eastward to Louisiana and southward into San Luis Potosí and Zacatecas, Mexico (Lahti and Leaché 2009). It occurs statewide in Oklahoma (Sievert and Sievert 2011). The purpose of this paper is to report, to my knowledge, the first information on

reproduction of S. consobrinus from Oklahoma as part of an ongoing survey of the reproductive cycles of lizards from western North America.

A sample of 89 S. consobrinus consisting of 40 adult males (mean SVL = 52.1 mm ± 5.7 SD, range = 38-63 mm), 46 adult females (mean SVL = 52.8 mm ± 6.5 SD, range = 41-66 mm) and 3 juveniles (mean SVL = 26.0 mm ± 2.6 SD, range = 23-28 mm) collected 1925 to 2010 and deposited in the Sam Noble Oklahoma

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Museum of Natural History (OMNH), University of Oklahoma, Norman was examined (Appendix).

A small incision was made in the lower part of the abdomen and the left gonad was removed for histological examination.

Histology slides were cut at 5 µm and stained with hematoxylin followed by eosin counterstain (Presnell and Schreibman 1997).

Histology slides were deposited at OMNH. Enlarged ovarian follicles (> 4 mm) or oviductal eggs were counted. An unpaired t-test was used to test for differences between male and female mean SVLs; linear regression analysis was used to test for the correlation between female body size (SVL) and clutch size. (Instat, vers. 3.0b, Graphpad Software, San Diego, CA).

There was no significant difference between mean SVLs of S. consobrinus male and female samples (t = 0.53, df = 84, P = 0.60). Three stages were present in the testicular cycle of S. consobrinus (Table 1): (1) regression: seminiferous tubules are at their smallest sizes and contain 2-3 layers of spermatogonia and Sertoli cells; (2) recrudescence: a proliferation of germ cells has commenced and is evidenced by the abundance of primary or secondary spermatocytes; (3) spermiogenesis: lumina of the seminiferous tubules are lined by sperm or clusters of metamorphosing spermatids. The period of sperm production encompassed March into July. The smallest reproductively active male (spermiogenesis) measured 38 mm SVL (OMNH 41608) and was collected in April.

Four stages were present in the monthly ovarian cycle of S. consobrinus from Oklahoma (Table 2): (1) quiescent, no yolk deposition; (2) early yolk deposition (basophilic vitellogenic granules in ooplasm); (3) enlarged ovarian follicles > 4 mm; (4) oviductal eggs.

Female reproduction occurred from May into July. I did not examine S. consobrinus from August, but the presence of females with enlarged follicles > 4 mm from July (Table 2) suggests some reproduction occurs in August. Mean clutch size (n = 16) was 7.0 ± 2.3 SD, range = 3-10. Linear regression analysis revealed a significant positive correlation between female body size (SVL) and clutch size. This relation is shown by the regression equation: Y = -9.6 + 0.29 X, r = 0.50, P = 0.05. One female from June (OMNH 11378) contained oviductal eggs and concomitant yolk deposition indicating S. consobrinus females may produce multiple clutches in the same reproductive season. This has been previously reported for S. consobrinus in Lahti and

Figure 1. Sceloporus consobrinus from Logan Co., KS. Photo by Danny Martin; www.dannymartinphotography.com.

Leaché (2009). The smallest reproductively active female (yolk deposition) measured 44 mm SVL (OMNH 29554) and was collected in April. Two slightly smaller females (OMNH 44015) SVL = 42 mm and (OMNH 30105) SVL = 41 mm, both collected in April were, subjectively, considered as adults that would have produced eggs later in the reproductive season. Sceloporus consobrinus of presumably hatchling size (OMNH 13208, SVL = 23 mm, OMNH 13211, SVL = 27 mm, OMNH 13210, SVL = 28 mm) were all collected in August. This is when hatchlings of S. consobrinus appeared in Illinois (Phillips et al. 1999). The period of reproduction presented herein from Oklahoma, April Table 1. Stages in the monthly testicular cycle of 40 adult male S. consobrinus from Oklahoma. n

Regressed

Recrudescence

March

5

0

0

5

April

16

0

0

16

May

4

0

0

4

June

8

1

0

7

July

6

4

1

1

Sept

1

1

0

0

Month

Spermiogenesis

Sceloporus consobrinus is found throughout the central United States; its northern range extends into South Dakota and eastward to southwest Illinois, while the southern range continues eastward to Louisiana and southward into San Luis Potosí and Zacatecas, Mexico.

Table 2. Stages in the monthly ovarian cycle of 46 adult female S. consobrinus from Oklahoma; * = one female with oviductal eggs and concomitant yolk deposition for a subsequent clutch. n

Quiescent

Early yolk deposition

Enlarged follicles > 4 mm

Oviductal eggs

March

3

3

0

0

0

April

15

6

6

0

3

Month

May

4

0

1

0

3

June

11

5

2

0

4*

July

13

6

1

3

3

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Table 3. Previous records in clutch sizes and reproductive period of S. consobrinus at different locations; * = based on two specimens.

Location

Clutch range

Multiple Clutches

Reproductive period

Colorado

8-12

Yes, 1-2

May to July

Source

Illinois

5-15

Not given

April to August

Phillips et al. 1999

Kansas

7

Yes, 2-3

May to August

Collins et al. 2010 Ballinger et al. 2010

Gillis and Ballinger 1992

Nebraska

4-6

Yes, 1-3

May into July

Nebraska

Not given

Not given

June into summer

Fogell 2010

Oklahoma

3-10

Yes

March into July

This paper

7

Yes, 2-3

May into July

Kiesow 2006

Texas*

6, 8

Not given

June

Smith 1946

Texas

Not given

yes

Spring through summer

Texas

7 to 9

Yes, 1-2

March into July

4-8

Yes, 2-3

May into July

South Dakota

Wyoming

into July is longer than that proposed (April to early May) by Lahti and Leaché (2009). Sceloporus consobrinus adheres to a reproductive strategy followed by 91% of North American lizards (Goldberg 2014) in which breeding occurs in spring, eggs are produced in late spring to summer; young emerge in late summer.

Previous clutch sizes from S. consobrinus by state are listed in Table 3. A clutch of three eggs from Oklahoma is a new minimum clutch size for S. consobrinus.

In view of the extensive range of S. consobrinus (Lahti and Leaché 2009), subsequent studies on S. consobrinus reproduction are warranted to ascertain geographic trends in the reproductive cycle.

Acknowledgments—I thank Cameron D. Siler (OMNH) for permission to examine S. consobrinus and Jessa A. Watters (OMNH) for facilitating the loan.

Literature Cited Ballinger, R.E., J.D. Lynch, and G.R. Smith. 2010. Amphibians and Reptiles of Nebraska. Rusty Lizard Press, Oro Valley, Arizona.

Collins, J.T., S.L. Collins, and T.W. Taggart. 2010.

Amphibians, Reptiles, and Turtles in Kansas. Eagle Mountain Publishing, Eagle Mountain, Utah.

Fogell, D.D. 2010. A Field Guide to the Amphibians and reptiles of Nebraska. Conservation and Survey Division, University of Nebraska, Lincoln.

Garrett, J.M., and D.G. Barker. 1987. A Field Guide to Reptiles and Amphibians of Texas. Texas Monthly Press, Inc. Austin.

Gillis, R., and R.E. Ballinger. 1992. Reproductive ecology of red-chinned lizards (Sceloporus undulates erythrocheilus) in Southcentral Colorado : comparisons

Garrett and Barker 1987 Vermersch 1992 Lewis 2011

with other populations of a wide-ranging species.

Oecologia 89:236-243.

Goldberg, S.R. 2014. Reproductive cycles of lizards from western North America. Sonoran Herpetologist 27:20-29.

Kiesow, A.M. 2006. Field Guide to Amphibians and Reptiles of South Dakota. South Dakota Department of Game, Fish and Parks. Pierre, South Dakota.

Lahti, M.E., and A.D. Leaché. 2009. Prairie lizard Sceloporus consobrinus Baird and Girard, 1853. Pages 210213 in: L.L.C. Jones and R.E. Lovich, eds. Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, Arizona, Leaché, A.D., and T.W. Reeder. 2002. Molecular systematics of the eastern fence lizard (Sceloporus undulatus): a comparison of parsimony, likelihood, and Bayesian approaches. Systematic Biology 51:44-68.

Lewis, D. 2011. A Field Guide to the Amphibians and Reptiles of Wyoming. The Wyoming Naturalist, Douglas, Wyoming.

Phillips, C.A., R.A. Brandon, and E.O. Moll. 1999.

Field Guide to Amphibians and Reptiles of Illinois.

Illinois Natural History Survey, Manual 8, Champaign.

Presnell, J.K., and M.P. Schreibman. 1997. Humason’s Animal Tissue Techniques. The Johns Hopkins Press, Baltimore.

Sievert, G., and L. Sievert. 2011. A Field Guide to Oklahoma’s Amphibians and Reptiles. Oklahoma Department of Wildlife Conservation, Oklahoma City.

Smith, H.M. 1946. Handbook of Lizards. Comstock Publishing Company, Ithaca, New York.

Vermersch, T.G. 1992. Lizards and Turtles of SouthCentral Texas. Eakin Press, Austin.

Four stages were present in the monthly ovarian cycle of S. consobrinus from Oklahoma: (1) quiescent, no yolk deposition; (2) early yolk deposition (basophilic vitellogenic granules in ooplasm); (3) enlarged ovarian follicles > 4 mm; (4) oviductal eggs. Female reproduction occurred from May into July.

Appendix: Sceloporus consobrinus from Oklahoma (by county) examined from the Oklahoma Museum of Natural History (OMNH), Norman.

Atoka County (n = 12) OMNH 38156-38158, 40090, 43028-43030, 43032-43034, 43036, 43037. Choctaw County (n = 5) OMNH 1825, 1894, 1896, 1898, 29554. Latimer County (n = 25) OMNH 10081-10085, 11600, 11602, 11604, 11606-11608, 11800-11807, 13208, 13210, 13211, 25711, 41606, 41608. Le Fiore County (n = 17) OMNH 16836, 38054, 38055, 41612, 41615-41618, 41664-41666, 41913-41917, 43041.

McCurtain County (n = 16) OMNH 1690, 1694, 1695, 1697, 29623, 30018, 30105, 30106, 30133, 30208, 30211, 38162, 38163, 38164, 38165, 40107. Pittsburg (n = 4) OMNH 38185-38188. Pushmataha County (n = 10) 8969, 8975, 11378, 23905, 28927, 28958, 44014-44016, 44018.

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RESEARCH ARTICLE

Effects of Buffelgrass Invasion on Density and Condition of Sonoran Desert Tortoises (Gopherus morafkai) Katherine M. Gray and Robert Steidl, The University of Arizona, School of Natural Resources and the Environment, Tucson, AZ. graykat@email.arizona.edu

H

umans have eliminated many biogeographic barriers that once isolated species, which has had profound ecological and economic consequences (D’Antonio and Vitousek 1992, Mack 2000).

Consequently, invasions by nonnative species threaten biodiversity worldwide (Hobbs and Mooney 2005), and have been identified as a threat to 50% of endangered species in the United States, the second most important cause of endangerment after habitat destruction and fragmentation (Wilcove et al. 2000).

Invasions by nonnative grasses are especially widespread and have had disproportionately strong effects on ecosystem structure and processes, especially fire (D’Antonio and Vitousek 1992, Lambrinos 2000, Litton et al. 2006). Buffelgrass (Pennisetum ciliare), a perennial, C4 bunchgrass native to Africa and Asia, was introduced to the southwestern United States in the 1930s and Mexico in the 1950s for erosion control and as cattle forage (Cox et al. 1998, Burquez-Montigo et al. 2002). Because of its ability to withstand hot and dry conditions, buffelgrass has invaded southwestern deserts, creating a dense grass layer in areas where ground cover was sparse historically (Rogstad et al. 2009). The marked increase in biomass and vegetation structure in areas invaded by buffelgrass is likely to increase frequency, size, and intensity of fire in those areas (Franklin et al. 2006, Franklin and Morina-Freaner 2010, McDonald and McPherson 2011). Additionally, buffelgrass can form dense stands and exclude native plants, potentially eliminating species that provide food and cover for native herbivores, and decrease open space, which could affect habitat quality for native animals (Burquez-Montijo et al. 2003, Olsson 2012a).

Sonoran Desert Tortoises (Gopherus morafkai) occur throughout southern Arizona and northwestern Mexico (Lamb et al. 1989, Barrett and Johnson 1990), a region where the distribution of buffelgrass has increased markedly in recent years (Franklin et al.

2006, Marshall et al. 2012). Recently, Sonoran Desert Tortoises were declared warranted for listing under the Endangered Species Act and are listed as a species of

Figure 1. One of fifty study plots invaded by buffelgrass. Saguaro National Park (Rincon District). Photo by Katherine Gray.

special concern by the Arizona Game and Fish Department (Arizona Game and Fish Department 1996, U. S. Fish and Wildlife Service 2010). Invasion by nonnative plants was identified as the most significant modification to habitat of Sonoran Desert Tortoises, and is thought to contribute to declines in tortoise populations (U. S. Fish and Wildlife Service 2010).

Changes in vegetation cover and structure resulting from invasions by nonnative plants can affect animal populations in multiple ways, including altering demography, distribution, and reproductive success (Litt and Steidl 2011). Most research on effects of buffelgrass invasion has focused on plants, with relatively little focus on animals (Marshall et al. 2012).

To evaluate effects of this invasion on desert tortoises, we characterized demography and physical condition of tortoises along a gradient of buffelgrass invasion in areas of high-quality habitat for tortoises. Our goal was to assess current effects of buffelgrass on density and condition of this rare, long-lived vertebrate to better understand potential future impacts as buffelgrass expands in distribution.

Humans have eliminated many biogeographic barriers that once isolated species, which has had profound ecological and economic consequences.

Consequently, invasions by nonnative species threaten biodiversity worldwide, and have been identified as a threat to 50% of endangered species in the United States.

Study Design We studied tortoises in southeastern Arizona, in both the Rincon and Tucson Mountain districts of Saguaro National Park. In 2010 and 2011, we identified areas with environmental features characteristic

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of high-quality habitat for desert tortoises, including areas with slopes between 20 and 30°, aspects between 150 and 260°, and elevations between 750 and 1200 m (Zylstra and Steidl 2009). We established 50 plots along a gradient of invasion by buffelgrass that were based on existing maps of buffelgrass distribution and field reconnaissance, ranging from locations with no buffelgrass to locations where buffelgrass was known to occur in relatively high densities. Plots were 4 ha (200 × 200 m), reflecting a Figure 2. An adult female Desert Tortoise eating buffelgrass at Sagauro National Park (Tucson District). Photo taken by Katherine Gray. compromise between the home-range size of desert tortoises (2-34 ha; Barrett 1990) and the amount of Berry and Christopher 2001): time required to survey a plot effectively. We surveyed mass /(length*width*height) 25 plots in 2010 and 25 different plots in 2011, 15 in the Tucson Mountains and 35 in the Rincon MounWe also quantified environmental covariates that tains, in southern Arizona (Fig. 1). could potentially explain the parameters we were interested in, such as the number of potential shelter sites, Methods topography, and vegetation.

We surveyed tortoises on each plot 4 times between Results and Discussion early July and October, a period that coincides with the monsoon season when tortoises are most active We generated a condition index for 131 adult desert (Woodbury and Hardy 1948). We surveyed during tortoises located across the gradient of buffelgrass mornings (0500-1000) and evenings (1600-2000), cover and found that condition of adults decreased alternating morning and evening surveys between by up to 10% as cover of buffelgrass increased, after successive surveys on each plot, which were separated accounting for sex. Densities of tortoises were not by at least 7 days. Across each plot, we established an affected by buffelgrass cover and were positively corarray of twelve 200-m parallel transects (2400 m total) related with the number of shelter sites. Reductions spaced at 16-m intervals to reflect the area within which most tortoises were detected in previous studies in body condition we observed in tortoises could be a of tortoises in the Sonoran Desert (Swann et al. 2002). result of reductions in the quantity or quality of forage plants in areas with relatively high buffelgrass cover.

On each survey, 2 to 3 surveyors aided by flashlights We found that cover of plants important to tortoises and mirrors, traversed transects and searched for for food decreased in areas invaded by buffelgrass, tortoises in the open, under vegetation, or in rock including grasses, desert vine, and prickly pear. Reduccrevices. We recorded the perpendicular distance from tions of succulents and forbs in areas invaded by the transect line to each tortoise encountered. buffelgrass, which are also important components in For each tortoise encountered, we classified age as diets of desert tortoises (Oftedal et al. 2002), also have adult (midline-carapace length [MCL] ≥180 mm) or been recorded in areas invaded by buffelgrass in the juvenile (MCL<180 mm), and classified sex of adults Sonoran Desert (Olsson et al. 2012). A reduction in based on concavity of the plastron, length of gulars, species diversity or abundance of forage plants could length of tail, and presence of swollen chin glands have adverse consequences on condition, because (Berry and Christopher 2001). We used a spring scale tortoises forage selectively due to their inability to to measure body mass (g), and used a dial caliper and excrete excess potassium (Oftedal et al. 2002). Also, tape measure to measure MCL, carapace width and home-range sizes of desert tortoises increase durheight, and width and length of plastron. To evaluate ing dry years in response to decreased availability of physical condition of the adult tortoises we captured, forage plants (Averill-Murray and Klug 2000); therewe generated an index of condition (Wallis et al. 1999;

SONORAN HERPETOLOGIST 27 (4) 2014

Densities of tortoises were not affected by buffelgrass cover and were positively correlated with the number of shelter sites.

Reductions in body condition we observed in tortoises could be a result of reductions in the quantity or quality of forage plants in areas with relatively high buffelgrass cover.

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fore, greater energy expenditures may be required by tortoises foraging in areas with high buffelgrass cover.

Site tenacity of desert tortoises in areas where forage has been reduced in response to buffelgrass could explain the reduced condition we observed in areas of relatively high buffelgrass cover, which could become more severe over time. Continued declines in condition could result in compromised immune systems, lower reproduction, lower growth rates, and ultimately death (Oftedal et al. 2002). Similar results have been observed in other species, where reproductive success and survival have been reduced in areas invaded by nonnative species, although animals continue to settle in and occupy these areas (Schmidt and Whelan 2000, Remes 2003, Lloyd and Martin 2005). Because desert tortoises are long-lived and site tenacious (O’Connor et al. 1994, Duda et al. 1999, Freilich et al. 2000), changes in plant community structure and composition in response to buffelgrass invasion may not cause sudden mortality nor trigger abandonment which could explain why density of tortoises did not vary across the gradient of buffelgrass we studied.

Density of tortoises in areas we surveyed with >15% cover of buffelgrass reflected densities of tortoises in areas where buffelgrass was absent, and also reflected estimates of tortoises reported from previous studies in this region (Swann et al. 2002, Averill-Murray and Averill-Murray 2005, Zylstra and Steidl 2010).

Within the areas we surveyed, tortoise density varied only with the number of potential shelter sites, which has been reported to influence density of tortoises in the Sonoran Desert consistently (Fritt and Jennings 1994, Averill-Murray et al. 2002, Reidle et al. 2008) and is likely a limiting factor for tortoise populations (Woodbury and Hardy 1948, McGinnis and Voigt 1971, Bailey et al. 1995, Rautenstrauch et al. 2002).

Sonoran Desert Tortoises use rock crevices for nesting, protection from predators and thermal extremes, and spend about 98% of their lives in these shelters (Nagy and Medica 1986). Because shelter sites are such an important factor in dictating carrying capacity and are not altered by the invasion of buffelgrass, potential effects of changes in the vegetation community on tortoises may be limited, at least in the initial stages of invasion.

Although we focused on the relatively immediate effects of buffelgrass on tortoises, buffelgrass has potential to influence tortoises over longer time horizons. Buffelgrass creates high fuel loads in areas that were once predominantly free of ground cover, which can increase the frequency, intensity, and extent of wildfires in areas where wildfire has been rare historically (McLaughlin and Bowers 1982, Brooks and Pyke 2002). If tortoises persist in areas invaded by buffelgrass, over the long term they likely will be exposed to more frequent wildfires that can cause high mortality.

For example, up to 11% of a population was estimated to have perished from the 1994 Mother’s Day fire

in the Rincon Mountains of Saguaro National Park (Esque et al. 2003). Wildfires might also perpetuate the spread of buffelgrass, as it thrives in post-fire conditions whereas native Sonoran desert vegetation does not (McDonald and McPherson 2011).

Acknowledgments—The project was made possible through the financial support of the Desert Southwest Ecosystem Cooperative Studies Unit, Friends of Saguaro National Park, T&E Incorporated, Tucson Herpetological Society, and Western National Parks Association. We thank Dana Backer and Don Swann from Saguaro National Park for their expertise on buffelgrass and desert tortoises, and their support during all aspects of the project. We appreciate the financial and administrative report (and patience) of Larry Norris of the Desert Southwest Ecosystem Cooperative Studies Unit. Joshua Ernst, Chris Magionami, Erin Zylstra, Bruce Weise, Steve Campbell, staff of Saguaro National Park, and many others who helped us in the field and contributed to our efforts in many others ways. Kevin Bonine and Bill Matter from the University of Arizona provided constructive reviews on an earlier draft of this report.

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2002. Distance sampling for Sonoran Desert tortoises. Journal of Wildlife Management 66:969-975.

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RESEARCH ARTICLE

Herpetofauna of the Canelo Arc

Robert L. Bezy, Natural History Museum of Los Angeles County, Los Angeles, CA 90007, USA; robertbezy@gmail.com Charles J. Cole, American Museum of Natural History, New York, NY 10024, USA This paper is dedicated to our friend and colleague, Erik F. Enderson, who introduced Kit Bezy and Bob Bezy to the Canelo area on 3 July 2000. During the ensuing decade, many wonderful days and nights were spent together observing and photographing the fauna and flora of the Canelos, Huachucas, and Patagonias. Erik’s magnificent images of the amphibians and reptiles in the region enliven these pages.

Situated in the 500 km gap between the northern terminus of the Sierra Madre Occidental and the

Mogollon Rim is a series of isolated mountain ranges known as the Madrean Archipelago (Lowe 1992). In Arizona and New Mexico, the 22 Madrean ranges appear to form three groups based on similarities among the herpetofaunas documented from montane biotic communities (Bezy and Cole 2014). The Canelo Hills were not analyzed as a separate range in that paper and here we focus on the Canelo arc of Madrean Evergreen Woodland that links the Huachuca Mountains of the southeastern group of ranges with the Patagonia Mountains of the southwestern group (Fig. 1).

For more than a century, the herpetofauna of the Huachuca Mountains has attracted attention (e.g., Stejneger 1902, Woodin 1953, Morrison et al. 1993). To the west, the fauna and flora of the Patagonia Mountains was surveyed recently (MABA 2013) and a list of the amphibians and reptiles of the Sonoita Creek area was published by Turner (2007). A thorough study of the Patagonia herpetofauna has yet to be conducted

and the amphibians and reptiles in the Canelo Hills linking these areas remain largely undocumented.

The Canelo Hills form a ridge extending ca 32 km from near Mount Hughes (1786 m elev., 31.6053°N, 110.68147°W, ca 8 km S Sonoita) southeast through Lookout Knoll (1880 m, 31.48275°N, 110.54053°W) to approximately Parker Canyon (1645 m, 31.42236°N, 110.45991°W) near the border between Santa Cruz and Cochise Counties, Arizona (Fig. 1). They are narrowly separated from the Santa Rita Mountains to the northwest by Sonoita Creek and merge with the Patagonia Mountains southwest of Corral Canyon and with the Huachuca Mountains southeast of Lyle Canyon. Although Brown and Lowe (1980) map the Canelo Hills as having a broad band of uninterrupted Madrean Evergreen Woodland (Figs. 1-3), our casual observations suggest that there is less woodland and more grassland (Semi-desert Grassland and Plains and Great Basin Grassland) than indicated by their coarsescale map.

We use “Canelo Hills” for the area as defined above in terms of topography and “Canelo arc,” or “the arc,”

SONORAN HERPETOLOGIST 27 (4) 2014

Situated in the 500 km gap between the northern terminus of the Sierra Madre Occidental and the Mogollon Rim is a series of isolated mountain ranges known as the Madrean Archipelago.

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for that part of the hills supporting Madrean Evergreen Woodland (=“woodland”).

Records of museum vouchers (specimens and images) for the arc demonstrate collecting and observational bias and are clustered in the northwest near Papago Springs (Fig. 2) and in the southeast near Canelo (Fig. 3). The records document a total of 22 species of amphibians and reptiles in Madrean Evergreen Woodland: 3 anurans (frogs and toads), 1 turtle, 6 lizards, and 12 snakes (Appendix 1). The arc shares a greater number of species with the woodlands of the high profile Huachuca (21) and Santa Rita (19) Mountains than with those of the low profile Patagonia Mountains (13). Clustering of similarities (Simpson’s S, UPGMA) among the herpetofaunas documented in woodland of the U.S. Madrean ranges (Bezy and Cole 2014) places the arc nearest the Huachuca Mountains with which they are connected via topography and biotic community. Note that every species documented for the arc also occurs in the woodland of the Huachuca Mountains. The one exception is the Yaqui Black-headed Snake (Tantilla yaquia) vouchered from the Canelo arc and the Santa Ritas but not the Huachucas.

The documented herpetofauna of the Canelo arc (Appendix 1) is composed primarily of species common to most Madrean ranges (e.g., Lithobates chiricahuensis, Chiricahua Leopard Frog, Fig. 4; Phryno-

soma hernandesi, Greater Short-horned Lizard, Fig. 5; Crotalus molossus, Black-tailed Rattlesnake, Fig. 6), with the exception of Hyla wrightorum (Arizona Treefrog, Fig. 7), Crotalus lepidus (Rock Rattlesnake, Fig. 8), and C. willardi (Ridge-nosed Rattlesnake) which have more restricted distributions in the area. Also present in the Canelo Hills region mapped by Brown and Lowe as Madrean Evergreen Woodland, but considered by us in on-site inspection to be grassland are: Scaphiopus couchii (Couch’s Spadefoot), Aspidoscelis uniparens (Desert Grassland Whiptail), Coluber flagellum (Coachwhip), Crotalus scutulatus (Mohave Rattlesnake), and Heterodon kennerlyi (Mexican Hog-nosed Snake).

Two species of montane rattlesnakes occur in the arc, Crotalus willardi near the Huachuca Mountains and C. lepidus near Sonoita. It would be of interest to compare genetic markers of the populations in the arc with those in the Santa Rita and Huachuca Mountains to clarify their historical relationships.

Hyla wrightorum (Fig. 7) is the only species of Madrean herpetofauna in southern Arizona that is uniquely shared by the Huachuca Mountains and the Canelo arc (Bezy et al. 2010). The frogs in these two ranges share a distinct mtDNA haplotype, but overall differentiation among the Mogollon Rim, HuachucaCanelo, and Sierra Madre Occidental populations is low for allozymes, mitochondrial DNA, and call parameters, suggesting that the three have become iso-

For more than a century, the herpetofauna of the Huachuca Mountains has attracted attention.

To the west, the fauna and flora of the Patagonia Mountains was surveyed recently (MABA 2013) and a list of the amphibians and reptiles of the Sonoita Creek area was published by Turner (2007).

Figure 1. Map of the Canelo Hills and adjacent ranges. Green denotes Madrean Evergreen Woodland and blue indicates Petran Montane Conifer Forest based on the map of biotic communities by Brown and Lowe (1980) as digitized by The Nature Conservancy in Arizona.

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lated relatively recently (Gergus et al. 2004). Considering the low level of differentiation and the occurrence at Turkey Creek, additional relictual populations of H. wrightorum may perhaps exist elsewhere in the arc.

The distribution of Sceloporus slevini (Slevin’s Bunchgrass Lizard) is enigmatic (Bezy 2004). It occurs in bunchgrass at high elevations in the Whetstone (Turner et al. 2003), Chiricahua, Huachuca, and Santa Rita Mountains, as well as at lower elevation in the grasslands of the San Rafael Valley, Sonoita Plain, and Empire Cienega. The species has been documented near the arc in the Huachuca Mountains, and recently was observed near Canelo Pass (Cole, pers. obs.).

Gastrophryne mazatlanensis (Mazatlan Narrowmouthed Toad) has been well-documented in the woodland of the Pajarito Mountains (Enderson 2002).

Lowe (1964) listed Gastrophryne as also occurring in the Patagonia Mountains, but no vouchers are available from the range. The listing may have been based on Campbell’s (1932) report of hearing the toads calling northeast of Patagonia. Searches have been made during the monsoons for narrow-mouthed toads in the Canelo arc but with no success to date. They may have occurred in the northern arc in the late Pleistocene (ca 42,000 ybp), as Czaplewski et al. (1999) tentatively referred a fossil humerus from Papago Springs Cave to Gastrophryne. Recently, Turner (2007) listed Gastrophryne for the Sonoita Creek area, increasing the likelihood that they eventually may be discovered in the distributional gap to the northeast. More than one species of Gastrophryne may occur in Arizona (Lowe 1964) and additional searches in the arc, as well as comparisons of mitochondrial and nuclear DNA sequences (e.g., Streicher et al. 2012) and morphology for all populations in the state would be of interest.

Although our discussion of the herpetofauna of Canelo arc is based on species, Ambystoma mavortium stebbinsi (Sonoran Tiger Salamander) is of considerable

interest due to its precarious genetic situation (Jones et al. 1988, 1995, Storfer et al. 2004). Individuals clearly assignable to this native subspecies on the basis of morphology and genetic markers persist primarily in the grasslands of the San Rafael Valley, but historically may have extended eastward to the Madrean Evergreen Woodland of the Canelo Hills. Fossil Ambystoma (tentatively referred to A. tigrinum) dating from the Late Pleistocene have been recovered from Papago Springs Cave (Czaplewski et al. 1999).

Many species found in the woodlands of the Huachuca and Santa Rita Mountains remain undocumented from the arc: Craugastor augusti (Barking Frog), Hyla arenicolor (Canyon Treefrog), Spea multiplicata (Mexican Spadefoot), Plestiodon callicephalus (Mountain Skink), Sceloporus jarrovii (Yarrow’s spiny Lizard), Sceloporus slevini (Slevin’s Bunchgrass Lizard), Lampropeltis pyromelana (Sonoran Mountain Kingsnake), and Tantilla wilcoxi (Chihuahuan Black-headed Snake).

It is notable that two species, Gyalopion quadrangulare (Thornscrub Hook-nosed Snake) and Oxybelis aeneus (Brown Vinesnake), found in semi-tropical habitats and extending north into the Pajarito and Patagonia Mountains and Sonoita Creek area (Van Devender et al. 1994, Turner 2007) are not known from the Canelo arc. Another snake wide-spread at lower latitudes and found in both the Patagonia and Santa Rita Mountains, Senticolis triaspis (Green Ratsnake), has not been documented from the arc nor from the Whetstone, Dragoon, Huachuca, and Mule Mountains.

For any of these species, it is doubtful that enough field work has been done in the Canelo arc to conclude with confidence that they do not occur there, especially as it would seem that they probably occurred there when the Madrean Evergreen Woodland may have been more widely distributed in the area several thousand years ago. If some of these species are truly absent from the arc, it might be because recently they

Clearly, the herpetofauna of the Canelo arc and the entire Madrean region merits additional field work yielding precisely georeferenced data for species occurrences and GIS-based analyses with fine scale vegetation layers.

Figure 2. Madrean Evergreen Woodland near Papago Springs, Santa Cruz Co., Arizona. Photo by Carol Townsend.

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Figure 3. Madrean Evergreen Woodland near Canelo Pass, Santa Cruz Co., Arizona. Photo by Carol Townsend.

Figure 4. Lithobates chiricahuensis (Chiricahua Leopard Frog) in the Canelo Hills. Photo by Erik F. Enderson.

were extirpated from natural causes, even if random.

Clearly, the herpetofauna of the Canelo arc and the entire Madrean region merits additional field work yielding precisely georeferenced data for species occurrences and GIS-based analyses with fine scale vegetation layers. These would facilitate the evaluation of ecological and evolutionary factors underlying montane species richness (see discussion in Graham et al. 2014). Ongoing inventories by MABA are laying a foundation for such an assessment.

At this point we can only speculate that the

Madrean Evergreen Woodland of the Canelo arc may have provided an important historical biogeographic connection between the Huachuca, Santa Rita, and Patagonia Mountains. Unfortunately, their herpetofauna remains poorly documented and the area often is viewed as uninteresting foothills traversed on the way to the higher profile Madrean ranges.

Acknowledgmentsâ&#x20AC;&#x201D;We thank Kit Bezy, Kathryn Bolles, Erik Enderson, and Carol Townsend for participation in field work; Erik Enderson and Carol

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Townsend for photography; Kit Bezy for illustration; and Kit Bezy, Kathryn Bolles, and Erik Enderson for helpful suggestions on a previous version of this paper.

The Nature Conservancy in Arizona provided the very useful digitized version of the Brown and Lowe (1980) map of biotic communities. We thank Jim Rorabaugh for bringing some records to our attention.

R.C.K. Bowie, C. Moritz, S.B. Baines, C.J. Schneider, J. VanDerWal, C. Rahbek, K.H. Kozak, and N.J. Sanders. 2014. The origin and maintenance of montane diversity: integrating evolutionary and ecological processes. Ecography 37:001-009.

Jones, T.R, J.P. Collins, T.D. Kocher, and J.B. Mitton.

1988. Systematic status and distribution of Ambystoma tigrinum stebbinsi Lowe (Amphibia: Caudata).

Copeia 1988:621-635.

Literature Cited Jones, T.R., E.J. Routman, D.J. Begun, and J.P. ColBezy, R.L. 2004. Slevinâ&#x20AC;&#x2122;s Bunchgrass Lizard Sceloporus lins. 1995. Ancestry of an isolated subspecies of slevini (Smith, 1966). Sonoran Herpetologist 17:62salamander, Ambystoma tigrinum stebbinsi Lowe: the 63. evolutionary significance of hybridization. MolecuBezy, R.L., and C.J. Cole. 2014. Amphibians and reptiles lar Phylogenetics and Evolution 4:194-202. of the Madrean archipelago of Arizona and New Lowe, C.H. 1964. Amphibians and reptiles of Arizona.

Mexico. American Museum Novitates 3810:1-23.

Pages 153-174 in C.H. Lowe (editor), The verteBezy, R.L., K.B. Bezy, K. Bolles, and E.F. Enderson. brates of Arizona. University of Arizona Press, 2011. Arizona Treefrog, Hyla wrightorum Taylor, Tucson.

1939. Sonoran Herpetologist 24:110-114.

Lowe, C.H. 1992. On the biogeography of the herBrown, D.E., and C.H. Lowe. 1980. Biotic communities petofauna of Saguaro National Monument. Pages of the Southwest. General Technical Report, Rocky 91-103 in C.P. Stone and E.S. Bellantoni (editors), Mountain Forest and Range Experiment Station, Proceedings of the symposium on research in SaUSDA Forest Service RM-78. guaro National Monument, Tucson, AZ: CooperaCzaplewski, J.A., J.I. Mead, C.J. Bell, W.D. Peachey, and tive Park Studies Unit, University of Arizona.

T.-L. Ku. 1999. Papago Springs Cave revisited, Part MABA. 2013. Madrean Archipelago Biodiversity AsII: vertebrate paleofauna. Occasional Papers of the sessment: Patagonia bioblitz. http://bit.ly/1rPWcCI Oklahoma Museum of Natural History 5:1-51.

Morrison, M.L., W.M. Block, L.S. Hall, and H.S. Stone.

Enderson, E.F. 2004. Great Plains Narrow-Mouthed 1995. Habitat characteristics and monitoring of amToad (Gastrophryne olivacea). Sonoran Herpetologist phibians and reptiles in the Huachuca Mountains, 15:80.

Arizona. Southwestern Naturalist 40:185-192.

Gergus, E.W., T.W. Reeder, and B.K. Sullivan. 2004.

Stejneger, L. 1902. The reptiles of the Huachuca Geographic variation in Hyla wrightorum: advertiseMountains, Arizona. Proceedings of the United ment calls, allozymes, mtDNA, and morphology.

States National Museum 25:149-158.

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Storfer, A., S.G. Mech, M.W. Reudink, R.E. Ziemba, Graham, C.H, A.C. Carnaval, C.D. Cadena, K.R. J. Warren, and J.P. Collins. 2004. Evidence for Zamudio, T.E. Roberts, J.L. Parra, C.M. McCain, introgression in the endangered Sonora Tiger Sala-

Figure 5. Phrynosoma hernandesi (Greater Short-horned Lizard) in the Canelo Hills. Photo by Erik F. Enderson.

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mander, Ambystoma tigrinum stebbinsi (Lowe). Copeia 2004:783-796.

Streicher, J.W., C.L. Cox, J.A. Campbell, E.N. Smith, and R.O. de Sรก. 2012. Rapid range expansion in the Great Plains Narrow-mouthed Toad (Gastrophryne olivacea) and a revised taxonomy for North American microhylids. Molecular Phylogenetics and Evolution 64:645-653.

Turner, D.S. 2007. Amphibians and reptiles of the Sonoita Creek State Natural Area, Arizona. Sonoran Herpetologist 20:38-42.

Turner, D.S., P.A. Holm, E.B. Wirt, and C.R. Schwalbe.

2003. Amphibians and reptiles of the Whetstone Mountains, Arizona. Southwestern Naturalist 48:347-355.

Van Devender, T.R., C.H. Lowe, and H.E. Lawler.

1994. Factors influencing the distribution of the Neotropical vine snake Oxybelis aeneus in Arizona and Sonora, Mexico. Herpetological Natural History 2:27-44.

Woodin, W.H. 1953. Notes on some reptiles from the Huachuca area of southeastern Arizona. Bulletin of the Chicago Academy of Science 9:285-296.

Figure 6. Crotalus molossus (Black-tailed Rattlesnake) in the Canelo Hills. Photo by Erik F. Enderson.

Figure 7. Hyla wrightorum (Arizona Treefrog) in the Huachuca Mountains. Photo by Erik F. Enderson.

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Figure 8. Crotalus lepidus (Rock Rattlesnake) in the Huachuca Mountains. Photo by Erik F. Enderson.

Appendix 1. Occurrence of native species of amphibians and reptiles in the Madrean Evergreen Woodland of the Canelo Hills and the Huachuca, Santa Rita, and Patagonia Mountains, Arizona, based on vouchers (specimens and images) in museum collections.

Taxon

English Name

Anura

Frogs and Toads

Anaxyrus punctatus

Red-spotted Toad

Anaxyrus woodhousii

Woodhouseâ&#x20AC;&#x2122;s Toad

Canelo X

Huachuca

Santa Rita

X

X

Patagonia

X

Craugastor augusti

Barking Frog

X

X

X

Hyla arenicolor

Canyon Treefrog

X

X

X

Hyla wrightorum

Arizona Treefrog

Incilius alvarius

Sonoran Desert Toad

Lithobates chiricahuensis

Chiricahua Leopard Frog

Lithobates tarahumarae

Tarahumara Frog

Lithobates yavapaiensis

Lowland Leopard Frog

Spea multiplicata

Mexican Spadefoot

Testudines

Turtles

Kinosternon sonoriense

Sonora Mud Turtle

Squamata

Lizards

Aspidoscelis sonorae

Sonoran Spotted Whiptail

X

X X

X

X

X X X

X X

X

X

X

X

X

X X

Aspidoscelis stictogramma

Giant Spotted Whiptail

X

Aspidoscelis tigris

Tiger Whiptail

X

Coleonyx variegatus

Western Banded Gecko

Crotaphytus collaris

Eastern Collard Lizard

X X

Elgaria kingii

Madrean Alligator Lizard

X

X

X

Holbrookia elegans

Elegant Earless Lizard

X

X

X

X

Phrynosoma hernandesi

Greater Short-horned Lizard

X

X

X

X

Phrynosoma solare

Regal Horned Lizard

X

Plestiodon callicephalus

Mountain Skink

X

X

Plestiodon obsoletus

Great Plains Skink

X

X

X

X

Sceloporus clarkii

Clarkâ&#x20AC;&#x2122;s Spiny Lizard

Sceloporus cowlesi

Southwestern Fence Lizard

X

X X

X

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Taxon

English Name

Sceloporus jarrovii

Yarrow’s Spiny Lizard

Sceloporus magister

Desert Spiny Lizard

Sceloporus slevini

Slevin’s Bunchgrass Lizard

Urosaurus ornatus

Ornate Tree Lizard

Squamata

Snakes

Coluber bilineatus

Sonoran Whipsnake

Crotalus atrox

Western Diamond-backed Rattlesnake

Canelo

Huachuca

Santa Rita

Patagonia

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

Crotalus lepidus

Rock Rattlesnake

X

X

X

Crotalus molossus

Black-tailed Rattlesnake

X

X

X

Crotalus pricei

Twin-spotted Rattlesnake

X

X

Crotalus willardi

Ridge-nosed Rattlesnake

X

X

X

Diadophis punctatus

Ring-necked Snake

X

X

Gyalopion canum

Chihuahuan Hook-nosed Snake

Gyalopion quadrangulare

Thornscrub Hook-nosed Snake

Hypsiglena chlorophaea

Desert Nightsnake

Lampropeltis pyromelana

Sonoran Mountain Kingsnake

Micruroides euryxanthus

Sonoran Coralsnake

Oxybelis aeneus

Brown Vinesnake

Pituophis catenifer

Gophersnake

X

X X

X

Rena dissectus

New Mexico Threadsnake Eastern Patch-nosed Snake

X

X

Salvadora hexalepis

Western Patch-nosed Snake

X

X

Senticolis triaspis

Green Ratsnake

Sonora semiannulata

Western Groundsnake Smith’s Black-headed Snake

X X

X

X

X

X

X

X

X

X

X

X

X

X X

X

Salvadora grahamiae

Tantilla hobartsmithi

X

X

X X

X

X

X

X

X

X X

Tantilla wilcoxi

Chihuahuan Black-headed Snake

Tantilla yaquia

Yaqui Black-headed Snake

X

Thamnophis cyrtopsis

Black-necked Gartersnake

X

X

Thamnophis eques

Mexican Gartersnake

X

X

Trimorphodon lambda

Sonoran Lyresnake

X

X

X

X

X

X

X

X

BOOK REVIEW

The Biology of Chameleons Review by Russell Ligon, Arizona State University

A s a biologist who has spent the last few years studying chameleon behavior, I had high hopes for the

beautiful new book, The Biology of Chameleons, edited by Krystal A. Tolley and Anthony Herrel. These hopes were, in almost every instance, exceeded. For many authors the work and information described are clearly labors of love, and the passion and knowledge of all of the authors of the 10 chapters that comprise this book come through beautifully, thanks in large part to the organization and editing by the book’s editors. I had a few minor issues (noted below), but on the whole I am confident that this book will become a valuable addition to the library of any scientist or lay-person with a dedicated interest in the biology of this strange and wonderful group of animals. Below, I

The Biology of Chameleons Krystal A. Tolley and Anthony Herrel (Editors) Hardcover, 288 pages ISBN: 9780520276055 November 2013 $65.00

SONORAN HERPETOLOGIST 27 (4) 2014

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have summarized the chapters included in this book to provide an overview for potential readers.

Chapter 1, “An Introduction” by Krystal A. Tolley and Anthony Herrel, sets the tone for this wonderful volume dedicated to increasing knowledge and making a trove of information available to a wider audience.

Chapter 2, “Chameleon Anatomy” by Christopher V. Anderson and Timothy E. Higham, provides a thorough description of chameleon skeletal, muscular, and anatomical features studied to-date. For nonanatomists, there is a lot of information to get through and, whether because more functional work has been completed on tongue extension or because it is one of Anderson’s specialties, the descriptions of the roles of different tongue muscles used in feeding and prey capture was much more approachable and engaging for me than some of pure descriptions of muscle groups and skeletal elements. This was true throughout the chapter – wherever functional explanations accompanied morphological descriptions I found I was better able to follow along and learn something new. One of the more interesting facts I gleaned from the chapter was the fact that chameleons exhibit a dramatic range in the number of caudal vertebrae they possess, ranging from 17 to 62 vertebrae! Though this was a very good chapter, I was disappointed when I tried to find the original source of information cited regarding the density of cones in chameleon retinas. In the chapter, the source of this information is cited from more recent sources which reference the older, original work (instead of citing the original work).

Chapter 3, “Chameleon Physiology” by Anthony Herrel, provides a nice mechanistic complement to the preceding chapter on morphology and anatomy.

This chapter is very well-written and easy to read, with descriptions of physiological phenomena that were easy to understand and interpret (though there was no introduction to some basic physiological terminology like “Q10”). Generally, this chapter provides the reader with a solid understanding of how chameleons are likely to sense the world, with clear descriptions of their visual and auditory systems. As a chameleon biologist, I did find something to quibble about regarding Herrel’s description of the factors controlling color change in chameleons. Herrel states that current consensus is that chameleon color change is under control of nervous system yet he excludes mention of a key study by Okelo (1986) definitively demonstrating that color change can be induced hormonally. Additionally, there a numerous references to stress-induced color change throughout chameleon husbandry literature, further suggesting the longer term hormonal control over color change in chameleons. This is probably a minor issue, but one worth mentioning.

Chapter 4, “Function and Adaptation of Chameleons” by Timothy E. Higham and Christopher V. Anderson, is an oddly titled (what is the function of a chameleon?) chapter about the functional adapta-

tions of chameleons. This chapter covers locomotory adaptations and those that function in feeding. In the first subsection on locomotion, focused on limb kinematics, the authors do a masterful job explaining the different ways that chameleon limbs contribute to their movement patterns and how these movements fit their specialized ecological niche. One of the key adaptations facilitating the thin-branch arboreality of chameleons is their fused digits, which allow chameleons to generate strong adduction forces on branches that consequently enable them to pull their bodies forward with their hind limbs in a manner unlike any terrestrial lizard. The section on mechanisms of projectile tongue use in prey capture was very thorough but, in my humble opinion, certainly could have benefitted by the inclusion of additional figures/images to illustrate key processes.

Chapter 5, “Ecology and Life History of Chameleons” by G. John Measey, Achille Raselimanana and Anthony Herrel, begins with detailed descriptions of habitat use among chameleons. Interestingly, field work on chameleons is so difficult that many inferences regarding where chameleons live and forage are apparently based on their nocturnal roosting locations, where they are easier for scientists to locate. Consequently, at the moment, the data required to split forest chameleons into more than two broad guilds (arboreal and leaf-litter) are lacking. In the section on chameleon life-history I learned that chameleon body size covers four orders of magnitude (across all species).

This chapter was interspersed with well-described case studies illustrating key points of chameleon ecology. These case studies/illustrative boxes were so effective and clear, I wish they were a part of every chapter in this book!

Chapter 6, “Chameleon behavior and color change”

by Devi Stuart-Fox gives a summary of the state of knowledge regarding the mechanisms of color change, stating that the control of color change is “poorly understood” but likely includes “a range of neurotransmitters and hormones…” Additionally, Stuart-Fox provides a very clear and easy to understand summary of the likely factors favoring the evolution of color change in chameleons, which makes sense because the work she summarizes is her own! The chapter also includes concise, yet comprehensive, reviews of chameleon territoriality, sperm storage, female reproduction tactics, male aggressive behaviors, sexual size dimorphism, with a larger section on anti-predator behaviors of these incredible animals. Overall, this was a fascinating and well-written chapter that made it easy for readers to grasp the current state knowledge regarding chameleon social behavior, as well as the control and numerous functions of rapid color change. I wish this had been available when I first began studying chameleons and I have no doubt it will be a boon to all future chameleon researchers.

Chapter 7, “Evolution and Biogeography of Cha

SONORAN HERPETOLOGIST 27 (4) 2014

Chapter 3, “Chameleon Physiology” by Anthony Herrel, provides a nice mechanistic complement to the preceding chapter on morphology and anatomy.

This chapter is very well-written and easy to read, with descriptions of physiological phenomena that were easy to understand and interpret.

103

meleons” by Krystal A. Tolley and Michele Menegon, provides a really nice complementary evolutionary and historical perspective regarding how and when chameleons came to be. What makes this chapter stand out to me was the detailed background Tolley and Menegon provide regarding climate and environmental events occurring during various stages of chameleon evolutionary history - they put splitting and diversification events into a logical and understandable context.

In addition to describing chameleon evolutionary events, this chapter also covers current chameleon diversity and distribution. Furthermore, this chapter describes locations and ranges of known chameleon groups: however, these sections could also serve as a roadmap for intrepid explorers who want to discover new chameleons species. Regions of great biodiversity, in addition to being prime locations for the discovery of new species, are also conservation hotspots- though this is a topic that Tolley and Menegon leave for another chapter. Unfortunately, as a non-biogeographer the sections labeled alpha- and beta-diversity did not help me, but the information included in those sections was still interesting and readily understandable.

Chapter 8, “Overview of Systmatics of the Chameleonidae” by Colin R. Tilbury, covers the systematics of chameleons. Tilbury begins with an interesting historical perspective of chameleon taxonomy and methodology, as well as a concise summary of the big-picture taxonomic issues (e.g., number of species, genera, changes over time) before delving into specific sub-familial groupings. These sections provide incredible details about the chameleons and will doubtless

prove to be a valuable resource and starting point for future comparative analyses.

Chapter 9, “Fossil History of Chameleons” by Arnau Bolet and Susan E. Evans, provides a comprehensive coverage of the fossil history of chameleons.

Unfortunately for those interested in chameleons, the fossil record of these animals is quite limited. However, the these authors do a commendable job summarizing the state of knowledge regarding chameleon fossils and placing major evolutionary events in chameleon history into a global perspective (i.e., what was happening in the world at that time). This includes a helpful table (table 9.1) and a figure linking evolutionary history and paleogeography that will undoubtedly be used in herpetology and evolutionary biology classes.

Chapter 10, “Chameleon Conservation” by Richard Jenkins, G. John Measey, Christopher V. Anderson, and Krystal A. Tolley, focuses on the myriad issues relating to chameleon conservation. The authors begin by summarizing the status of the most heavily threatened species (critically endangered by the IUCN Red List), and work through several other categories of risk in detail. The status of chameleons is troubling (a huge understatement on my part); but the authors do a commendable job describing both common and specific threats, outlining high-value habitats and areas, and going into great detail on chameleon-specific threats of trade/export/importation of chameleons.

The authors end with specific priorities for future research and regulatory actions, and with a sense that more data on diverse taxa will aid the conservation of more species.

Information for Contributors

Remember the THS in Your Will

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.

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.

Including the THS in your will is an excellent way A uthors should submit original articles, notes, book reviews to the editor, either via email using an attached to support the value of this organization and the

Sonoran Herpetologist Natural History Observations

The Tucson Herpetological Society invites your contributions to our Natural History Notes section.

We are particularly interested in photographs and descriptions of amphibians and reptiles involved in noteworthy or unusual behaviors in the field. Notes can feature information such as diet, predation, community structure, interspecific behavior, or unusual locations or habitat use. Please submit your observations to Howard Clark, editor.sonoran.herp@gmail.com.

Submissions should be brief and in electronic form.

The status of chameleons is troubling (a huge understatement on my part); but the authors do a commendable job describing both common and specific threats, outlining high-value habitats and areas, and going into great detail on chameleonspecific threats of trade/export/importation of chameleons.

Local Research News

T

he Sonoran Herpetologist welcomes short reports for our Local Research News, a regular feature in our journal. We are interested in articles that can update our readers on research about amphibians and reptiles in the Sonoran Desert region. These articles need be only a few paragraphs long and do not need to include data, specific localities, or other details. The emphasis should be on how science is being applied to herpetological questions. Please submit your materials to Howard Clark, editor.sonoran.herp@gmail.com.

Submissions should be brief and in electronic form.

SONORAN HERPETOLOGIST 27 (4) 2014

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TAXO N O M Y

New tree frog named for Ozzy Osbourne

Submitted by Suman Pratihar, Associate editor, Sonoran Herpetologist; pratihar_vu@rediffmail.com

B

ack in 1981, Black Sabbath frontman Ozzy Osbourne bit the head off a bat while performing on stage. More than 25 years later, a newly discovered species of tree frog has been named in his honor. No, the tiny Amazonian Dendropsophus ozzyi does not prey on bats, but it does sound like them. The males of the species have especially large vocal sacs that they use to produce high-pitched noises and communicate with females, National Geographic reports.

Read more here: http://bit.ly/1zFlLbS

Dendropsophus ozzyi, seen here in its Amazon habitat, has an unusually high-pitched, chirpy call. Photograph by Pedro Peloso.

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 ___________________________________________________________

SONORAN HERPETOLOGIST 27 (4) 2014

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Sonoran Herpetologist (ISSN 2333-8075) is the newsletter-journal of the Tucson Herpetological Society, and is Copyright Š 1988-2015. 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 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 Societyâ&#x20AC;&#x2122;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 Robert Villa, cascabel1985@gmail.com Vice President Krista Schmidt, turtlerad@hotmail.com Secretary Don Swann, donswann@dakotacom.net Treasurer Margaret Fusari, maggiefusari@gmail.com Directors: John Ginter, jginter@gsrcorp.com Don Moll, donandjanice@q.com Jim Rorabaugh, jrorabaugh@hotmail.com Walter Merker, walter.merker@gmail.com Steven Condon, sjcondon2001@gmail.com Cody Hurlock, codycha@msn.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 Jim Rorabaugh, Program Chair 2nd Wednesday, 7:15 PM Board of Directors Meeting Last Tuesday of each month (except December), 7:00 PM TBD Speakers Bureau (scheduled presentations) Robert Villa & Ed Moll Conservation Committee Dennis Caldwell Herpetological Information Hotline Bob Brandner, (520) 760-0574 Jarchow Conservation Award Open 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 Webpage http://tucsonherpsociety.org Heidi Flugstad, Webmaster, heidi_flugstad@hotmail.com

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

tucsonherpsociety.org

SONORAN SONORAN HERPETOLOGIST HERPETOLOGIST 25 (1)272012 (4) 2014

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