Sh vol 271 2014

25 Volume 27

January 2012 March 2014

THIS MONTH’S PROGRAM

John C. Murphy

FUTURE SPEAKERS

A Second Look at the Trinidad & Tobago Herpetofauna 7:15 PM; Tuesday, 18 March; NEW MEETING PLACE:

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15 April 2014: Philip A. Medica

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20 May 2014: B. H. Brattstrom

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

“Recital Hall” of Pima Community College West Campus, 2202 W. Anklam Road, Tucson, AZ 85709

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J

ohn C. Murphy was a high school teacher in the Chicago suburbs for most of his teaching career, and for most of those years he also had a research associate position in the Division of Reptiles and Amphibians at the Field Museum. In the early 1990s while he was finishing the Trinidad and Tobago book, he started to work with Harold Voris and Daryl Karns on homalopsids snakes, a project that spanned more than two decades and involved a considerable number of field trips to Southeast Asia. Daryl Karns died suddenly, Harold Voris retired, and Murphy also retired from his day job. He now spends most of his time studying herps, particularly snakes. In 2010 he and several others decided to take a second look at the Trinidad and Tobago herpetofauna. It turns out that like most other places on the planet there are more amphibian and reptile species there than previously thought and the species known from the islands are not necessarily the same ones thought to be present 25 years ago. This presentation is an overview and update on the fauna since the 1997 publication the Amphibians & Reptiles of Trinidad and Tobago.

Trinidad and Tobago are continental islands (or at least Trinidad is a continental island) and Tobago may be an oceanic island with a continental fauna. While geographers consider T&T the most southerly islands in the Lesser Antilles, they are better regarded as outliers of the South American continent with a fauna made up mostly of species from the northern Andes of Venezuela and the Guiana Shield. While many of the species on the islands have been considered populations of widespread species, there are more

Number 1

ISSN 2333-8075

Patronyms of the Pioneer West (Part XIX) “Scaphiopus couchii – Gnome of the Desert” by Edward O. Moll

100-MILE CIRCLE 7

“Desert Night Lizard, Xantusia vigilis Baird, 1859” by Robert L. Bezy

ANNOUNCEMENT 11

GeckoWatch: Evolution on Your Porch

IN MEMORIAM 12

“Bob Stebbins: inspiration for herpers of all ages” by Brian K. Sullivan

RESEARCH 21

“Reproductive Cycles of Lizards from Western North America” by Stephen R. Goldberg

BOOK REVIEW 30 John Murphy avoiding the strike of a Python bivittatus in Thailand. Photo by Daryl R. Karns.

endemic and near endemic species than previously thought. The presentation will summarize what has been reported in the literature and what we have discovered in the lab and field in terms of the systematics and natural history of a wide range of species. Some of the findings include the discovery of Scarthyla vigilans on Trinidad, a species previously known from Venezuela, and found calling in

“Harmony Hollow” reviewed by Howard O. Clark, Jr.

CURRENT RESEARCH NEWS 31

“When Snakes Fly” by David Bertelsen

31

“Variable Venom (Why it’s best not to be bitten in the first place)”

by David Bertelsen

SONORAN HERPETOLOGIST 27 (1) 2014

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brackish water habitats; the recently described skink Copeoglossum aurea is quite arboreal; Tobago’s glassfrog uses male parental care, and a single male may guard as many as four or five egg masses at once that are all attached to the underside of a leaf; there is a new species of parrot snake (Leptophis) from Tobago known from a single specimen, and the widespread tropidurid lizard formerly known as Plica plica is in fact a complex of at least eight different species, with the one on Trinidad

and Tobago now called P. caribeana. We are closer to understanding the nature of the Tupinambis teguixin species Group, and the Trinidad Bothrops remains a mystery for the movement. Trinidad has three species of aquatic dipsadid snakes that will use brackish water habitats and this provides a clue for a biogeographic hypothesis regarding the T&T fauna. Within the next two years we plan to have completed a field guide to the herps of the islands.

FUTURE SPEAKERS

15 April 2014: Philip A. Medica Biologist/Herpetologist USGS

Herpetological Studies in Nevada: A review of the Herps of Nevada and Desert Tortoise Ecology 7:15 PM; Location to be Determined

P hil’s fascination with wildlife began as a high school student collecting garter snakes and painted and spot-

ted turtles in nearby swamps. Simultaneously, in 1956 he joined a reptile club at the New York Zoological Park where monthly meetings chaired by Dr. James A. Oliver presented interesting talks and films on various herpetological subjects including Ditmars Horned Lizard and King Cobras. This introduction to herpetology followed by a 3 month collecting trip to Mexico with a member of the American Museum of Natural History clinched his interest in herps and the deserts of the southwestern U. S. Departing for college in New

Mexico in 1959 and mentored by Dr. James R. Dixon, Dr. Walter G. Whitford, and Dr. Ralph J. Raitt, along with encouragement to collect specific reptiles for Dr. Robert C. Stebbins to paint for his new field guide, ultimately lead to a professional career in herpetology.

Studies in Nevada pertaining to the long-term growth and natural history of the Desert Tortoise as well as population dynamics of lizards impacted by gamma radiation, and the effects of rainfall and drought were pursued with Dr. F. B. Turner and Dr. K. A. Nagy with UCLA at the Nevada Test Site. Numerous other herpetological studies pertaining to the natural history of reptiles were undertaken in Arizona, California, and Mexico with noteworthy ecological and behavioral observations reported in peer-reviewed literature. He joined the federal service in 1992 as an ecologist with the Bureau of Land Management, and subsequently served as a Research Wildlife Biologist with the U. S. National Biological Survey, Desert Tortoise Coordinator with the U. S. Fish & Wildlife Service, and biologist with the U. S. Geological Survey, becoming emeritus in May 2013. Over the past 50 years numerous students were under Phil’s supervision conducting field studies on reptiles as well as small mammals; many of which have continued in the field of biological sciences.

Phil’s fascination with wildlife began as a high school student collecting garter snakes and painted and spotted turtles in nearby swamps.

Simultaneously, in 1956 he joined a reptile club at the New York Zoological Park where monthly meetings chaired

by Dr. James A. Oliver presented interesting talks and films on various herpetological subjects including Ditmars Horned Lizard and King Cobras.

Phil Medica observing behavior of Marine Iguanas in the Galapagos Islands.

Photo by C. M. Stibor.

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20 May 2014: Bayard H. Brattstrom Professor of Zoology, California State University, Fullerton, now living at Horned Lizard Ranch, Wikieup, Arizona

Social behavior of Desert Lizards 7:15 PM; Location to be Determined

B ayard H. Brattstrom is Professor of Zoology, Emeritus, in the Department of Biology, California

State University, Fullerton, California. He currently lives on a 640-acre Horned Lizard Ranch, south of Wikieup, Arizona. He received his B.S. in Zoology from San Diego State University, and both his M.A. and Ph. D. in Zoology from the University of California, Los Angeles. He has received the Distinguished Teaching and Outstanding Research Awards from California State University, Fullerton and has been a Visiting Professor at Monash University, University of Sydney, University of Queensland, and a Visiting Researcher at James Cook University in Australia.

He is a fellow of the Herpetologists League and the American Association for the Advancement of Science. He is primarily a herpetologist with interest and numerous publications on thermoregulation, social behavior, ecology, and paleontology. He has also published on fish, birds, mammals, cacti, “warm-blooded� plants as well as on the repopulation of San Benedicto Island, Mexico after a 1953 eruption. He has served as a consultant to local, state and federal agencies, private corporations, and non-profit environmental groups, mostly on rare and endangered species. He has worked on several films, Disney’s Epcot Center, and testified, based on his herpetological research, in several criminal cases including one attempted-murder trial involving a rattlesnake. He is also the author of books of poetry, Australian slang and teaching, and has had photographs published in books, magazines and textbooks.

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Presentation Abstract

tudies on the social behavior of lizards have shown them capable of a variety of behavioral postures, sequences, and sociality, that exceed that found in some birds and mammals. While many lizards are territorial, others are hierarchical, and some have harems.

Fossorial and leaf-litter lizards, on the other hand, may just show simple, often size-related, dominance during chance encounters.

Much of lizard behavior is simple and genetic (displays, courtship), while other aspects of behavior are learned. There are, however, strong density and resource-related determinants of social behavior (food, hiding and egg-laying placa, and display sistes).

With an increase in population numbers (an increase in density), aggressive interactions increase and behavior may switch from territory to hierarchy. Most lizards

Bayard Brattstrom with model of Long-nosed Leopard Lizard, Gambelia wislizenii. This exquisite replica was created by local artist Sandra Lee. Photo by Mary Shipman.

in most situations know their social status by virtue of size, color, odors, sounds, or behavior of dominants. Subordinates respond by behaviors or postures (escape, submission, waves) to indicate submission, thereby avoiding energetically expensive or risky social interactions.

Behavior observed in the laboratory or other crowded situations allows the observer to see many if not all possible postures; behaviors, and social interactions that may be in the behavioral make-up of any given species. It does not mean that the lizard has to, or does utilize all of them in the wild, only that they are available if it needs them. While elaborate malemale displays are known for quite a few species of lizards, complete behavioral inventories (ethograms) are known for only a few species. I report here on postures, positions, ethograms for nine species of southwestern United States lizards: Coleonyx variegatus, Calliasaurus draconoides, Uta stansburiana, Sceloporus occidentalis, Phrynosoma coronatum, Heloderma suspectum, Dipsosaurus dorsalis, Uma scoparia, and Aspidoscelis hyperythrus, to species differences and the complexity of lizard behavior.

SONORAN HERPETOLOGIST 27 (1) 2014

Studies on the social behavior of lizards have shown them capable of a variety of behavioral postures, sequences, and sociality, that exceed that found in some birds and mammals.

While many lizards are territorial, others are hierarchical, and some have harems.

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PAT R O N Y M S O F T H E P I O N E E R W E S T ( PA R T X I X )

Scaphiopus couchii – Gnome* of the Desert Edward O. Moll, Tucson Herpetological Society, Tucson, AZ, e.o.moll@gmail.com

R oused by monsoon thunder, a 3 inch,

yellow-green gnome emerges from his subterranean chamber, concluding a 10 month siesta in order to make his annual visit to the world above. Wasting no time, for time is an enemy to his kind, this robust male spadefoot begins searching for a sizable puddle to serve as a temporary domain and nursery. Upon finding one of adequate size he begins emitting his characteristic, mournful “ waaaaaaaa.” Such sad bleating is understandable, considering it has been a multitude of months since his last sexual encounter and the impending breeding may well be completed in only a day or two.

This turns out to be his lucky night as a large female who has found his “waaaaaaaa” to be a cut above the pond’s other male bleaters, is approaching. The ensuing one-night stand is consummated with amplexus that stimulates the female to exude some 3000 eggs while the male swathes them with spermatozoa.

Their annual obligation complete, the proud parents abandon their potential progeny to the elements. Haste is of the essence, as these temporary ponds are already shriveling due to evaporation. In the warm water, the eggs hatch within a day and tadpoles metamorphose into froglets in just over a week.

Their reproduction completed for another year, the spadefoots transform into eating machines, gorging on sufficient arthropods to survive through the next protracted aestivation. Fortunately swarms of termites, a spadefoot delicacy, are emerging in profuse numbers at this time. The frogs gorge themselves with this high energy repast to over 50 percent of their body weight in a single evening. Following this mandatory period of gluttony and the departure of the monsoon rains, the satiated spadefoots use the keratinized spades on their hind feet to dig backward into the ground where they scrape out and enlarge aestivation chambers. As a final touch of housekeeping, each will cast off layers of sticky skin to line and waterproof its boudoir, thus assuring that humidity will remain adequate ‘til the next monsoon.

Figure 1. Couch’s Spadefoot (Scaphiopus couchii). Photo by author.

The life history depiction above is patterned after that of Couch’s Spadefoot, Scaphiopus couchii, one of the largest and best known of the American spadefoots (Fig. 1). The generic name of this frog, Scaphiopus, incorporates the root Scaph meaning spade and pus referring to foot. The specific epithet, couchii, is a patronym referring to Darius Nash Couch, its collector. For much of their history, spadefoots have been called spadefoot toads. This has created some confusion, as spadefoots and true toads reside in different families. Spadefoots differ by having vertical instead of round pupils, no parotoid glands, comparatively smooth skin and one keratinized spade instead of two.

Thus the term spadefoot is considered superior to spadefoot toad.

For the majority of years since they were first described as Scaphiopus by John Holbrook (Moll 2007) in 1836, spadefoots resided in the family Pelobatidae, along with their old world cousins, the Eurasian spadefoots. In 1866 E. D. Cope split the new world species into Scaphiopus and Spea, based on what might appear to be rather nebulous characteristics: eyelids wider than the space between them and a wedge-shaped spade instead of a sickle-shaped one. Although rejected by some authors (e.g., Wright and Wright 1949, Schmidt 1953), Spea seems to have survived the test of time and is used in most of the recent literature. In regard to the family, however, recently a small band of

Wasting no time, for time is an enemy to his kind, this robust male spadefoot begins searching for a sizable puddle to serve as a temporary domain and nursery. Upon finding one of adequate size he begins emitting his characteristic, mournful “ waaaaaaaa.”

*Referring to spadefoots as gnomes is a tribute to the Arthur Bragg’s 1965 classic work on the group, “Gnomes of the Night.”

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ever-vigilant molecular biologists (Garcia-Paris et. al.

2003) were stunned to discover that the taxon, Pelobatidae, was not monophyletic. To rectify this heinous situation, the new world spadefoots were banished from the Pelobatidae and consigned to the occasionally used taxon, Scaphiopodidae. Hopefully this classification will escape notice of future molecular biologists at least for awhile. It is hard for us elder ecologists to keep pace with current taxonomic volatility.

Scaphiopus couchii ranges across the southwestern U.S. with populations in Oklahoma, Texas, New Mexico, Arizona, and California. A small isolated population also occurs in southeastern Colorado. In addition, it has an extensive range in Mexico extending southward to Nayarit, Zacatecas, and Queretaro. The species also ranges along the entire east-side of Baja California. In Arizona the family Scaphiopodidae is represented by Scaphiopus couchii and three members of Spea (bombifrons, intermontana, and multiplicata).

Couch’s Spadefoot might well be considered the champion of xeric-adapted herps. W. W. Mayhew (1962) reported that in California’s Colorado Desert during super-drought times rainfall was insufficient to stimulate emergence of spadefoots from their subterranean chambers for up to three years. This feat makes pikers out of reptile Van Winkles such as the Arizona Mud Turtle (Kinosternon arizonense) which is known to aestivate for only a little more than a year waiting for its temporary water habitat to refill. Or consider the Gila Monster (Heloderma suspectum) which spends 92 percent of its life inactive in deep crevices or underground shelters. Nevertheless, the monster surfaces for a relatively small amount of time each year, seeking the requisite 3 to 5 rodent and avian repasts necessary to support its extended period of inactivity. Obviously neither can hold a candle to the mind-boggling abilities of the spadefoot.

Scaphiopus couchii inherited its name from a career military officer and closet naturalist Darius Nash Couch (Fig. 2; unless indicated otherwise, the following biographical material is taken from Gambone 2000). Couch was born in Putnam County, New York in 1822 to Jonathan and Betsey Couch. The name Darius came from Betsey’s first husband, as did a readymade family of three sisters. At least one or two additional sisters resulted from Jonathan’s first marriage, and an additional 6 children were born to Jonathan and Betsey. Being farmers, they lived by the farmer’s credo: the more cheap help the better. Farming life did not appeal to Darius, and in 1842 he entered West Point, where he graduated in 1846. In the same year he was promoted in the army to brevet second lieutenant 4th Artillery.

Shortly after graduation he fought in the Mexican American War (1846-1847), and was brevetted First Lieutenant for gallant and meritorious conduct. He participated in the Battle of Buena Vista (February 22-23, 1847), and later participated in the Seminole

Figure 2. Darius Nash Couch. Photo from Brady National Photographic Art Gallery (Washington, D.C.).

war in 1849 and 1850. On the negative side during the Mexican–American War, Couch developed health problems (possibly rheumatic fever and dysentery) that affected him periodically throughout his life.

Due in part to this illness and part to a fondness that he had developed for Mexico, Couch decided to take a year (1853 and 1854) off to explore the country at his leisure. The only problem was how to finance the trip. Darius had heard from colleagues that a man named Spencer Baird at the Smithsonian Institution had financed trips for other military men willing to collect animals for the museum’s expanding collection.

This sounded like a perfect solution to Couch, as he had always had an interest in nature and this would pay for his travels.

Lieutenant Couch signed on with Baird and early on in his trip provided a windfall for the Smithsonian. John Louis Berlandier (Moll 2003), a biologist, pharmacist, physician, and naturalist living in Matamoros, died prematurely in 1851—drowning while trying to cross a swollen stream. Berlandier left a large collection of archeological and natural history specimens along with natural history notes concerning the region, in the care of his wife. Couch learned of this impressive collection in 1853, when he visited Matamoros, and wrote to Baird, urging the Smithsonian to purchase it. Surprisingly, Baird could not raise enough funds from either the Smithsonian or the government for the purchase. However, Couch was so impressed with the collection, that he bought it with his own money. By agreement with Baird he sent the collection to the Smithsonian for safekeeping. The SONORAN HERPETOLOGIST 27 (1) 2014

Couch’s Spadefoot might well be considered the champion of xeric-adapted herps. W. W. Mayhew (1962) reported that in California’s Colorado Desert during superdrought times rainfall was insufficient to stimulate emergence of spadefoots from their subterranean chambers for up to three years.

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Smithsonian eventually did purchase some items from the collection, but not enough to reimburse Couch for his investment. However, Baird did help Couch find purchasers among other museums and private collectors. Two specimens of the little Texas Tortoise were in this collection, and subsequently were sent to Jean Louis Rodolphe Agassiz at Harvard (Moll 2004), who described them as Xerobates berlandieri (now Gopherus berlandieri).

Couch also collected several new species of his own on this expedition. Between Santa Catarina and Monterey, he found a tiny, inch-long nondescript fish, inhabiting a spring. Several specimens were pickled and sent to Smithsonian ichthyologist and Baird’s right hand man, Charles Girard, who named it for the young lieutenant, Limia couchianus. This fish, whose name was later changed to Xiphophous couchianus proved to be the most northern population of the platyfish group. One of its values is that it has proven useful in the study of melanomas. Unfortunately, the wild population is currently on the verge of extinction.

Among the birds Couch collected was an oriole that he named Icterus scottii. In his dedication Couch wrote: “I have named this handsome bird as a slight token of my high regard for Major Winfield Scott, Commander in Chief of the U.S. Army” (Evans 1993).

As it turns out, the oriole had been named previously, so its scientific name could not be replaced, but the common name of Scott’s Oriole has endured.

Another bird captured on the trip was named by Baird as Couch’s Kingbird. Finally, last but best, a group of spadefoots collected near the Rio Nasas, in Matamoros, Tamaulipas, were new, and described by Baird in Couch’s honor as Scaphiopus couchii. A subspecies of a pocket gopher and a rock squirrel were also named in his honor.

One final species bearing Couch’s name is the Sierra Gartersnake, Thamnophis couchii. The original description was provided by Robert Kennicott, one of a number of young biologists Spencer Baird took under his wing. When Baird wrote up the reptile section of the Williamson Route Railroad Survey from the Mississippi River to the Pacific Ocean, he asked Kennicott to write up the description of one of the gartersnakes collected on the survey, Eutaenia couchii (Thamnophis couchii; Kennicott in Baird 1859). Quite possibly Kennicott never met or even knew of Couch. Baird may have suggested the name because he still felt indebted to Couch for purchasing the Berlandier collection.

Following the Mexican trip, Couch’s tenure as a naturalist ended. Upon returning to the United States, Couch married Mary Crocker, the daughter of a congressman, in 1854. Then he resigned from the Army in 1855, took up residence in Taunton, Massachusetts, and became a merchant in his wife’s family business.

He and Mary had two children, a daughter named Alice, born in 1855 and a son named Leonard Crocker, in 1856.

With the onset of the Civil War, duty called once more and Darius agreed to be the Colonel of the 7th Regiment of Massachusetts Volunteers that entered the war on the Union side. Couch was involved with many major battles: Yorktown, Second Bull Run, Antietam, and Chancellorsrville to mention a few. By the end of the war he had been promoted to Major General, and had a Pennsylvania fort (Fort Couch) and a monument built in his honor (For a biography and complete history of his battles and wartime experiences, consult Gambone 2000).

Couch resigned from the army in May, 1865, and ran for Governor of Massachusetts. Despite being a war hero, he was defeated. For the remainder of his life, Couch held a variety of occupations including: Collector of the Port of Boston; President of the Virginia Mining and Manufacturing company; Quarter Master General of the state of Connecticut; and Adjunct General of the State of Connecticut.

Major General Darius N. Couch died in Norwalk, Connecticut in 1897 (Fig. 3). Today his name is perpetuated in zoological circles by a tiny fish, a kingbird, a gartersnake, and a robust gnome-like spadefoot. Not bad for a non-biologist Aknowledgements—K. L. Lance, D. L. Moll, and J. K. Moll read early drafts of this manuscript and offered many useful suggestions.

Literature Cited Bragg, A.N. 1965. Gnomes of the night. The spadefoot toads. University of Pennsylvania Press. 127 p.

Cope, E.D. 1866. On the structures and distribution of the genera of arciferous Anura. Journal of the Academy of Natural Science (Philadelphia) 6(1):67112.

Among the birds Couch collected was an oriole that he named Icterus scottii. In his dedication Couch wrote: “I have named this handsome bird as a slight token of my high regard for Major Winfield Scott, Commander in Chief of the U.S. Army” .

Figure 2. Stone marking the graves of Darius and Mary Couch.

Photo by author.

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Evans, H.E. 1993. Pioneer Naturalists. Henry Holt and Co. New York. 294 pp.

Gambone, A.M. 2000. Major-General Darius Nash Couch. Enigmatic Valor. Army of the Potomac Series. Butternut and Blue, Baltimore, MD. 335 p.

Garcia-Paris, M., D.R. Buchholz, and G. Parra-Olea.

2003. Phylogenetic relationships of Pelobatoidea re-examined using mtDNA. Molecular Phylogenetic Evolution 1:12-23.

Kennicott, R. 1859 in: S.F. Baird (ed.). Reptiles. Exploration & survey R.R. route Mississippi River to Pacific Ocean (Williamson Route). Washington D.C. 10 Vol. 4(4):9-13.

Mayhew, W.W. 1962. Scaphiopus couchi in California’s Colorado Desert. Herpetologica 18(3):153-161.

Moll, E.O. 2003. Gopherus berlandieri (Agassiz, 1857)—

Texas Tortoise. Patronyms of the Pioneer West II. Sonoran Herpetologist 16(6):38-39.

Moll, E.O. 2004. Gopherus agassizi (Cooper, 1863)— Desert Tortoise. Patronyms of the pioneer west VIII. Sonoran Herpetologist 17(5):50-53.

Moll, E.O. 2007. Holbrookia maculata (Girard, 1851) —Common lesser earless lizard. Patronyms of the pioneer west XV. Sonoran Herpetologist 20(3):2630.

Schmidt, K.P. 1953. A checklist of North American amphibians and reptiles. 6th ed. American Society of Ichthyologists and Herpetologists. University of Chicago Press, Chicago. viii + 280p.

Wright, A.H., and A.A. Wright, 1949. Handbook of Frogs and Toads of the United States and Canada.

Comstock Publ., Ithaca, New York xxi + 640 p.

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

Desert Night Lizard, Xantusia vigilis Baird, 1859

Robert L. Bezy, Herpetology, Natural History Museum of Los Angeles County; robertbezy@gmail.com

F or three decades following Baird’s 1859 original description, Xantusia vigilis was considered one of the

rarest North American lizards (Zweifel and Lowe 1966). This changed dramatically in 1893. During a field trip from Stanford University to southern California’s Mohave Desert, John Van Denburgh and Charles Gilbert glimpsed a small lizard deep in the crevice of a Joshua Tree (Yucca brevifolia). They managed to extract the reptile and discovered it was the rare X. vigilis (Fig. 1). Reacting in grand style, they searched though the fallen Joshua Tree limbs in the area, finding a large number of night lizards (Jennings 1997). In 1895 Van Denburgh described the event: The key to their home once discovered, the collection of a large series of these lizards was merely a matter physical exertion. Every fourth or fifth stem that was examined gave up its Xantusia, and in one instance five, as many

as were previously known to collections, were found under a single tree.

The association of Desert Night Lizards with Joshua Trees grew into an eco-legend and many a naturalist passing though the Mohave Desert stopped, turned a fallen branch or two, and quickly became a believer.

The lizards were easy to collect in large numbers and became one of the favored research animals of California’s rapidly growing universities. The extensive literature on the species can be covered only briefly here, and the reader is referred to additional summaries in Zweifel and Lowe (1966) and Bezy (1982).

Early on, Malcolm Miller authored a series of papers providing the results of remarkably detailed studies of seasonal variation and histology of the gonads and endocrine glands (e.g., Miller 1948a, 1948b) and in the process worked out some of the basic aspects of reproduction and ecology of the species (Miller 1951,

The association of Desert Night Lizards with Joshua Trees grew into an eco-legend and many a naturalist passing though the Mohave Desert stopped, turned a fallen branch or two, and quickly became a believer.

Figure 1. Desert Night Lizard (Xantusia vigilis) in the Mohave Desert, Arizona. The species has vertically elliptical pupils and lacks movable eyelids. Photo by Kit Bezy.

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Figure 2. Richard G. Zweifel (left; 1957, after completion of the Xantusia vigilis field study) and Charles H. Lowe (right; 1955, recording data at the study site near Llano, California). Photos courtesy of Richard Zweifel.

1954). He concluded that in southern California’s Antelope Valley, mating takes place in the last week of May and in the first week of June. Ovulation occurs in June and early July, gestation lasts about three months, and one to three young are born in September and October. Timing of these events was found to vary slightly from year to year.

The species has one of the lowest reproductive potentials among lizards. The average litter size for 148 broods was found to be 1.87 (calculated by Zweifel and Lowe 1966 from data in Miller 1954). However, in a dry year Miller found only 2 of 36 large adult females carried developing embryos. Reproduction by only a portion of the adult females in a given year is a phenomenon that appears to be widespread among night lizards (e.g., Island Night Lizard, X. riversiana, Goldberg and Bezy 1974; Pajapan Night Lizard, Lepidophyma pajapanense, Méndez-de la Cruz et al. 1999; Granite Night Lizard, X. henshawi, Goldberg 2013).

The embryos of the Desert Night Lizard have a well-developed placenta capable of exchanging amino acids (Miller 1951, Yaron 1977, Stewart and Presch 1992) and nearly half of the weight gain of the embryo appears to be derived from material transported across the oviduct wall to the placenta. At parturition, the neonates average 22-23 mm SVL and weigh 0.23 g (Miller 1951), and the mothers have been observed to consume the fetal membranes (Cowles 1944).

The habitat disruption required to find Desert Night Lizards makes it difficult to study their life history and ecology. In 1949 Richard Zweifel and Charles Lowe (Fig. 2) took on this thorny challenge. Zweifel (pers. com., 2012) recalled: “I was an undergraduate at UCLA and Lowe a graduate student. He asked if I would like to join him on the proposed research and I accepted immediately. (Chuck was and remained a close friend and a better source of advice than any of my formal advisors.)” Conducting eight annual winter surveys in the Antelope Valley of southern California, they balanced the necessity of capturing and marking

lizards with the need for minimal habitat disruption.

The results comprise the first and only long-term study of life history of the Desert Night Lizard (Zweifel and Lowe 1966).

In the first three years of life the lizards grew from a mean of 23 mm to a 35.1 mm SVL. After three years the females grew at a faster rate, eventually reaching 50 mm, compared to a maximum of 42 mm for males.

The number of neonates in the population was found to be significantly correlated with winter rainfall, varying from 0.6 to 1.6 per adult female.

During their seven year study, 197 of the 216 recaptures (91%) occurred at stations where the lizards had been captured previously. Only 19 “travelers” were found, and these moved a distance of 11.0 m to 320.0 m, with a mean of 43.6 m. Zweifel and Lowe (1966)

The species has one of the lowest reproductive potentials among lizards.

The average litter size for 148 broods was found to be 1.87 (calculated

by Zweifel and Lowe 1966 from data in Miller 1954).

However, in a dry year Miller found only 2 of 36 large adult females carried developing embryos.

Figure 3. Geographic distribution of the Desert Night Lizard (Xantusia vigilis) in Arizona and Sonora. Map by Kit Bezy.

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established that the species is highly philopatric and has a low reproductive potential, slow growth, and long life expectancy, achieving a population density of 47 lizards per hectare.

The social and genetic underpinnings of philopatry in the Desert Night Lizard recently have been examined in detail. Davis et al. (2011) studied kin selection in the species by combining a mark-recapture study with micro-satellite DNA analyses. A total of 2,120 lizards were captured of which 265 were recaptured.

Home range size varied from 0.1 - 53 m2 with a mean of 6.05 m2 (mean diameter, 5.59 m). Kinship groups Figure 4. Habitat of the Desert Night Lizard (Xantusia vigilis) in the Hualapai Valley, were formed through the delayed dis- Arizona, with Joshua Trees (Yucca brevifolia) and Mohave Yuccas (Yucca schidigera; left persal of offspring. Groups containing foreground). Photo by Kathryn Bolles. nuclear family members were found to be more stable than groups containing less related lizards. Kin-based social groups in the species may be a by-product of viviparity promoting delayed dispersal. Conducting crossfostering experiments in the field and laboratory, Rabosky et al. (2012) found kin presence actively promotes both philopatry and winter aggregation. Philopatry is further promoted by direct fitness benefits to adults and particularly neonates resulting from winter aggregation and delayed dispersal (Rabosky et al. 2012).

Figure 5. Habitat of the Desert Night Lizard (Xantusia vigilis) in the Poachie Mountains, Brattstrom (1953) found that Arizona, with Bigelow’s Nolinas (Nolina bigelovii) and Banana Yuccas (Yucca baccata; far left foreground). Photo by Kathryn Bolles. the most frequent items in the diet are ants and beetles. Goldberg et al.

János Xantus, the Hungarian American naturalist who (1993) recovered two species of gastrointestinal paracollected the type specimens of Xantusia vigilis at Fort sites from X. vigilis both of which have been found Tejon, California (Moll 2003). only in xantusiid lizards. Nightsnakes (Hypsiglena) As presently delineated, the species occurs in Caliare observed in the same plant litter as Desert Night fornia, Nevada, Utah, Arizona, and Sonora (Sinclair et Lizards and have been considered to be important al. 2004, Leavitt et al. 2007, Noonan et al. 2013). Since predators. However, Rodríguez-Robles et al. (1999) Klauber’s 1939 report of Desert Night Lizards in found only three X. vigilis among the 92 prey items in Arizona, our knowledge of the distribution of the spethe stomachs of the 397 nightsnakes examined. cies in the state has grown steadily. It is now known at Desert Night Lizards have a low preferred body many localities in western Arizona scattered from Utah temperature (30°C, Cowles and Bogert 1944, Brattstrom 1965), and exposure to 36.5°C for 7 days caused to Sonora (Fig. 3; Klauber 1940; Bezy 1967, 1982, 2005; Lowe 1964; Tomko 1975; Jones et al. 1981, 1982; sterility in males (Cowles and Burleson 1945). Oxygen Bezy and Sites 1987; Smith et al. 2008). consumption is only half that of other lizards and Decaying branches and trunks of Joshua Trees peaks during the day, suggesting a diurnal activity cycle (Yucca brevifolia) and Mohave Yuccas (Y. schidigera; Fig. (Mautz 1979). Field measurements indicate that night 4) comprise the principal habitat in Arizona’s Mojave lizards in their natural habitats have extraordinarily Desert from north of the Colorado River, extending low energy, food, and water requirements (Mautz and south to the Hualapai Valley and Bill Williams River.

Nagy 2000).

Along the Hualapai and Poachie mountains, the habitat The species has vertically elliptically pupils. It lacks includes Bigelow’s Nolinas (Nolina bigelovii) and Banana movable eyelids, likely the reason for the name vigilis, Latin for awake. The genus and family were named for Yuccas (Yucca baccata; Fig. 5). In the Arizona’s Sonoran SONORAN HERPETOLOGIST 27 (1) 2014

Kinship groups were formed through the delayed dispersal of offspring. Groups containing nuclear family members were found to be more stable than groups containing less related lizards.

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Desert they have been found in decaying Desert Agaves (Agave deserti; Fig. 6) and Saguaros (Carnegiea gigantea) in the Estrella, Harquahala, Kofa, Castle Dome, Cabeza Prieta, and Tinajas Altas mountains (see also Bezy 2005 and Smith et al. 2008). Packrat (Neotoma spp.) nests provide important habitat throughout the range (Zweifel and Lowe 1966). Slevin (1949) found a Desert Night Lizard active at night in a Creosote Bush (Larrea tridentata) flat in the Gran Desierto of Sonora.

Results of phylogenetic analyses of DNA sequences from nuclear and mitochondrial genes (Sinclair et al.

2004, Leavitt et al. 2007, Noonan et al. 2013) indicate that X. vigilis is most closely related to a clade containing the Arizona Night Lizard (X. arizonae) and Cole’s Night Lizard (X. jaycolei). These studies also indicate that populations living in rock-crevices in the Cerbat, Cottonwood, and Hualapai mountains of Arizona are X. vigilis rather than X. arizonae to which they previously had been assigned by Klauber (1940) and Bezy (1967).

The picture emerges that the Desert Night Lizard is a highly successful habitat specialist. Its low energy and water requirements are met within the protection of decaying yuccas, agaves, and other plants, where the combination of low predation, high life expectancy, slow growth, kin-based social groups, and low reproductive potential sustain dense populations (Zweifel and Lowe 1966, Mautz and Nagy 2000, Davis 2012).

Acknowledgments—I thank Kit Bezy, Kathryn Bolles, Thomas Brennan, Jay Cole, Erik Enderson, Stephen Goldberg, Clayton May, Edward Moll, Michael Robinson, and Wade Sherbrooke for participation in field work; Kit Bezy and Kathryn Bolles for photography; Kit Bezy for illustration; Richard Zweifel for photos and informative discussions; and Kit Bezy, Kathryn Bolles, and Stephen Goldberg for helpful suggestions on a previous version of this paper.

Literature Cited Baird, S.F. 1859. Description of new genera and species of North American lizards in the Museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences Philadelphia 1858:253-256.

Bezy, R.L. 1967. Variation, distribution, and taxonomic status of the Arizona Night Lizard (Xantusia arizonae). Copeia 1967:653-661.

Bezy, R.L. 1982. Xantusia vigilis. Catalogue of American Amphibians and Reptiles 302.1-302.4.

Bezy, R.L. 2005. The night lizards (Xantusia) of Arizona. Sonoran Herpetologist 18:14-19.

Bezy, R.L., and J.W. Sites, Jr. 1987. A preliminary study of allozyme evolution in the lizard family Xantusiidae. Herpetologica 43:280-292.

Brattstrom, B.H. 1952. The food of the night lizards, genus Xantusia. Copeia 1952:168-172.

Brattstrom, B.H. 1965. Body temperatures of reptiles.

American Midland Naturalist 73:376-422.

Figure 6. Habitat of the Desert Night Lizard (Xantusia vigilis) in the Sonoran Desert, Arizona, with Desert Agave (Agave deserti).

Cowles, R.B. 1944. Parturition in the yucca night lizard.

Copeia 1944:98-100.

Cowles, R.B., and C.M. Bogert. 1944. A preliminary study of the thermal requirements of desert reptiles. Bulletin of the American Museum of Natural History 83:261-296.

Cowles, R.B., and G.L. Burleson. 1945. The sterilizing effect of high temperature on the male germ-plasm of the yucca night lizard, Xantusia vigilis. American Naturalist 79:417-435.

Davis, A.R. 2012. Kin presence drives philopatry and aggregation in juvenile Desert Night Lizards (Xantusia vigilis). Behavioral Ecology 23:18-24.

Davis, A.R., A. Corl, Y. Surget-Groba, and B. Sinervo.

2011. Convergent evolution of kin-based sociality in a lizard. Proceedings of the Royal Society B 278:1507-1514.

Goldberg, S.R. 2013. Reproduction in the Granite Night Lizard, Xantusia henshawi (Squamata, Xantusiidae), from southern California. Sonoran Herpetologist 26:9-11.

Goldberg, S.R., and R.L. Bezy. 1974. Reproduction in the Island Night Lizard, Xantusia riversiana. Herpetologica 30:350-360.

Goldberg, S.R., C.R. Bursey, and R.L. Bezy. 1993.

Gastrointestinal helminths of night lizards, genus Xantusia (Xantusiidae). Journal of the Helminthological Society of Washington 60:165-169.

Jennings, M.R. 1997. John Van Denburgh (1872-1924): Pioneer herpetologist of the American West. Pages 323-350 in: T.W. Pietsch and W.D. Anderson, Jr. (editors). Collection Building in Ichthyology and SONORAN HERPETOLOGIST 27 (1) 2014

Results of phylogenetic analyses of DNA sequences from nuclear and mitochondrial genes (Sinclair et al. 2004, Leavitt et al.

2007, Noonan et al. 2013) indicate that X. vigilis is most closely related to a clade containing the Arizona Night Lizard (X. arizonae) and Cole’s Night Lizard (X. jaycolei).

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Herpetology. American Society of Ichthyologists and Herpetologists, Special Publication 3.

Jones, K.B., D.B. Abbas, and T. Bergstedt. 1981. Herpetological records from central and northeastern Arizona. Herpetological Review 12:16.

Jones, K.B., L.M. Porzer, and K.J. Bothwell. 1982. Herpetological records from west-central Arizona. Herpetological Review 13:54.

Klauber, L.M. 1939. Studies of reptile life in the arid Southwest.

Bulletin of the Zoological Society of San Diego 14:1-100.

Klauber, L.M. 1940. Notes from a herpetological diary, II. Copeia 1940:15-18.

Leavitt, D.H., R.L. Bezy, K.A. Crandall, and J.W. Sites, Jr. 2007.

Multi-locus DNA sequence data reveal a history of deep cryptic vicariance and habitat-driven convergence in the Desert Night Lizard Xantusia vigilis species complex (Squamata: Xantusiidae). Molecular Ecology 16:4455-4481.

Lowe, C.H. 1964. Amphibians and reptiles of Arizona, pages 153-174 in: C.H. Lowe (editor). The vertebrates of Arizona.

Univ. Arizona Press, Tucson.

Mautz, W.J. 1979. The metabolism of reclusive lizards, the Xantusiidae. Copeia 1979:577-584.

Mautz, W.J., and K.A. Nagy. 2000. Xantusiid lizards have low energy, water, and food requirements. Physiological and Biochemical Zoology 73:480-487.

Méndez-de la Cruz, F.R., M. Villagran-Santa Cruz, O. Hernandez-Gallegos, N.L. Maniquez-Moran, and F.J. RodriguezRomero. 1999. Reproductive cycle of the tropical night lizard Lepidophyma pajapanensis from Veracruz, Mexico. Journal of Herpetology 33:336-339.

Miller, M.R. 1948a. The seasonal histological changes occurring in the ovary, corpus luteum, and testis of the viviparous lizard, Xantusia vigilis. University of California Publications in Zoology 47:197-224.

Miller, M.R. 1948b. The gross and microscopic anatomy of the pituitary and the seasonal histological changes occurring in the pars anterior of the viviparous lizard Xantusia vigilis. University of California Publications in Zoology 47:225-246.

Miller, M.R. 1951. Some aspects of the life history of the yucca night lizard, Xantusia vigilis. Copeia 1951:114-120.

GeckoWatch: Evolution on Your Porch

G eckoWatch is a citizen science project to map the

fine-scale distribution of nonnative geckos in the United States. The project was developed by herpetologists from the Natural History Museum of Los Angeles County and California State University, Northridge, in collaboration with more than two dozen other partner institutions. The primary interest is in mapping the rapidly increasing range of the Mediterranean House Gecko, Hemidactylus turcicus. However, we are interested in all nonnative gecko species.

There are at least 18 species of nonnative geckos that have established populations in the United States. Seven species

Miller, M.R. 1954. Further observations on reproduction in the lizard Xantusia vigilis. Copeia 1954:38-40.

Moll, E.O. 2003. Patronyms of the pioneer West: János Xántus and Fort Tejon or how the night lizards got their name.

Sonoran Herpetologist 16:26-29.

Noonan, B.P., J.B. Pramuk, R.L. Bezy, E.A. Sinclair, K. de Queiroz, and J.W. Sites, Jr. 2013. Phylogenetic relationships within the lizard clade Xantusiidae: Using trees and divergence times to address evolutionary questions at multiple levels. Molecular Phylogenetics and Evolution 69:109-122.

Rabosky, A.R.D., A. Corl, H.E. Liwanag, Y. Surget-Groba, and B. Sinervo. 2012. Direct fitness correlates and thermal consequences of facultative aggregation in a desert lizard. PLOS ONE 7(7):1-8 e40866.

Rodríguez-Robles, J.A., D.G. Mulcahy, and H.W. Greene. 1999.

Feeding ecology of the desert night snake, Hypsiglena torquata.

Copeia 1999:93-100.

Sinclair, E.A., R.L. Bezy, K. Bolles, J.L. Camarillo R., K.A. Crandall, and J.W. Sites. 2004. Testing species boundaries in an ancient species complex with deep phylogeographic history: genus Xantusia (Squamata: Xantusiidae). American Naturalist 163:396-414.

Slevin, J.R. 1949. Range extension of Xantusia vigilis. Herpetologica 5:148.

Smith, E.L., R.L. Bezy, and T.C. Brennan. 2008. Night lizards in the Sierra Estrella. Sonoran Herpetologist 21:50-53.

Stewart, J.R., and W. Presch. 1992. Placental development of the Desert Night Lizard, Xantusia vigilis. American Zoologist 32:153A. Tomko, D.S. 1975. The reptiles and amphibians of the Grand Canyon. Plateau 47(4):161-166.

Van Denburgh, J. 1895. The species of the genus Xantusia.

Proceedings of the California Academy of Sciences (Ser. 2) 5:523-534.

Yaron, Z. 1977. Embryo-maternal interrelations in the lizard Xantusia vigilis, pages 271-277 in: J.H. Calaby and C.H. Tyndale-Briscoe (editors). Reproduction and evolution.

Australian Acad. Sci., Canberra.

Zweifel, R.G., and C.H. Lowe. 1966. The ecology of a population of Xantusia vigilis, the Desert Night Lizard. American Museum Novitates 2247:1-57. are established in Florida alone and four are established only in Hawai’i. The remaining seven species are found in multiple states.

The Mediterranean House Gecko has been the most successful invader, with established populations in at least 21 states.

To undertake any research on these nonnative geckos, scientists must first understand where these geckos occur. As we learn about the rapidly changing distributions of these nonnative geckos, we can then ask: (1) What are the impacts of these nonnative geckos on our native species?; (2) What makes some species successful colonizers?; and, (3) What are the likely routes of colonization? Observations from citizen scientists are essential to answering these questions and allowing us to learn about the invasion biology of these nonnative geckos.

To learn more about the project, visit the GeckoWatch webpage at www.nhm.org/geckowatch.

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IN MEMORIAM

Bob Stebbins: inspiration for herpers of all ages

Brian K. Sullivan, School of Mathematics & Natural Sciences, Arizona State University, Phoenix, AZ; bsullivan@asu.edu

R obert Cyril Stebbins passed away this past year (23 September 2013), at the age of 98, not long after

his last field guide appeared (Stebbins and McGinnis 2012; see Clark 2013 and Sweet 2013 for reviews).

Like all good herpers, I got to know Stebbins through his first Peterson field guide (to western reptiles and amphibians), published in 1966, and reissued in 1985 and 2003. This book achieved legendary status among naturalists in general, and herpetologists of the western United States in particular, in part because Stebbins was an accomplished illustrator (I would say “artist”, but he eschewed this label). He provided outstanding illustrations and paintings for all his books, in addition to detailed natural history about each and every species included in his many exceptional books, from 1951 through 2012. He was good friends with Chuck Lowe, an irascible fellow natural historian and herpetologist who was his polar opposite: Stebbins was an affable, soft-spoken individual whereas Lowe was a bear of man in personality and physical stature.

Lowe infamously stepped from a vehicle in the middle of nowhere (100 miles from Tucson) and informed the group of graduate students from Phoenix that they “…need to step away from the snake!” as it “… belongs to me!”, though they had stopped ten minutes earlier when Lowe was nowhere to be seen. He approached as they began to photograph the large Western Diamond-backed Rattlesnake (Crotalus atrox) crossing the paved road, just after midnight one summer night over 40 years ago. By contrast, when I showed Stebbins a uniquely patterned Western Banded Gecko (Coleonyx variegatus) that I had collected just moments before, he was so considerate of my feelings that he could hardly muster the gumption to ask if I would consider letting him keep it; I had to convince him that it was a pleasure on my part. My purpose here is to convey the depth of passion Stebbins manifested not just for herpetology, but for herpetologists—especially young and impressionable ones, mere kids who contacted him motivated only by their curiosity and keen interest in wildlife.

My life with Bob Stebbins began when I was just ten years old, and searching for guidance with respect to my interests in herpetology. In the fall of 1967, I was enrolled in a local wildlife education program, and as a result, I picked up my first copy of Stebbins (always referred to as “Stebbins”, never “Peterson guide” or “the field guide” or some such). The program, taught by the curator of vertebrates at the Alexander Lindsay Junior Museum in the East Bay Area, was my first exposure to Stebbins. The instructor of both “Reptiles” and “Raising Wild Pets”, two courses I repeated in 5th and 6th grade, conveyed an admiration for Stebbins

that bordered on rock star adoration. I was rapt as he spoke of one of the many serpents brought to the museum previously, one that—quite surprisingly in my view—he couldn’t identify, which led to him approaching Stebbins for help with identification. When he described the reaction of Stebbins as he brought the serpent forth from the standard issue cloth snake bag, it was the fact that Stebbins instantly grasped that the snake was a little-known, rear-fanged Boomslang, from Africa, that was so impressive. My instructor had been unknowingly free-handling a snake already responsible for the death of a famous herpetologist. I instantly

My purpose here is to convey the depth of passion Stebbins manifested not just for herpetology, but for herpetologists—especially young and impressionable ones, mere kids who contacted him motivated only by their curiosity and keen interest in wildlife.

Figure 1. Alameda Striped Racer (Masticophis lateralis euryxanthus), top (photo by Brian Sullivan); Gilbert’s Skink (Plestiodon gilberti), middle (adult; photo by Randy Babb) and Western Skink (Plestiodon skiltonianus), bottom (photo by Randy Babb); information on these species in Contra Costa County, California, provided anecdotal information conveyed to Stebbins in the early 1970s.

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concluded I was listening to the wrong teacher. I needed to meet Stebbins, and given my new job as a museum volunteer, prior to the arrival of any more African snakes at the museum’s “in box” (where locals could place unwanted pets and uninvited houseguests for the museum staff to contend with, one way or another). Shortly thereafter, based on the recommendation of my now disgraced instructor, I was off to the bookstore when field guides could be had for ~$7.

“Making Captures” was just what I needed to read: I spent more time in this section than any other; I’ve even written scientific manuscripts elaborating and outlining the improvements to some of the methods he described so many years ago. Years later, I was impressed with his extraordinary patience in explaining himself and this practice (making captures) when a student argued that the subjects should be released.

I’ve known very few people that would qualify, but Stebbins was “saintly” in many respects. Soft-spoken, yet with bull-dog tenacity on any topic near and dear to his heart, such as placing limits on off-road vehicle activity which was rapidly eroding the desert landscape he loved.

Some few years later, the spring of 1971, I was the proud owner of two impressive spiny lizards of the desert Southwest: a pair of male and female Granite Spiny Lizards, Sceloporus orcutti. The male of this species is perhaps the most gaudy of all North American lizards, with a suffusion of purple, blue, and turquoise across not only its ventral but all dorsal surfaces as well. The drab female, brown and tan, pales by comparison, but it was her egg-laying one night as summer began that was to provide my first opportunity for direct interaction with Stebbins. From his larger book on amphibians and reptiles of the western United States (Stebbins 1954), I knew that there had only been two clutches described for S. orcutti. Hence, I knew that any information I gathered would be “important”, and I dutifully recorded the details of each egg produced by this female, placing the length and width of each in two columns in my “field note book” (which I constructed following the guidelines in Stebbins 1966).

The next day I enlisted the help of—yes, mom; she assisted in the typing of my draft, exercising especial skill in “spell-checking” (she served in this capacity all through my high school experiences), and then we shipped it off, having no idea what, if any, response there might be.

I was not shocked as much as over-joyed when a reply from R. C. Stebbins arrived; I was struck by the heavy stock of the quality letter-head, but most compelling of all was the content. It was clear from that point on that I was “in”, as he not only complemented me on my “carefully recorded data” but expressed his desire to make use of them in his revision of the 1954 book on which I had staked my claim to significance!

The clincher, however, was the precise manner of how he concluded his request: “…I would like to make use

of these in my book, which you may know I am in the process of revising”. Well, yes, of course I knew exactly that, and thus, I realized, if only via the awkward line of an over-worked professor, he had implied that I was a member of the academy. He treated me as if I was aware of the fact that he was working on his revision. Cool.

Given the encouragement, and the feeling I was now a part of the West Coast herpetological scene, I spent considerable time concerned with how my fieldwork could provide the most useful information, though in hindsight, it was more a matter of how it might be best used to impress Stebbins. My time in the chaparral biome surrounding Mt. Diablo in the East Bay provided two such opportunities: first, in the spring of 1972 I found an Alameda Striped Racer, Masticophis lateralis euryxanthus (Fig. 1), a most impressive and it turned out, federally protected snake (one of the few). Later that year I turned up a second individual, and then, as I continued with my interests in Corral Hollow, the road-riding mecca for herpers in the Bay Area (due to the high diversity of snakes, many of which reach the northern limits of their distribution thereabouts), I found individuals there that were intermediate between the protected subspecies, and the nominate form, M. l. lateralis, that occurs throughout chaparral elsewhere in California. I knew these snakes would be of interest to Stebbins; I began gathering data on the color pattern (knowing it was key to their ID from Stebbins, the book) of all striped racers I encountered. It was 1974 before I gathered sufficient information to contact him; by that time I had learned that he had once captured an Alameda Striped Racer in the East Bay Area by leaping after one as it rapidly crawled away, successfully grabbing it by the long, thin tail. I fancied that I might have captured almost as many individuals of this snake species as he had; I was certain it would be useful to him if I could find out more.

While searching for more racers in the spring of 1972, I found a Western Skink, on the western edge of the San Joaquin Valley (east slope of North Peak) in April, and shortly thereafter, captured a Gilbert’s Skink to the west, near Lime Ridge close to my High School, De La Salle (Fig. 1). This contradicted a widely known herpetological factoid for the Bay Area: Western Skinks occur on the coast, and Gilbert’s Skinks occur in the drier interior. This was depicted not only in my larger Stebbins, but in the smaller, and much more personable guide to reptiles and amphibians of the San Francisco Bay Area, copies of which I had obtained on multiple occasions since the early to mid 60s (a common gift from relatives knowing of my predilections). I knew this was important; moreso than the length and width, and overall number of eggs produced by my Granite Spiny lizard: Stebbins made mention of the fact that these two lizard species were separated by habitat preferences in his books.

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I’ve known very few people that would qualify, but Stebbins was “saintly” in many respects.

Soft-spoken, yet with bulldog tenacity on any topic near and dear to his heart, such as placing limits on off-road vehicle activity which was rapidly eroding the desert landscape he loved.

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So, another letter—another rapid reply; not only did Bob offer praise for my work on the distribution of skinks to date, he outlined a project for me!

I was clearly moving up in the herpetological world.

Specifically, based on my highly simplistic geographical descriptions—long before the advent of GPS apps on cellular phones—he suggested I should, with the proper amount of “good, hard fieldwork”, be able to document areas of sympatry, a place where the two skinks might co-occur. It is unimportant that I never did find such a site (perhaps because small but significant differences in habitat preferences keep the two forms separate), what mattered was the degree to which we had formed a professional relationship—at least in my mind it was clear that he expected me to continue to work on this issue, and provide him with updates; I couldn’t have asked for more!

As my high school years unfolded, my trips to the desert via relatives transitioned into those I could undertake with my closest friends, all of whom became familiar with Stebbins. High school also provided the opportunity to improve my writing abilities, something Stebbins encouraged. My fall semester of junior year found me in a course otherwise completely foreign to my prior academic endeavors: advanced composition (our text was Strunk and White, The Elements of Style, the most important guide, next to Stebbins, that I ever owned, and like Stebbins, the only one for which I have had or currently retain multiple copies). For this high school course, unsurprisingly, one had to develop a single, all consuming term paper. My choice of topic?

A comparison of the Chuckwalla and Clark’s Spiny Lizard. With my instructor’s encouragement, I composed yet another letter to Stebbins, but now I pushed the newly evolving boundaries of our relationship: rather than enclosing data of some small but potential use to him, I now was unashamedly simply writing to ask a favor: could he help me in providing literature that I might use in my manuscript?

Imagine: the hero of your childhood called upon to help with your homework; the surprising thing is

that I didn’t think it was at all odd, nor did I feel in any way awkward about it. Nor did Bob; in replying, he treated me as he would a colleague, providing details on a number of contributions that could help (because he kept information on herps of the western US organized by species [individual file folders for each species], it was relatively easy for him to do so).

In this fashion I was introduced to the dissertation of his most recent student, Kristin Berry, and its focus on the Common Chuckwalla, including its social behavior, impacted me deeply (I returned to study this lizard exactly 20 years later, and have spent the last 20 years analyzing its behavior and status in the face of urbanization in the Sonoran Desert). I marvel looking back, for it was still three years before we were to meet face to face. The reply was also unique in another respect: it was two full pages in length—Bob was taking my request very seriously.

That following spring, 1974, while driving in the extreme southeastern Bay Area, I came upon a series of striped racers expressing characters suggestive of intergradation between the rare and protected Alameda subspecies and the form found throughout the rest of California. Some of these had been found DOR (“dead on the road”), and were collected. They served as the basis for the first of my “collections” to be deposited at the MVZ (Museum of Vertebrate Zoology, Cal), as important a step as I could imagine short of actually meeting Stebbins in the process!

Imagine my disappointment when he set it up so that one of his grad students would handle the duty. Ah well, I gained entry and spent a considerable amount of time in his lair, and even learned the complex system of cards for recording data of specimens to be entered into the pickled collection, and looked up a number of his specimens. This visit also led me to establish relationships with his students, and with the backing of Stebbins, the ability to visit the world renowned MVZ collection, and even gather data on all of the Alameda Striped Racers that were preserved in the collection, ones that I examined to produce two

Imagine: the hero of your childhood called upon to help with your homework; the surprising thing is that I didn’t think it was at all odd, nor did I feel in any way awkward about it.

Figure 2. Corral Hollow Road, famed road riding locale, central California, left; a road-killed King Snake (Lampropeltis californinae), taken from roadway to fence post for feeding by a raven, right. Photos by the author.

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manuscripts during college on geographic variation in this federally protected snake. Having pitched an idea—that the southeastern Bay Area, and my favorite snake hunting road, Corral Hollow, could be critical to this endangered species, I was set. Who could deny access to a herpetologist with such a project in mind?

Though I was crest-fallen each visit when he was not “in the building”, it led to my making his acquaintance in perhaps the most appropriate setting of all, entirely by chance.

Night driving at Corral Hollow was a regular pastime once I was old enough to drive (Fig. 2). Each spring that he taught Vertebrate Natural History, Stebbins organized a caravan of vehicles to load up late in the afternoon, and with TAs driving, 30-40 students would descend on Corral Hollow to see what level of diversity and abundance of herps they could muster in one evening’s worth of road riding. In the spring of 1977 I was now a regular at Corral Hollow; I would entice a friend, or two—even ask a girl on a date—and spend it cruising the road at 15 mph or so from dusk ‘til midnight. The snakes, of eleven different species, were abundant: my “best” night, 30 snakes of seven different species, stands as an all-time personal best in spite of my lifetime of efforts on multiple continents under a variety of conditions (Sullivan 1981a, b).

Given how rarely snakes are encountered while hiking, even when searching for them, road-riding is a wellunderstood method of snake collecting, and one that Stebbins championed regularly (it has served me well; I have published more papers on North American snake communities using road-riding survey methods the last 40 years than just about anyone, I would venture— save L. Klauber, a San Diego area herpetologist that Stebbins introduced me to the next year, by directing me to his many publications on this technique).

Back to my story: May, 1977—driving Corral Hollow Road that night, I noticed more than one “competitor” (any car driven as slow as mine must be up to something; 9 times out of 10 they are collectors, or drunks). More “natural” competitors, such as ravens, I tolerate—they appear to survey roads regularly looking for roadkill; I watched one swoop down and grab a snake that had just been hit by a vehicle before I could stop to identify it. Fortunately, it paused to consume its free lunch on the roadside fence, where I could sneak up to photograph it (Fig. 2). For the human variety (of potential competitors), I generally play it dumb, and stop to ask how they are doing, without giving them any encouragement (the last thing I want is for folks to think the road is “good”, and to return regularly, or talk it up as a snake hunting destination, my worst nightmare). On this warm spring night in 1977, after two cars passed me going the opposite direction, a third one approached, and slowed: I slowed to a stop, rolled down my window, and to my surprise, came face to face with the most attractive young snake chaser I’d met: a TA for Stebbins! She asked if I was having any

luck, and outlined that they were not. By that time, I’d seen four snakes on the road already; she admitted that they were doing especially poorly because there were five vehicles in their group. She indicated they were on a fieldtrip from Cal; I name-dropped a bit on Stebbins—she said, “oh yeah? Well, professor Stebbins is in that car right there!” as she motioned over her shoulder toward the car well down the roadway.

I was taken aback, to be sure, but she quickly concluded I was harmless, not so obnoxious as my comments about who I knew might have indicated, and we parted, with a plan to continue snake hunting and compare notes later. I’d not driven more than a mile or so and the car she had originally pointed out, having stopped next to her vehicle in the interim, pulled up next to mine, and the window was lowered: a quiet voice asked “Brian Sullivan?” to which I replied, “Dr. Stebbins, I presume?” (no one laughed). “Yes, it’s me”…we chatted briefly about the bad luck they were having, and how you just can’t account for variation among vehicles. We didn’t get out, we didn’t shake hands, but somehow, it all seemed very appropriate.

When I arrived at Cal in the fall of 1977, just a few months after our chance meeting, I became aware that Stebbins was retiring in the spring of 1978, and yet, would teach both of his classes, Vertebrate Natural History (known then as “the 107 course”), and Herpetology in the spring. That fall, however, my established relationship with Stebbins paid off yet again: having obtained work-study funds, I was pouring over all the job listings, and having focused on those listings with any mention of biology, zoology, or animals of any kind, I had a handful of postings to pursue. First impression? There are more rats than students at UC Berkeley.

I had just about had my fill of reading over the expected “duties” associated with the first three dozen jobs I perused (to wit: clean, wash, restock with shavings, food and water associated with the care of thousands of rats) when I saw the last listing, for a professor named Wake, Marvalee. Instead of any mention of “small mammals”, which I now took to mean “white rodents”, it simply stated: “care of amphibians”; little did I know that I was perhaps the only undergrad that arrived at her office for an interview that afternoon that could actually spell “caecilian.”

Caecilians, or apodans, are little-known, limbless amphibians that Marvalee worked on at the time (and continues working on to this day); that fall day at Cal I was ecstatic to find she was offering to pay me to act as their care taker (the first of what would be a string of dream jobs!). I had barely described my long and established relationship with Stebbins when she excused me, leaving me with the distinct impression the job was mine, and I have to admit, I was confident as well. Just tossing his name around within 48 hours of arriving on campus had paid dividends; not bad.

Tragically, I only worked with those lovely and mysteriSONORAN HERPETOLOGIST 27 (1) 2014

...the window was lowered: a quiet voice asked “Brian Sullivan?” to which I replied, “Dr. Stebbins, I presume?” (no one laughed).

“Yes, it’s me”… we chatted briefly about the bad luck they were having, and how you just can’t account for variation among vehicles.

15

ous creatures for two weeks before I had (apparently) single-handedly wiped out the largest (~100 individuals) and only known colony of the limbless amphibians in the US, all dead due to an unknown pathogen! Marvalee did not hold it against me. Subsequently, it was determined that our overly-hygenic approach to their care was the problem, and when placed in moist earth they did just fine. Having killed off the animals I was to be paid to care for over the next 12 months, Marvalee gave Figure 3. Herpetology field trip, April, 1978. Stebbins bends over to photograph me a variety of jobs to allow anuran larvae near Skagg Springs, northwest of Cal. Photo by Craig Benkman. me first-hand experience as a novice grad student in her growing and active lab. complete my independent project, but to extract two As I learned what it was like to be a graduate publications for my efforts that subsequently appeared student in Marvalee’s lab, I soon enjoyed Stebbins in in the Journal of Herpetology (Sullivan 1981a, b). More the 107 class (known by its number alone to many Cal importantly, ten years after I graduated from Cal I graduates), a field-oriented course covering the tetrawas in the running for my first full-time, assistant pods. At the time, it was a two quarter class, for which professor position: one of the three members of the Stebbins was responsible for 50% (amphibians and search committee pushed me to the top of the list reptiles), and in my case for considerably more: my based in part on my having had vertebrate natural independent project (IP; a massive undertaking that history (“107”) under Stebbins. Knowing that was could make or break your grade in this whopping 10 enough to convince him I was going to do a good job credit course). I was off to see Stebbins to pitch my IP (which entailed teaching a similar vertebrate biology in the first week of class, and it consisted of: a roadclass). Twenty years later I became the editor of the riding study of the snakes of Corral Hollow. Driving same journal that published my snake papers in part my way to my first real herpetological research, and the because early on another Cal graduate serving as editor over-the-top enthusiasm of Stebbins was a welcome vouched for my overall breadth of knowledge based in support mechanism—he suggested so many themes no small part to my undergraduate training overseen for me to pursue while undertaking my project that

by Stebbins, Harry Greene, and Marvalee Wake. after about 30 minutes worth, he looked at me, with It was on our first herpetology fieldtrip to the his sympathetic grin, and said, “oh…well, I think that northern coast of California that I became acquainted for this class, you could just try to gather as much with Bob’s amazing stamina in the field. On the field information as you can about each snake”. trip to the coast, the TA asked me to drive with StebAs we continued our discussion, he excitedly sugbins as others were self-conscious about stepping gested I draw a picture of each snake in my field note forward, and I loved it; but, his eyesight was failing (he book (I did this for the first one, but never again; it provided cars behind us with two DOR specimens; took too much time away from sampling!), record the no one ever told him the truth…at the first stop, I compass direction it was headed (yes), degree to which offered to drive for the rest of the trip). Having piled it was pressed against the pavement (yes), the time into 5 passenger cars, we arrived along the beautiful (yes), temperature of the air, road, and snake (yes, yes, coast road in the early afternoon, looking for small and yes!), and a number of other possible parameters. streams draining across the legendary State Route 1, I knew he’d like it, but I had no idea how much. Even to meet the Pacific Ocean after a rapid descent from though it was his last quarter at Cal, he remained the adjacent forests. Having found an area to pull off, engaged and excited, meeting with me whenever time the assembled group listened intently as Bob described permitted (I found that most other students appeared the fact that though we might find California Giant overly sensitive to his stature, or perhaps age, and soft- Salamanders (Dicamptodon ensatus) near the roadway, spoken demeanor, and I rarely had competition during the real treat was far upstream, over boulders and office hours). Stebbins, along with Harry Greene (the under vines, through stinging nettles and icy water: the professor that replaced Stebbins) the following year, rarely seen Southern Torrent Salamanders (Rhyacotriton provided me with the guidance sufficient not only to variegatus) found in the headwaters of small creeks such SONORAN HERPETOLOGIST 27 (1) 2014

As we continued our discussion, he excitedly suggested I draw a picture of each snake in my field note book (I did this for the first one, but never again; it took too much time away from sampling!), record the compass direction it was headed (yes), degree to which it was pressed against the pavement (yes), the time (yes), temperature of the air, road, and snake (yes, yes, and yes!)...

16

Figure 4. Photographs from the herpetology field trip to the Mohave Desert, May, 1978 (last class field trip with Stebbins). Left: Bob Stebbins speaks to students, including Jeff Kauffman whose field guide was thrown into the desert in west Texas, about a Mohave Fringe-toed Lizard (Uma scoparia) near Pisgah Crater, east of Barstow (photo by C. Benkman); upper right and right center: students with Stebbins (center, blue shirt, long white pants) at an ORV park north of Trona; bottom right: the author; bottom right-center: mottled Banded Gecko (Coleonyx variegatus) collected by the author near Pisgah Crater that was retained by Stebbins and subsequently used to illustrate pattern variation, matched with the original used in Stebbins, 1966, shown immediately above.

Gecko images modified from Stebbins (2003).

as the one near our parking area. As I peered into the primeval forest covering the steep slopes rising behind our cars, I wondered how a professor on the verge of retirement could still conduct rigorous fieldwork, and then realized he was gone—he’d plunged into the bramble and disappeared. He simply pushed through the vines, waded in the icy cold creek, and in no time at all disappeared into the darkness of the ancient coastal redwood forest. My only thought: there was no way this old geezer was going to leave me behind.

We were huffing and puffing before we had rounded the first bend of the creek, and it was 20 minutes of steady uphill slogging before he finally paused and started turning over small rocks in the trickle that remained of the creek. As I caught my breath it became clear—we were entirely alone; he didn’t even notice that no one else had made it. We could sometimes hear the others, far below, but while I gasped for air, I realized they would not be taking part in this “on the spot” lecture session.

At the end of that spring term, I was finishing my 107 final along with 120 other students, and Stebbins had been the proctor. I approached, handing in my exam, but then asked if he had finished grading the tests and term papers for my other class with him,

herpetology. Ah, yes—he distinctly remembered that he and the TA had finished up with a standard graph paper depiction of the scores of all the students in the class, and he asked her: “what’s this over here? This stray mark is too far off the chart, isn’t it?” to which the TA replied “…oh, no—that’s a STUDENT… Brian is so far above all the rest, that’s his cumulative score!” As so often was the case, he grinned, and chuckled quietly as he told me the story, taking some small satisfaction, I think, in knowing that I was absolutely over-joyed at the outcome.

The following weekend we had our “end of term” herpetology party at the TA’s house, which also served as a student-based retirement party for Stebbins. The students arrived early and stayed late, but Stebbins was there to see many of the photographs and slides that various students had taken during our two primary fieldtrips, one to the north coast of California (Fig. 3), and the other to the western and central Mohave Desert (Fig. 4; see Fig. 5 for Bob during an earlier fieldtrip of his own). The highlight for me was the autograph session the TA arranged: all the students brought their copy of Stebbins, and he sat in a centrally located chair and signed each one as it was handed to him in turn. I had never had a book signed, and haven’t SONORAN HERPETOLOGIST 27 (1) 2014

We were huffing and puffing before we had rounded the first bend of the creek, and it was 20 minutes of steady uphill slogging before he finally paused and started turning over small rocks in the trickle that remained of the creek.

As I caught my breath it became clear— we were entirely alone; he didn’t even notice that no one else had made it.

17

done it since (save for having Bob sign my second edition of Stebbins in 1985), but the look on his face when I handed him my copy was priceless. He initially looked surprised, as if I’d handed him something else by mistake, but as he opened it, and began to carefully turn the pages, he said with great sincerity and depth of feeling “Oh my, now this is what I love to see!

Look at the wear and Figure 5. Stebbins taking a break, 1964, Galapagos (photo courtesy of Ted Papenfuss). tear—look at the soiling of these pages! This book has been used as I like to kill every gawd damn one of em!”…he failed; intended!” As he continued gushing I am sure my star hats off to Houghton-Mifflin. He threw it into the never shone so bright, and the 30 or so students that desert, and as he did, his wad of wet tobacco fell from were surrounding us patted me on the back repeatedly, his mouth into Jeff ’s lap; we departed in a rage of have long ago relinquished all thoughts of competing spinning wheels, flying rocks, and the drunken cowboy with me for title of “herp nerd, first class”. The tape stumbling back to his fully loaded truck…Ed Abbey I used to repair and replace the binding after 12 years would’ve been proud: we were fleeing away from the of continuous use was barely holding it together; I see scene of the attack on Stebbins when I realized we now that he had to sign on the all important section of were going south on a road that dead-ended at the Rio Making Captures (Fig. 6) as so much of the introducGrande River…after a quick glance at the rear-view tion was missing (this copy has since been rebound, mirror, acceleration over a small hill, an abrupt turn and repaired; all but 5 pages accounted for). I was into a wash, driving just far enough off the road that impressed that he took such obvious pride and satishe passed by without seeing us (whew!), followed by faction in knowing that I was such a serious student of a u-turn, quick stop and search: Yes! We found Jeff ’s herpetology. Something akin to a father of sports icon. Stebbins! We left for Arizona the next morning.

My classmates and I left for a midnight screening of After I received my Bachelor’s degree from Cal in Rocky Horror Picture Show, complete with bikers (in the 1979, I headed to ASU in the Sonoran Desert solely theatre; ah, life at Cal, post-60s), but Stebbins declined. because of the herpetological diversity; I picked After herpetology and natural history classes were Phoenix over Tucson solely because Bob warned me over, the spring semester turned to summer, and I off UA—he thought Lowe was just too cantankerous. took one of my longest herping trips to the desert Prior to leaving Berkeley, I stopped by Bob’s office, Southwest, with Jeff Kauffman, a fellow student from and spoke with him about my career options. It was herpetology. With my well-used Stebbins in hand, I then I realized something terrible had happened: made at least one, and sometimes two trips to the desbiologists like Bob had seen the writing on the wall, ert Southwest each year (until I arrived “permanently” and recognized that the golden era of the 50s, 60s, and in 1979). On this trip, we made it to the Big Bend of 70s was over. No longer would a major professor have Texas, and it was there one night while road-riding that a stable full of young studs, multiple graduate students a drunken local, complete with cowboy hat, tobaccopursuing various projects; the crashing job market juice stained chin, and gun rack, grabbed Stebbins—the made it clear that at most we might move to a “equibook—by reaching his weathered arm through the librium” steady-state model in which a professor might passenger-side window of our small sedan. We had literally expect, if honest with themselves, to train a just pulled over, and while Jeff was thumbing through single replacement in their lifetime. That spring of my his copy of Stebbins, the stranger had pulled up, departure I had seen Stebbins brought to tears when stumbled out, and approached. While interrogating us he realized that his last three students would not be about what we were up to, he suddenly wrenched the stepping into academic positions as herpetologists— book away from Jeff, and attempted to rip it in half not one of them. He told me in no uncertain terms just like a phonebook while exclaiming that “Snakes! I that when I had first contacted him, just ten short SONORAN HERPETOLOGIST 27 (1) 2014

He initially looked surprised, as if I’d handed him something else by mistake, but as he opened it, and began to carefully turn the pages, he said with great sincerity and depth of feeling “Oh my, now this is what I love to see! Look at the wear and tear—look at the soiling of these pages!

This book has been used as I intended!”

18

Figure 6. “Stebbins” (the field guide), 1966 (1st edition; left), 1985 (2nd edition; top), 2003 (3rd edition; right), and two letters received from Stebbins during the early 1970s (lower right).

years before, he had been confident and absolutely correct when advising “all that you need is a Ph.D.; that is job security for a herpetologist!” But now, now he literally threw his hands in the air and said “I have no idea what to tell you is necessary, Brian—I cannot advise you on a fool proof plan for success as a herpetologist in academia…” I asked if perhaps simply publishing a few papers prior to graduation would help, by establishing my reputation early… “well, no, publications, even grants—there are plenty of folks out there with accomplishments along those lines that are out of work…I really don’t know what to tell you”.

I had heard a bit of this from the graduate students I’d met over the previous two years while working in Marvalee’s lab; with my whole life having been dedicated to an academic career in herpetology, there was no derailing it now, but it did make me realize it was not going to be as easy as I had once imagined.

It was with this backdrop that I met with Bob again, six years after my departure from Cal. I had indeed stuck with the academic roadmap, and obtained my Ph.D. from ASU. Just as Bob predicted, I wasn’t able to step into a position as a herpetologist; I ended up taking various post doctoral positions while hunting for that elusive tenure track position. I was in the middle of this search when I was invited to participate in a symposium on desert ecosystems at the Palm Springs Living Desert Musuem in 1985. This was where I met

Bob as less of a student, and as more of a colleague.

Almost as exciting to me was that the second edition (Fig. 6, Stebbins 1985) of “Stebbins” was introduced at those Palm Springs meetings; imagine my delight when I thumbed through the pages to see the banded gecko I collected during the 1978 herpetology class field trip, wonderfully painted in living color! Yes, the very same individual I found while night driving: the importance?

A degree of spotting he was unfamiliar with, and felt required illustration. I had to agree.

In the mid-90s, I returned to California, Berkeley, to conduct an “update” study of the Corral Hollow snake community (comparing data of the 1970s with the 1990s; he loved it!). It was my first academic sabbatical, one of those events that makes the relatively low pay of my dream job all the more palatable: a year of snake chasing on partial salary. I imagined that I would find the snakes of Corral Hollow few and far between, and my doom and gloom follow-up study would be front page news in the conservation journals (I was never so pleased to be disappointed: all the snakes found there originally are still present, and in roughly similar numbers, with one—the Pacific Rattlesnake—having increased dramatically due to the heavy rains of the early 90s). As I began my studies I dropped by Cal to call on Bob, and caught up with him at an art show, featuring his work. We chatted briefly and he told me, in hushed tones as if he didn’t SONORAN HERPETOLOGIST 27 (1) 2014

He told me in no uncertain terms that when I had first contacted him, just ten short years before, he had been confident and absolutely correct when advising “all that you need is a Ph.D.; that is job security for a herpetologist!” But now, now he literally threw his hands in the air and said “I have no idea what to tell you is necessary, Brian— I cannot advise you on a fool proof plan for success as a herpetologist in academia…”

19

want others to hear (he spoke this way so often it was tough to tell), that “I am taking art lessons; I want to learn to really paint— I’ve only ever been an illustrator, really…”

I was struck dumb; not an “artist”? In the parlance of the day, really?!

In 2000 I was preparing for the graduation of my third Ph.D. candidate, Matthew Kwiatkowski, who had worked on the social behavior of Chuckwallas, much like the Stebbins student that had proFigure 7. Field notes of Stebbins (1949) in which he first detected the similarities between larvae (and vided so much matetoads) from the Mojave and Virgin rivers (left; with the permission of the Museum of Vertebrate rial for my high school Zoology, University of California, Berkeley); range maps adopted from Stebbins (2003) for the Arizona composition course in Toad (Bufo microscaphus; lower right), and Woodhouse’s Toad (Bufo woodhousii upper; used with permission); a hybrid toad (top right) and Arizona Toad (right) are shown next to the maps.

1973. He had found the target journal, Herpetological Conservation and Biology, that females are very seunlike so many outlets limited by publication costs, lective in their choice of mates, preferring to live near encouraged figures and photographs. I decided that it and mate with males with brightly colored tails. These would be informative to publish the high quality range tails conveyed information about males: that their termaps from Bob’s 3rd edition of the field guide (Stebritories were rich in good plant foods (high in carotbins 2003) and called him to discuss the prospect. He enoids), and that they were efficient (“good”) foragers immediately agreed, and our last conversation confor such plants. I thought that one of the paintings by cerned his early work on those two toads: he collected Bob of a male chuckwalla would be a perfect gradua large number of both species and their hybrids from ation gift—and it was, but I had to get a copy first. the precise site described in my paper (the lower Virgin I called the MVZ, and eventually heard back from River) exactly 60 years prior (Fig. 7). More critically, at Bob directly: he was in his office from time to time, the time there was some confusion (controversy even) almost 25 yrs since he retired. He was so pleased that regarding the taxonomic status of these toads; some his painting was to serve the purpose of a graduating gift for someone that had furthered our understanding considered the two taxa highly divergent species (Bufo compactilis and B. woodhousii), but Bob recognized that of one of his favorite lizards, the Chuckwalla. He also mentioned that it would have been ideal if a male with one of the toads in southwestern Utah was closely a brightly colored orange tail was his subject, like those related to B. californicus from southern California, not B. woodhousii, much less B. compactilis (he made the studied by Matt, rather than one from the northern case clearly in his 1951 volume on amphibians of the Mohave Desert, studied by his student, Kristin. His western United States). Decades later I marveled at son, John, was handling sales of his artwork, and it was with him that I worked out the details of shipping the clarity of his thinking—though we differed in our taxonomic outlooks, he was exceptionally adept at and obtaining a second print for myself. The world had come full circle: thirty years later, here I was work- grasping relationships at the population level. Taxonomically, he was conservative, a “lumper” who saw ing on Chuckwallas, contacting Bob for copies of his great value in the polytypic species, and was always work, rather than advice on a high school term paper, slow—careful and cautious, he always told me—in acand he was all too willing to help out. cepting suggestions on species status for a number of My last interaction with Bob concerned the use of western amphibians we discussed over the years (e.g., his beautiful range maps for two toads I had worked although he had accepted the status of Bufo californicus on for decades: Woodhouse’s Toad (Bufo [Anaxyrus] and B. microscaphus by the time of his third edition woodhousii), and the Arizona Toad (Bufo [Anaxyrus] [2003], he still synonomized Hyla wrightorum with Hyla microscaphus; Fig. 7). In 2010 I was publishing a manueximia, though the work of one my students showed script on hybridization between these two toads, and SONORAN HERPETOLOGIST 27 (1) 2014

We chatted briefly and he told me, in hushed tones as if he didn’t want others to hear (he spoke this way so often it was tough to tell), that “I am taking art lessons; I want to learn to really paint—I’ve only ever been an illustrator, really…” I was struck dumb; not an “artist”?

In the parlance of the day, really?!

20

these two treefrogs are distinct).

The amazing fact of it all is my story is not unique; Bob Stebbins influenced countless students of all ages and all levels. Over the years, as I bored one colleague after another with stories about my first letter to Stebbins (Fig. 6), his rapid response, and consistent encouragement, more than once the listener waited patiently or interrupted quickly to let me know that they too had been provided with similar encouragement. At first, I was taken aback, imagining that this somehow diminished my relationship with Bob, but the more I heard, the more impressed I became with just how many individuals he had influenced over this lifetime—many younger herpetologists long after he had retired (in 1978). Accomplished scientists across a variety of disciplines, united by their interest in herpetology and exposure to Stebbins in their formative years include: Steve Arnold, Kristin Berry, Steve Busack, Bruce Bury, John Endler, Gary Fellers, Bob Hansen, Lynne Houck, Ray Huey, Dave Morafka, Ted Papenfuss, Eric Pianka, Dick Sage, Brad Shaffer, and Dick Zweifel to name those I am familiar with. It is impossible for me to imagine anyone coming along that can fill his shoes.

Acknowledgments—I thank John Alcock, Bob Hansen, and M. A. Kwiatkowski for comments on this manuscript; G. Rice shared her insights on the influence of Bob Stebbins. I especially thank Craig Benkman, a fellow graduate of herpetology with Bob in 1978; he graciously provided photos used herein.

Jeff Kauffman, another fellow graduate, traveled to

the desert Southwest; without him, the tales of that trip might have ended very differently.

Literature Cited Clark, H.O., Jr. 2013. Review of: Field guide to amphibians and reptiles of California. Western North American Naturalist 73:258-259.

Stebbins, R.C. 1951. Amphibians of Western North America. University of California Press, Berkeley.

Stebbins, R.C. 1954. Amphibians and Reptiles of Western North America. McGraw-Hill Press, New York.

Stebbins, R.C. 1966. A field guide to Western Reptiles and Amphibians. Houghton-Mifflin Co., Boston.

1st Edition.

Stebbins, R.C. 2003. A field guide to Western Reptiles and Amphibians. Houghton-Mifflin Co., Boston.

3rd Edition.

Stebbins, R.C., and S.M. McGinnis. 2012. Field Guide to Amphibians and Reptiles of California. Revised Edition. California Natural History Guides. University of California Press, Berkeley, CA. Sullivan, B.K. 1981a. Observed differences in body temperature and associated behavior of four snake species. Journal of Herpetology 15:245-246.

Sullivan, B.K. 1981b. Distribution and relative abundance of snakes along a transect in California.

Journal of Herpetology 15:247-248.

Sweet, S.S. 2013. Review of: Field guide to amphibians and reptiles of California. Revised Edition. Copeia 2013:785-786.

RESEARCH ARTICLE

Reproductive Cycles of Lizards from Western North America Stephen R. Goldberg, Whittier College, Department of Biology, Whittier, CA 90608, sgoldberg@whittier.edu

S

ome early excellent works on North American lizard reproduction are by Blount (1929) Phrynosoma solare, Altland (1941) Sceloporus undulatus, Reynolds (1943) Eumeces fasciatus, Woodbury and Woodbury (1945) Sceloporus graciosus, Miller (1948) Xantusia vigilis, Fitch (1954) Eumeces fasciatus, Wilhoft and Quay (1961) Sceloporus occidentalis, Asplund and Lowe (1964) Uta stansburiana and Urosaurus ornatus. There was heightened interest and a dramatic increase in studies of lizard reproductive cycles in the 1970s and 1980s.

An example of significant works were

by Ballinger (1973), Vitt (1973, 1986), Tinkle (1972, 1973), Mayhew (1963, 1965, 1966a, b), Goldberg (1974), and Goldberg and Bezy (1974).

Fitch (1970) summarized previous studies on lizard reproduction and provided information on geographic variation (Fitch 1985). Fitch (1980) discussed various aspects of reptile reproduction. The purpose of this

Over the years, as I bored one colleague after another with stories about my first letter to Stebbins, his rapid response, and consistent encouragement, more than once the listener waited patiently or interrupted quickly to let me know that they too had been provided with similar encouragement.

Western Fence Lizard (Sceloporus occidentalis), San Luis Obispo Co., California. Photo by Howard Clark.

paper is to summarize the knowledge on reproduction of lizards from western North America (Table 1) and to suggest future investigations. Taxonomy follows Jones and Lovich (2009).

SONORAN HERPETOLOGIST 27 (1) 2014

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Table 1. Lizards of western North America with sources on their reproduction. F = fall breeder, S = spring breeder, B = fall and spring breeder, U = breeding season unknown.

Crotaphytidae Great Basin Collared Lizard

Crotaphytus bicinctores

Goldberg and Mahrdt 2011

S

Eastern Collared Lizard

Crotaphytus collaris

Fitch 1956, Trauth 1979

S

Sonoran Collared Lizard

Crotaphytus nebrius

Babb 2009

S

Baja California Collared Lizard

Crotaphytus vestigium

Goldberg and Mahrdt 2010

S

Cope’s Leopard Lizard

Gambelia copei

Goldberg et al. 2010

S

Blunt-nosed Leopard Lizard

Gambelia sila

Tollestrup 1982

S

Long-nosed Leopard Lizard

Gambelia wislizenii

Tanner and Krough 1974, Tollestrup 1982, Goldberg 2013c

S

Desert Iguana

Dipsosaurus dorsalis

Mayhew 1971

S

Common Chuckwalla

Sauromalus ater

Johnson 1965, Nagy 1973, Prieto 1977

S

Zebra-tailed Lizard

Callisaurus draconoides

Vitt and Ohmart 1977

S

Greater Earless Lizard

Cophosaurus texanus

Ballinger et al. 1972, Goldberg 2013a

S

Elegant Earless Lizard

Holbrookia elegans

Goldberg 2012a

S

Common Lesser Earless Lizard

Holbrookia maculata

Jones and Ballinger 1987

S

Banded Rock Lizard

Petrosaurus mearnsi

Hain 1965, Cozens 1974, Goldberg 2011a

S

Blainville’s Horned Lizard

Phrynosoma blainvillii

Goldberg 1983

S

Texas Horned Lizard

Phrynosoma cornutum

Ballinger 1974

S

Pygmy Short-horned Lizard

Phrynosoma douglasii

Lahti and Beck 2009

S

Goode’s Horned Lizard

Phrynosoma goodei

Mulcahy 2009

S

Greater Short-horned Lizard

Phrynosoma hernandesi

Goldberg 1971b

S

Flat-tailed Horned Lizard

Phrynosoma mcallii

Howard 1974, Goldberg 2011b

S

Round-tailed Horned Lizard

Phrynosoma modestum

Howard 1974, Goldberg 2012b

S

Desert Horned Lizard

Phrynosoma platyrhinos

Pianka and Parker 1975, Goldberg 2012c

S

Regal Horned Lizard

Phrynosoma solare

Blount 1929, Parker 1971

S

Dunes Sagebrush Lizard

Sceloporus arenicolus

Fitzgerald and Painter 2009

S

Twin-spotted Spiny Lizard

Sceloporus bimaculosus

Webb 2009

S

Clark’s Spiny Lizard

Sceloporus clarkii

Tinkle and Dunham 1986

S

Prairie Lizard

Sceloporus consobrinus

Jones and Ballinger 1987

S

Southwestern Fence Lizard

Sceloporus cowlesi

Babb and Leaché 2009

S

Common Sagebrush Lizard

Sceloporus gracious

Goldberg 1975a, Woodbury and Woodbury 1945

S

Yarrow’s Spiny Lizard

Sceloporus jarrovii

Goldberg 1970, 1971a, Ballinger 1973

F

Desert Spiny Lizard

Sceloporus magister

Parker and Pianka 1973, Vitt and Ohmart 1974

S

Canyon Lizard

Sceloporus merriami

Grant and Dunham 1990

S

Western Fence Lizard

Sceloporus occidentalis

Wilhoft and Quay 1961, Goldberg 1973, 1974, 1975b, Jameson and Allison 1976

S

Texas Spiny Lizard

Sceloporus olivaceus

Blair 1960

S

Granite Spiny Lizard

Sceloporus orcutti

Mayhew 1963

S

Crevice Spiny Lizard

Sceloporus poinsettii

Ballinger 1973

F

Slevin’s Bunchgrass Lizard

Sceloporus slevini

Newlin 1976, Ballinger and Congdon 1981

S

Plateau Fence Lizard

Sceloporus tristichus

Tinkle 1972, Tinkle and Dunham 1986

S

Yellow-backed Spiny Lizard

Sceloporus uniformis

Goldberg 2012d

S

Striped Plateau Lizard

Sceloporus virgatus

Vinegar 1975, Smith et al.1995

S

Coachella Fringe-toed Lizard

Uma inornata

Mayhew 1965

S

Colorado Desert Fringe-toed Lizard

Uma notata

Mayhew 1966a

S

Yuman Fringe-toed Lizard

Uma rufopunctata

Goldberg 2012e

S

Iguanidae

Phrynosomatidae

SONORAN HERPETOLOGIST 27 (1) 2014

22

Mohave Fringe-toed Lizard

Uma scoparia

Mayhew 1966b

S

Long-tailed Brush Lizard

Urosaurus graciosus

Vitt and Ohmart 1975, Vitt et al. 1978

S

Baja California Brush Lizard

Urosaurus nigricaudus

Goldberg 1996

S

Ornate Tree Lizard

Urosaurus ornatus

Asplund and Lowe 1964, Michel 1976, Ballinger 1977, Tinkle and Dunham 1983, Van Loben Sels and Vitt 1984, Goldberg 2013e

S

Common Side-blotched Lizard

Uta stansburiana

Asplund and Lowe 1964, Tinkle 1961, Parker and Pianka 1975, Goldberg 1977, Nussbaum 1981, Goldberg 2012h

S

Texas Banded Gecko

Coleonyx brevis

Goldberg 2012i

S

Reticulate Banded Gecko

Coleonyx reticulatus

Dixon 2009a

S

Switak’s Banded Gecko

Coleonyx switaki

Grismer 2002

S

Western Banded Gecko

Coleonyx variegatus

Parker 1972, Goldberg 2012f

S

Peninsular Leaf-toed Gecko

Phyllodactylus nocticolus

Dixon 2009b

S

Arizona Striped Whiptail

Aspidoscelis arizonae

Sullivan 2009a

S

Canyon Spotted Whiptail

Aspidoscelis burti

Goldberg 1987

S

Gray Checkered Whiptail

Aspiodoscelis dixoni

Walker et al. 1991, 1994

S

Chihuahuan Spotted Whiptail

Aspidoscelis exanguis

Schall 1978

S

Gila Spotted Whiptail

Aspidoscelis flagellicauda

Stevens 1980

S

Common Spotted Whiptail

Aspidoscelis gularis

Ballinger and Schrank 1972

S

Little White Whiptail

Aspidoscelis gypsi

Rosenblum and Burkett 2009

S

Orange-throated Whiptail

Aspidoscelis hyperythra

Bostic 1966, Goldberg 2013d

S

Little Striped Whiptail

Aspidoscelis inornata

Lowe and Goldberg 1970, Christiansen 1971, Stevens 1983

S

Marbled Whiptail

Aspidoscelis marmorata

Dixon 2009c

S

New Mexico Whiptail

Aspidoscelis neomexicana

Christiansen 1971

S

Colorado Checkered Whiptail

Aspidoscelis neotesselata

Livo 2009

S

Pai Striped Whiptail

Aspidoscelis pai

Sullivan 2009b

S

Plateau Spotted Whiptail

Aspidoscelis scalaris

Leavitt 2009

S

Six-lined Racerunner

Aspidoscelis sexlineata

Fitch 1958, Hoddenbach 1966

S

Sonoran Spotted Whiptail

Aspidoscelis sonorae

Routman and Hulse 1984

S

Common Checkered Whiptail

Aspidoscelis tesselata

Jennings 2009

S

Tiger Whiptail

Aspidoscelis tigris

Goldberg and Lowe 1966, Pianka 1970, Goldberg 1976

S

Desert Grassland Whiptail

Aspidoscelis uniparens

Hulse 1981

S

Plateau Striped Whiptail

Aspidoscelis velox

Persons and Wright 2009

S

Red-backed Whiptail

Aspidoscelis xanthonota

Rosen 2009

S

Arizona Night Lizard

Xantusia arizonae

Bezy 2009a

S

Bezy’s Night Lizard

Xantusia bezyi

Bezy 2009b

U

Sandstone Night Lizard

Xantusia gracilis

Lovich 2009

S

Granite Night Lizard

Xantusia henshawi

Lee 1975, Goldberg 2013f

S

Island Night Lizard

Xantusia riversiana

Goldberg and Bezy 1974

S

Sierra Night Lizard

Xantusia sierrae

Bezy 2009c

U

Desert Night Lizard

Xantusia vigilis

Miller 1948, Zweifel and Lowe 1966

S

Wiggins’ Night Lizard

Xantusia wigginsi

Bezy 2009d

U

Mountain Skink

Plestiodon callicephalus

Swann et al. 2009

S

Gilbert Skink

Plestiodon gilberti

Goldberg 2009

S

Many-lined Skink

Plestiodon multivirgatus

Goldberg 2012g

S

Gekkonidae

Teiidae

Xantusiidae

Scincidae

SONORAN HERPETOLOGIST 27 (1) 2014

23

Great Plains Skink

Plestiodon obsoletus

Hall 1971, Goldberg 2013b

S

Western Skink

Plestiodon skiltonianus

Tanner 1957, Goldberg 2005

S

Four-lined Skink

Plestiodon tetragrammus

Fitzgerald 2009

S

Little Brown Skink

Scincella lateralis

Brooks 1967, Kwiatkowski 2009

S

California Legless Lizard

Anniella pulchra

Goldberg and Miller 1985

B

Northern Alligator Lizard

Elgaria coerulea

Vitt 1973

S

Madrean Alligator Lizard

Elgaria kingii

Goldberg 1975c

F

Southern Alligator Lizard

Elgaria multicarinata

Goldberg 1972

S

Panamint Alligator Lizard

Elgaria panamintina

Goldberg and Beaman 2003

S

Texas Alligator Lizard

Gerrhonotus infernalis

Greene et al. 2009

F

Heloderma suspectum

Goldberg and Lowe 1997

S

Anguidae

Helodermatidae Gila Monster

Discussion The reproductive cycles of lizards from western North America are typical of temperate zone lizards (both in the northern and southern hemisphere) which breed in spring. Eggs are deposited in spring, neonates emerge in summer and have several months to feed and grow before ceasing activity in autumn. Young of the year are typically active during autumn by which time adults have entered hibernation. Approximately 87/96 (91%) of North American lizards follow this regimen of spring reproduction. A few produce sperm in fall (Goldberg 1971a, Ballinger 1973, Goldberg 1975c, Mount 1963, Goldberg and Beaman 2004) and the enigmatic Anniella pulchra produces sperm in both fall and spring (Goldberg and Miller 1985).

One, however, must exercise caution in characterizing the precise timing of spring reproductive cycles as there is considerable year to year variations due to prevailing climactic conditions, see for example, Xantusia vigilis (Zweifel and Lowe 1966), Urosaurus ornatus, (Martin 1973, 1977, Ballinger 1977), Sceloporus occidentalis (Goldberg 1975b), Aspidoscelis hyperythra (Romero-Schmidt and Ortega-Rubio 2000). During years of reduced rainfall, there may be no reproduction (Mayhew 1963, 1966a, b, Zweifel and Lowe 1966, Nagy 1973; Romero-Schmidt and Ortega-Rubio 2000).

Timing of events in the reproductive cycles may vary in members of the same genus. This was noted in lizards that breed in autumn like Sceloporus jarrovii (Goldberg 1971a), Sceloporus poinsettii (Ballinger 1973) which live at high elevation and whose range extends south into Mexico. Some of the Mexican sceloporine lizards ares also known to reproduce in autumn (Guillette and MĂŠndez-de la Cruz 1993) while most species of sceloporine lizards reproduce in spring, see for example, Sceloporus occidentalis (Goldberg 1973, 1974) and Sceloporus graciosus (Goldberg 1975a). Spring breeding occurs in E. multicarinata from southern California (Goldberg 1972), whereas fall breeding occurs in E. kingi (Goldberg 1975c) which occurs at high elevations in mountains of the Madrean Archipelago,

Gerrhonotus infernalis (Greene et al. 2009) and Elgaria paucicarinata (Goldberg, and Beaman 2004) which is restricted to the Cape Region of Baja California Sur (Grismer 2002). Most scincids are spring breeders, see for example, P. obsoletus (Hall 1971, Goldberg 2013b), however, Plestiodon egergius (Mount 1963) from southeastern North America breeds in the fall as do the Mexican high-elevation skinks, Plestiodon copei (Guillette 1983) and Plestiodon brevirostris (Goldberg 2002).

While most members of the genus Phrynosoma breed in the spring, see for example P. cornutum (Ballinger 1974, 1977), however, P. ditmarsi (Goldberg 1999) from Sonora, Mexico is a fall breeder.

Viviparity is occasionally noted in lizards living in high altitudes or high latitudes as opposed to oviparous congeners in warmer climates. North American anguids exhibit oviparity (Elgaria multicarinata; Goldberg 1972), whereas, viviparity occurs in Elgaria coerulea (Vitt 1973) which may be a response to the cooler enviroments it inhabits.

Studies of the same species at different elevations have revealed differences in reproduction, presumably related to the low temperatures of the high elevation environments. This was noted in Sceloporus occidentalis and Uta stansburiana in the San Gabriel Mountains, Los Angeles County, California (Goldberg 1974, 1977).

Both species initiated testicular recrudescence (renewal of germinal epithelium for the next period of sperm formation) earlier at higher elevations than at lower elevations of the same species. This presumably reflected their shortened activity seasons. Saint Girons and Duguy (1970) reported late summer testicular recrudescence began earlier in the mountains then at sea level in Lacerta muralis in France. Yolk deposition in high elevation S. occidentalis was delayed until late March as opposed to an earlier start (February) in low elevation females (Goldberg 1974). This is presumably due to cooler temperatures at high elevations which limit lizard activity.

Lizards living at higher latitudes have been shown to produce a smaller number of clutches than their conspecifics living at lower latitudes. McCoy and SONORAN HERPETOLOGIST 27 (1) 2014

The reproductive cycles of lizards from western North America are typical of temperate zone lizards (both in the northern and southern hemisphere) which breed in spring. Eggs are deposited in spring, neonates emerge in summer and have several months to feed and grow before ceasing activity in autumn.

24

Hoddenbach (1966) reported Cnemidophorus tigris from Colorado lay one clutch, whereas females from Texas produced two clutches. The single Colorado clutch was larger (3.4 eggs) than both Texas clutches (2.2 eggs, each). Geographic differences in the timing of reproduction of Urosaurus ornatus are summarized in Goldberg (2013e).

Topics of interest for future studies might include comparisons of timing of reproductive cycles of exotic lizards in their country of origin versus newly colonized locations. Kraus (2009) provided an extensive list of exotic lizard introductions. Norval et al. (2012) reported timing of the reproductive cycle of the invasive lizard, Norops sagrei in Taiwan was similar to that reported from Belize, Cuba, Florida and Hawaii. Goldberg and Kraus (2011a) found male Chamaeleo jacksonii join the breeding population at a smaller size than their native Kenya suggesting C. jacksonii has a higher reproductive potential in Hawaii as opposed to Kenya. Carlia ailanpalai, invasive in the Mariana Islands, Oceania has been implicated in transmitting Salmonella to humans and stealing food of native lizard species (Goldberg and Kraus 2012). Judging by the success of Phelsuma laticauda in colonizing diverse habitats in the Hawiian Islands, (Goldberg and Kraus 2011b) subsequent colonizations by this species should be discouraged.

Conclusions In view of the difficulty in obtaining collecting permits for large monthly samples, utilization of existing collections in museums has become increasingly important. Because there are only limited holdings of large samples of a particular species in museums and many museums prohibit invasive procedures, future lizard reproductive studies will utilize capture, palpation and release of females. Further information on the reproductive condition may be obtained through biochemical and molecular analysis from a drop of blood with the lizard being subsequently released.

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Blount, R.F. 1929. Seasonal cycles of the interstitial cells in the testis of the horned toad (Phrynosoma solare). Journal of Morphology 48:317-343.

Bostic, D.L. 1966. A preliminary report of reproduction in the teiid lizard, Cnemidophorus hyperythrus beldingi. Herpetologica 22:81-90.

Brooks, G.R., Jr. 1967. Population ecology of the ground skink, Lygosoma laterale (Say). Ecological Monographs 37:71-87.

Christiansen, J.L. 1971. Reproduction of Cnemidophorus inornatus and Cnemidophorus neomexicanus (Sauria, Teiidae) in northern New Mexico. American Museum Novitates 2442:1-48.

Cozens, T.R. 1974. The behavioral ecology of the banded rock lizard: Petrosaurus mearnsi (Sauria: Iguanidae). M.S. thesis, San Diego State University, San Diego, CA. Dixon, J.R. 2009a. Reticulate banded gecko Coleonyx reticulatus Davis and Dixon, 1958. Pages 304-307 in: L.L.C. Jones SONORAN HERPETOLOGIST 27 (1) 2014

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and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Dixon, J.R. 2009b. Peninsular leaf-toed gecko Phyllodactylus nocticolus Dixon, 1964. Pages 318-321 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest.

A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Dixon, J.R. 2009c. Marbled whiptail, Aspidoscelis marmorata (Baird and Girard, 1852). Pages 362-365 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Fitch, H.S. 1954. Life history and ecology of the five-lined skink, Eumeces fasciatus. University of Kansas Publications Museum of Natural History 8:1-156.

Fitch, H.S. 1956. An ecological study of the collared lizard (Crotaphytus collaris). University of Kansas Publications Museum of Natural History 8:213-274.

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Fitch, H.S. 1980. Reproductive strategies of reptiles. Pages 25-31 in: Reproductive Biology and Diseases of Captive Reptiles. J.B., Murphy and J.T. Collins (editors). Society for the Study of Amphibians and Reptiles, Ithaca, NY. Fitch, H.S. 1985. Variation in clutch and litter size in New World reptiles. University of Kansas Museum of Natural History Miscellaneous Publication 76:1-76.

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A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Fitzgerald, L.A., and C.W. Painter. 2009. Dunes sagebrush lizard Sceloporus arenicolus. Pages 198-201 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Goldberg, S.R. 1970. Seasonal ovarian histology of the ovoviviparous iguanid lizard Sceloporus jarrovi Cope. Journal of Morphology 132:265-276.

Goldberg, S.R. 1971a. Reproductive cycle of the ovoviviparous iguanid lizard Sceloporus jarrovi Cope. Herpetologica 27:123-131.

Goldberg, S.R. 1971b. Reproduction in the short-horned lizard Phrynosoma douglassi in Arizona. Herpetologica 27:311-314.

Goldberg, S.R. 1972. Reproduction in the southern alligator lizard Gerrhonotus multicarinatus. Herpetologica 28:267273.

Goldberg, S.R. 1973. Ovarian cycle of the western fence lizard, Sceloporus occidentalis. Herpetologica 29:284-289.

Goldberg, S.R. 1974. Reproduction in mountain and lowland populations of the lizard Sceloporus occidentalis.

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Goldberg, S.R. 1975a. Reproduction in the sagebrush lizard, Sceloporus graciosus. American Midland Naturalist 93:177-187.

Goldberg, S.R. 1975b. Yearly variations in the ovarian cycle of the lizard Sceloporus occidentalis. Journal of Herpetology 9:187-189.

Goldberg, S R. 1975c. Reproduction in the Arizona alligator lizard, Gerrhonotus kingi. Southwestern Naturalist 20:412-413.

Goldberg, S.R. 1976. Reproduction in a mountain population of the coastal whiptail lizard, Cnemidophorus tigris multiscutatus. Copeia 1976:260-266.

Goldberg, S.R. 1977. Reproduction in a mountain population of the side-blotched lizard, Uta stansburiana (Reptilia, Lacertilia, Iguanidae). Journal of Herpetology 11:31- 35.

Goldberg, S.R. 1983. Reproduction of the coast horned lizard, Phtrynosoma coronatum, in southern California.

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Goldberg, S.R. 1987. Reproductive cycle of the giant spotted whiptail, Cnemidophorus burti stictogrammus. Southwestern Naturalist 32:510-511.

Goldberg, S.R. 1996. Urosaurus microscutatus (Small-scaled Lizard) Testis Cycle. Herpetological Review 27:202.

Goldberg, S.R. 1997. Phyllodactylus xanti (Leaf-toed gecko).

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Goldberg, S.R. 1999. Phrynosoma ditmarsi (Rock Horned Lizard). Testis Cycle. Herpetological Review 30:167.

Goldberg, S.R. 2002. Eumeces brevirostris (Shortnose skink).

Reproduction. Herpetological Review 33:134.

Goldberg, S.R. 2005. Reproductive cycle of the western skink, Eumeces skiltonianus (Sauria: Scincidae) in southern California. Texas Journal of Science 57:295-301.

Goldberg, S R. 2009. Reproductive cycle of Gilbert’s skink, Plestiodon gilberti (Squamata: Scincidae) from California.

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Goldberg, S.R. 2011a. Reproduction in the banded rock lizard. Petrosaurus mearnsi (Squamata: Phrynosomatidae).

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Goldberg, S.R. 2011b. Notes on reproduction of the flattailed horned lizard, Phrynosoma mcallii (Squamata: Phrynosomatidae). Bulletin of the Chicago Herpetological Society 46:30-31.

Goldberg, S.R. 2012a. Reproduction of the elegant earless lizard, Holbrookia elegans (Squamata: Phrynosomatidae).

Texas Journal of Science 62:281-286.

Goldberg, S.R. 2012b. Notes on reproduction in roundtail horned lizards, Phrynosoma modestum (Squamata: Phrynosomatidae) from New Mexico. Bulletin of the Chicago Herpetological Society 47:34-36.

Goldberg, S.R. 2012c. Notes on reproduction of desert horned lizards, Phrynosoma platyrhinos (Squamata: Phrynosomatidae) from southern California. Bulletin of the Chicago Herpetological Society 47:89-91.

Goldberg, S.R. 2012d. Reproduction of the yellow-backed SONORAN HERPETOLOGIST 27 (1) 2014

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spiny lizard, Sceloporus uniformis (Squamata: Phrynosomatidae) from California. Bulletin of the Southern California Academy of Sciences 111:25-28.

Goldberg, S.R. 2012e. Reproduction in the Yuman fringetoed lizard, Uma rufopunctata (Squamata: Phrynosomatidae). Southwestern Naturalist 57:457-458.

Goldberg, S.R. 2012f. Reproduction in the western banded gecko, Coleonyx variegatus (Squamata: Eublepharidae), from California. Sonoran Herpetologist 25:65-67.

Goldberg, S.R. 2012g. Reproduction of the many-lined skink, Plestiodon multivirgatus (Squamata: Scincidae) from New Mexico. Sonoran Herpetologist 25:74-76.

Goldberg, S.R. 2012h. Reproduction of the side-blotched lizard, Uta stansburiana (Phrynosomatidae) from Baja California Sur, Mexico. Sonoran Herpetologist 25:97-99.

Goldberg, S.R. 2012i. Notes on reproduction of the Texas banded gecko, Coleonyx brevis (Squamata: Gekkonidae).

Gekko 6:29-32.

Goldberg, S.R. 2013a. Reproduction in the granite night lizard, Xantusia henshawi (Squamata: Xantusiidae), from southern California. Sonoran Herpetologist 26:9-11.

Goldberg, S.R. 2013b. Reproduction of the Great Plains skink, Plestiodon obsoletus (Squamata: Scincidae) from New Mexico. Sonoran Herpetologist 26:33-35.

Goldberg, S.R. 2013c. Notes on reproduction in longnose leopard lizards, Gambelia wislizenii (Squamata: Crotaphytidae) from southern California. Bulletin of the Chicago Herpetological Society 48:79-82.

Goldberg, S.R. 2013d. Notes on reproduction of orangethroated whiptail lizards, Aspidoscelis hyperythra (Teiidae) from California. Bulletin of the Chicago Herpetological Society 48:107-108.

Goldberg, S.R. 2013e. Reproduction in the ornate tree lizard, Urosaurus ornatus (Squamata: Phrynosomatidae), from Sonora, Mexico. Sonoran Herpetologist 26:49-51.

Goldberg, S.R. 2013f. Reproduction in the granite night lizard, Xantusia henshawi (Squamata: Xantusiidae), from southern California. Sonoran Herpetologist 26:9-11.

Goldberg, S.R.,and K.R. Beaman. 2003. Elgaria panamintina (Panamint Alligator Lizard). Reproduction. Herpetological Review 34:143.

Goldberg, S.R., and K.R. Beaman. 2004. Reproduction in the San Lucan Alligator lizard, Elgaria paucicarinata (Anguidae) from Baja California Sur, Mexico. Bulletin of the Southern California Academy of Sciences 103:144-146.

Goldberg, S.R., and R.L. Bezy. 1974. Reproduction in the island night lizard, Xantusia riversiana. Herpetologica 30:350-360.

Goldberg, S.R., and F. Kraus. 2011a. Notes on reproduction of Jackson’s chameleon Chamaeleo jacksonii (Squamata, Chamaeleonidae), from Hawaii. Herpetological Bulletin 115:1-3.

Goldberg, S R., and F. Kraus, F. 2011b. Notes on reproduction of the gold dust day gecko, Phelsuma laticauda (Gekkonidae) from Hawaii. Current Herpetology 30:79-81.

Goldberg, S.R., and F. Kraus. 2012. Reproduction in the invasive lizard, Carlia ailanpalai (Squamata: Scincidae) from Oceania. Russian Journal of Herpetology 19:199-202.

Goldberg, S.R., and C.H. Lowe. 1966. The reproductive cycle of the western whiptail lizard (Cnemidophorus tigris) in southern Arizona. Journal of Morphology 118:543548.

Goldberg, S R., and C.H. Lowe. 1997. Reproductive cycle of the Gila monster, Heloderma suspectum Cope (Helodermatidae) in southern Arizona. Journal of Herpetology 31:161-166.

Goldberg, S.R., and C.R. Mahrdt. 2010. Reproduction in the Baja California Collared lizard, Crotaphytus vestigium (Squamata: Crotaphytidae). Bulletin of the Southern California Academy of Sciences 109:153-156.

Goldberg, S.R. and C.R. Mahrdt. 2011. Reproduction in the Great Basin collared lizard, Crotaphytus bicinctores (Squamata: Crotaphytidae). Bulletin of the Southern California Academy of Sciences 110:189-192.

Goldberg, S.R. and C.M. Miller. 1985. Reproduction of the silvery legless lizard, Anniella pulchra pulchra (Anniellidae) in southern California. Southwestern Naturalist 30:617-619.

Goldberg, S.R., K.R. Beaman, and C.R Mahrdt. 2010. Reproduction in the Baja California Leopard Lizard, Gambelia copeii (Squamata: Crotaphytidae). Bulletin of the Southern California Academy of Sciences 109:15-17 Grant, B.W., and A.E. Dunham. 1990. Elevational covariation in environmental constraints and life histories of the desert lizard Sceloporus merriami. Ecology 71:17651776.

Greene, H.W., P.M.Ralidis, and E.W. Acuña, 2009. Texas alligator lizard Gerrhonotus infernalis Baird, 1859 “1858).

Pages 492-495 in: L.L.C. Jones and R.E. Lovich (eds.).

Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Grismer, L.L. 2002. Amphibians and reptiles of Baja California including its Pacific islands and the islands in the Sea of Cortés. University of California Press, Berkeley, CA. Guillette, Jr. L.J. 1983. Notes concerning reproduction of the montane skink, Eumeces copei. Journal of Herpetology 17:144-148.

Guillette, L.J., and F.R. Méndez-de La Cruz. 1993. The reproductive cycle of the viviparous Mexican lizard Sceloporus torquatus. Journal of Herpetology 27:168-174.

Hahn, W.E. 1964. Seasonal changes in testicular and epididymal histology and spermatogenic rate in the lizard Uta stansburiana stejnegeri. Journal of Morphology 115:447-460.

Hain, M.L. 1965. Ecology of the lizard Uta mearnsi in a desert canyon. Copeia 1965:78-81.

Hall, R.J. 1971. Ecology of a population of the Great Plains skink (Eumeces obsoletus). University of Kansas Science Bulletin 49:357-388.

Hoddenbach, G.A.1966. Reproduction in western Texas Cnemidophorus sexlineatus (Sauria: Teiidae). Copeia 1966:110-113.

Howard, C.W. 1974. Comparative reproductive ecology of horned lizards (Genus Phrynosoma) in southwestern United States and northern Mexico. Journal of the SONORAN HERPETOLOGIST 27 (1) 2014

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Arizona Academy of Science 9:129-116.

Hulse, A.C. 1981. Ecology and reproduction of the parthenogenetic lizard Cnemidophorus uniparens (Teiidae). Annals of Carnegie Museum 50:353-369.

Jameson, Jr., E.W., and A.Allison. 1976. Fat and breeding cycles in two montane populations of Sceloporus occidentalis (Reptilia, Lacertilia, Iguanidae). Journal of Herpetology 10:211-220.

Jennings, R.D. 2009. Common checkered whiptail (unisexual) Aspidoscelis tesselata (Say, 1823). Pages 390-393 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ Johnson, S.R. 1965. An ecological study of the chuckwalla, Sauromalus obesus Baird in the western Mojave desert.

American Midland Naturalist 73:1-29.

Jones, S.M., and R.E. Ballinger. 1987. Comparative life histories of Holbrookia maculata and Sceloporus undulatus in western Nebraska. Ecology 68:1828-1838.

Jones, L.L.C., and R.E. Lovich (eds.). 2009. Lizards of the American Southwest. A photographic field guide. Rio Nuevo Publishers, Tucson, AZ. Kraus, F. 2009. Allien reptiles and amphibians. A scientific compendium and analysis. Springer Science, New York.

Kwiatkowski, M.A. 2009. Little brown skink Scincella lateralis (Say in James, 1823). Pages 466-469 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest.

A Photographic Field Guide. Rio Nuevo Publishers: Tucson, AZ. Lahti, M.E., and D.D. Beck. 2009. Pygmy short-horned lizard Phrynosoma douglasii (Bell, 1829). Pages 170-173 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers: Tucson, AZ. Leavitt, D.J. 2009. Plateau spotted whiptail Aspidoscelis scalaris (Cope, 1892). Pages 378-381 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers: Tucson, AZ. Lee, J.C. 1975. The autecology of Xantusia henshawi henshawi (Sauria: Xantusiidae). Transactions of the San Diego Society of Natural History 17:259-278.

Livo, L.L. 2009. Colorado checkered whiptail (unisexual) Aspidoscelis neotesselata (Walker, Codes and Taylor, 1997).

Pages 370-373 in: L.L.C. Jones and R.E. Lovich (eds.).

Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers: Tucson, AZ. Lovich, R.E. 2009. Sandstone night lizard Xantusia gracilis Grismer and Galvan, 1896. Pages 420-423 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers: Tucson, AZ. Lowe, C.H., and S.R. Goldberg. 1970. Reproduction in the little striped whiptail. Journal of the Arizona Academy of Science 6:162-164.

Martin, R F. 1973. Reproduction in the tree lizard (Urosaurus ornatus) in central Texas: drought conditions. Herpetologica 29:27-32.

Martin, R.F. 1977. Variation in reproductive productivity of range margin tree lizards (Urosaurus ornatus). Copeia 1977:83-92.

Mayhew, W.W. 1963. Reproduction in the granite spiny lizard, Sceloporus orcutti. Copeia 1963:144-152.

Mayhew, W.W. 1965. Reproduction in the sand-dwelling lizard, Uma inornata. Herpetologica 21:39-55.

Mayhew, W.W., 1966a. Reproduction in the arenicolous lizard Uma notata. Ecology 47:9-18.

Mayhew, W.W. 1966b. Reproduction in the psammophilous lizard Uma scoparia. Copeia 1966:114122.

Mayhew, W.W. 1971. Reproduction in the desert lizard Dipsosaurus dorsalis. Herpetologica 27:57-77.

McCoy, C.J. and G.A. Hoddenbach. 1966. Geographic variation in ovarian cycles and clutch size in Cnemidophorus tigris (Teiidae). Science 154:1671-1672.

Michel, L. 1976. Reproduction in a southwest New Mexican population of Urosaurus ornatus. Southwestern Naturalist 21:281-299.

Miller, M.R. 1948. The seasonal histological changes occurring in the ovary, corpus luteum and testis of the viviparous lizard, Xantusia vigilis. University of California Publications in Zoology 47:197-224.

Mount, R.H. 1963. The natural history of the redtailed skink, Eumeces egregius Baird. American Midland Naturalist 70:356-385.

Mulcahy, D.G. 2009. Goode’s horned lizard Phrynosoma goodei Stejneger, 1893. Pages 174-177 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide.

Rio Nuevo Publishers: Tucson, AZ. Nagy, K.A. 1973. Behavior, diet and reproduction in a desert lizard, Sauromalus obesus. Copeia 1973:93102.

Newlin, M.E. 1976. Reproduction in the bunch grass lizard, Sceloporus scalaris. Herpetologica 32:171-184.

Norval, G., S.R. Goldberg, and J. Mao. 2012. The reproductive cycle of the brown anole (Anolis sagrei) an introduced lizard species in Taiwan. Russian Journal of Herpetology 19:75-81.

Nussbaum, R.A. 1981. Seasonal shifts in clutch size and egg size in the side-blotched lizard, Uta stansburiana. Oecologia 49:8-13.

Parker, W.S. 1971. Ecological observations on the regal horned lizard (Phrynosoma solare) in Arizona.

Herpetologica 27:333-338.

Parker, W.S. 1972. Aspects of the ecology of a Sonoran desert population of the western banded gecko, Coleonyx variegatus (Sauria, Eublepharinae).

American Midland Naturalist 88:209-224.

Parker, W.S., and E.R. Pianka. 1973. Notes on the ecology of the iguanid lizard, Sceloporus magister.

Herpetologica 29:143-152.

Parker, W.S. and E.R. Pianka. 1975. Comparative ecology of populations of the lizard Uta stansburiana.

Copeia 1975:615-632.

Persons, T.B. and J.W. Wright. 2009. Plateau striped SONORAN HERPETOLOGIST 27 (1) 2014

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whiptail (unisexual) Aspidoscelis velox (Springer, 1928).

Pages 402-405 in: L.L.C. Jones and R.E. Lovich (eds.).

Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Pianka, E.R. 1970. Comparative autecology of the lizard Cnemidophorus tigris in different parts of its geographic range. Ecology 51:703-720.

Pianka, E.R. and W.S. Parker. 1975. Ecology of horned lizards: a review with special reference to Phrynosoma platyrhinos. Copeia 1975:141-162.

Prieto, A.A. 1977. Reproduction in the Arizona chuckwalla, Sauromalus obesus tumidus. American Midland Naturalist 98:463-469.

Reynolds, A.E. 1943. The normal seasonal reproductive cycle in the male Eumeces fasciatus together with some observations of the effects of castration and hormone administration. Journal of Morphology 72:331-377.

Romero-Schmidt, H.L. and A. Ortega-Rubio. 2000. Reproduction of the Cape orange-throat whiptail, Cnemidophorus hyperythrus hyperythrus. Herpetological Natural History 7:1-7.

Rosen, P.C. 2009. Red-backed whiptail, Aspidoscelis xanthonota (Duellman and Lowe 1953. Pages 406-409 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Rosenblum, E.B., and D. Burkett. 2009. Little white whiptail Aspidoscelis gypsi. Pages 350-353 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest.

A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Routman, E.J., and A.C. Hulse. 1984. Ecology and reproduction of a parthenogenetic lizard, Cnemidophorus sonorae. Journal of Herpetology 18:381-386.

Saint Girons, H. and R. Duguy. 1970. Le cycle sexuel de Lacerta muralis L.en plaine et en montagne. Bulletin du MusÊum National d’Histoire Naturelle 42:609-625.

Schall, J.J. 1978. Reproductive strategies in sympatric whiptail lizards (Cnemidophorus) two parthenogenetic and three bisexual species. Copeia 1978:108-116.

Smith, G.R., R.E. Ballinger, and B.R. Rose.1995. Reproduction in Sceloporus virgatus from the Chiricahua Mountains of southeastern Arizona with emphasis on annual variation. Herpetologica 51:342-349.

Stevens, T.P. 1980. Notes on thermoregulation and reproduction in Cnemidophorus flagellicaudus. Journal of Herpetology 14:418-420.

Stevens, T.P. 1983. Reproduction in an upper elevation population of Cnemidophorus inornatus (Reptilia, Teiidae).

Southwestern Naturalist 28:9-20.

Sullivan, B.K. 2009a. Arizona striped whiptail Aspidoscelis arizonae (Van Denburgh, 1896). Pages 326-329 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Sullivan, B.K. 2009b. Pai striped whiptail Aspidoscelis pai (Wright and Lowe, 1993). Pages 374-377 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American South-

west. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Swann, D.E., M.D. Caron, and T. Edwards. 2009.

Mountain skink Plestiodon callicephalus Bocourt, 1879. Pages 446-448 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest.

A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Tanner, W.W. 1957. A taxonomic and ecological study of the western skink (Eumeces skiltonianus). Great Basin Naturalist 17:59-88.

Tanner, W.W., and J.E. Krough. 1974. Ecology of the leopard lizard, Crotaphytus wislizenii at the Nevada Test Site, Nye County, Nevada. Herpetologica 30:63-72.

Tinkle, D.W. 1961. Population structure and reproduction in the lizard Uta stansburiana stejnegeri. American Midland Naturalist 66:206-234.

Tinkle, D.W. 1972. The dynamics of a Utah population of Sceloporus undulatus. Herpetologica 28:351359.

Tinkle, D.W. 1973. A population analysis of the sagebrush lizard, Sceloporus graciosus in southern Utah.

Copeia 1973:284-296.

Tinkle, D.W, and A.E. Dunham. 1983. Demography of the tree lizard, Urosaurus ornatus, in central Arizona. Copeia 1983:585-598.

Tinkle, D.W., and A.E. Dunham. 1986. Comparative life histories of two syntopic sceloporine lizards.

Copeia 1986:1-18.

Tollestrup, K. 1982. Growth and reproduction in two closely related species of leopard lizards, Gambelia silus and Gambelia wislizenii. American Midland Naturalist 108:1-20.

Trauth, S.E. 1979. Testicular cycle and timing of reproduction in the collared lizard (Crotaphytus collaris) in Arkansas. Herpetologica 35:184-192.

Van Loben Sels, R.C., and L.J. Vitt. 1984. Desert lizard reproduction seasonal and annual variation in Urosaurus ornatus (Iguanidae). Canadian Journal of Zoology 62:1779-1787.

Vinegar, M.B. 1975. Demography of the striped plateau lizard, Sceloporus virgatus. Ecology 56:172-182.

Vitt, L.J. 1973. Reproductive biology of the anguid lizard, Gerrhonotus coeruleus principis. Herpetologica 29:176-184.

Vitt, L J. 1986. Reproductive tactics of sympatric gekkonid lizards with a comment on the evolutionary and ecological consequences on invariant clutch size. Copeia 1986:773-786.

Vitt, L.J., and R.D. Ohmart. 1974. Reproduction and ecology of a Colorado River population of Sceloporus magister (Sauria: Iguanidae). Herpetologica 30:410-416.

Vitt, L.J., and R.D. Ohmart. 1975. Ecology, reproduction, and reproductive effort of the iguanid lizard Urosaurus graciosus on the Lower Colorado River.

Herpetologica 31:56-65.

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Vitt, L.J., and R.D. Ohmart. 1977. Ecology and reproduction of lower Colorado River lizards: I. Callisaurus draconoides (Iguanidae). Herpetologica 33:214-222.

Vitt, L. J., R.C. Van Loben Sels, and R.D. Ohmart.

1978. Lizard reproduction: annual variation and environmental correlates in the iguanid lizard Urosaurus graciosus. Herpetologica 34:241-253.

Walker, J.M., J.F. Scudday, R.V. Kilambi, and C.C. Cohn. 1991. Activity, temperature, age, size and reproduction in the parthenogenetic whiptail lizard Cnemidophorus dixoni in the Chinati Mountains in Trans-Pecos Texas. American Midland Naturalist 126:256-268.

Walker, J.M., J.E. Cordes, C.C. Cohn, H.L.Taylor, R.V. Kilambi, and R.L. Meyer. 1994. Life history characteristics of 3 morphotypes in the parthenogenetic Cnemidophorus dixoni complex (Sauria, Teiidae) in Texas and New Mexico. Texas Journal of Science 46:27-33.

Webb, R.G. 2009. Twin-spotted spiny lizard Sceloporus bimaculosus Phelan and Brattstrom, 1955. Pages 202205 in: L.L.C. Jones and R.E. Lovich (eds.). Lizards of the American Southwest. A Photographic Field Guide. Rio Nuevo Publishers, Tucson, AZ. Wilhoft, D. C. and W.B. Quay. 1961. Testicular histology and seasonal changes in the lizard Sceloporus occidentalis. Journal of Morphology 108: 95-106.

Woodbury, M. and A.M. Woodbury. 1945. Life history studies of the sage brush lizard, Sceloporus g. graciosus with special reference to cycles in reproduction.

Herpetologica 2:175-196.

Zweifel, R.G., and C.H. Lowe. 1966. The ecology of a population of Xantusia vigilis, the desert night lizard.

American Museum Novitates 2247:1-57.

BOOK REVIEW

Harmony Hollow Howard O. Clark, Jr., Editor, Tucson Herpetological Society, Tuscon, AZ; editor.sonoran.herp@gmail.com

By Donald G. Kramer Published: 1/17/2006 AuthorHouse, Bloomington, IN Format: Perfect Bound Softcover Pages: 96 Size: 5 × 8 ISBN: 978-1-42590-045-8 Print Type: B/W

A s a product of the 1970s, I oftentimes recall a simple childhood where I’d roam the countryside and local orchards in search of wildlife and adventure.

On occasion I would encounter a tree frog or snake or study the birds as they fly over. Once the mosquitos made their appearance I would head indoors for dinner. Childhood freedom as described here is rare nowadays and the lackadaisical days of yesteryear are likely extinct for most kids.

Donald G. Kramer has captured this exploratory spirit in his chapter book Harmony Hollow, aimed for young readers. I read the entire book in one setting.

I was quickly reminded of my adventurous past and could readily relate to the three main characters—life friends aged 12 to 13, set out for an adventure.

The boys were set to explore a nearby deep valley set in the mountains of Washington state. Their fathers work for the Forest Service for part of the year. They initially planned to camp at a designated campsite but all along they were scheming to spend a few days in the Harmony Hollow.

The adventure they embark upon is more than they imagined—with a bit of heroism and cryptozoology.

The book is a delightful and quick read—perfect for any young boy or girl ready to lose themselves on a rainy afternoon. Books that capture the free spirit of the child are rare, and I believe Kramer has written a winner.

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CURRENT RESEARCH NEWS

When Snakes Fly David Bertelsen, member, Tucson Herpetological Society, Tuscon, AZ; david.bertelsen8@gmail.com

T he genus Chrysopelea is a colubrid group of five species found from western India to Indonesia.

Known as “flying snakes,” these beautiful reptiles, ranging in length from 0.6 to 1.2 m, have rear fangs but are no threat to humans. Socha (2011) described behavior in the genus, and Holden et al. (2014) analyzed the aerodynamics of Chrysopelea paradise, the most proficient glider of the genus (it can produce glide ratios up to 4.5 and can actively turn in flight). These are typical snakes without body appendages such as skin flaps yet they can produce lift, minimal drag, and vortex-shedding by shaping their bodies, creating an airfoil by splaying their ribs. Both launch and landing are different from other gliding animals, not surprising given the absence of limbs. Although they sometimes simply fall or dive, individuals usually take off by dangling from a branch in a J-shape, then thrust their bodies upward and forward before releasing, probably to increase velocity (and lift) for the flight and to clear nearby obstacles such as branches. As it launches, the snake flattens its body from just behind its head to its vent into a low triangle with a concave base, the lower

ends of the triangle forming a distinct lip that helps provide lift. With the body angled upward in relation to the flight path, the snake undulates its head from side to side, forming an S-shape that in effect acts as fore and aft wings. The posterior portion of the body oscillates in a vertical plane and may facilitate turning.

To land on a branch, the snake forms a hook with its body, while the tail and posterior portion of the body touch down first and the head last when landing on the ground. A great National Geographic video showing Socha and the snake at work can be viewed at http:// on.natgeo.com/1im5rBn. Literature Cited Socha, J.J. 2011. Gliding flight in Chrysopelea: Turning a snake into a wing. Integrative and Comparative Biology 51:969-982.

Holden, D., J.J. Socha, N.D. Cardwell, and P.P. Vlechos.

2014. Aerodynamics of the flying snake Chysopelea paradise: how a bluff body cross-sectional shape contributes to gliding performance. The Journal of Experimental Biology 217:382-394.

Variable Venom (Why it’s best not to be bitten in the first place) David Bertelsen, member, Tucson Herpetological Society, Tuscon, AZ; david.bertelsen8@gmail.com

C

rotalus oreganus helleri, the Southern Pacific Rattlesnake, lives in diverse habitats from sea level to over 3000 m in northwestern Baja California, southwestern California, and Santa Catalina and Coronado del Sur islands. Adults range in length from 0.6 to 1.4 m and eat a variety of prey species. Coloration ranges from browns to grays and blacks (see http://bit.ly/1e4pQGk); the distinctive characteristic of this subspecies is the terminal tail band which is at least twice as wide as the other bands. Not surprising given the diversity of habitats and prey, C. o. helleri is known for its extreme variation in venom, resulting in considerable difficulty in determining appropriate treatment and management of bites.

Using individuals from four habitat types in California, Sunagar et al. (2014) investigated the toxin diversity of C. o. helleri venom using a proteomics-transcriptomics approach. The objective was to analyze the molecular evolution and diversification within toxin types and the expression of these types within the venom of individuals in different parts of the subspecies’ range. The analysis showed significant differences in the venom composition of the four sub-populations,

with some being hemorrhagic and others neurotoxic.

Sunagar et al. (2014) concluded that venom differences resulted from differential evolutionary selection pressures and that the rates of molecular change varied both between toxin types and within the venom of the subpopulations. The authors argued that the results support the hypothesis that the underlying cause of venom variation was adaptation to particular prey.

Literature Cited Sunagar, K., E.A.B. Undheim, H. Scheib, E.C.K. Gren, C. Cochran, C.E. Person, I. Koludarov, W. Kelln, W.K. Hayes, G.F. King, A. Antunes, and B.G. Fry. 2014. Intraspecific venom variation in the medically significant Southern Pacific Rattlesnake (Crotalus oreganus helleri): Biodiscovery, clinical and evolutionary implications. Journal of Proteomics, in press. http://dx.doi.org/10.1016/j.jprot.2014.01.013.

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Amphibian & Reptile Conservation (ISSN: 1083-446X; eISSN: 1525-9153) (ARC) is an open-access international journal that publishes original peer-reviewed research, reviews, short communications, new species descriptions, and country reviews that focus on a variety of topics related to the biology, conservation, natural history, and diversity of amphibians and reptiles. ARC is now published semi-annually and each issue typically focuses on one country or subject but not exclusively, which general issues being produced as well. The most recent issue (September 2013) focuses on Mexico and includes seven papers. An upcoming issue on the country of Colombia is being developed for 2014.

Potential author(s) are encouraged to submit papers directly to the editor at: arc.publisher@gmail.com and/or for further discussion and consideration.The NEW journal website is now located at: amphibian-reptile-conservation.org and is the offical journal destination for news, updates, and current and past papers published in the journal. All current and previously published papers of ARC have been added to ZenScientist.com and can be downloaded for free there as well on the official ARC website: http://bit.ly/1lyUsKc MEETING MINUTES BOD minutes can be found here: www.dropbox.com/sh/bxl4xdoyxc1x4bx/s4fCPAHXgS MEMBERSHIP

Membership Information Individual Family Student

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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.

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.

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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, Sarah Ashby by email at halfaussie@gmail.com or by phone at (520) 396-9832.

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

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$ _______ 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 Sarah Ashby at halfaussie@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.

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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 Roy Averill-Murray, royaverillmurray@charter.net 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.

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

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

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.

Officers 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 Sarah Ashby, halfaussie@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 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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