Tlaconete Pinto
Arizona ’s Lost Salamander

Originally published in the Sonoran Herpetologist "Herpetofauna of the 100-mile Circle" 17 (12) 2004

Robert L. Bezy 1, Erik F. Enderson 2, and Kevin E. Bonine 3

1 Natural History Museum of Los Angeles County, Los Angeles, California 90007, USA.
2 2627 S. Jaguar Rd, Tucson, Arizona 85730, USA.
3 Department of Ecology and Evolutionary Biology and School of Natural Resources, University of Arizona, Tucson, Arizona 85721, USA.

Much water has flowed under many bridges since the January of the year of our Lord one thousand nine hundred and thirteen, when a college sophomore opened a letter and read a sentence, the results whereof are the following pages. It has been thirteen years since the eighteen-year-old boy read those few words and in those thirteen years for him the surf has whitened the shores of Caribbean islands; the slopes of the Balsams have been blue in the distance; hawks have soared a thousand feet below the naked peak of Sharp Rock; the iridescent wings of Morpho have fluttered through glades in the rain forest and in the mist of the cataract at Xico; from the mooring at Vera Cruz tall Orizaba has stood against the western sky; the low sun has shone on the ice of Ixtaccihuatl, most unforgettable of mountains; step after step, for him, far out at sea, Chirripo Grande, "nie von Menschenfuss betreten," has climbed aloft; and from shaken Irazu, while the ash cloud of the eruption rose above his head and floated, a black pall beaten by the fierce wind, he has seen through a break in the clouds, far to the Southwards, the sheer and menacing shaft of rock which is Cerro de la Muerte. And in that letter and in that sentence there was no hint of high peaks or of distant places, of mountains seen and unclimbed and forever remembered, of the coughing of tigers by night in the black forest, there were only these words in that letter from Leonhard Stejneger "I would like to point out to you, however, that Herpetology offers fields more in need of investigation than the snakes, viz., the salamanders."

Emmett Reid Dunn 1926

This passage from the forward to Dunn's (1926) classical monograph of the salamanders of the family Plethodontidae stands as perhaps the most poetic description of herpetological field work ever penned. In the grandeur of this forward, the insightful discussion of the evolution of the Plethodontidae, and the nearly 400 pages of systematic accounts of the species, it is easy to overlook the one line in this monograph that is, perhaps, of most significance to Arizona herpetology. This occurs, without discussion, on p. 360 in the account of Oedipus bellii: "Specimens seen 50, as follows: ….United States: Arizona: Ft. Whipple 3 (U.S.N.M., uncatalogued, now lost)."

Dunn’s citation of these three specimens from Fort Whipple (near Prescott, Yavapai Co., Arizona) is highly significant in terms of salamander biogeography. The gigantic red-blotched salamander now known as Pseudoeurycea bellii (Fig. 1, 4) was described by Gray in 1850 (as Spelerpes bellii) from Mexico and is a member of the Plethodontidae, the Lungless Salamanders. Plethodontids comprise the largest salamander family with over 350 species found from British Colombia and Nova Scotia south to Brazil and Bolivia and in a small, disjunct portion of southern Europe (Dunn 1926; Frost 2004; Wake 1966; Wake and Lynch 1976). Arizona lies in a distributional gap between three geographic clusters of plethodontid species: (1) California and the Pacific Northwest; (2) the eastern U.S.; and (3) tropical Mexico and Central and South America (Fig. 2). This immense void (the Plethodontid Gap) in the distribution of Lungless Salamanders extends throughout the major North American deserts (Sonoran, Chihuahuan, Mojave, and Great Basin) and into much of the Great Plains, Mexican Plateau, and northwest Mexico. The absence of Lungless Salamanders from these arid and semi-arid regions is perhaps to be expected for amphibians that absorb oxygen through moist skin and lay eggs in damp soil.

What is surprising is that three species of plethodontid salamanders persist in isolated mountain ranges within this vast arid gap. Two of these are found in New Mexico: Aneides hardii (Sacramento Mountains Salamander; Fig. 3) found only in the high elevation wet coniferous forests of the Sacramento, Sierra Blanca, and Capitan Mts. (Lincoln and Otero Cos.); and Plethodon neomexicanus (Jemez Mountains Salamander) found in rock slides high in the Jemez Mts. (Sandoval and Los Alamos Cos.; Stebbins 2003; Wake 1965; Williams 1973). Studies of allozymes, albumin immunology (Larsen et al. 1981) and mitochondrial DNA (Mahoney 2001) support earlier morphological work (Lowe 1950; Wake 1963, 1966) that concluded these two New Mexico species are ancient geographic relicts that diverged from other species of Lungless Salamanders in the Miocene, over 20 million years ago.

The third species that persists within the Plethodontid Gap is Tlaconete Pinto (Pseudoeurycea bellii). This magnificent plethodontid is widely distributed in southern and central Mexico, extending north along the Sierra Madre Oriental to the northern-most cloud forests in Tamaulipas (Martin 1958). In western Mexico, Dunn’s (1926) distribution map of the species shows it no further north than Nayarit, and Taylor (1938, 1941) concluded that the lost specimens cited by Dunn from Fort Whipple, Arizona, were questionable. However, when Lowe (1955) reviewed the salamanders of Arizona, he considered that field work to that date had been inadequate to rule out the possibility that plethodontids may occur in moist coniferous forests at high elevations in the state (especially in the Pinaleño and White Mountains) as they do in New Mexico.

The possibility of an Arizona Lungless Salamander received a great boost by the events of 1964. In September of that year, a field party of four (Thomas J. Cox, John W. Wright, Kenneth L. Hale, and Clyde J. Jones) attempted to reach Yecora, Sonora, by truck, but were forced to proceed on foot. During the return hike back to the truck, Clyde Jones collected a small black salamander which was brought to the University of Arizona on 5 September. On the following morning a second field party (Charles H. Lowe, Clyde J. Jones, and John W. Wright) were flown to Yecora by David Vactor in a Cessna 180, where they proceeded (during a lengthy rainstorm) via pack animals to the locality where the first salamander had been taken. On 7 September they succeeded in collecting six additional salamanders (details as reported by Lowe et al. 1968). The seven salamanders were found to be Pseudoeurycea bellii. This locality in the northern Sierra Madre Occidental of Sonora is ca 880 km (550 mi) airline distance northwest of the nearest known localities for the species in Nayarit and 770 km (480 mi) SE of Dunn’s lost USNM specimens from Fort Whipple, Arizona. In addition to representing a spectacular range extension, the population was described as a separate subspecies, P. b. sierraoccidentalis, characterized by having dorsal red blotches that are few in number and irregular in shape and position (Lowe et al. 1968).

In the subsequent 40 years since this discovery, only one additional locality has been reported for Pseudoeurycea bellii sierraoccidentalis. On 24 June 1987, Tom Van Devender and Peter Holm observed four adults in the Ocampo area, providing the first record of a plethodontid salamander from the state of Chihuahua and the second known locality for the subspecies (Van Devender et al. 1989).

The discovery of the population of Pseudoeurycea bellii in the northern Sierra Madre Occidental of Sonora led Lowe et al. (1968) to call for a renewed search for relictual populations of plethodontid salamanders in the Southwest in “ranges such as the Bradshaws, Chiricahuas, Pinaleños, and Mogollons, which are Cordilleran outliers positioned between the Sierra Madre and the Rockies.”

In spite of all the water that has flowed under the bridges since Dunn (1926) cited the three lost uncatalogued specimens of Pseudoeurycea bellii from Fort Whipple, no additional Lungless Salamanders have been found in Arizona. We are left with many unanswered questions: What happened to the specimens Dunn cited? Were they actually collected at Fort Whipple and by whom? Were they Pseudoeurycea bellii? Do plethodontid salamanders remain hidden in some remote Arizona mountain range? The mystery remains unresolved, but Dunn has left us with an enduring vision of what he may have intended by “mountains seen and unclimbed and forever remembered.”

Acknowledgments. We are grateful to Chuck Lowe for inspiration; Kit Bezy and Kate Bolles for participation in field work in New Mexico and for review of an earlier draft of this paper; Richard Felger, George Ferguson, and Tom Van Devender for invaluable information about Sonora; George Bradley for access to specimens in the University of Arizona Herpetological Collection; Darrel Frost and Dave Wake for information on nomenclature; and Kit Bezy for production of the map.

Literature Cited

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