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Full Issue, Vol. 65 No. 2 Western North American Naturalist Volume 65 Number 2 Article 21 4-29-2005 Full Issue, Vol. 65 No. 2 Follow this and additional works at: https://scholarsarchive.byu.edu/wnan Recommended Citation (2005) "Full Issue, Vol. 65 No. 2," Western North American Naturalist: Vol. 65 : No. 2 , Article 21. Available at: https://scholarsarchive.byu.edu/wnan/vol65/iss2/21 This Full Issue is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Western North American Naturalist 65(2), © 2005, pp. 145–152 NOTES ON THE BIOGEOGRAPHY AND DORSAL COLORATION OF CICINDELA AMARGOSAE DAHL (COLEOPTERA: CARABIDAE) Michael G. Kippenhan1,2 ABSTRACT.—The widely distributed and fragmented populations of the tiger beetle Cicindela amargosae are docu- mented for dorsal coloration, elytral maculation, habitat, and adult escape behavior. Currently, there are 2 recognized subspecies, C. a. amargosae and C. a nyensis. The analysis of populations indicated that the variation in dorsal coloration did not coincide with the accepted subspecific criteria for this species, thus illustrating the difficulty in applying a sub- specific category unequivocally to tiger beetles. Key words: Cicindela amargosae, tiger beetle, subspecies, phenotypic variation, habitat. Cicindela (Cicindelidia) amargosae Dahl is 1982, Werner 1994); however, Leffler (1979) a polytypic species associated with grass mar- presented distinguishing morphological and gins of moist, often alkali-encrusted areas of geographical attributes for the 2 species. Apart drainages in southern Oregon, western Nevada, from studies by Dahl (1939) and Rumpp (1956), and eastern California (Fig. 1, Table 1). There are C. amargosae has received little attention in 2 recognized subspecific forms: Cicindela entomological literature except for Leffler (1979), a. amargosae and C. a. nyensis Rumpp. Dahl who included this species as part of the Pacific (1939) described C. willistoni amargosae from Northwest tiger beetle fauna, and Freitag (1999), “four miles north of Furnace Creek, Death who listed all populations for this species out- Valley, Inyo County California.” This subspecies side Death Valley as C. a. nyensis. was separated from other geographical forms In a geographic outline of the populations of the C. willistoni group by the combination of C. amargosae, Rumpp (1956) found C. a. of bright blue-green dorsal coloration, macula- amargosae, along with C. w. pseudosenilis and tion pattern reduced to an apical lunule, and C. californica pseudoerronea Rumpp, isolated elytral punctation (Dahl 1939). Interestingly, along the natural springs associated with the Dahl (1939) believed this species to be a sub- alkali flats north of Furnace Creek, Death Valley, species of C. willistoni even though C. willis- California. Cicindela a. amargosae was not found toni pseudosenilis W. Horn was sympatric at downstream of Furnace Creek at Saratoga the type locality. In a study of the Death Val- Springs, even though C. w. pseudosenilis, C. c. ley, California, tiger beetles, Rumpp (1956) pseudoerronea, and a 3rd species, C. n. nevadica elevated C. amargosae to specific rank based LeConte, were present (Rumpp 1956). Inter- on the observations that it did not interbreed estingly, Rumpp’s tiger beetle collection at the with C. w. pseudosenilis at the type locality California Academy of Sciences includes a and there was a lack of hybrids. Utilizing color series of C. a. amargosae collected at Saratoga and body length as diagnostic criteria, Rumpp Springs in 1963. Rumpp (1956) found C. a. (1956) described C. amargosae nyensis from nyensis associated with the intermittent chan- “1.6 miles south of Springdale, Nye County, nels of the Amargosa River near Springdale, Nevada.” In addition to the allopatry from the Nye County, Nevada. Although the Death Val- nominate form, this subspecies was character- ley and Springdale populations are in close ized by its matte-black dorsal coloration, “softer” proximity to one another (<80 km), small elytra relative to the nominate form, and mountain ranges apparently are geographical smaller overall body length. Various authors barriers between the 2 type localities. Rumpp have considered C. amargosae to be a sub- (1956) believed that populations connected to specific form of C. senilis G. Horn (Boyd et. al the Springdale and Furnace Creek populations 11425 S.E. Claybourne St., Portland, OR 97202. 2C.P. Gillette Arthropod Biodiversity Museum, Colorado State University, Fort Collins, CO 80523. 145 146 WESTERN NORTH AMERICAN NATURALIST [Volume 65 (1956) assertion that dorsal color variation was due to hybridization. Because most cicindelid taxonomy relies solely on morphological char- acters to determine subspecific status, the con- sideration of ecophenotypic characters and their role in cicindelid color expressions (Pearson Lk. Alvord Lk. Warner and Vogler 2001, Schultz 2001) is often left un- 1 explored. The objective of this study is to review 2 the current distribution and habitat of C. amar- gosae populations throughout its range while 3 evaluating the correlation between distribu- Lk. Surprise tion and dorsal coloration and maculation. 4 METHODS AND MATERIALS 5 Lk. Lahontan 6 I reevaluated the criteria utilized by Rumpp (1956) to differentiate the subspecific forms of 7 C. amargosae. These include (1) geography 8 and allopatry, (2) dorsal coloration and elytra Lk. Rennie 10 maculation, (3) habitat and adult escape be- 9 havior, and (4) total body length. To evaluate 11 each of these criteria, I collected individuals AmargosaAmargosae R. R. of C. amargosae from locations throughout the 12 species range. In addition to specimens cap- 13 tured in the field, pinned material including the type of C. a. nyensis and paratypes of C. a. amargosae were examined from the California Academy of Sciences Collection (San Fran- cisco, California). RESULTS AND DISCUSSION Fig. 1. Known populations of Cicindela amargosae and pluvial lakes (numbers correspond to Table 1). Geography and Allopatry Vestiges of the ancient lakes that once occu- by the Amargosa River would have individuals pied the Great Basin offer an understanding of expressing a variety of dorsal coloration indica- the present-day distribution of C. amargosae tive of hybridization. Accordingly, the popula- (Fig. 1). Leffler (1979) believed that C. amar- tions downstream of Springdale at Ash Mead- gosae inhabited the shores of pluvial Lake ows, Nye County, Nevada, were categorized as Lahontan and associated lake basins. The “hybrid,” inasmuch as individuals matched the reduction of these pluvial lakes in post-Pleis- description of both subspecific forms (Rumpp tocene times to smaller remnant lakes (Reheis 1956; Table 2). In addition, Rumpp (1956) con- 1999) can be considered a valid explanation for sidered as “hybrids” populations from north- the widespread and fragmented distribution of western Nevada and adjacent California that current populations in the northern half of the exhibited dorsal coloration described as “black, species range. Historically, it is likely the local- green and bronze.” Rumpp (1956) apparently ized distribution of the Death Valley, Califor- was unaware of C. amargosae populations out- nia, populations had a much more widespread side California and Nevada; however, Leffler range along the shores of pluvial Lake Manly, (1979) examined populations from Lake and which at one time was close to 161 km long Harney Counties, Oregon, all of which were (Sharp and Glazner 1997). Populations close to placed as C. a. nyensis. Tecopa, Inyo County, California, are associated During a study of the current distribution with remnants of ancient Lake Tecopa, which of C. amargosae in 2002 and 2004, I made num- occupied the area south of Death Valley until erous observations that contradicted Rumpp’s 500,000 years ago (Sharp and Glazner 1997). 2005] BIOGEOGRAPHY OF CINCINDELA AMARGOSAE 147 TABLE 1. Known populations of Cicindela amargosae (arranged north to south). Location Source 1. OR: Harney Co., Alvord Hot Springs MGKC, CSUCa 2. OR: Lake Co., Crumb Lake Leffler 1979b 3. CA: Modoc Co., Surprise Lake CASCc 4. NV: Wascoe Co., Gerlach CASC, MGKCd 5. CA: Lassen Co., Honey Lake CASC, MGKC 6. NV: Washoe Co., Truckee Meadows LaRivers 1946 7. NV: Esmeralda Co., Fish Lake CASC 8. NV: Nye Co., Springdale CASC, MGKC 9. CA: Inyo Co., Death Valley Nat. Mon., Furnace Creek CASC, MGKC 10. NV: Nye Co., Ash Meadows NWR CASC, MGKC 11. CA: Inyo Co., Shoshone CASC 12. CA: Inyo Co., Tecopa Hot Springs CASC 13. CA: Inyo Co., Death Valley Nat. Mon., Saratoga Springs CASC aColorado State University Collection, Fort Collins, CO bLiterature sources only cCalifornia Academy of Sciences, Golden Gate Park, San Francisco, CA dMichael G. Kippenhan Collection, Portland, OR The Death Valley and Tecopa populations, along implications of these 2 color forms. In an eco- with populations not directly associated with logical role, color, elytral maculation, and ven- pluvial Lake Lahontan, such as Springdale and tral setae function as the primary mechanisms Ash Meadows, Nye County, Nevada, may have by which adult tiger beetles regulate body arrived via watercourses originating from the temperature (Schultz and Hadley 1987, Pear- shores of pluvial lakes and are currently asso- son and Vogler 2001). While C. a. amargosae
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