Great Basin Naturalist

Volume 47 Number 2 Article 2

4-30-1987

Zoogeography of Great Basin : patterns of distribution and differentiation

George T. Austin Nevada State Museum and Historical Society, Las Vegas

Dennis D. Murphy Stanford University

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Recommended Citation Austin, George T. and Murphy, Dennis D. (1987) "Zoogeography of Great Basin butterflies: patterns of distribution and differentiation," Great Basin Naturalist: Vol. 47 : No. 2 , Article 2. Available at: https://scholarsarchive.byu.edu/gbn/vol47/iss2/2

This Article 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 Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. ZOOGEOGRAPHY OF GREAT BASIN BUTTERFLIES: PATTERNS OF DISTRIBUTION AND DIFFERENTIATION

George T. Austin and Dennis D. Murphy"

Abstract. —The butterflies of the Great Basin exhibit general patterns of distriljution and speciation similar to those found for other taxa, particularly birds. Two major centers of infraspecific differentiation and coinciding distribution limits of taxa are identified, each with three subregions. Great Basin butterflies are characterized by pallidity and substantial endemism below the level.

The Great Basin of western North America Materials and Methods is a huge area, nearly 520,000 square kilome- Distribution maps for taxa and ters, of largely internal drainage between the other distinct phenotypes occurring within Rocky Mountains to the east and the Sierra and on the margins of the Great Basin were Nevada to the west. It includes Utah west of constructed from a variety of sources. Nevada the Wasatch Plateau, extreme southwestern data are drawn primarily from the collections Idaho and southeastern Oregon, California and field notes at the Nevada State Museum, east of the Sierra Nevada, and nearly all of Carson City, the senior author's personal col- Nevada (Fig. 1). Elevations range from lection, and collections made by the Center l,000-m lowlands dominated by sagebrush for Conservation Biology at Stanford Univer- {Artemisia) and saltbush {Atriplex) to numer- sity. Eastern California data were obtained ous, mostly north-south oriented mountain from the notes and collections of a number of ranges which may exceed 3,000 m. These private collectors. Southern Oregon records mountain ranges, most of which are forested are from Dornfeld (1980), and Rocky Moun- only at the higher elevations, constitute is- tain and eastern Great Basin records are from lands of boreal habitat. Lowland wet areas are Ferris and Brown (1981). Some Sierra Nevada similarly islandlike. The area is largely unin- data were obtained from Shapiro et al. (1979) habited by and is relatively undis- humans and the collections of the Nevada State Mu- livestock has turbed except for grazing which seum. Numerous other literature sources had substantial impact on the composition of were consulted. the vegetation, especially at lower elevations The maps thus prepared were examined to (e.g., 1982, 1983). Rogers Thomas determine patterns of distribution within the Studies of the distribution and biogeogra- Great Basin and adjacent areas. Attention was phy of Great Basin biota have dealt largely paid to the absence or presence of species with vertebrates (e.g., Behle 1963, 1978, within the Great Basin and the extent of their Brown 1971, 1978, Grayson 1982, 1983, John- apparent distributions and differentiation in son 1975, 1978, Smith 1978) and plants (e.g., the Great Basin.

Billings 1978, Harper et al. 1978). Here we present information on the distribution of Taxa and Distribution Great Basin butterflies, paying particular at- tention to the distributional limits of species, The 155 butterfly species occurring in the subspecies, and well-differentiated segre- Great Basin include some 240 subspecies and gates, and to centers of infraspecific differenti- well-differentiated segregates. More than half ation. Additionally, we discuss the role of "is- the species are geographically polytypic in land" effects in shaping local species richness. this and adjacent regions, including the Rocky

Nevada State Museum and Historical Society, 700 Twin Lakes Drive, Las Vegas, Nevada 89107. Department of Biological Sciences, Stanford University, Stanford, California 9430.5.

186 April 1987 Austin, Murphv: Great Basin Butterflies 187

OREGON IDAHO [ NEVADA f WARNER Jarbidge Mountains

I SUBREGION Independence Mountains

JARBIDGE SUBREGION •KEast Humboldt Mountains

^ i CENTRAL IF Ruby Mountains SUBREGION

Virginia ^[/ ^ Range

, Pine Nut 3|l Mountains jl A Snake Range

Wassuk

?V,^ Range

. TOIYABE \ SUBREGION INYOV^ SNAKE ^^ SUBREGION SUBREGION White Mountains

• ..••' ^-1 \

ARIZONA MOJAVE SUBREGION

Spring Range ^ \ /

Fig. 1. The Great Basin showing subregions and locations mentioned in the text.

Mountains and Sierra Nevada. No species are stricted to the Great Basin. A number of addi- endemic to the Great Basin, consistent with tional groups of populations within the region previous findings for birds (Behle 1963). show some measurable diflerentiation. The About 50 subspecies and other well-differen- distributions of these taxa and segregates by tiated infraspecific segregates (distinct groups geographic affinities are summarized in Table 1. of phenotypically similar, but unnamed, pop- Nearly 90% of all Great Basin butterfly spe- ulations) of butterflies, however, are re- cies are also found in the Rockv Mountains. 188 Great Basin Naturalist Vol. 47, No. 2

Table 1. AflFinities of the Great Basin butterfly fauna. Taxa include subspecies and distinct unnamed segre- gates.

Species April 1987 Austin. Murphy: Great Basin Butterflies 189

Table 3. Distribution of Great Basin endemic butter- Table 3 continued. flies by region.' Speyeria zerene gtindeh Eastern Region Limenitis archipptis lahontani Jarbidge sthenele ))atilus Ochlodcs sijlvatwides bonneviUa Neominois ridingsii stretchii *Lycaeua eclitha nevadensis Here and in subsequent tables, seg. (segregate) is used to denote distinct * ritti mattonii sets of phenotypically similar populations which are as yet unnamed. *Speycha atlantis urcyi *Narrowl\ distributed ta.\on *Spcycria atlantis clko Speijcria mormonia ai-tonis Phyciodes campestris seg. est extant populations apparently are now Etiphydryas colon nevadensis well to the north. Papilio indra, in addition, Snake exists as a relatively isolated endemic subspe- Polites sabtileti seg. cies in the general area, Plebejus Satyritun sacpium seg. same and

*Incisalia au^ttsttts (?) seg. lupini occurs as widely scattered populations *Euphilotcs hattoides seg. across central Nevada. Several of these same Toiyabe species also extend into montane areas south Polites sabtileti seg. of the Sierra Nevada cordillera (Emmel and Papilio indra nevadejisis Emmel 1973), indicating an ability to survive *Speyeria e^leis toiyabe in more xeric conditions than those at their * pallesccns distribution centers. Widespread Euphydryas editha lehmani Etiphydryas editha koreti Centers of Differentution Western Region A number ol Great Basin species are com- Inyo paratively unvarying in phenotype over a mexicana blanca broad area from the Rocky Mountains or east- *Hesperia miriamae seg. *Polites sabtileti seg. ward, west to the Sierra Nevada or beyond. Ltjcaena rtibidtis seg. Others exhibit considerable regional differen- langstoni tiation and may include one or more pheno- *Plebejiis icarioides seg. types restricted to the Great Basin. The large *Plebejtis saepiohis seg. Coenonympha ochraceae mono number of phenotypic endemics suggests that *Cercyonis pegala tvheeleri the Great Basin is at least a moderately active Neominois ridingsii seg. area of infraspecific differentiation. Examina- Central tion of the distributions of subspecies and seg- *Pseiidocopaeodes eitntis seg. regates of polytypic species in the Great Basin *Polites sabtileti genoa and adjacent butterfly faunas shows rather *Etiphilotes rita seg. Speyeria zerene malcolmi well defined distribution patterns suggesting Speyeria callippe nevadensis "centers of differentiation. ' Similar to Behle s Etiphydryas editha monoensis (1963) findings for birds, these centers are *Cercyonis pegala seg. bounded by areas where numerous range lim- Warner its coincide, further suggesting that the Great Polites sabtileti seg. Basin consists of definable biogeographical *PoUtes sabtileti seg. units (Fig. 1, Table 3). These regions gener- *Pieris napi seg. rtibidiis rtibidtis ally coincide with distril)utional limits or more Cercyonis pegala stephensi widespread butterfly taxa and are strikingly Widespread similar to distributional centers found for Hesperia tineas macstvaini birds (Behle 1963, 1978).

Widespread in Gre.at Basin Eastern Region Colias alexandra edicardsii Lycaena arota virginiensis The Great Basin may be viewed as two Mitotira siva chalcosiva distinct centers of butterfly distribution and baiieri differentiation (Fig. 1). The first is the Eastern Etiphilotes rita pallesccns Glaticopstjche piasiis ncvada Region bounded by the Wasatch Front in the Speyeria nokomis apacheana east, to and including the Reese River Valley —

190 Great Basin Naturalist Vol. 47, No. 2 and from the northern Hmits of the Mojave Subregion (southwestern Lahontan basin) Desert in the south, north to southern Idaho including the White Mountains and adjacent and southeastern Oregon. The area inchides areas, Wassuk and Sweetwater mountains, the Pleistocene Lake Bonneville basin, east- and adjacent east slope of the Sierra Nevada of ern portions of the Pleistocene Lake Lahontan Nevada and California; (2) the Warner Subre- basin, the Ruby group of drainages, and the gion (southern Oregon Lakes drainage)—in- southern portion of the Snake River drainage cluding northeastern California, northwest- group (see Smith 1978). This region is com- ern Nevada, and south central Oregon; and (3) prised of three subregions: (1) the Jarbidge the Central Subregion (west central and Subregion (southern Snake, northern Bon- northwestern Lahontan basin)—the area be- neville, and northeastern Lahontan drain- tween the above (Fig. 2). Behle (1963) ex- ages)—including the area north of the Hum- cluded, but later included (Behle 1978), the boldt River to central Humboldt County in Inyo and Warner subregions in the Western Nevada, adjacent southeastern Oregon, Region and discussed them as separate bio- southern Idaho, and northwestern Utah; (2) geographic entities (see also Miller 1941, the Snake Subregion (Ruby and southern Johnson 1970). Bonneville drainages)—including the remain- The Inyo Subregion is the most well de- der of the Great Basin along the eastern bor- fined center of butterfly differentiation in the der of Nevada and western Utah; and (3) the Western Region (Table 3). Speciation in this

Toiyabe Subregion (southeastern Lahontan area is greatest in the White Mountains where drainage)—including the central portion of at least one endemic subspecies and four en- Nevada. The Eastern Region loosely corre- demic segregates are recognizable. Another sponds to Behle's (1963) concept; however, he subspecies is restricted to the Owens Valley. did not subdivide the region, and he included An additional four subspecies are more widely more of Idaho. distributed in the subregion. The most clearly defined center of differen- The Warner Subregion has at least two en- tiation in the Eastern Region is the Jarbidge demic subspecies and three endemic segre-

Subregion. Three subspecies are narrowly re- gates. The Central Subregion is geographi- stricted to the Jarbidge and Independence cally broad and not sharply defined. There are ranges and another to the Ruby and East three restricted segregates and one restricted

Humboldt ranges (Table 3). Three other sub- subspecies and three more widely ranging species and one segregate are distributed subspecies, some of which extend for varying more broadly in the subregion. The Snake distances into the Inyo and/or Warner subre-

Subregion has two apparent narrowly dis- gions. One additional subspecies is relatively tributed segregates in the vicinity of the widespread throughout the Western Region. Snake Range and two others more widely dis- Finally, 11 endemic Great Basin subspecies tributed. The Toiyabe Subregion has two nar- occur in at least one (usually more) subregion rowly distributed subspecies and one subspe- of both the Eastern and Western regions. cies and one segregate more widespread. Two other endemic Great Basin subspecies are Speciation Phenomena more widely distributed in the Eastern Re- gion, ranging into two or more of the subre- Zones in which subspecies or segregates gions. interface are found throughout the Great Basin. Some, however, emerge prominently Western Region as areas of intergradation for a wide variety of This region includes the area from the west- species when distribution and differentiation ern edge of the Eastern Region (defined patterns are examined. Similar phenomena above) to the east slope of the Sierra Nevada, were identified and discussed for birds by north from the Mojave Desert to south central Johnson (1978). Here we follow that presenta-

Oregon (Fig. 1). The area includes the west- tion for butterflies. Areas where speciation ern portion of the Pleistocene Lake Lahontan appears to be less obvious coincidently have basin and the southern Oregon Lakes been less well studied. But, while further drainage group (Smith 1978). Again three sub- knowledge may somewhat alter the details, regions may be discriminated: (1) the Inyo the overall definition of these zones and the April 1987 Austin. Murphy: Great Basin Butterflies 191

MOJAVE ZONE 6

02

\ 3o

( \.v O^ \ ^

\ \

Fig. 2. Areas of Interaction among Great Basin butterflies. Numbers refer to species pairs in Table 4 (Sierra Nevada), Table 5 (northeastern Nevada), Table 6 (eastern and central Nevada), and Taljle 7 (Mojave). Solid symbols refer to interspecific hybridization; open symbols refer to intraspecific intergradation. 192 Great Basin Natur.\list Vol. 47, No. 2 interactions described appear to be sound. The east slope of the Sierra Nevada and the Note also that there are some recent records adjacent western Great Basin, as a conse- of shifts in the ranges of birds in these same quence, might be expected to be an area of geographic areas of interaction (e.g., Johnson vigorous interaction among distinguishable and Johnson 1985). butterfly taxa which may have only recently

Pairs of related taxa or segregates are cate- come into contact. This is the single most gorized by the type and degree of interaction. active area of intergradation for Great Basin In the following sections we discuss areas birds (Johnson 1978, Johnson and Johnson where subspecies or segregates and closely 1985), and the same appears to be true for related species come into contact. A number butterflies (Table 4). These interactions in- of closely related species hybridize in these clude many species and involve a wide variety areas. Intergradations between subspecies or of types and degrees of differentiation and/or segregates include primary intergradations disjunct distributions within this geographic

(those between phenotypically similar sub- area (Fig. 2). species or segregates such as those along a Geographic areas of contact within the cline) and secondary intergradations (those Sierra Nevada Zone are generally narrow. between phenotypically more dissimilar sub- Sierran Speyeria zerene zerene and a western species segregates such as "internal contact" Great Basin subspecies, S. -.. malcohni, for of those from distant points on a ring or example, intergrade (a primary intergrada-

rassenkreis) . Also, in the same geographic tion) only in the vicinity of Carson Gity areas some species and subspecies (or segre- (Moeck 1957, Grey and Moeck 1962). The gates) exhibit range disjunctions. These zones more widespread Great Basin subspecies, S. of allopatry may be geographically wide (such z. gunderi, intergrades with S. z. conchyliattis as across the Lahontan Basin) or narrow (be- in the Granite and Warner mountains on the tween adjacent mountain ranges). Taxa also northern Nevada-California border (a sec- can be separated by elevation (high and low ondary intergradation between Rocky Moun- populations in the same mountain range) or tain and Sierra Nevada subspecies [Grey and time (flying at different seasons). Moeck 1962, Grey 1972]). Two Speyeria cal-

lippe phenotypes (S. c. nevadensis and S. c. Sierra Nevada Zone near semivirida ) intergrade in this same area. Perhaps the most striking element of Great A population of Neominois ridingsii, appar-

Basin biogeography is the predominance of ently intermediate between N. r. stretchii Rocky Mountain and closely related Great and an as yet undescribed Sierra Nevada seg- Basin taxa in relative proximity to the east regate, occurs here as well. Euphydnjas ani- slope of the Sierra Nevada. The occurrence of cia wheeleri and E. chalcedona macglashanii Sierra Nevada biotic elements east into the produce an apparently intermediate popula-

Great Basin, conversely, is rare (Behle 1978, tion in the Sweetwater Mountains (Murphy

Harper et al. 1978, Johnson 1978, Tanner and Ehrlich 1983), while E. anicia macyi and 1978). For instance, in the Snake Range, cen- E. a. veazieae intergrade across a broad area trally located in the Great Basin, 54% of 86 along the Oregon and Nevada border. species are represented by subspecies that are The extreme western Great Basin ranges either shared with the Rocky Mountains or additionally have some Sierra Nevada- are Great Basin subspecies or segregates most derived taxa or segregates which are pheno- similar to Rocky Mountain ta.xa. Only 3% of typically distinct. Thoryhcs mexicana blanca species are of Sierra Nevada affinity. In the of the White, Wassuk, and Sweetwater moun- Toiyabe Range in the west central Great tains and undescribed Hesperia miriamae and Basin, 50% of 92 species have Rocky Moun- Lycaena ruhidus segregates restricted to the tain affinities, while 4% are similar to Sierra White Mountains are examples of populations Nevada taxa. And, in the White Mountains, closely related to Sierra Nevada taxa. The lat- just several dozen kilometers from and in di- ter appears related to the Sierran subspecies, rect sight of the Sierra Nevada crest, 34% of 79 L. r. monochensis, and is replaced elsewhere species are most similar to Rocky Mountain in the Great Basin by the widespread L. r. taxa, and only 17% are of Sierra Nevada affin- sirius. ity. Some species or subspecies (or segregate) April IBS- Austin, MuHPHY: Great Basin Butterflies 193

Tablp: 4. Taxa pairs of butterflies that sliow various Table 4 eontinui'd. speciation phenomena in the western Great Basin/east slope Sierra Nevada region (most widespread Great Basin 40. Euphydryas editha lehmani and £. c. monoensis taxon hsted before Sierra Nevada or other taxoii). 41. Cocnonymp}ui ochracea mono and C. ochraceae b rendu interspecific hybridiza- Narrow zone of sympatry and AlLOCHRONIC SYMP.\TRY BETW EEN REPRESENT.\TI\'ES of DI- tion: VERGENT SUBSPECIES: A/, nelsoni nclsoni 1. Mitoura siva chalcosiva and 42. Euphilotcs battoides baucri and E. b. glaucon 2. Etiplujdryas anicia whecleri and E. ciuilccdona mac- 43. Euphilotcs enoptes ancilla and E. e. enoptes glashanii 3. Limenitis weidemeyerii latifascia and L. lorquini Narrow zone of sympatry and intergradation be- pairs are narrowly synipatric, or nearly so, tween REPRESENTATIVES OF DINERGENT SUBSPECIES: with little or no hybridization or intergrada- 4. Hesperia comma harpahts and H. comma ijosemite tion in this zone. The closely related Chlosyne 5. Anthochahs sara thoosa and A. s. stclla palla and C. acastus appear to be sympatric at 6. Lycacna arota vir

12. Spcycria zcrcnc ^tindcri and S. z. conchyliatit.s may be different species). Another species 13. Spcycria callippc ncvadcnsis and S. c. scmivirida pair, Limenitis wcidemcijehi and L. lorquini, 14. Phyciodcs campestris campestris and P. c. montana hybridizes in a very narrow zone just east of 15. Ncominois riding.^ii stretcliii and N. ridingsii seg. the Sierra Nevada (Perkins and Perkins 1967) zone of allopatry between closely related Narrow with extensions northward into Idaho and SPECIES: southwestern Alberta. Yet another pair, Mi- 16. Clilosync acasttts acastus and C. palla seg. 17. Coenonympha ochracea mono and C. ampclo.s am- toura siva and M. nelsoni, have long been pclos considered distinct species. They, however, Narrow zone of allop.iiTRY between represent.atines hybridize in a broad region in the western of divergent SUBSPECIES: Great Basin and hence may be one species. 18. Thorybcs mcxicana blanca and T. mcxicana nciada Isolated high-elevation populations of at 19. Hesperia miriamac seg. and H. m. miriamac least two species, Polites sabuleti and Phy- 20. Politcs .mbulcti ncnoa and P. s. tccum.sch (ele\ational)

21. Pontia .sisymbrii elivata and P. s. sisyvd)rii ciodcs campestris, exist in the Sierra Nevada 22. Euchloe hyantis lotta and E. hyantis ssp. bounded on both the east and west by more 23. Lycacna cditlia nevadensis and L. c. editha widespread, lower-elevation subspecies. Two 24. Lycacna rubidus sirius and L. rubidus ssp. (ele\a- other species, Euphydnjas editha and E. tional) exist as a series of elevational sub- 25. Satyrium fulininosum scmihina and S. fuUfiinosum chalcedona, ssp. species (perhaps ecotypes) on the west slope 26. Satyrium californica seg. and S. c. cyfinus to the crest of the Sierra Nevada and as a

27. Satyrium sylvinus seg. and S. .s-. syJvinus single middle-elevation subspecies on the 28. Strymon niclinus pudica and S. m. setonia east slope and into the western Great Basin. 29. Glaucopsyche piasus nevada or G. /;. toxcuma and G. p. piasus Numerous Great Basin subspecies (or segre- 30. Plebejus melissa melissa and P. m. fridayi or P. m. gates) are "replaced" by Sierra Nevadan taxa paradoxa (elevational) between the western portion of the Great 31. Plebejus sacpiolus sacpiohis and P. sacpiobis seg. Basin and the crest of the Sierra Nevada (elevational) (Table 4). There is usually narrow elevational 32. Plebejus shasta minnehaha and P. s. sha.sta 33. Euphydryas editha monoensis and P. p. aurilacus or allopatry and/or allochrony (imposed by ele-

P. t'. nubig,ena (elevational) vational differences in phenology) between 34. Euphydryas anicia uhceh^ri and £. a. vcazicac or £. these phenotypes, but intergradation occurs fl. nmcyi in some. Furthermore, both Euphilotcs 35. Cercyonis pegala seg. and C. p. stcphcn.'ii enoptes and E. battoides are represented by Broad zone of allopatry between representatix es of divergent subspecies: sympatric allochronic "populations." These

36. Polites sonora iitahensis and P. s. sonora distinct univoltine populations fly at single 37. Spcyeria cijbele letona and S. c. /t'fo locations at different times of the year and thus 38. Spcyeria egleis toiyabe and S. e. cgleis thus are reproductiveK' isolated temporally 39. Spct/cnV; >»or»i()n!« fl/toHi.S' and S. ;/i. mormonia (hence should constitute "allochronic species"). 194 Great Basin Naturalist Vol. 47, No. 2

Table 5. Pairs of butterfly taxa showing various specia- Table 6. Pairs of butterflies taxa showing various spe- tion phenomena in the northeastern Great Basin (most ciation phenomena in central (C) and eastern (E) Great widespread Great Basin taxon hsted first). Basin (the most widespread Great Basin taxon is Hsted

first). Narrow zone of sympathy between closely related SPECIES without HYBRIDIZATION: Narrow zone of sympathy and interspecific hybridiza-

1. Euphydryas anicia wheeleri and E. colon nevadensis tion: 1. Plehejus acmon texanus P. htpini lupini (C) Narrow zone of sympatry and interspecific hybridiza- and tion: Narrow zone of symp.\try and intergrad.\tion be- 2. ancilla and £. hattoides ^laitcon tween REPRESENT.ATIVESOF divergent SUBSPECIES: 3. Coenonynipha ochracea hrenda and C. ampelos elko 2. Pontia sisymbrii elivata and P. sisymbrii seg. (E, C) 3. Satyrium behrii crossii and S. b. behrii (C) Narrow (usually) zone of sympatry and intergrada- 4. Cehistrina ladon echo and /. cinerea (G) tion between representatives of divergent subspe- C. 5. Glaucopsyche piasus nevada and G. piasus daunia CIES: (E) 4. Colias alexandra edwardsii and C. a. astraea 6. Plebejus acmon texanus and P. a. acmon (G) 5. Plehejus acmon acmon and P. a. lutzi 7. Speyeria zerene gunderi and S. ~. platina (E) 6. Speyerianokornis apacheana iin(\S. n. nokomis 8. Phyciodes campestris campestris and P. c. camillus 7. Speyeria egleis titahensis and S. e. linda iC) 8. Phyciodes campestris campestris and P. c. camillus 9. Limenitis wcidemeyerii latifascia and L. ic. angusti- 9. Euphydryas editha lehmani and E. e. Iwtchinsi fascia (E) 10. Limenitis archippus lahontani and L. a. archippus Narrow zone of allopathy between hepresentatin'Es Narrow zone of allop.atry between closely related OF divergent SUBSPECIES: SPECIES: 10. Lycaena arota virginiensis and L. a. schellhachi (E) 11. Papilio bairdii and P. oregonius (may be conspecific) 11. Euphilotes battoides baueri and E. battoides seg. or Narrow zone of allopathy between representatives £. battoides nr. bernadino (C) OF divergent SUBSPECIES; 12. Euphilotes enoptes ancilla and E. enoptes seg. (G) 12. Anthocharis sara thoosa and A. sara hrowningi 13. Plebejus saepiohis saepiohis and P. s. gertschi (E) 13. Satyrium sylviniis seg. S. s. putnaini and (elevational) 14. pallescens and E. r. mattunii 14. Euphydryas editha lehmani and £. e. koreti (E, G) 15. atlantis greyi S. atlantis elko Speyeria and (elevational)

Broad zone of allopathy between representatives of 1.5. Neominois ridingsii stretchii and N. r. dionysus (G) dinergent subspecies: 16. Satyrium saepium provo and S. saepium seg. 17. Lycaena nivalis browni and L. n. nivalis southern Idaho, and southeastern Oregon, but for these latter areas few pertinent data Disjunctions between distinct species and exist. Information does exist for much of Elko between subspecies or segregates within the and Humboldt counties and northern Eureka same species are manifest in both narrow and and Lander counties in Nevada. This area is wide zones of allopatry in the Sierra Nevada considerably smaller in extent and lacks the Zone (Table 4). Some of these "gaps" are just a abrupt topographical and ecological disconti- few miles wide, such as between the eastern- nuity of the Sierra Nevada-Great Basin inter- most margin of the Sierra Nevada and the face. Nonetheless, some combination of fac- westernmost Great Basin mountain ranges. tors there promotes diflferentiation and But other gaps include much of the broad replacement. The region also marks the west- expanse between the eastern Sierra Nevada ern or southernmost extent of the distribu- and the mountains of central Nevada. Many tions of many species in the Great Basin (see species that range continuously across the re- below). gion north of the Great Basin are also absent in As in the Sierra Nevada Zone, there are this same broad area. Note that many gaps in replacements (specific and subspecific) with distribution more or less coincide with re- or without hybridization or intergradation and gions of intergradation and of overlap be- some, mostly narrow, allopatries (Table 5). tween pairs of taxa discussed above. While the zone of interaction along the Sierra

Nevada is east/west in orientation, that in Northeastern Nevada Zone northeastern Nevada is more complicated

Another area of substantial apparent incipi- (Eig. 2). The majority of interactions there ent speciation in the Great Basin is the north- involve east/west replacements of Rocky eastern portion of Nevada (Fig. 2). This area Mountain taxa with those of the Great Basin or should probably include northwestern Utah, Sierra Nevada. There are, however, several .

April 1987 Austin, Murphy: Great Basin Butterflies 195

Table 7. Pairs of butterfly taxa showing various specia- Eastern Nevada-Western Utah Zone tion phenomena at the transition Ix'tween the Great Basin and Mojave Desert (Great Basin taxon Hsted first). This region, which includes White Pine and Lincoln counties in Nevada and parts of adja- Narrow zone of sympathy and intergradation be- cent Utah, is a comparatively minor area of tween representatives of divergent subspecies: speciation and faunal replacement (Table 6, 1 Pyrfius communis communis and P. c. albescens (par-

tial elevational allopatr\', possil)l\ different species) Fig. 2). The apparent subspecific endemics

2. Hespcropsis lihija lena and H. /. Uhya are shown in Table 3. Most phenotypically 3. Anthocharis cethura cethuni and A. c. pinui identifiable replacements consist of Great 4. Mitoura siva chalcosiva and M. s. rhodope Basin subspecies or segregate replacing 5. Glaucopsijche hm.damus oro and G. hj^damus seg. (partial elevational allopatry) Rocky Mountain subspecies with minor in- 6. Euphtjdnjas anicia wheeleri and E. a. idcna tergradation. There is, in addition, some re- 7. pauhis and C. s. miisoni placement of desert subspecies or segregates Narrow zone of allopatry between closely related with subspecies or segregates which range SPECIES: widely north of this zone. This portion of the 8. Chlosyne acastus acastus and C. neutnoc^cui ncu- Great Basin is most noteworthy as a northern moegeni or western limit of the distributions of a num- Narrow zone of allopatry between representatives below). OF divergent SUBSPECIES: ber of taxa (see

9. Polites sabuleti sabuleti and P. s. chusca 10. Papilio indra nevadensis and P. indra n^aiiini or P. Central Nevada Zone indrci seg. This area includes the central Nevada 11. Euphilotes battoides baueri and E. b. imn-tini 12. Plebejus melissa melissa and P. mclissa seg. mountains and valleys and is another compar- 13. Apodemia monno mormo and P. mornw seg. (partial atively minor area of interaction among phe- elevational and seasonal allopatry) notypes (Table 6, Fig. 2). Many of the interac- Broad zone of allop.\try between closely related tions discussed for the previous two zones SPECIES: extend for varying distances into the Central 14. chlosyne acastus acastus and C. palla vallismoi-tis Nevada Zone. Both east/west and north/south Broad zone of allop.-vtry between represent.\ti\'es of interactions are involved. A particularly inter- divergent subspecies: 15. Plebejus icarioides ardea and P. icarioides seg. esting feature in this zone, and in other areas

16. Plebejus shasta minnehalui and P. s. charlestonensis to the north as well, is the apparent hybridiza- 17. Speyeria zerene gunderi or S. ;. malcolmi and S. ;. tion between two species of blues, Plebejus carolae acmon and P. lupini (Goodpasture 1973). The 18. Euphydryas anicia wheeleri and E. a. morandi zone, in part, forms the eastern edge of a 19. Eiincnitis archippus lahontani and L. a. obsoleta 20. Liinenitis weidemeyerii latifascia and L. w. nevadae broad gap or zone of allopatry between spe- cies which are present between here and the north/south replacements of taxa from Oregon Sierra Nevada (see above). or Idaho with generally widespread Great Mojave Desert-Great Basin Zone Basin taxa. One subspecies each of both Speyeria e^leis and S. atlantis extends into This area, including parts of Lincoln, Nye, this zone from the north and another from the and Esmeralda counties, Nevada, and Wash- east (Austin 1983). Furthermore, intergrada- ington County, Utah, is recognized as the tion of phenotypes occurs among at least northern limit of Mojave Desert plants (Beat- seven other subspecies pairs. For some of ley 1975, Meyer 1978) and birds (Behle 1978, these (e.g., Speyeria nokomis, Swisher and Johnson 1978), hence the southern limit of the Morrison 1969) this blending occurs over a Great Basin. Mammalian and herpetological broad area of the eastern Great Basin and distributions also support this as a distinct northwestern Colorado; for others (e.g., Eu- area of biological discontinuity (Hall 1946,

phydryas editha) the cline is quite narrow. Banta 1965a, b). Several butterfly species oc- Finally, hybridization apparently occurs be- curring widely in both areas exhibit different tween Euphilotes battoides and E. enoptes phenotypes on either side of this transition, (Shields 1977) and between the semispecies while others intergrade across this area (Table Coenonympha ampelos and C. ochracea of 7, Fig. 2). There is a zone of allopatry for some the C. tullia superspecies complex in this taxa and segregates between the Great Basin area. and Mojave Desert, but this zone is generally 196 Great Basin Naturalist Vol. 47, No. 2

Table 8. Rocky Mountain butterfly species extending that occur to the western limits of the Rocky west to the Sierra Nevada across the Great Basin. Mountains, (2) extreme western taxa extend- ing Hesperopsis libya no further east than the east slope of the Hesperia tineas Sierra Nevada, (3) taxa of mainly Rocky Moun- Colias (ilexandra tain affinity that occur to the eastern borders Lijcaena ruhidus of the Great Basin, then north across Idaho Mitoura siva and Oregon and, in numerous cases, south Speyeria nokomis Chlosync acastus into the Sierra, and (4) southern taxa that oc- Eiiphydryas anicia cur north to southern Nevada and/or south- Limenitis weidcmeycrii western Utah. Coenonympha ochracea Other species reach the limits of their Neominois ridingsii ranges somewhere within the Great Basin re- gion. This includes a number of butterfly taxa narrow. Only for Limenitis archippus is there that enter only the eastern portion of the a broad zone of allopatry; several hundred Great Basin and otherwise possess a distribu- kilometers separate L. a. obsoleta in the Colo- tional pattern like the species in (3) above. rado River drainage and L. a. lahontani along The limits of these latter two groups coincide the Humboldt River. closely with the zone boundaries discussed in the previous section on speciation. Wasatch Front Zone Few Sierra Nevada species extend into the The interface of the western escarpment of Great Basin and only Plehejus hipini, as men- the Rocky Mountains with the Great Basin in tioned above, for a substantial distance. The central Utah superficially presents topograph- remainder occur, for the most part, only in the ical and ecological contrasts comparable to western Great Basin ranges. Of the two appar- that of the Sierra Nevada zone. Nevertheless, ent endemic species of butterflies in the faunal replacement in this zone is not as strik- Sierra Nevada, Hesperia miriamae and Colias ing as along the western edge of the Great hehrii, only H. miriamae extends its distribu- Basin. Some endemic subspecies (or segre- tion into the Great Basin as a phenotypically gates) occur in this zone, and there is replace- distinct isolate found solely in the White ment of some Rocky Mountain taxa with those Mountains. Endemic Sierra Nevada subspe- of the Great Basin. A sizable number of Rocky cies also have made few inroads into the Great Mountain subspecies as discussed below, Basin. Among the approximately 20 primarily however, extend past this area well into the alpine or subalpine taxa, only Plehejus Great Basin. Widespread Great Basin butter- fraukUnii podarce (one record from the Vir- flies such as Hesperia comma harpahis, Pontia ginia Range) and Pohtes sabuleti tecumseh, sisymbrii elivata, Euchloe hyantis lotta (this Chlosyne w. whitneyi, and Euphydryas taxon may be a species in itself, separate from editha nuhigena (Sweetwater Mountains) ex- £. hyantis fide P. A. Opler), Lycaena ruhidus tend east into the Great Basin. The east slope sirius, Plehejus icarioidesfuUa, P. shasta min- of the Sierra Nevada, in turn, is the western nehaha, Speyeria coronis snyderi, and S. cal- distribution limit of at least 11 Rocky Moun- lippe harmonia range west from the Wasatch tain species (Table 8). Front across virtually the entire Great Basin, A number of Rocky Mountain species (some some as far as the east slope of the Sierra of which also occiu" in the Sierra Nevada) enter Nevada. the Great Basin only in northeastern Nevada

(Table 9). Most of these species have re- stricted Great Basin distributions and occur in Distributional Limits both the Sierra and Rocky Mountains. Nu- Distributional limits of butterflies in the merous additional species occur as isolates on Great Basin and adjacent areas exhibit repeat- many of the Great Basin ranges. ing patterns of particular interest. Some spe- Three species with primarily southern dis- cies, as mentioned, totally avoid the Great tributions, Hesperopsis alpheus, Anthocharis

Basin, occurring solely at its borders. This cetJiura, PhilotieUa speciosa, occur through- overall situation essentially results from four out much of the western Great Basin but not distinct distribution patterns: (1) eastern taxa the eastern. Several others extend to the east- April 1987 Austin, Muhfhy: Great Basin Butterflies 197

Table 9. Widespread butterfly species eTiterinti tlie Tabi.K 10. List and general distribution of Great Basin Great Basin only in tlie northeastern portion. pallid iiutterfly taxa.

Hesperia nevada Western Great Basin Parnasshis phocbus Thonjhi's mcxicana hlanca Papilio eurijmcdon Euphilotes rita svg. Pieris napi Speyeria zerene tmdcolmi ("zerene" ssp. group) Lijcacna cupreus Speyerid callippe nevadensis ("nevadensis" ssp. group) Lijcaena dorcas Coenonympha ochracca mono Speyeria ctjhele Cercyoni.s pc^ala steplwnsi Spcijcria atlantis Neominois ridingsii sag. Spctjehd inonnonia Phijciodes tharos Central Great Basin Polites sabideti seg. Speyeria egleis toiyahe Cercyonis oetus pallescens ern and central regions. None, however, Northeastern Great Basin reach northeastern Nevada except as strays or Ochlodes sylvanoidcs honncvilla Lycaena editha nevadensis nonpermanent populations. A number of spe- Speyeria atlantis greyi cies reach their northern distributional limits Speyeria atlantis elko {"irene" ssp. group) in southern Nevada, south of the Mojave Speyeria mormonia artonis Desert/Great Basin transition (Austin and Pliyciodes campestris seg. Austin 1980). Likewise, numerous Great Coenonymplia antpelos elko Basin species have their southern distribu- General Great Basin tional limits near that boundary. Nonetheless, Hesperia uncas lasus Incisalia eryphon seg. more than 10% of the butterfly species in the Speyeria nokomis apacheana Spring Mountains in extreme southern Ne- Speyeria zerene ounderi vada are of Great Basin affinity, and several Limenitis nrchippus lahontani endemic subspecies and segregates in this Cercyonis sthenele paulus range appear to be closely related to Great Basin taxa (Austin 1981). This suggests a more extensive southern distribution for much of taxa and segregates are restricted to the north- the Great Basin fauna in the past and agrees eastern region, seven are in western Nevada, with our knowledge of the vicissitudes in three are in central Nevada, and six are more Pleistocene climate (e.g., Martin and generally distributed. Some pallid subspecies Mehringer 1965, Wells 1983). Taxa with pri- and segregates are extremely restricted geo-

marily northern distributions (e.g. , alpine Co- graphically, such as Cercyonis oetus palles- lias, Boloria, Erebia, Oeneis), on the other cens, found only in small areas of the Reese hand, contribute very little in general to the River and Big Smoky valleys, and an unde- Great Basin fauna. However, three putative scribed Euphilotes rita segregate, found only "species," Papilio oregonius, Euphydnjas at Sand Mountain east of Fallon. White alka- colon, and Coenonympha ampclos (each con- line or other pale soil was suggested as the key specific with or siblings of more widespread to predator-mediated selection for a pale Great Basin species), enter the northeastern ground color for many of these species

region. One, C. ampelos , extends the furthest (Emmel and Emmel 1969, 1971, Emmel and south, well into western Nevada to the Carson Mattoon 1972, Wielgus and Wielgus 1974). River basin. This may be true for some nondesert species as well (e.g., Hovanitz 1940, 1941, Bagdonas

and Harrington 1979) and is supported by the PALLIDITi presence of extreme dark phenotypes of some At least 20 butterfly species exhibit their species in dark-background, marshy areas of most pallid phenotype in the Great Basin the Great Basin (e.g., Polites sabuleti in east- (Table 10). An additional three butterfly sub- ern Nevada). The presence of pallid pheno- species groups reach their extreme in pallidity types in much of the Great Basin, of course, is in the region. Linsdale (1938) and Hall (1946) also consistent with Watts (1968) findings as- noted a similar phenomenon in Nevada birds sociating lighter basal wing color with warmer and mammals. Seven of the pallid l:)utterfly thermal regimes. 198 Great Basin Naturalist Vol. 47, No. 2

Discussion now restricted to areas far north and west of that range. Furthermore, pika (Ochotona The Great Basin butterfly fauna substanti- princeps) remains have been recovered more generalities previ- ates many zoogeographic than 1,000 m lower in elevation than known at ously drawn for other taxonomic groups, par- present. Grayson (1982) implies that: (1) bo- ticularly birds, in this region. Foremost, there real mammals were widely distributed across is a general impoverishment of species rich- the lowlands, (2) extinction led to the present ness inward from the peripheries, especially absence of certain species on certain montane the Mountains westward. This from Rocky islands, (3) certain species became extinct on would be predicted from the similar distribu- all montane islands, and (4) there was no tion patterns recorded for plants (Billings Holocene recolonization. 1978, Harper et al. 1978), in light of the close For butterflies, we have only present-day association of butterflies and their larval host distributions to examine. Butterflies, like plants. Nevertheless, suitable habitat (includ- birds, are considerably more vagile than most ing adequate specific host plant availability) mammals; thus, it is not surprising that they appears to exist for many butterfly species show less-dramatic effects of island size and missing from portions of the Great Basin. isolation. That butterflies are more mobile

This impoverishment, as well as the previ- than mammals (but less so than birds) is re- ously noted endemism, presence of "relict" flected in the comparatively low slope associ- populations, and indications of recent extinc- ated with the species-area curves for butter- tions (Austin 1985), is consistent with an "is- flies from Great Basin mountain ranges land effect" (MacArthur and Wilson 1967). (Murphy et al. 1986, Wilcox et al. 1986). This situation in the Great Basin largely arises Hence, rates of interrange (interisland) dis- from the sequestering of biotic diversity in persal should be higher, and recolonization comparatively small and isolated patches of after extinction more frequent, in butterflies montane habitat surrounded by sagebrush- than in mammals. Nonetheless, a significant dominated desert. The insular biogeography, area effect is found for butterflies. But, sup- particularly area effects and immigration- porting the notion that rates of extinction ex- extinction dynamics, of the montane Great ceed that of colonization in at least some but-

Basin mammals, birds, and butterflies has terfly species, Wilcox et al. (1986) have shown been discussed previouslv (Brown 1971, that the numbers of "sedentary" butterfly 1978, Austin 1981, Murphy and Wilcox 1985, species are better correlated with area than Murphy et al. 1986, Wilcox et al. 1986). The are "vagile butterflies. Less-mobile taxa same relationships are seen in fish and land (e.g., montane land snails and lowland fish) snails (Smith 1978, Pratt 1985). exhibit an even greater effect of isolation and Montane or boreal biotic elements in the extinction in this region (Smith 1978, Pratt Great Basin appear to exhibit relictual distri- 1985). butions. This is best substantiated by mam- Note that islandlike effects of area and isola- malian distributions since they include both tion are not restricted to montane or boreal recent (Brown 1971, 1978) and fossil (Grayson elements in the Great Basin. Lowland ripar- 1982, 1983) evidence. These data indicate that ian butterflies appear to be equally isolated, present-day boreal mammalian faunas are not and the faunas of these communities exhibit at equilibrium (that is, they lack balanced similar effects (Austin 1985). Riparian butter- rates of extinction and of colonization) but are fly species richness decreases from the Golo- largely the result of range constriction and rado River Valley northward (upstream) into subsequent extinction (without recoloniza- the central Great Basin. In the northern Great tion) of a once widespread Pleistocene fauna. Basin, species richness decreases from the Fossil evidence from the central Great Basin relatively rich upper river valleys (Humboldt, reinforces the popular view that boreal habi- Garson, Walker) downstream toward the cen- tat, extensive in the Pleistocene, withdrew tral Great Basin. northward and contracted toward montane Given the high number of phenetically dis- summits. Grayson (1983) reports the fossil tinct, geographically restricted endemic but- presence of the vole Phenacomys cf inter- terfly subspecies and segregates, it is of inter- medius in the Toquima Range. This species is est to note patterns of differentiation in other ,

April 1987 Austin, Murphy: Great Basin Butterflies 199 taxa within the Great Basin. Speciation in all exists between subspecies in the extreme taxa is most striking along the western and southern or extreme western Great Basin for northeastern edges of the Great Basin. Bnt, several mammal species. Subspecific en- differentiation certainly is not limited to these demics largely follow the patterns described areas. Stutz (1978), for instance, identified above for butterflies. One species, Mi- several rich evolutionary sites for Atriplcx in crodipodops pallidus, in fact, is a Great Basin the Great Basin, similar to those found for endemic. The distributions of mammals at the birds (Behle 1963, Johnson 1978), and corre- species level (Hagmeier 1966) are consistent sponding to centers of differentiation and lim- with our butterfly data; and more fine- its of distributions of plants in the Great Basin grained, below-the-species-level studies may as outlined by Cronquist et al. (1972). These well further strengthen this comparison with studies and our butterfly data clearly indicate the our findings. existence of distinct areas of interaction and spe- ciation within the whole of the Great Basin. Acknowledgments As we mentioned in several sections above, butterflies and birds are extremely similar in Numerous people made their records and/ their patterns of distribution and differentia- or specimens of Great Basin butterflies avail- tion within the Great Basin. This similarity able to us: R. Albright, D. E. Allen, R. also extends to other taxa including reptiles Bailowitz, D. L. Bauer, J. Brock, F. M. and amphibians (Stebbins 1954) and mammals Brown, J. M. Burns, C. Gallaghan, H.

(Hall 1946, Hall and Kelson 1959). Am- Clench, J. T. Cooper, C. Crunden, T. E. hijstoma tigrinum and Bufo woodhousei are Dimock, D. Eff", J. F. Emmel, T. C. Emmel, Rocky Mountain amphibian species not oc- C. D. Ferris, C. F. Gillette, R. E. Gray, L. P. curring in the Great Basin but extending west Grey, D. Guiliani, C. Hageman, K. Hansen, along its northern margin. A far-western G. Harjes, C. Henne, P. J. Herlan, J. Lane, Great Basin subspecies of Bufo boreas re- R. L. Langston, C. S. Lawson, A. Ludke, W. places a widespread interior subspecies in the W. McGuire, C. D. MacNeifl, J. Masters, S. western Great Basin, and an isolated subspe- O. Mattoon, D. MuUins, J. S. Nordin, P. A. cies occurs in the Inyo Region. Reptiles that Opler, E. M. Perkins, Jr., A. Pinzl, F. W. and

avoid the Great Basin but occur along its bor- J. D. Preston, R. Robertson, K. Roever, R. C. ders include Phnjnosoma douglasii and Rosche, R. W. Rust, F. Ryser, P. and S. Sav-

Thamnophis sirtalis, the latter occurring in age, J. A. Scott, C. Sekerman, O. E. Sette, A. both the Rocky Mountains and the Sierra Ne- Shapiro, A. O. Shields, D. Shillingburg, R.

vada. Subspecific intergradation occurs along Skalski, M. Smith, N. J. Smith, R. E. Stan- the Sierra Nevada-Great Basin interface ford, G. B. Straley, W. Swisher, K. B. Tid-

{Sceloporus graciosus, S. occidentalis well, J. W. Tilden, D. Thomas, W. Whaley, Thamnophis elegans, Crotahis viridus) and R. E. Wells, B. A. Wilcox, and D. Young. We near the Mojave Desert-Great Basin transi- are grateful to all. We thank the curators of tion {Callisaurus draconoides, Phnjnosoma various collections who made specimens platyrhinos, Uta stansburiana). Extension of available for examination: F. H. Rindge primarily southern species northward in the (American Museum of Natural History), L. D.

western Great Basin east of the Sierra Nevada and J. Y. Miller (Allyn Museum of Entomol-

is relatively common. Tanner (1978) com- ogy), J. E. Rawlins and C. W. Young

mented on the absence in the Great Basin of (Carnegie Museum of Natural History), J. P. expected montane species or of endemic spe- Donahue (Los Angeles County Museum), C. cies of amphibians and reptiles. Murvosh (University of Nevada, Las Vegas),

Numerous examples of similar phenomena F. Ryser (University of Nevada, Reno), and J. exist among mammals. Species such as Lepus Burns, C. F. G. Clarke, and R. K. Robbins americanus, Eutamias amoenus, Tamiasciu- (National Museum of Natural History). We

rns douglasii, and Maries americana are also thank S. R. Naegle, P. A. Opler, J. A. found in both the Rocky Mountains and Sierra Scott, A. M. Shapiro, R. E. Stanford, and B. Nevada but not the Great Basin. Others ex- A. Wilcox for helpful comments on drafts of tend northward from the southern deserts the manuscript, and P. Church for typing onlv in the western Great Basin. An interface earlv drafts. 200 Great Basin Naturalist Vol. 47, No. 2

Funding for some survey work on which Ferris, C. D., and F. M. Brown. 1981. Butterflies of the Rocky Mountain this discussion was based was provided by states. University of Oklahoma Press, Norman. grant DAR 8022413 from the National Science Goodpasture, C. 1973. Biology and systematics of the Foundation. The junior author has been sup- Plebejus (Icaricia) acmon group (: Ly-

caenidae). I. ported by a grant from the Koret Foundation Review of the group. J. Ento- of San Francisco. mol. Soc. 46: 468-485. Grayson, D K. 1982. Toward a history of Great Basin mammals during the past 15,000 years. Pages 82-101 in D. B. Madsen and F. Literature Cited J. O'Gonnell, eds., Man and environment in the Great Basin. Soc. Amer. Archaeol. Papers, No. 2. Austin, G T 1981. The montane butterfly fauna of the 1983. Paleontology of Gatecliff Shelter—small Spring Range, Nevada. J. Lepid. Soc. .35; 66-74. mammals. Chapter 6 in D. H. Thomas, O. 1983. A new subspecies of Speijeria atlantis (Ed- J. Davis, D. K. Grayson, W. N. Melhorn, T. wards) (Nymphahdae) from the Great Basin of Thomas, and D. Tre.xler, The archaeology of Mon- Nevada. J.' Lepid. Soc. 37: 244-248. itor Valley. 2. GatecliffShelter. Anthropol. Papers 1985. Lowland riparian butterflies of the Great Amer. Mus. Nat. Hist., Vol. 59. Basin and associated areas. J. Res. Lepid. 24: Grey, L P 1972. Notes on Speijeria zerene populations 117-131. in

Modoc Co. , California. News Lepid. Soc. 1972(6): 2. Austin, G T., and A T Austin 1980. Butterflies of Clark Grey, L. P., and A. H. Moeck. 1962. Notes on overlap- County, Nevada. J. Res. Lepid. 19: 1-63. ping subspecies. I. An example in Speijeria zerene Bagdonas, K., and M Harrington 1979. Variant adap- (Nymphahdae). J. Lepid. Soc. 16: 81-97. tations in moths in Grouse Canvon, Utah. News H.agemeier, E M 1966. A numerical analysis of the dis- Lepid. Soc. 1979(5): 6 (abstract).' tributional patterns of North American mammals. Banta, B H. 1965a. A distributional check list of the II. Re-evaluation of the provinces. Svst. Zool. 15: recent reptiles inhabiting the state of Nevada. 279-299. Occ. Pap. Biol. Soc. Nevada, No. 5. Hall. E R 1946. Mammals of Nevada. University of 1965b. A distributional check list of the recent California Press, Los Angeles. amphibians inhabiting the state of Nevada. Occ. Hall, E R , and K. R Kelson. 1959. The mammals of Pap. Biol. Soc. Nevada, No. 7. North America. 2 vols. Ronald Press, New York. Beatley, J. G. 1975. Glimates and vegetation pattern Harper, K T , D C Freeman, W K. Ostler, and L. G. across the Mojave/Great Basin Desert transition of Klikoff. 1978. The flora of Great Basin mountain southern Nevada. Amer. Midi. Nat. 93: 53-70. ranges: diversity, sources, and dispersal ecology. Behle, W H. 1963. Avifaunistic analysis of the Great Great Basin Nat. Mem. 2: 81-103. Basin region of North America. Proc. 13th Intern. Holdren, C E., and p. R. Ehrlich 1982. Long range OrnithoL Gongr.: 1168-llSl. dispersal in checkerspot buttei-flies: transplant ex- 1978. Avian biogeography of the Great Basin and periments with Eupluidn/as gillctti. Oecologia 50: intermountain region. Great Basin Nat. Mem. 2: 125-129. 55-80. Hovanitz, W 1940. Ecological color variation in a butter- Billings, W. D. 1978. Alpine across phytogeography the fly and the problem of "protective coloration." Great Basin. Great Basin IMat. Mem. 2: 105-117. Ecology 21:.371-,380. Brown, H. 1971. J. Mammals on mountaintops: nonequi- 1941. Parallel ecogenotypical color variation in librium insular biogeography. Amer. Nat. 105: buttei-flies. Ecology 22: 259-284. 467-478. Johnson, N K. 1965. The breeding avifaunas of the Sheep 1978. The theory of insular biogeography and the and Spring ranges in southern Nevada. Condor 67: distribution of boreal birds and mammals. Great 93-124. Basin Nat. Mem. 2: 209-227. 1970. The affinities of the boreal avifauna of the Gronquist, A. M. a, Holmgren N H Holmgren, and J. Warner Mountains, California. Occ. Pap. Biol. L Reveal. 1972. Intermountain flora, vascular Soc. Nevada, No. 22. plants of the intermountain west, U.S.A. Vol. 1. 1975. Controls of number of bird species on mon- Hafner Publ. Co., New York. tane islands in the Great Basin. Ecologv 29:

Dornfeld, E. J 1980. The buttei-flies of Oregon. Timber .545-567. Press, Forest Grove, Oregon. 1978. Patterns of avian geography and speciation Emmel.T. G.andJ. F Emmel. 1969. A new subspecies of in the intermountain region. Great Basin Nat.

the Cercyonis meadi group (Satyridae). J. Lepid. Mem. 2: 137-1.59. Soc. 23: 161-164. Johnson, N. K , and C B Johnson. 1985. Speciation in 1971. An extraordinary new subspecies oiCercij- sa.\)sudkers {SpJnjrapicus). II. Sympatry and mate onis oetus from central Nevada (Lepidoptera, preference in S. ruber daf^etti and S. nuchalis. Satyridae). Pan-Pacific Entomol. 47: 1,55-157. Auk 102: 1-15. 1973. The butterflies of southern California. Nat. LiNSDALE, J. M 1938. Bird life in Nevada with reference Hist. Mus., Los Angeles Co., Sci. Series 26. to modifications in structure and behavior. Con- Emmel, T. G, and S O. Mattoon. 1972. Cerctjonis pe- dor 40: 137-159. gala blanca, a "missing type" in the evolution of MacArthur, R. H , AND E. O Wilson. 1967. The theory

the genus Ccrci/oru.s(Satvridae). J. Lepid. Soc. 26: of island biogeography. Princeton University 140-149. Press, Princeton, . April 1987 Austin, Murphy: Great Basin Butterflies 201

1986. butterflies of North America, a Martin, P S., and P. J Mehringer. 1965. Pleistocene Scott. J A The pollen analysis and biogeography of the south- natural history and field guide. Stanford Univer- west. Pages 433-452 in H. E. Wright and D. G. sity Press, Stanford, California. Fray, eds., The Quaternary of the United States. Shapiro. A. M., C. A. Palm, and K L Wcislo 1979. The Princeton University Press, Princeton, New ecology and biogeography of the butterflies of the Jersey. Trinitv Alps and Mount Eddy, northern Califor-

nia. J.' Res. Lepid. 18;69-15L Meyer, S. E. 1978. Some factors governing plant distribu- Shields, O 1977. Studies on North American Philotes tions in the Mojave-Intermountain transition (). V. Taxonomic and biological, notes zone. Great Basin Nat. Mem. 2: 197-207. continued. J. Res. Lepid. 16: 1-67. Miller, A. H 1941. A review of centers of differentiation Smith, GR. 1978. Biogeography of intermountain fishes. for birds in the western Great Basin region. Con- Great Basin Nat. Mem. 2: 17-42. dor 43: 257-267. Sterbins. R. C 1954. Amphibian and reptiles of western MOECK, A. H. 1957. Geographic variability in Speijeria. North America. McGraw-Hill, New York. Comments, records and description of a new sub- Stutz. H. C. 1978. Explosive evolution of perennial

species. Milwaukee Ent. Soc. , Special Paper. Atriplex in western America. Great Basin Nat. Murphy, D.. and P R. Ehrlich. 1983. Biosystematics of Mem. 2: 161-168. the Euphijdnjas of the Great Basin with the de- Swisher. W L. and A L Morrison. 1969. News, 22; scription of a new subspecies. J. Res. Lepid. notes—one desirable species. News Lepid. Soc. 254-261. 1969(3): 4. Murphy, D. D.. and B A Wilcox. 1985. Buttei-fly diver- Tanner, W W. 1978. Zoogeography of reptiles and am- sity in natural habitat fragments: a test of the valid- phibians in the intermountain region. Great Basin 43-53. ity ofvertebrate-based management. In]. Verner, Nat. Mem. 2: Thomas. D H 1983. Large mammals. In D. H. Thomas, M. L. Morrison, C. J. Ralph, and R. H. Barrett, ed.. archaeology of Monitor Valley. 2. Gate- eds.. Modeling habitat relationships of terrestrial The Nat. Hist. Anthrop. Pap. vertebrates. University of Wisconsin Press. cliffShelter. Amer. Mus. 59: 126-129. Murphy, D D , B A Wilcox, G. T. Austin, and P R Watt, W B. 1968. Adaptive significance of pigment poly- Ehrlich 1986. Butterflies of the Great Basin morphisms in Colias butterflies. I. Variation of ranges. J. Res. Lepid., in press. melanin pigment in relation to thermoregulation. Perkins, S. P., and E. M Perkins, Jr 1967. Revision of Evolution 22: 437-458. the Limenitis weidemeijerii complex, with descrip- Wells, P V 1983. Paleobiogeography of montane islands tion of a new subspecies (). Lepid. J. in the Great Basin since the last glacialpluvial. Soc. 21:213-234. Ecol. Monogr. 53: 341-382. of central Great Pratt. W L. 1985. Insular biogeography WiELGUS, R. S,. and D. Wielgus 1974. A new sandy- Basin land snails: extinction without replacement. desert subspecies of Megathymus coloradensis

J. Arizona-Nevada Acad. Sci. (1985 Proc. Suppl.) (Megathymidae) from extreme northern Arizona. 20: 14 (abstract). Bull. Allyn Mus., No. 17. his- T. Rogers, G. F. 1982. Then and now. A photographic Wilcox, B. A , D D Murphy. P R Ehrlich, and G tory of vegetation change in the central Great Austin. 1986. Insular biogeography of the montane Basin Desert. University of Utah Press, Salt Lake butterfly faunas in the Great Basin: comparison with Citv. birds and mammals. Oecologia69: 188-194.