Speyeria nokomis ( Edwards, 1863b )

Dunford, James C., 2009, Taxonomic overview of the greater fritillary genus Speyeria Scudder and the atlantis - hesperis species complexes, with species accounts, type images, and relevant literature (Lepidoptera: Nymphalidae), Insecta Mundi 2009 (90), pp. 1-74 : 15-17

Speyeria nokomis ( Edwards, 1863b)

( Figure 10-11 )

Argynnis Nokomis Edwards, 1863b: 221 .

Acidalia Semnopsyche nokomis form valesinoides-alba Reuss, 1926: 69.

Speyeria nokomis (Edwards) [ dos Passos and Grey 1945a].

Common names. Nokomis fritillary, western seep fritillary.

Type deposited. Neotype (male) designated by dos Passos and Grey (1947) at American Museum of Natural History ( Figure 10 ).

Type locality. Rocky Mountains and mountains of California. Neotype (male) ( Figure 10 ) defined by dos Passos and Grey (1947) as Mount Sneffels, Ouray County, Colorado; however, dos Passos and Grey (1965) reconsidered this designation and Brown (1965) noted that this locality was an unlikely habitat for S. nokomis and that the specimen does not fit the original description. Ferris and Fisher (1971) revised the type locality and designated a lectotype (male) ( Figure 11 ) taken from Colorado for S. nokomis . Ferris and Fisher (1971) discuss the likelihood that the type locality for S. nokomis nokomis was probably somewhere in eastern Utah; however, the specimen they designated as lectotype is taken from Mesa County, Colorado (see below). Grey (1989) later noted that S. nokomis does occur at Mt. Sneffels, based on collection records located at the AMNH. Although the true type locality and type specimen will likely remain obscure or missing, the neotype designation provided by dos Passos and Grey (1947) is reaffirmed by Grey (1989).

Type label data. Taken from dos Passos and Grey (1947): “ Oslar Sneffels Mts Ouray Co Col Aug 9000 Ft.; A. nokomis ; Ex Coll. Wm. C. Wood Acc 36915; NEOTYPE, Argynnis Nokomis [male], Edwards” .

Identification, taxonomy, and variation. There are approximately 7 described subspecies in the nokomis complex. Adult wingspan ranges from 63-80 mm. Male dorsal wing coloration is orange with sparse black dots while females are black basally and whitish outwardly with many black spots. The dorsal submarginal dark chevrons do not touch the black marginal line. Forewings are pinkish-orange ventrally with white spots. Discal coloration is variable in both sexes depending on the geographic location. The ventral hindwing disc on males and females is light to dark brown with submarginal band tan in many regions; females bear a gray-green disc with the submarginal band yellow-green in California and Nevada populations. Eastward populations tend to have darker hindwing discs. The hindwings on both sexes have relatively small silver spots and they typically bear black edges. Most forms of S. nokomis exhibit sexual dimorphism.

The uncus on the male genitalia is hooked and similar to those of S. idalia and the Semnopsyche group; however, the female has only a single bursal sac. The eye coloration in living specimens is yellowgreen ( Glassberg 2001b). The egg is cream colored when laid and becomes tan after a few days. Detailed egg morphology is included in Scott and Mattoon (1981). Larvae typically bear a yellow to orange dorsal stripe and yellow to orange transverse stripes with rows of yellow- orange or black spines. Black patches surround spines dorsally and laterally. Female larvae typically feed ten days longer than do males ( Allen et al. 2005). Detailed larval descriptions, including setal maps, are included in Scott and Mattoon (1981). Pupae are black with center of wing cases orange, and bear orange stripes on the abdomen. Pupae vary in coloration throughout the range of S. nokomis .

Range. Speyeria nokomis is presently known from eastern California to western Colorado, south through eastern Arizona and western New Mexico, with populations as far south as Mexico. Tuttle (2007) noted that R. Holland reported specimens located in the CMNH were taken in the 1930’s from the Mescalero Apache Reservation in the Sacramento Mountains and represent the southern-most occurrence of nominate ‘ nokomis ’. Known localities are widely separated due to restricted habitat. Many populations are declining because of capping of springs and other habitat modifications caused by human disturbances such as livestock grazing ( Hammond and McCorkle 1984).

Life History. Habitats include Upper Sonoran to Canadian Zone moist meadows near streams, permanent spring fed meadows, marshlands, boggy streamsides, and seeps; S. nokomis can be found in canyons with pinyon pines and junipers. Britten et al. (1994) studied the isozyme variability of S. nokomis populations in the Great Basin and noted that there was little gene flow between populations, further confirming that S. nokomis is confined to mesic seep habitats with great expanses of unsuitable, xeric habitat isolating populations. Eggs are laid singly and haphazardly near host plants. Unfed first instar larvae hibernate, and some later instars may also aestivate during drought conditions from April through June ( Scott 1986b). Larvae overwinter in grass stems after emerging ( Pyle 1995). Males patrol all day in meadows or along streams seeking females, especially in valley bottoms if the spring is a on a slope. Scott (1975) noted that males would fly about 1/ 2 meter off the ground and dip into to hollows to search. This species tends to fly on the average later than most other Speyeria . Flight period is usually from late July to mid September or mid August to mid September in the southern part of its range. The range of S. nokomis was likely more continuous during moister climatological times. Populations are now separated by vast desert landscapes. Scott (2008a) estimates that 50% of S. nokomis sites were lost by 1900 due to water diversion. A population [ S. nokomis coerulescens (W. J. Holland) ] that once flew high in the Santa Catalina Mountains north of Tucson, Arizona has not been seen since 1938 and presumably has been extirpated ( Glassberg 2001b). Fleishman et al. (2001b) note that extinction and colonization events for S. nokomis populations in the Great Basin are related to multiple aspects of habitat quality, such as extreme climatic events and grazing-mediated availability of nectar. The results from Britten et al. (1994) indicate there is little gene flow among S. nokomis populations in the Great Basin, and that these populations have lost genetic variability as the result of small effective populations sizes and genetic drift; thus, conservation of individual colonies may be important for the evolutionary potential of this species. Results from mark and recapture studies conducted by Britten et al. (2003) indicate that suitable but vacant habitat patches should be maintained for potential recolonization by S. nokomis apacheana (Skinner) in the central Great Basin.

Larval host plants. Viola sororia ( Emmel et al. 1970; Scott and Mattoon 1981; Scott 1986a; Robinson et al. 2002).

Adult food resources. Thistles ( Scott 1986b), especially Arizona red thistle (Cliff Ferris, pers. comm.).

Figures

Figure 6-10. Type images for Speyeria spp.; dorsal, ventral, and label images. 6) Speyeria idalia neotype, described as Papilio Nymphalis Phaleratus Idalia, male, AMNH. 7) Speyeria diana holotype, described as Papilio Nymphalis Phaleratus Diana, male, BMNH. 8) Speyeria cybele neotype, described as Papilio Nymphalis Phaleratus Cybele, female, BMNH. 9) Speyeria aphrodite neotype, described as Papilio Nymphalis Phaleratus Aphrodite, male, AMNH. 10) Speyeria nokomis neotype, described as Argynnis Nokomis, male, AMNH. Images 6 and 9-10 by James C. Dunford; 7-8 by Kim Goodger Buckmaster.

Figure 11-15. Type images for Speyeria spp.; dorsal, ventral, and label images. 11) Speyeria nokomis lectotype, described as Argynnis Nokomis, male, AMNH. This specimen is not recognized as the name bearing type. 12) Speyeria edwardsii lectotype, described as Argynnis edwardsii, male, FMNH. 13) Speyeria coronis neotype, described as Argynnis Coronis, male, CMNH. 14) Speyeria zerene lectotype, described as Argynnis Zerene, male, NMNH. 15) Speyeria carolae holotype, described as Argynnis coronis carolae, male, CMNH. Images 11-13 and 15 by James C. Dunford; 14 by Robert Robbins.