Speyeria hesperis (Edwards, 1864)

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 : 29-30

Speyeria hesperis (Edwards, 1864)

( Figure 4C , 25-46 )

Argynnis Hesperis Edwards, 1864a: 502 .

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

Speyeria hesperis (Edwards) [ Scott et al. 1998].

Common names. Hesperis fritillary, western fritillary.

Type deposited. Neotype (male) designated by Brown (1965; see also dos Passos and Grey 1965) at Carnegie Museum of Natural History ( Figure 25 ).

Type locality. Defined by dos Passos and Grey (1947) based on lectotype (= neotype from Brown 1965; see also dos Passos and Grey 1965) as Turkey Creek Junction, Jefferson County, Colorado.

Type label data. “ Hesperis [male] type Colo; Neotype, Argynnis hesperis [male], W. H. Edwards designated by dos Passos and Grey 1965 ” .

Identification, taxonomy, and variation. Formerly considered a subspecies of S. atlantis , S. hesperis is variable throughout its range and 21 subspecific taxa are currently recognized. Speyeria hesperis and S. atlantis occur together or in close proximity, especially in cool, forested habitats, in many areas of western North America. Dos Passos and Grey (1965) noted that S. hesperis represented an unsilvered subspecies of S. atlantis along the Front Range in Colorado. Tebaldi (1982) (also see Ferris 1983) used starch gel electrophoresis of six enzymes to analyze the relationships among three phenotypes of S. atlantis and found that the phenotypes could be considered “semispecies.” Scott et al. (1998) divided S. atlantis into a distinct species based on wing coloration and wing pattern, hesperis having mostly unsilvered or cream colored ventral hindwing spots and S. atlantis always silvered. However, this situation still needs to be addressed in greater detail. The original description for S. hesperis identifies it as having opaque/ cream colored spots (also see Figure 25 ). Several ‘ hesperis ’ subspecies (i.e., S. h. electa , S. h. ratonensis, S. h. schellbachi, S. h. dorothea, S. h. nausicaa, S. h. capitanensis) are apparently always silvered. It may be that S. electa is a valid species and represents the silver-spotted western ‘species’ in the atlantis species complex (Cliff Ferris, pers. comm.). Adult wingspan ranges from 50-68 mm. The ventral hindwing disc is red-brown to orange-brown and can be silvered or unsilvered. Scott et. al. (1998) split S. atlantis and S. hesperis forms based on wing pattern and coloration and a few larval characters. Adult eye coloration in living individuals is blue-gray in some populations, and this may help separate some populations from similar S. aphrodite populations, which bear dull, yellow-green eyes ( Glassberg 2000). Larvae are generally solid black and bear orange spines with black tips. There are two brown lines located dorsomedially. The pupa is similar to S. atlantis in shape, but stouter; the color is brown on the head and wing cases. The abdomen is brown with some areas yellow-brown. Larval and pupal coloration varies throughout the range of S. hesperis (see Scott et al. 1998) due to various local climatic conditions.

Range. Speyeria hesperis occurs from Alaska, central Yukon and southwestern Northwest Territory, south through Canada east to western Manitoba, and in the western United States along the Rocky Mountains, to central California, northeastern and central Arizona, and central New Mexico.

Life history. Speyeria hesperis forms occur in moist meadows, gulches, and along cool slopes ( Scott 1986b). Scott (2006b) observed females laying eggs on pine needles, Quercus leaves, grasses, and various other plants near Viola spp. Edwards (1888c) described the morphology of the egg, larval instars, and pupal stage and provided the phenology of each stage. Flight period is from early June to late October.

Larval host plants. Viola canadensis var. scopulorum , V. adunca , V. sororia affinis , V. rydbergii , V. adunca bellidifolia , V. nuttallii , V. purpurea ( Scott 1992, 2006b; Scott et al. 1998).

Adult food resources. Yellow composites, mints ( Opler and Wright 1999); especially Monarda (Cliff Ferris, pers. comm.).

Figures

Figure 4. Male genitalic armature. A) Speyeria idalia (locality unknown). B) Speyeria cybele (Nebraska). C) Speyeria hesperis (aedeagus removed) (Idaho). D) Digitus located on left valva, Speyeria edwardsii (Nebraska). Images by James C. Dunford.

Figure 21-25. Type images for Speyeria spp.; dorsal, ventral, and label images. 21) Speyeria atlantis canadensis lectotype, described as Argynnis atlantis canadensis, male, AMNH. Now synonymized under Speyeria atlantis (Scott et al. 1998). 22) Speyeria atlantis hollandi holotype, described as Argynnis atlantis Hollandi, male, CNIC. 23) Speyeria atlantis sorocko holotype, described as Speyeria atlantis sorocko, male, AMNH. 24) Speyeria atlantis pahasapa holotype, described as Speyeria atlantis pahasapa, male, AMNH.25) Speyeria hesperis neotype, described as Argynnis Hesperis, male, CMNH. Images 21 and 23-25 by James C. Dunford; 22 by Norbert Kondla.

Figure 26-30. Type images for Speyeria spp.; dorsal, ventral, and label images. 26) Speyeria hesperis helena lectotype, described as Argynnis lais, male, CMNH. 27) Speyeria atlantis dennisi holotype, described as Argynnis lais form dennisi, male, AMNH. Now synonymized under Speyeria hesperis helena (Scott et al. 1998). 28) Speyeria hesperis beani holotype, described as Dryas atlantis beani, male, NMNH. 29) Speyeria hesperis brico holotype, described as Speyeria hesperis brico, male, CNIC. 30) Speyeria hesperis ratonensis holotype, described as Speyeria atlantis ratonensis, male, LACM. Images 26-27 by James C. Dunford; 28 by Robert Robbins; 29 by Norbert Kondla; 30 by Weiping Xie.

Figure 31-35. Type images for Speyeria spp.; dorsal, ventral, and label images. 31) Speyeria hesperis greyi holotype, described as Speyeria atlantis greyi, male, AMNH. 32) Speyeria hesperis lurana holotype, described as Speyeria atlantis lurana, male, AMNH. 33) Speyeria hesperis irene lectotype, described as Argynnis irene, female, NMNH. 34) Speyeria hesperis hanseni holotype, described as Speyeria atlantis hanseni, male, LACM. 35) Speyeria hesperis dodgei holotype, described as Argynnis dodgei, male, AMNH. Images 31-32 and 35 by James C. Dunford; 33 by Robert Robbins; 34 by Weiping Xie.

Figure 36-40. Type images for Speyeria spp.; dorsal, ventral, and label images. 36) Speyeria hesperis viola holotype, described as Speyeria atlantis viola, male, AMNH. 37) Speyeria hesperis elko holotype, described as Speyeria atlantis elko, male, NVMC. 38) Speyeria hesperis tetonia holotype, described as Speyeria atlantis tetonia, male, AMNH. 39) Speyeria hesperis wasatchia holotype, described as Speyeria atlantis wasatchia, male, AMNH. 40) Speyeria hesperis chitone lectotype, described as Argynnis chitone, male, CMNH. Images 36 and 38-40 by James C. Dunford; 37 by George Baumgartner and Scott Klette.

Figure 41-45. Type images for Speyeria spp.; dorsal, ventral, and label images. 41) Speyeria hesperis electa lectotype, described as Argynnis Electa, male, CMNH. 42) Speyeria hesperis nikias lectotype, described as Argynnis nikias, male, CMNH. Now synonymized under Speyeria hesperis electa (Scott et al. 1998). 43) Speyeria hesperis schellbachi holotype, described as Speyeria atlantis schellbachi, male, LACM. 44) Speyeria hesperis nausicaa lectotype, described as Argynnis Nausicaa, male, CMNH. 45) Speyeria hesperis dorothea holotype, described as Speyeria atlantis dorothea, male, AMNH. Images 41-42 and 44-45 by James C. Dunford; 43 by Weiping Xie.

Figure 46-48. Type images for Speyeria spp.; dorsal, ventral, and label images. 46) Speyeria hesperis capitanensis holotype, described as Speyeria atlantis capitanensis, male, AMNH. 47) Speyeria hydaspe lectotype, described as Argynnis Hydaspe, male, NMNH. 48) Speyeria mormonia lectotype, described as Argynnis Mormonia, male, NMNH. Image 46 by James C. Dunford; 47-48 by Robert Robbins.