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---------- 1969. A revised synonymic list of the Nearctic Melitaeinae with taxonomic
notes (Nymphalidae). J. Lepid. Soc. 23: 115-125.
---------- 1970. A revised synonymic catalog with taxonomic notes on some Nearctic
Lycaenidae. J. Lepid. Soc. 24: 26-38.
McAlpine, W. S., 1939. A new Metal Mark (Calephelis) from Texas. Bull. Brooklyn Ent. Soc. 34: 75-80.
McDunnough, J. 1938. Checklist of the Lepidoptera of Canada and the United States of America, Part 1, Macrolepidoptera. Mem. So. Calif. Acad. Sci. 1 (1).
---------- 1939. Checklist of the Lepidoptera of Canada and the United States of
America, Part 2, Microlepidoptera. Mem. So. Calif. Acad. Sci. 2(1).
A REVISION OF SPEYERIA NOKOMIS (NYMPHALIDAE)1
Clifford D. Ferris
University of Wyoming, Laramie, Wyoming
Mike Fisher
1200 Summit Road, Ponderosa Hills, Parker, Colorado
It is proposed that Speyeria nokomis nitocris (Edwards) be elevated from the synonymy as placed by dos Passos (1964), to subspecific rank for the reasons which are delineated in the paragraphs below.
In an earlier work (dos Passos and Grey 1947), Speyeria nokomis nitocris was treated as a valid subspecific taxon, but later (1964), dos Passos treated this taxon as synonymous with S. nokomis nokomis (Edwards). For the past several years, we have made a study of this insect in Colorado (M. F.) and in Arizona and New Mexico (C. D. F.). The Colorado insect, S. nokomis nokomis is quite distinct in habitat and fascia from the Arizona-New Mexico insect, S. nokomis nitocris,
Speyeria nokomis nokomis (Edwards)
Original Description: "Descriptions of certain species of diurnal Lepidoptera found within the limits of the United States and British America." No. 3. Proc. Acad. Nat. Sci. Philadelphia 14: 221 (1863). Figured by Edwards, B. N. A. pi. Argynnis4 (1868).
Type Locality: This was fixed by dos Passos and Grey (1947) as Mount Sneffels, Ouray Co., Colorado. This location has been questioned by Brown (1957b, p. 335) and does not appear consistent with presently known habitats for n. nokomis. The neotype male (AMNH) bears the label "Oslar Sneffels Mts Ouray Co Col Aug 9000 Ft.," "A. nokomis." and "Ex Coll. Wm. C. Wood Ace. 36915."
Discussion: In the plate which accompanies this paper, three subspecies of Speyeria nokomis are figured. Dorsally the males are similar
1 Published with the approval of the Director, Wyoming Agricultural Experiment Station, as Journal Paper No. 434.
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but distinct differences are evident on the ventral surfaces. In S. n. nokomis, the discal area of the hind wing is a pale cinnamon brown. The silver spots are large, brilliant, and moderately edged with black. The species as a whole is strongly dimorphic (excepting n. wenona) and the females dorsally tend almost to black basally. In n. nokomis, dorsally the females show a blue-green overscaling in the dark areas with a light buff submarginal band. On the primary, the band shows a yellow-orange flush; on the secondary the flush is bluish. Ventrally, the females are similar to the males as far as the primaries are concerned, except that the colors are more intense and the ground color is a darker red. On the secondary, the submarginal band is a light buff and the ground color of the discal area is buff overscaled with a greenish-black dusting. Females from the Paradox Valley colony in Colorado tend to be more blue dorsally than specimens from other localities.
This insect is currently known from several locations in Colorado, from Uintah Co. in eastern Utah, and in a blend with n. apacheana from more westerly areas of Utah. The Colorado locations are Unaweep Canyon, Mesa Co., 6000' and Paradox Valley, Montrose Co., 5200'. The Colorado habitats are nearly neutral seep areas where there is a constant flow of water and are surrounded by willow thickets. The violets which serve as the larval foodplant grow in the understory of the seeps and are found only sparsely in the thickets. The surrounding region, except in the vicinity of other seeps and washes is relatively arid. The sites of both colonies are unusual with respect to their geology. The usual rock formation in this portion of Colorado is Mancos shale. The Unaweep Canyon seep emerges along a spring line in granite and flows over what appears to be stream alluvium. The canyon is unusual in that it is mainly composed of granite. The Paradox Valley colony is in a blowsand area. The Uintah Co., Utah area is a rather unrewarding looking meadow, but with the necessary prerequisites of willow, violets, and a constant flow of water. The presence of willow is probably only incidental to the presence of nokomis. Both willows and violets are frequently found in the same location in arid regions.
There is one additional recent record of S. nokomis from Colorado. This is a worn female taken by Scott Ellis at Rogers Mesa Delta Co., 5850', 26 August 1958. The source of this specimen has not yet been determined, although it perhaps came from an undiscovered Colorado colony.
Dos Passos and Grey (1947) fixed the type locality for S. nokomis nokomis as Mount Sneffels, Ouray Co., Colorado. We feel that we must take exception to this fixation for several reasons. Collecting in recent
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years in this area (Brown, in litt.) has failed to turn up this butterfly. The ecology of the fixed type locality is not consistent with currently known collection areas. Furthermore the neotype does not fit the original description (Brown, 1965). There is also a disparity in dates. Dos Passos (1964) indicates an 1862 ("1861-3") date for the Edwards publication of the description of n. nokomis. If this is the correct date, it would not seem possible for the specimens to have come from Ouray Co., as the first whites to enter this area came in 1871. Oslar is known to have been quite inaccurate in his data. As Brown has mentioned (1965), the Macomb Expedition entered the area west of Paradox Valley in 1859 (Macomb and Newberry 1876). It is possible that the material ascribed to Edwards 1862 was actually collected on this expedition. Brown placed the probable type locality for S. nokomis nokomis (Edwards) (and not for the neotype S. nokomis nokomis dos Passos and Grey) as Ojo Verde, San Juan Co., Utah. The location Ojo Verde is mentioned in the Macomb/Newberry report. The topographical coordinates are given in the report as lat. 38° 14' 50" and long. 109° 26' 40". Present United States Coast and Geodetic Survey maps do not show Ojo Verde, but the coordinates would indicate it to be in the vicinity of Hatch Wash. In August, 1969, the senior author attempted to locate this site. He was unsuccessful as that portion of Utah is now posted as an impact area for test missiles launched from Green River, Utah. The warning notices and travel directions were sufficiently ambiguous that he did not wish to risk entry into the area.
The comments in the Macomb/Newberry report would appear to indicate a possible habitat for n. nokomis. Macomb and his associates camped at Ojo Verde in mid-August and would thus have been there at the correct time of the year for the insect's flight. Of the area, the following comments are made: "The Ojo Verde is a copious spring in a caiion cut out of the red sandstone, ten miles west of La Teneja. The surrounding country is very sterile, sparsely set with sage bushes and small cedars, but about the spring the bottom of the canon is covered with the greenest and most luxuriant grass."
It would appear then that the type locality for S. nokomis nokomis (Edwards) is most probably somewhere in eastern Utah, and not in Ouray Co., Colorado, although L. P. Grey now favors the Rio Hondo area in Lincoln Co., New Mexico.
There also exists a record of a single male and two females of S. n. nokomis taken September 15, 1877 by Lt. C. A. H. McCauley at the crossing of the Lower Rio Florida, La Plata, Co., Colorado 6500-7000' some
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twenty miles east of Durango. These specimens are now part of the Strecker Collection at the Chicago Natural History Museum. The male was apparently misidentified at one point in time and appears to be S. cybele carpenteri (Edwards) (see Brown 1957a). It was originally recorded in the McCauley report (1879) as Argynnis cybele. The female records were checked by L. P. Grey and one is figured in the report (Plate I).
The flight span of S. n. nokomis is generally during the first three weeks in August, although it has been taken from mid-July (Unaweep Canyon) into September.
Speyeria nokomis nitocris (Edwards)
Original Description: "Description of new species of diurnal Lepicloptera found in North America." Trans. Amer. Ent. Soc. 5: 15-16 (1874). Figured by Edwards, B. N. A. 3: [91-92], Argynnis I; Holland, B. B., p. 85, pi. 13. (type).
Type Locality: White Mountains, Arizona. Based upon the discussion in Brown (1965), and the study by the senior author of the distribution of nokomis in Arizona, the type locality is probably east of Ft. Apache, along the East Fork of the White River, Apache Co., Arizona. The lectotype (designated by dos Passos and Grey 1947) (CM) bears the label "Nitocris $ W. Mtns. Ariz, type" and is the specimen figured by Holland.
Discussion: The accompanying plate clearly shows the differences between n. nokomis and n. nitocris. Generally the colors are much more intense in n. nitocris. In the males, ventrally, the discal area of the secondary is a deep cinnamon brown, the silver spots are quite brilliant and heavily margined with black scales. The submarginal band is darker than in n. nokomis and shows an orange flush over the lighter buff ground color. In the females, dorsally, the dark colored areas are more extensive than in n. nokomis and the blue-green overscaling is more prominent. Ventrally, the discal area of the secondary has a cinnamon ground color which is heavily overscaled with velvety black. The submarginal band is yellow-buff but overscaled with dark scales.
The habitat of S. n. nitocris lies in lush Canadian Zone meadows or along the mountain streams which feed such meadows. The adults show a preference for feeding upon red thistles of various species. S. nokomis nitocris generally flies at higher elevations that does S. n. nokomis. It has been taken from 5400' to 8500' and appears to be more common at higher elevations. It has been recorded from Arizona: Gila, Coconino, Greenlee, Navajo, Apache Cos.; from New Mexico: Catron, Valencia, San Miguel, Taos Cos. S. n. nitocris is found in the region of the Mogollon Rim and White Mountain country in Arizona, and in the extension of this terrain into New Mexico (Mogollon Mountains and eastward). It then
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Fig. 1. Probable distribution for S. nokomis. Solid dots indicate S. nokomis nitocris; open dots indicate S. nokornis nokomis. Not all collection sites are shown. The dotted line which encloses each series of dots indicates the probable range of the given subspecies where suitable habitats exist. Mt. Sneffels is represented by the open triangle and Ojo Verde by the solid triangle.
occurs in pockets at various other locations in New Mexico. The flight period extends from mid-August into early September.
Of the two subspecies, nitocris is the more common. Both are quite local, but nitocris has been taken in numerous localities and is sometimes found in high concentration. In both subspecies the males appear to outnumber the females by a ratio of more than 5:1. The females appear less active than the males and keep to areas with dense vegetation. In nitocris, the males appear as soon as the dew starts to dry off in the
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morning and they may be seen feeding or patrolling their territories. The females appear more toward noon and except when feeding, are generally flushed from the tall grass and willows which border the streams associated with their habitat. Mating behavior has been described by Ferris (1969).
It is interesting to note that as one examines western populations of S. nokomis through Utah and into California, the ratio of the sexes becomes nearly equal.
From a somewhat limited study, it would appear that both subspecies appear in strength in any given colony on an alternate year basis.
For comparison, also illustrated in the plate are specimens of S. nokomis apacheana (Skinner) from California. Ventrally these are quite distinct from the Rocky Mountain Region-Northern Arizona-New Mexico material. In the males the discal area is almost the same color as the sub-marginal band. In the females, the discal area is concolorous with the submarginal band, but is overscaled with greenish-black.
We interpret the range of S. nokomis nokomis as being restricted to southwestern Colorado, and to Uintah and San Juan Cos., Utah. It would appear from preliminary studies that n. nokomis and n. apacheana inter-grade in Utah. An interesting note on the blend zone situation was presented by Swisher and Morrison (1969). They collected nokomis from California to Colorado. Based upon a somewhat limited number of specimens, they found n. apacheana to be the only form in California. In Washington Co., Utah (SW Utah at Leeds), they found 24% intermediate between n. apacheana and n. nokomis. Collecting at Glendale, Kane Co. (40 miles east of Leeds) yielded 17% intermediates and 15% referable to n. nokomis. In northeastern Utah at Vernal, Uintah Co., they found 20% intermediate and 44% were referable to nokomis. Colorado specimens (Montrose Co.) were all n. nokomis. The remaining percentage figures at each site apply to n. apacheana. The sex ratio, males to females, are reported as 3:2 for n. apacheana and 9:1 for n. nokomis as an average over the sites visited. Females of n. nokomis are quite scarce in Colorado.
The range of S. nokomis nitocris is restricted to northern Arizona, New Mexico and south-central Colorado along the New Mexico border. We feel that S. nokomis nigrocaerulea (Cockerell and Cockerell) should be treated as a synonym of S. nokomis nitocris because it evidently represents an eastern outpost of the latter subspecies. Occasional New Mexico specimens are larger than material from the White Mountains of Arizona, and sometimes more darkly marked, but we do not feel this varia-
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Fig. 2. a-d. S. nokomis nokomis, Unaweep Canyon, Mesa Co., Colo., 6000'. b, c, Female, 20-viii-1967; c, d, male, 18-viii-1967.
e-h. S. nokomis nitocris. e, Female, Willow Creek, Catron Co., New Mex., 25-viii-67, 8100'; f, female, Pine Creek, Gila Co., Ariz., 7-viii-67, 5400'; g, male, Alpine, Apache Co., Ariz., 17-viii-67, 8300'; h, male, Pine Creek, Gila Co., Ariz., 7-viii-67, 5400'.
i-1. S. nokomis apacheana, Round Valley, Inyo Co., Calif., 5000'. i, j, Female, 24-viii-1968; k, 1, male, ll-viii-1963.
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tion sufficient to warrant nomenclatural distinction. The New Mexico insect was originally described from Beulah. This has created some confusion as Beulah does not currently exist as a town. It is a mail route in the Sapello Valley, San Miguel Co., New Mexico. Farming and ranching have essentially wiped out the habitat for nokomis. We would include here the specimen taken by Rotger in Conejos Co., Colorado and described in Brown (1957b).
In some species of Speyeria, there is considerable change in discal color intensity as a function of altitude. S. morraonia is a good example. We considered this to be the situation with S. nokomis. Examination of collection sites indicates that both n. nokomis and n. nitocris are found at comparable altitudes although the latter extends into higher altitudes. The ecologies of the respective habitats are quite different. It is on basis of fascia, habitat, and geographical location that we have separated the two subspecies. It should be noted that the underside color of n. nitocris fades quite rapidly on the wing, and this may have been the reason for confusing the two subspecies in the past. The very dark color of the disc in fresh n. nitocris is most distinct from the color of the same area in fresh n. nokomis.
We would thus propose a revised treatment of S. nokomis as follows:
SPEYERIA (SPEYERIA) Scudder, 1872
Type: Papilio Nymphalis Phaleratus idalia Drury, "1770" [1773] Argynnis Auctorum XDryas Hiibner, [1806] (Opinion 278, name 75) (partim) Neoacidalia Reuss, 1926 (partim) 612 nokomis (Edwards), "1861-3" (1862)
a n. nokomis (Edwards), "1861-3" (1862)
form valesinoidesalha (Reuss), 1926 (emendatio) b n. nitocris (Edwards), "1874-6" (1874)
nigrocaerulea ( Cockerell and Cockerell), 1900 ab. rufescens (Cockerell), 1909
nitrocris dos Passos, 1964 (lapsus calami) c n. ccerulescens (Holland), 1900'
cserulescens McDunnough, 1938 (lapsus calami) d n. apacheana (Skinner), 1918
nokomis (Edwards) 1873 (nee Edwards, "1861-3" [1862]) e n. wenona dos Passos and Grey, 1945
Acknowledgments
The authors would like to thank the following collectors who supplied specimens for examination, and information on habitats and ecology: J. D. Eff, Bruce Harris, J. H. Masters, and R. E. Stanford. Scott Ellis provided extensive habitat notes for Colorado. Special thanks are due F. Martin Brown of Colorado Springs, Colorado and L. Paul Grey of Lin-
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coin, Maine who provided collection data, reviewed the manuscript, and made many pertinent comments during the preparation of this paper. Mr. W. H. Shaughnessy of the Academy of Natural Sciences, Philadelphia kindly provided data for specimens collected by Skinner in New Mexico. Mr. Kilian Roever of Phoenix, Arizona kindly provided additional Arizona collection records and made some additional comments upon the habits of n. nitocris. He noted that females of this species generally oviposit in shaded or semi-shaded open woods where there are seeps, although one female was observed ovipositing in an open meadow formed by an old beaver dam. Also supplied were records of specimens referable to n. apacheana from Garfield, Iron, Kane, and Washington Cos., Utah. The intermediate situation between n. nokomis and n. apacheana was noted as discussed above.
Literature Cited
Brown, F. M. 1957a. The McCauley expedition to the San Juan region of Colorado in 1877. J. N.Y. Ent. Soc. 65: 139-145.
---------- 1957b. Colorado Butterflies. Denver.
---------- 1965. The types of the Nymphalid butterflies described by William Henry
Edwards Part I. Argynninae. Trans. Amer. Ent. Soc. 91: 233-350.
dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. Lep. Soc. Mem. 1.
dos Passos, C. F. and L. P. Grey. 1947. Systematic catalogue of Speyeria (Lepi-doptera, Nymphalidae) with designations of types and fixations of type localities. Novitates 1370.
Edwards, W. H. 1868-1872. The Butterflies of North America. Volume 1. American Ent. Soc. Reprinted 1888 by Houghton Mifflin and Company, Boston.
---------- 1874-1884. The Butterflies of North America. Volume 2. Houghton
Mifflin and Company.
---------- 1887-1897. The Butterflies of North America. Volume 3. Houghton
Mifflin and Company.
Ferris, C. D. 1969. Some additional notes on mating behavior in butterflies. J.
Lepid. Soc. 23: 271-272. Holland, W. J. 1898. The Butterfly Book. Doubleday, Page and Co., New York. Macomb, J. N. and J. S. Newberry. 1876. Report of the exploring expedition
from Santa Fe, New Mexico, to the junction of the Grand and Green rivers of
the great Colorado of the West, in 1859. Washington. McCauley, C. A. H. 1879. Report on the San Juan Reconnaissance of 1877, by
Lieutenant C. A. H. M'Cauley, Third Artillery, in charge. In Index to the
Executive Documents of the House of Representatives for the Third Session of
the Forty-fifth Congress, 1878-'79. In 18 volumes. Volume V. Report of the
Chief of Engineers, Part III. Washington. Swisher, W. L. and A. L. Morrison. 1969. News of the Lepid. Soc, 15 April,
p. 4.