AMERICAN MUSEUM NOVITATES Published by Number 1054 THE AMERICAN MUSEUM OF NATURAL HISTORY March 22, 1940 New York City

NEW BUTTERFLY SUBSPECIES FROM WYOMING (NYMPHALIDAE, PIERIDAE) BY ALEXANDER B. KLOTS1 Boloria2 pales halli, new subspecies luster, if any. The spot at the end of the cell is consistently white in color and only faintly This is a very distinct subspecies of pale8 from pearly, never silvery, being thus less lustrous high altitudes in the Wind River Range, Sub- than in any of the other Nearctic subspecies lette Co., Wyoming, the recording of which ex- and most of the Palaearctic ones. Especially tends the known range of pales in North America noteworthy is the considerable reduction of the many hundreds of miles to the southward. submarginal row of red, triangular spots which Hallt may be characterized as follows: in other subspecies form the inner borders of the MALE.-Average length of fore-wing, 18.3 marginal row of silvery spots. In halli most of mm. these red spots are reduced in size; this, in con- Ground-color of upper side of wings bright junction with the above-mentioned almost total orange-brown, very lightly tinged with pinkish absence of white, pearly or silvery, results in a or reddish. The usual transverse rows of spots very characteristic obliteration of the marginal and irregular lines are all present. Those in the row of spots. basal half (or slightly more than half) of the The males show comparatively little variation wings are somewhat narrower than in most of in color and markings, far less than would be the Palaearctic forms of the species, as is also expected in a species occupying a habitat that the case with B. pales alaskensis (Holland). is subject to such extreme climatic changes. The fore-wings are dusted with fuscous scaling FEMALE.-Average length of fore-wing, 19.02 at the bases, somewhat more broadly along the mm, inner margin than along the costa. The hind- Ground-color of most of the area of upper side wings are more heavily shaded basally than the of both wings light gray, tinted with light brown- fore-wings; but this shading almost always fails ish yellow. The color tends to be noticeably to reach the end of the discal cell. lighter, brighter and more yellow in the costal, Ground-color of under-side of fore-wing some- subcostal and apical light areas and spots of the what duller, and with a slightly more pinkish fore-wing, and in the anal light spots of the hind- tinge than that of the upper side. The spots and wing. In sharp contrast to the general ground- bands of the upper side "show through" very color, the costal and subapical region of the bind- faintly, those in and at the end of the discal cell wing has the ground-color bright orange-brown, more strongly. The apex and a narrowing as bright as the ground-color of the male. Dark border that does not reach the anal angle are markings much heavier and more suffused than yellow, slightly tinged with greenish. In this in the male. yellow area some of the spots of the upper side Ground-color of under-side of fore-wing a are repeated, of a brownish-red color; but these somewhat darker, more reddish brown than in are of much smaller extent than the correspond- the male. The black markings of the upper side ing spots on the upper side. are nearly all repeated here, though much smaller Ground-color of under-side of hind-wing yel- than on upper side. The apical and outer- low slightly tinged with greenish; dark markings marginal light area is of a definitely more green- brownish red to bright red. In the great ma- ish yellow than in the male. The dark red jority of the specimens the dark markings are so markings in this area are as in the male. The considerably diluted by dusting of the greenish- black dots at the ends of the veins are heavier yellow ground-color that the entire wing shows than in the male. a noticeable lack of contrast between the light Ground-color of under-side of hind-wing a and dark areas. With the exception of the large more greenish yellow than in the male, the green- spot at the end of the cell, all of the markings, ish tinge being more pronounced in the areas which in other subspecies of pales (especially which on the upper side of the wing are black or most of the Palaearctic ones) are bright silvery fuscous. The dark red- markings average only white, here show only faint traces of a pearly very slightly heavier than in the male; but 1 College of the City of New York. since their hue is in greater contrast with the 2 Boloria Moore, genotype pales (Denis and ground-color, the wing has a more contrasty Schiffermueller) should, in the author's estimation, As in the male the be recognized as a valid genus. Pales has usually and variegated appearance. been placed in the genus Brenthis Huebner. See pearly or silvery white markings are all very also note under the next subspecies here described. much reduced in size and luster. 2 AMERICAN MUSEUM NOVITA TES [No. 1054

There is a great deal of variation' in the ex- only *one specimen, a male from Point tent of the black markings and fuscous dusting on the upper side of the wings of the females, Barrow, Alaska (dos Passos collection), to proportionately more than in the males. There which this name would be applied. It evi- is also considerable variation in the intensity of dently represents an extreme Arctic sub- the dark red markings of the under-side of the species. hind-wings. Below are listed the chief characteristics TYPE LOT.-Holotype male, alIotype fe- that serve to differentiate halli from these male, 139 male paratypes and 57 female three named forms. In the case of paratypes, vicinity of Green River Pass, alaskensis this is based on comparison Wind River Range, Sublette Co., Wyom- with the material listed above; with re- ing, July 18-31, 1939, collected by David gard to reiffi and nearctica it is based mostly Bigelow and the author. The holotype, on the descriptions of these forms by Reuss, allotype and a series of paratypes have Verity and Lehmann. been deposited in the collection of The From alaskensis, halli is distinguished by American Museum of Natural History. the following: The remaining paratypes are in the collec- (1) The deeper orange-brown ground-color tions of Mr. Bigelow and the author. of the upper side of the wings of the males. (2) The much lighter, yellow-brown tinged, Series of them will be distributed to numer- gray ground-color of the upper side of the wings ous museums. of the female, which contrasts strongly with the COMPARISON WITH OTHER NORTH AMERI- bright yellow-brown, costal and apical region of CAN SUBSPECIES OF pales.-Three other the hind-wing. In alaskensis females the ground-color of most of the area of the wings is a names have been applied to forms of pales much darker, duller yellow-brown; tbat of the in North America; these are: costal and apical region of the hind-wing is not as bright as in haili; and so the general appear- (1) Brenthis pales var. alaskensis HOLLAND, ance of alaskensis females is much duller and less 1900, Ent. News, XI, p. 383. contrasty. (2) Boloria reiffl REUSS, 1925, Intern. Ent. (3) The reduction in size, on the upper side Zeitung, XIX, pp. 279-280. of both wings of the males, of the submarginal (3) Boloria pales nearctica VERITY, 1932, row of black spots. In alaskensis these tend to Deutsch. Ent. Zeitschr., Iris, XLVI, pp. 104 be more produced inwardly, forming inward- and 108. pointing triangles. In halli they tend to be less triangular in shape. These spots also "show Of these I have been able to study satis- through" less on the under-side in halli, a factory material of alaskensis only for characteristic which accentuates the lack of comparison with halli. Of this subspecies marginal spots in this form (see no. 7 ff.). I have studied critically 30 males and 23 (4) The slightly greater reduction of the basal fuscous dusting on the upper side of both females from various southern and western wings of the males. This is more noticeable localities in Alaska, and Atlin, B. C. on the hind-wings, for in haili the dusting almost These are in the American Museum, dos always fails to reach the end of the discal cell, Passos and Klots collections. while in alaskensis it nearly always does so. (5) The reduction of the dark, red-brown, Reiffi was described from a single male subapical markings on the under-side of the fore- from an unknown locality in British Colum- wings of both sexes. bia. Reuss mentions particularly the bril- (6) The reduction and dilution by the liance and contrast of the red and silver ground-color of the dark, red-brown markings of the under-side of the hind-wings of both sexes. markings of the under-side, a characteriza- (7) The reduction of the white, pearly or tion that applies to one of the Atlin, B. C., silvery markings of the under-side of the hind- specimens studied by me but not to the wings of both sexes, especially the almost total other. I suspect that reiffi will eventually loss of the marginal row of pearly or silvery spots. In halli there are at most only faint prove to be a synonym of alaskensis. suggestions of these spots in all but a few speci- Nearctica was described by Verity on the mens. An exception is the spot in cell Cu, basis of Lehmann's description (Seitz, which is usually fairly well defined and slightly Macrolepidoptera.., V, p. 423, 1913), as pearly. In alaskensis this row of spots is nearly always well defined, being definitely silvery or Argynnis pales, of a number of specimens pearly, and margined inwardly with red-brown. from extreme northwestern Alaska, Lat. (8) The brighter, slightly more greenish- 69° 40' N., Long. 1410 W.). I have seen yellow color of the apex and margin of the under- 1940] NEW BUTTERFLIES FROM WYOMING 3 side of the fore-wing and the ground-color species occurs over an area of two hundred of the under-side of the hind-wing. or more square miles in the Wind Rivers, From reiffi, halli is distinguished by the for this mountain range has a very large following: area above timberline. This being so, I (1) The lack of contrast on the under-side am very much surprised that I failed to of the hind-wings of both sexes, between the take the species in 1935 on the mountains greenish-yellow ground-color and the dark, around Green River Lake, for on at least red-brown markings; this is due to the reduction in extent and dilution in hue of the latter. two of these I collected far above timber- Reiffi is stated in the original description to line. It is possible that some temperature have the yellow ground-color strong and out- factor limits the species to the interior standing ("stark, vortretende, gelber Grund- areas of the range in proximity to the farbe"). high peaks. (2) The great reduction of the white, silvery glaciers and the very markings of the under-side of the hind-wings. Females were proportionately much less Reiffi is stated in the original description to have often observed visiting flowers than were the silver marks strong and normal. the males; but individuals were frequently From nearctica, halli is distinguished by observed resting in the low willows (Salix) that form dense mats in the Hudsonian the following: Zone bog areas. (1) The greater reduction in the males of Specimens were very rare in the higher the basal fuscous dusting on the upper side. Nearetica was described (by Lehmann, whose meadows of the true Arctic-Alpine Zone description forms the basis for Verity's applica- above 11,000 ft. alt., which is surprising. tion of the name) as having the fore-wings Only occasional strays from theAtower heavily dusted basally, and the hind-wings levels were seen. This is in direct c6ntrast broadly shaded with greenish black as far as the apex of the cell. In practically all speci- to some of the other butterflies such as mens of halli the black basal shading definitely Euphydryas, Aglais milberti, Plebejus saepi- does not extend to the end of the cell, a character- olus, Erebia theano ethela, E. tyndarus istic which also as noted serves to differentiate callias, etc., which, although they occur it from ala8keris%. (2) The lack of contrast and variegation of most abundantly in Hudsonian Zone, the markings of the under-side of the hind- frequently fly up into even extreme Arctic- wings. Lehmann calls nearctica "uncommonly Alpine Zone and are often common there. variegated" in this respect. On the wing, halli males appear to have (3) The reduction of the white, silvery mark- a somewhat lighter, more rapid, and more ings of the under-side of the hind-wings. darting flight than the species of Brenthis FIELD NOTES, ETC.-Halli was very taken in the same localities. The slightly common in Hudsonian Life Zone meadows, pinker hue of the ground-color of the wings at and above timberline, which in the vi- of the male is more noticeable in specimens cinity of Green River Pass is at about in flight in the sunshine than in dried 10,00-10,500 ft. altitude. The males specimens. On the wing the females re- were everywhere, visiting flowers of Arnica, sembled very strongly the dark, worn Erigeron, Polygonum and Eriogonum. The specimens of Phyciodes that were flying females were much less common, although with them. The flight of the females is the presence of many rather worn indi- much less swift and less graceful than that viduals toward the end of our collecting of the males. period showed that we had collected ACKNOWLEDGMENTS.-The first speci- through the height of the halli season. mens of halli collected are said to have been We took specimens everywhere that we taken in the Wind River Range many collected in Hudsonian Zone, i.e., in Stroud years ago by Mr. Carl Rungius, the well- Creek Canyon, around Peak Lake, along known artist. Later, Mr. Gaylord C. the upper reaches of Trail Creek, around Hall of New York City, who has done Summit Lake and the Elbow Lakes, in much collecting in this range, took a series. the upper parts of Pine Creek Canyon, On hearing that I intended collecting at and all along the trail from Summit Lake high altitudes in the Wind Rivers, Mr. to Palmer Lake. Without a doubt the Hall generously placed his knowledge of 4 AMERICAN MUSEUM NOVITATES [No. 1054

the region, and of collecting and conditions specimen in the type series of laddi (17 there, at my disposal. So it is with great specimens) is as light-colored beneath as pleasure that I name this subspecies for even darker than average specimens of him. caelestis; and of the caelesti8 available for comparison (24 topotypical specimens, Hall Clossianal aphirape laddi, Valley, Colo., and 7 specimens from other Colorado localities) only two, both females new subspecies (which sex averages darker beneath), are In a previous paper (Journ. N. Y. Ent. as dark as any of the laddi. Soc., 1937, XLV, p. 328) the author re- As mentioned in a previous paper (loc. corded the capture of four specimens of cit.) the Snowy Range is not very definitely aphirape in the Snowy Range, Wyo., separated from the Colorado mountain and commented upon their apparent differ- ranges, so that the presence there of differ- ence from C. aphirape caelestis (Hemming) ent subspecies is a bit surprising. Study (alticola (Barnes and McDunnough)), the of series of other Snowy Range butterflies Colorado subspecies. In July, 1939, an seems to show, however, that in a number additional series of the Snowy Range form of the species there are slight but constant was collected by Anthony Ladd of New differences from the corresponding Colo- York City and the author. These agree rado forms. so consistently with the previously col- Aside from the darker coloring beneath, lected specimens that the author has no laddi shows no other points of difference hesitation in naming the Snowy Range from caelestis and similarity to dawsoni or population as a new subspecies which is other Canadian subspecies. This particu- dedicated to the co-collector. larly refers to the silvery hue of the white From C. aphirape caelestis (Hemming), spots and the ocellate spot in the dark, its nearest relative, laddi is distinguished basal spot in the cell, both on the under- by the following characteristics: side of the hind-wing, which were dis- (1) The very considerably darker, red- cussed in my previous paper (loc. cit.) brown color of the subapical patch on the under- as points of difference between caelestis side of the fore-wing, and the similarly darker, and dawsoni. red-brown color of the submarginal, median and post-basal bands of the under-side of the TYPE SERIEs.-All specimens of type hind-wing. In caelestis these dark markings series from Snowy Range, Albany Co., are of a much lighter, more yellow-brown. Wyo. Holotype e, 9 cl and 3 9 para- (2) A tendency for the submarginal rows of types, July 7, 1939, collected by Ladd and round, black spots on both upper and under- sides of the fore-wing to be somewhat sinuate, Klots in willow bog, Hudsonian Zone, alts. bending proximally below M2 and distally again 10,400 to 10,800 ft. Allotype 9, 1 d, and below Cu2. In caelestis this row of spots tends 1 9 paratype, August 13, 1935, collected to be straighter. by Klots, in willow bog, near Class (Lewis) Laddi is not an extremely distinct sub- Lake, Hudsonian Zone, alt. 10,900 ft. species in the sense of every specimen being 1 e paratype (with wings incompletely distinguishable from any specimen of any expanded) July 17-23, 1935, collected by other subspecies. Very few subspecies are. Klots, in willow bog near Class (Lewis) But that there is a very considerable and Lake, Hudsonian Zone, alt. 10,900 ft. almost constant difference between it and Holotype, allotype, 3 ci and 1 9 para- caelestis is evident at a glance upon com- types in The American Museum of Natural parison of series of the two forms. No History. The remainder of the paratypes are in the author's collection. Some will 1 Clossiana Reuss, genotype gelene (Denis and Schiffermueller), should be recognized as the generic be distributed to museums. name for those North American species other than Boloria pales which have heretofore usually been Colias gigantea harroweri, placed in Brenthis Huebner. Brenthis should be re- stricted to the Palaearctic species hecate (Esper) the new subspecies genotype, ino (Rottemburg) and daphne (Denis and Schiffermueller). Reuss has erected the subgenus For many years this, as well as other Proclo8siana for aphirape; but the validity of this seems a bit dubious. yellow Colias from western North America, 1.940] NEW BUTTERFLIES FROM WYOMING 5 has been passing under the name of Colias heavier. Discal spot prominent. Hind-wing A small pale yellowish, evenly but not very heavily christina form astraea Edwards. dusted with fuscous scales. Discal spot large, series taken by the author in 1935 (now a more or less double, only slightly pearly in- part of the present type lot) was thus tenta- ternally, very diffusely ringed with light purplish tively named (Journ. N. Y. Ent. Soc., brown of a rather more dilute hue than in male. more broadly pink. 1937, XLV, p. 321). Further collecting Fringes as in male, in 1939 in the same locality in which this TYPE LOT.-AII are from Clear Creek, 1935 material was taken has furnished addi- vicinity Lower Green River Lake, Sublette tional specimens and evidence which not Co., Wyo., alt. 8400 ft. Holotype male, only warrants the description of this form three male and two female paratypes, as new but also shows that in all probability August 3, 1935, collected by Klots; allo- it is a southern subspecies of Colias gigantea type female, nine male and one female para- Strecker. The type locality of gigantea types, July 16, 1939, collected by Klots, is in the vicinity of Churchill, Manitoba. except the female paratype which was col- lected by David Bigelow. MALE-Average length of fore-wing, 24.5 FIELD NOTES.-The allotype female was mm. Ground-color of upper side of wings yellow, ovipositing on willow (Salix) when cap- slightly greenish tinged. Black border of nor- tured. All the specimens were taken in a mal width, tending to be rather smooth in- somewhat acid, moss (Hypnum)-willow wardly, somewhat produces basally along inner margin of fore-wing. Veins strongly yellow in bog area that lies in the Clear Creek black borders. Black discal spot of fore-wing "bottom" about three-quarters of a mile present in all specimens studied, usually strongly below the "natural bridge" which is shown defined, sometimes large, reduced to a trace in on the Fremont Peak quadrangle of the only one of the twelve males of the type series, U. S. Topographic Map. A number of and very small in one other. Discal spot of hind-wing "showing through" from under-side other females were seen dropping into the as a very pale orange spot. vegetation of the bog as if to oviposit. General color of under-side somewhat lighter Willow is probably the only possible food- yellow than that of upper side, paler on the inner for the species in this bog. The margin, and deeper, almost orange, at the plant extreme base of the fore-wing. Fore-wing species is very strongly limited to the bog fairly heavily dusted with fuscous scales along environment; of all those seen in 1935 and costa, more lightly dusted on disc beyond end of 1939, including at least a dozen and a half cell and on apex. Discal spot of fore-wing that were not taken, none were outside prominent, with many yellow and some yellow- brown scales internally. of the limits of the bog area. Under-side of hind-wing rather greenish DISCUSSION.-As mentioned previously, yellow, evenly but not very heavily dusted this form has for many years been passing with fuscous scales; this dusting tends to be lighter toward the outer margin. Discal spot under the name of Colias astraea Edwards, of hind-wing large, often double, pearly in- of which the type locality is Yellowstone ternally, diffusely ringed with purplish brown National Park, Wyoming. The name which frequently invades and tints the central, astraea should be used only for pale orange pearly region. or specimens of the species christina Fringes pink, more yellowish above than be- yellow low. Fringe of fore-wing becomes yellow below Edwards, although the type specimen of vein Cu2 of 2dA. On the under-side the costal astraea (in the Carnegie Museum) has margin of both wings is distinctly purplish pink; considerable orange on the upper side of and the outer margin of the fore-wing and the It be noted that Edwards outer and anal margins of the hind-wing are the wings. may more narrowly pink. described astraea from a single male only, FEMALE.-Upper side: Ground-color bright so that the male and female figured as yellow. Black border absent, or present as only "types" by Holland (Butterfly Book, re- a trace in the subapical region. Discal spot of 1931, Pi. LxvIII, fig. 26, 27) fore-wing present, large, fuscous or black. vised edition, Discal spot of hind-wing large, double, solid, are not types at all, but merely specimens bright orange. from the Edwards collection. The male Under-side: Fore-wing. Basal and discal thus illustrated is considerably less orange parts of wing yellow, somewhat orange tinted; apical, terminal and inner-marginal parts pale than the true type. yellow, fuscous dusting as in male somewhat McDunnough (Canadian Entomologist, 6 AMERICAN MUSEUM NOVITATES [No. 1054

1922, LIV, p. 135, and 1928, LX, p. 270) (2) g. gigantea, 52 males and 12 females, all has called attention to the essential differ- topotypes from Churchill, Manitoba; 19 males ences between gigantea and the yellow and 17 females from Alaska. forms of christina, pointing out the food- Because of the variation shown by plant difference between the two species. harroweri it has seemed best to limit the From these observations and my own it type series to the specimens from the seems that the forms of the christina- Clear Creek colony for which there is emilia-alexandra complex are legume feed- habitat data. ers, while gigantea feeds exclusively on COMPARISON WITH g. gigantea.-From willow. Correlated with this is a marked g. gigantea, harroweri is distinguished by the difference in habitat: christina ffies in the following: dry open country, while meadows and (1) The black borders of the upper sides of gigantea is limited in the southern part the wings of the males, which are somewhat of its range to willow bogs and in the north wider than in g. gigantea. to tundra. (2) The color of the apex of the under-side of Mr. of the fore-wing and of the entire under-side of In this respect some observations the hind-wing of the males. In harroweri these Jack May, the experienced Canadian lepi- areas are of a greenish yellow which does not dopterist, are pertinent. In a letter to contrast strongly with the ground-color of the me a number of years ago Mr. May stated rest of the fore-wing. In g. gigantea these an for areas are of a richer, somewhat orange-yellow that he had found excellent locality which contrasts with the paler, more dilute hue gigantea, namely, in the large swamps in of the rest of the fore-wing. the interior of the Riding Mountains, Mani- (3) The fuscous dusting of the under-side toba. I have received a considerable of the hind-wings of both males and females. In harroweri this is much sparser than in g. series of these specimens from him. gigantea. Accordingly I have no hesitation in as- (4) The ground-color of the females. In signing this Wyoming form to gigantea, harroweri this is usually bright yellow, as in all of which species it constitutes the most four specimens of the type series, three out of The name harroweri four specimens from Yellowstone Park, and southern subspecies. thirteen out of seventeen specimens from should certainly be applied to Wyoming, southern Manitoba and Alberta. Of the five Idaho and Montana specimens of gigantea; exceptions one from Yellowstone and one from and it probably should also be used for soutbern Manitoba are white, and the other those from the southern regions of Mani- three are pale yellow. Of twelve female topo- types of g. gigantea six are white and six are toba, Alberta and British Columbia. very pale yellowish. The southern Canadian specimens are not (5) In a number of other characters there typical, however, showing in some respects seem to be minor differences; these are: size, and in other in which harroweri seems to average very slightly intergradations to g. gigantea, smaller than gigantea; the black discal spot of respects intergrading to christina, emilia, the fore-wing, which seems to average slightly alexandra and occidentalis in a very larger in harroweri; and the border of the puzzling way. fore-wing of the female, which seems to be has been studied slightly less reduced in gigantea. The number The following material of specimens available for study is not great critically in preparing these diagnoses enough, however, to warrant any assurance with (American Museum and Klots collections): regard to these. (1) harroweri, in addition to the type series, I take pleasure in naming this subspecies 75 males and 21 females from various localities K. Harrower of the U. S. in Wyoming, Montana, Idaho, Manitoba, Al- for Mr. James berta and British Columbia. Forest Service, Pinedale, Wyoming.