SKIPPERS AND OF A DISJUNCT ASPEN PARKLAND AREA IN ALBERTA

NORBERT G. KONDLA, Parks Division, Alberta Recreation and Parks, Rimbey, Alberta TOC 2J0.

, Accounts of faunas and The study area was situated in 28- habitat use of the Alberta aspen 26-18-W4, about 11 km south of East parkland are scarce. Case and Bird Coulee, on the highest portion of the' provide some information on the hills where elevation ranges from 945 Peace River parkland5 and Thormin m to 1037 m. Here the escarpment is deals with the Beaverhill Lake area.24 dissected by ravines, with local relief However, other accounts treat in¬ varying from gently rolling to steeply dividual species (e.g. Bird3) or tax¬ sloping. Figure 1 shows the general onomic matters (e.g. Ehrlich9) rather appearance of the study area. , j than the typical fauna of a given About 50% of the study area is habitat. This paper reports on the covered by Aspen Poplar forest with a most detailed study of a local aspen moderately well-developed low shrub parkland butterfly fauna undertaken understory of Rose and Buckbrush. to date in Alberta. The herbaceous component of the The central aspen parkland section forests is rich and includes such is a fairly distinctive zone in south- species as Sprengel’s Sedge, Wild central Alberta. Two outliers of aspen parkland are present near Drumheller, on the Wintering Hills and on the Hand Hills. The Wintering Hills, an upland rising more than 120 m above the plains southwest of Drumheller, are a generally east-west oriented ridge, the northeastern es¬ carpment of which supports a sub¬ stantial area of aspen parkland which is separated by a distance of about 80 km from the central aspen parkland. For this study, we selected a representative area of about 260 ha for collection and observation of butterflies. All habitats were intensive¬ ly searched by the writer and S. Harris from 1976 to 1980 inclusive on 20 different days with a seasonal spread from 1 May to 3 September. Visits were timed to minimize the possibility of missing any species because of its flight period. Figure 1. General view of the study area.

4 Blue Jay /etch, Peavine, Canada Anemone, Hesperlldae ind Western Canada Violet. TAWNY-EDGED SKIPPER (Polites About 30% of the study area con- themistocles) — a worn specimen ;ists of grassland, most of which was was found 1-8-77 in low shrub habitat lominated by Rough Fescue prior to near aspen forest; early to mid¬ he era of intensive cattle grazing, summer. /luch of this grassland has been al- LONG DASH (Polites mystic dacotah) — ered to the point where Porcupine uncommon in grassland; 13-7-79; 5rass and June Grass are now flight period not known. lominant. Certain slopes with a south UNCAS SKIPPER (Hesperia uncas uncas) md west exposure support an open — one female was found 29-7-78 on a irass-forb vegetation typical of west-facing hillside in open Por¬ :oulees in the region. cupine Grass-June Grass vegetation; flight period not known. The remainder of the study area is ASSINIBOIA SKIPPER (Hesperia comma >ccupied by shrubiands and mis- assiniboia) — locally abundant 29-7- :ellaneous local vegetation types. 78, 20-8-79, 3-8-80; mid to late >mall stands of Hawthorn, summer; this was easily ob¬ ^hokecherry, and Saskatoon are served on flowers of Liatris punctata (Blazing Star) and showed pro¬ >resent, as are larger areas of nounced hilltopping behavior; up to 8 Jeaked Willow. Low shrub thickets of individuals would frequently engage Juckbrush, Rose, and Gooseberry in rapid, erratic flights in tightly ire prevalent. Miscellaneous spaced groups. egetation types are Bearberry mats, i White birch stand, and two small NEVADA SKIPPER (Hesperia nevada) — eepage areas. very locally distributed on hilltops and ridge crests with short grass; 23-6-79. The area is floristically interesting, GARITA SKIPPER (Oarisma garita) — ^specially from a biogeographic abundant in grasslands; 19-6-77, 13- >erspective. In a small area one can 7-79; late spring to mid-summer; ind such unlikely associates as more abundant in less heavily grazed leart-leaved Arnica (a mountain grasslands. pecies), White birch (a boreal forest ARCTIC SKIPPER (Carterocephalos pecies), and Pincushion Cactus (a palaemon mandan) — occasional in •rairie-badlands species). and near aspen forest; 19-6-77; mid to late spring. Annotated List COMMON CHECKERED SKIPPER (Pyrgus communis communis) — abundant in In the following list the term spring grassland and low shrub habitat; 3-9- efers to May and June while summer 78, 23-6-79, 13-7-79; late spring to efers to July, August and the first few late summer. ays of September. Butterfly DREAMY DUSKY WING (Erynnis icelus) — henology is described with terms occasional in and near aspen forest; ke “early spring to mid-summer.” 21-5-78, 23-6-79; mid to late spring. lates indicate days for which PERSIUS DUSKY WING (Erynnis persius) pecimen records are available and — fairly common in grassland near re written in an abbreviated format aspen forest; 30-5-76, 21-5-78, 9-5- 3.g. 29-7-78 means 29 July 1978). 80, 18-5-80; early to late spring. lost of the common names are from AFRANIUS DUSKY WING (Erynnis looper12. Scientific names follow dos afranius) — fairly common in assos7 except where more recent grasslands; 29-7-78, 9-5-80, 11-5-80; rork indicates that a change is war- flies in two broods, early to mid spring anted. and mid to late summer.

arch, 1981. 39(1) 5 NORTHERN CLOUDY WING (Thorybes along the Red Deer River.18 Hooper pylades) — uncommon at the edge of use of the name Cypress Hills Ol aspen forest; 30-5-78, 23-6-79; mid to World Swallowtail is misleading.12 Th late spring. taxon dodi was first described fror badlands and the greatest portion c SILVER-SPOTTED SKIPPER (Epargyreus its range comprises badlands/prairi| clarus clarus) — occasional in habitats. Hence the name Badlandl clearings in aspen forest and along Old World Swallowtail seems morj edges of forest; 23-5-76; mid to late appropriate. spring. CANADIAN TIGER SWALLOWTAi ji (Papilio glaucus canadensis) - abundant in and along edges c aspen forest; most frequently see patrolling the edge of the aspen fores at heights of 2 to 5 m; 30-5-76, 23-6 79; mid to late spring.

Silver-spotted Skipper. C. Wallis

Canadian Tiger Swallowtail. C. Wallh

Papillonldae Plerldae ZELICAON SWALLOWTAIL (Papilio WESTERN CHECKERED WHITE (Pierii\\ zelicaon nitra) — follows occidentalis occidentalis) — fairh Fisher11; males of the black abundant in all non-forest areas; fre phenotype were found 8-5-77 and 23- quently seen hilltopping; the spring 6-79 on grassland hilltops; males and females of the more abundant yellow brood is dramatically smaller anc| phenotype were also found hilltop¬ darker than later broods; 1-5-77, 6-5 [ ping 1-5-77, 14-5-78, 29-7-78, 23-6- 78, 3-9-78, 23-6-79, 20-8-79, 4-8-80 i 79, 9-5-80, 11-5-80; early spring to early spring to late summer in multi- mid summer; this lengthy range of pie broods. dates combined with the fresh con¬ CABBAGE BUTTERFLY (Pieris rapae) - dition of the 1978 record suggests occasional in open areas; 1-5-78, 3-9- either an extremely staggered 78, 2-9-79, 9-5-80; early spring to late emergence or a small second brood. summer. BADLANDS OLD WORLD SWALLOWTAIL YELLOW ALFALFA BUTTERFLY (Co//as| (Papilio machaon dodi) — only found philodice eriphyle) — abundant in all on a grassy hilltop; 23-6-79, 20-8-79, non-forested areas; albinistic females) 9-5-80, 11-5-80, 18-5-80, 3-8-80. This were found fairly frequently; 30-5-761 interesting swallowtail was described 23-5-77, 1-8-77, 16-8-77, 29-7-78, 3-j in 1939 by J. McDunnough primarily 9-78, 20-8-79, 2-9-79, 18-5-80; early on the basis of specimens collected spring to late summer.

Blue Jayl 6 I .EXANDRA SULPHUR (Colias alexandra forewing, occurs only about 100 km to alexandra) — fairly abundant in native the north at Buffalo Lake. When the grasslands; 29-7-78, 23-6-79, 20-8- zone of intergradation between 79, 18-5-80; mid spring to late fletcheri and aliparops is summer. documented in this region, it may well All females found to date have been be a very narrow zone similar to the albinistic, a situation that appears to situation reported by Clench for two be normal for Alberta populations of other subspecies of the Striped this taxon. Ferris assigned Alberta Hairstreak in the southeastern USA.® prairie material to subspecies astraea'0 but since then research has BROWN ELFIN (Callophrys augustinus) — very locally distributed; found only in shown the presence of pure yellow a Bearberry patch at the top of the es¬ populations in the prairie region of carpment; 1-5-77, 6-5-78; early to mid Alberta. The name used here for these small, yellow prairie Is spring; insufficient voucher material used provisionally as the most logical is available to assign a subspecies available name. In fact there is no name with confidence; geographic published research to show that these character variation of this species in populations are the same as the Alberta needs to be investigated. southern C. a. alexandra. Further GRAY HAIRSTREAK (Strymon melinus) — study may even show that these pop¬ only one specimen found on 11-5-80. ulations represent a distinct species. PURPLISH COPPER (Epidemia helloides) This insect was not reported from the — found at a seepage area on 23-6- prairies of southern Saskatchewan by 79, 2-9-79; late spring to late summer. Hooper12 although it probably occurs there. Here is an opportunity for MELISSA BLUE (Lycaeides melissa naturalists in the prairies of southern melissa) — uncommon in dry Saskatchewan to provide specimens grassland; 23-6-79, 2-9-79, 18-5-80, for taxonomic and biogeographic 4-8-80; mid spring to late summer. study. [There are two known GREENISH BLUE (Plebejus saepiolus specimens for southern arnica) — fairly common in grassland Saskatchewan, -both from the and low shrub areas; 23-5-76, 4-6-77, Cypress Hills. One was collected 18- 5-80; mid to late spring. several years ago by Donald Hooper, the other in July 1977 by W. Harris ACMON BLUE (Plebejus acmon lutzi) — and S. Lamont. — Editors] very local in dry grassland with bare soil and umbrellaplant (Eriogonum .YMPIA MARBLE (Euchloe Olympia) — flavum); 23-6-79. only one specimen found on 18-5-80; RUSTIC ARCTIC BLUE (Agriades glandon the most northwesterly reported rustica) — locally abundant in dry locality for this species in North grassland; fond of nectaring at America. flowers of Silverberry (Elaeagnus comutata)-, 23-6-79, 2-9-79; this late caenldae record plus another from Rumsey in August 1979 suggests a small second )RAL HAIRSTREAK (Harkenclenus titus brood, apparently an unreported immaculosus) — males were found phenomenon. hilltopping on 29-7-78 and 20-8-79; mid to late summer. WESTERN-TAILED BLUE (Everes amyn- DCKY MOUNTAIN STRIPED tula albrighti) — fairly common in and HAIRSTREAK ( liparops near aspen forest; 23-5-76, 30-5-76, aliparops) — this is an inconspicuous 19- 6-77; mid to late spring. and apparently uncommon butterfly SILVERY BLUE (Glaucopsyche lygdamus in Alberta; one female was found 16- couperi) — common in grassland and 7-77, perched on a Buckbrush leaf at low shrub habitats; 23-5-76, 30-5-76, the edge of aspen forest. The sub¬ 1-5-77, 8-5-77, 23-5-77, 4-6-77, 14-5- species fletcheri, characterized by 78, 9-5-80, 11-5-80, 18-5-80; early to large orange patches on the dorsal mid spring.

arch, 1981. 39(1) 7 SPRING AZURE (Celastrina argiolus lucia) — occasional in clearings, edges of forest, and more open portions of the aspen forests; 1-5-77; early to mid spring. Nymphalldae WHITE ADMIRAL (Limenitis arthemis rubrofasciata) — occasional in aspen forest, clearings, and tall shrub thickets; 16-7-77, 13-7-79; late spring to mid-summer.

Satyr Angle Wing. C. Wall

23-5-77, 11-5-80; previous literatui referred Alberta material to the sul species satyrus but dos Passos refe Alberta populations to neomarsyai White Admiral. C. Wallis this situation needs further study. GRAY COMMA (Polygonia progne) ■ RED ADMIRAL (Vanessa atalanta rubria) found in aspen forest on 13-5-79 an — occasional in or near forest and tall 3-7-79; the latter record is of the di; shrub habitat; 19-6-77; flight period tinctive summer form, l-argenteum\ not known. PEARL CRESCENT (Phyciodes tharos) | PAINTED LADY (Cynthia cardui) — oc¬ abundant in grassland and low shrul casional in most habitats and areas; mid spring to mid-summer; 1$ sometimes abundant on hilltops; late 6-77, 16-7-77, 29-7-78, 18-5-80. one afternoon about 50 to 60 males SILVER-BORDERED FRITILLARY (Bolon and females were found on a 50 m by selene) — uncommon and local in c 20 m section of grassy hilltop; 23-6- near sedge-rich seepage meadow; 79, 20-8-79; only found in 1979, a year 13-7-79. in which the species was generally abundant in southern Alberta. MILBERT’S TORTOISE SHELL (Nymphalis milberti furcillata) — occasionally seen in all habitats; 1-5-77, 19-6-77, 13-5-79; flight period not clear. MOURNING CLOAK (Nymphalis antiopa antiopa) — occasional in aspen forest and tall shrub habitat; worn, overwintered individuals were fre¬ quently seen in early spring while fresh adults were seen from mid¬ summer to late summer; 3-9-78. SATYR ANGLE WING (Polygonia satyrus) — uncommon in aspen forest; 8-5-77, Silver-bordered Fritillary.

8 Blue Ja^ EADOW FRITILLARY (Boloria bellona areas; early to late summer; 16-7-77,

jenistai) — abundant in rich grassland 29- 7-78, 3-9-78, 20-8-79, 2-9-79, 3-8- and low shrub areas; mid spring to 80, 4-8-80; Kondla has identified the mid-summer, in at least two broods; need for research to clarify variation 23-5-76, 30-5-76, 23-5-77, 4-6-77, 16- at the subspecies level in Alberta.13 7-77, 21-5-78, 29-7-78, 18-5-80. VARIEGATED FRITILLARY (Euptoieta DWARD’S FRITILLARY (Speyeria claudia) — one female was found edwardsii) — known only from one hilltopping on 20-8-79. fresh male on 30-5-76 at the edge of aspen forest; an exceptionally early Satyrldae record for this uncommon but distinc¬ RINGLET (Coenonympha inornata ben- tive species. jamini) — abundant in richer ^LLIPPE FRITILLARY (Speyeria callippe grasslands and low shrub habitats; calgariana) — locally abundant in mid-spring to early summer; 23-5-76, grassland, especially on hilltops; late 30- 5-76, 23-5-77, 4-6-77, 23-6-79,18- spring to mid-summer; 19-6-77, 29-7- 5-80. 78, 23-6-79, 13-7-79. RIDING’S SATYR (Neominois ridingsii ridingsii) — rare on grassy slope with considerable proportion of bare ground; 13-7-79. COMMON WOOD NYMPH (Cercyonis pegala ino) — abundant in richer grassland and low shrub habitats; early to late summer; 16-7-77, 1-8-77, 29-7-78, 13-7-79, 4-8-80. VARUNA ARCTIC ( uhleri varuna) — fairly abundant in grasslands; early to late spring; 30-5-76, 8-5-77, 23-5- 77, 4-6-77, 23-6-79, 9-5-80, 11-5-80, 18-5-80. ALBERTA ARCTIC (Oeneis alberta alber¬ ilippe Fritillary. C. Wallis ta) — locally abundant in grasslands; early to mid spring; 1-5-77, 8-5-77, ’LANTIS FRITILLARY (Speyeria atlantis) 14-5-78, 21-5-78, 13-5-79, 9-5-80, 11- — very abundant in open habitats; 5-80, 18-5-80; emergence varies by 2 late spring to mid-summer; 19-6-77, to 3 weeks depending on spring 16-7-77, 13-7-79; although typically weather; in late springs the main flight variable, this population is period of the Alberta Arctic overlaps predominantly of the darker beani that of the Varuna Arctic. phenotype rather than the paler RED-DISKED ALPINE (Erebia discoidalis helena phenotype. mcdunnoughi) — fairly common but locally distributed in fescue RMONIA FRITILLARY (Speyeria mor- grasslands; early to mid-spring; 1-5- monia eurynome) — abundant in 77, 14-5-78, 11-5-80. most open areas; late spring to mid¬ summer; 16-7-77, 29-7-78. COMMON ALPINE (Erebia epipsodea) — fairly common in richer grasslands 3 EAT SPANGLED FRITILLARY and low shrub areas; mid-spring to (Speyeria cybele pseudocarpenteri) early summer; 4-6-77, 23-6-79; — very abundant, especially at the Ehrlich shows the study area to be in edges of aspen forest; females were a zone of intergradation between E. e. frequently noted well within the freemani and E. e. epipsodea.9 He woods; early to mid-summer; 16-7- also indicates that no abrupt boun¬ 77, 29-7-78, 13-7-79, 4-8-80. daries exist between the subspecies 5HRODITE FRITILLARY (Speyeria and that considerable variation within aphrodite) — very abundant in open a population masks geographic arch, 1981. 39(1) 9 variation unless longer series of Alberta, i.e. Calgary2, Plateau Mtn. specimens are available for study. Kananaskis Provincial Park14, Star When more material is available this dard13. In the present study aree! population will likely be assignable to species diversity rises rapidly t the subspecies freemani. almost maximum by mid spring with poorly defined peak in late spring an then a slight drop followed by hig Discussion diversity until midsummer. Only i A total of 57 species have been late summer is there a substantia found, 55 of which are probably reduction in species diversity. resident in the study area. The fauna According to the account of mat' is comprised of: Hesperiidae 13 locating behavior given by Scott, th (23%), Papilionidae 3 (5%), Pieridae 5 two major methods used to locafr (8%), 12 (21%), mates are: perching behavior (male 17 (29%), Satyridae 7 rest at characteristic sites and in (12%). Except for a substantially vestigate passing objects by flying ou enriched Hesperiidae fauna and at them) and patrolling behavio depauperate Pieridae fauna, this is (males fly almost continuously ii representative of the situation in search of females).18 , Alberta as a whole. As in most prairie areas, the skipper fauna here is rich in Information provided by Scott19 an< comparison to the boreal and cor- personal observations show that th' dilleran regions of the province. For a fauna of the study area is made up c non-cordilleran area, the Speyeria 31 species which predominantly us patrolling behavior and 24 specie fauna is surprisingly rich, in both which predominantly use perchin* number and abundance of species. The violet flora (food plants of the behavior to locate mates. Th greater fritillaries) is equally rich, in- Hesperiidae are predominant! cluding Viola adunca, V. perchers while the Papilionidae an nephrophylla, V. nuttallii, V. inveterate patrollers. Some specie pedatifida, V. rugulosa. peculiarities that were noted are: th< Assiniboia Skipper and Nevada The Satyridae are predominantly Skipper characteristically perch oi spring fliers here. The few Pieridae of hill and ridge tops; the Zelicaoi the study area are found throughout Swallowtail and Badlands Old Worl< the spring and summer. Most of the Swallowtail patrol less than 2 metre Lycaenidae, especially the blues, are above the ground on grassy hilltop spring fliers although the hairstreaks while the Tiger Swallowtail patrol are distinctly summer fliers. The mostly at heights of 2-5 metres at th< Nymphalidae fly most abundantly edge of aspen forest. The Pieridae an from late spring to midsummer. all patrollers, as are most of th< Skippers include spring flying species Lycaenidae, although the hairstreak: with legumes and poplars for larval and coppers are perchers. food plants, and summer flying Most of the Nymphalidae are species with grasses for larval food patrollers, especially the greate plants. Overall abundance and diver¬ fritillaries which are very active. The sity was greatest in the spring in the Satyridae are interesting because the grassland and in summer in early flying species use perchine shrubland and woodland habitats. behavior supplemented by patrolline Butterfly phenology data does not while the later flying species are show a pronounced short term peak patrollers. The visibility of individua in species diversity as other areas in butterflies is correlated with the

10 Blue Jay nount of time they spend flying. The ct that patrolling males spend more ne flying than do perching males is a definite influence on population itimates based on visual estimates flying butterflies. Since the study area is the highest II in the region with many local, nailer scale topographic highs, this scussion would not be complete thout treating the phenomenon of Itopping. In the broadest sense the 'm refers to the congregation of in- cts on various kinds of topographic 3h points in the landscape.20 The pic has generated substantial dis- ssion and Shields presents a com- ehensive literature review.21 Blazing Star. Bernard de Vries. Many explanations or theories have *en advanced but it is overly riplistic to suggest a single best ex- sources abundant on hilltops rather anation for the phenomenon. In the than the hilltop situation. ore narrow, behavioral definition of Alberta Arctic behavior fits the Itopping17 it seems most widely mate-locating definition of hilltopping cepted that hilltops are meeting quite nicely. Males perch on the ridge eas for the sexes to mate. In the top, periodically fluttering up several oader definition there are most like- feet, apparently surveying the slope a variety of factors and suitable ex- below them for females, and dashing anations to suit the variety of downslope to investigate butterflies at ecies, ecological circumstances, lower elevations. This behavior has d geographic locations involved. been previously described in Oeneis Twelve of the species found in the uhleri uhleri in Colorado.15 esent study area were very ticeably restricted to or were more The aggressive, and more or less continuous hilltop flights of the undant on local topographic highs, Zelicaon Swallowtail, Badlands Old two cases, the Brown Elfin and World Swallowtail, Western :mon Blue, the restricted dis- Checkered White, Coral Hairstreak, aution of the larval food plants is Painted Lady, and Callippe Fritillary jarly the causative factor. Such is are also good examples of mate- t the case for the Melissa Blue or locating hilltopping behavior. e grass feeders (Assiniboia ipper, Nevada Skipper, Alberta The study area has a variety of ctic). Adults of the Assiniboia butterfly habitats due to differing ipper are strongly attracted to slope exposures, groundwater con¬ wers of Blazing Star (Liatris punc- ditions, and grazing history. The la) while adults of the Nevada grassland and shrubland habitats are ipper were seen to show a similar much more productive for butterflies traction to flowers of Prairie than the aspen forest. Naturalists oundsel (Senecio canus) and a visiting areas like this will find the lall yellow mustard. Thus, these in- shrubby edges of aspen forest and cts may be attracted to nectar grassy hilltops to be superior places irch, 1981. 39(1) 11 to observe a variety of butterflies and 12HOOPER, R. R. 1973. The butterflies c interesting behavior. Saskatchewan. Saskatchewai Museum of Natural History. 216 pp. 13KONDLA, N. G. 1979. Skippers anc butterflies of a prairie farm. Alberti Naturalist 9:71-75. ^IRD, C. D. 1975. A calendar of the butterflies and skippers of the alpine 14KONDLA, N. G. and C. D. BIRD. 1979 area of Plateau Mountain. Alberta The skippers and butterflies o Naturalist 5:26-28. Kananaskis Provincial Park, Alberta Blue Jay 37:73-85. 2BIRD, C. D. 1975. A revised calendar of the butterflies and skippers of "MASTERS, J. H. and J. T. SORENSON Calgary. Calgary Field Naturalist 1969. Field observations on fores 6:312-314. Oeneis (Satyridae). Journal of th< Lepidopterists’ Society 23:155-161. 3BIRD, C. D. 1979. The Question Mark — another “eastern” butterfly new to "McDUNNOUGH, J. H. 1939. A new race Alberta. Alberta Naturalist 9:66. of Papilio belonging to the machaof complex. Canadian Entomologis 4BOWMAN, K. 1944. Additions and correc¬ 71:216-217. tions to checklist of the macro- of Alberta. Canadian En¬ "SCOTT, J. A. 1968. Hilltopping as a tomologist 76:191-192. mating mechanism to aid the survival “CASE, J. W. and C. D. BIRD. 1977. of low density species. Journal ol Butterflies and skippers of west- Research on the Lepidoptera 7:191- central Alberta. Blue Jay 35:208-219. 204.

“CLENCH, H. K. 1972. The boundary 18SCOTT, J. A. 1974. Mate locating between Satyrium liparops and its behavior of butterflies. America^ subspecies strigosum (Lepidoptera: Midland Naturalist 91:103-117. Lycaenidae). Annals of the Carnegie Museum 44:11-24. 10SCOTT, J. A. 1975. Mate locating behavior of western North American 7dos PASSOS, C. F. 1964. A Synonymic list butterflies. Journal of Research on of the Nearctic Rhopalocera. the Lepidoptera 14:1-40. Lepidopterists’ Society Memoir 1.145 20SHEPARD, J. H. 1966. A study of the PP- hilltopping behavior of Pieris oc\ “dos PASSOS, C. F. 1968. A name for cidentalis Reakirt (Lepidoptera; Polygoni a satyr us marsyas Pieridae). Pan-Pacific Entomologist (Lepidoptera: Nymphalidea). Trans¬ 42:287-291. actions of the American En¬ tomological Society 95:153-159. 21SHIELDS, O. 1967. Hilltopping: An ecological study of summit con¬ “EHRLICH, P. R. 1955. The distribution and gregation behavior of butterflies on a subspeciation of Erehia epipsodea southern California hill. Journal of1 Butler (Lepidoptera: Satyridae). Research on the Lepidoptera 6:69- University of Kansas Science Bulletin 178. 37:175-194. 22STALKER, A. M. 1973. Surficial Geology 10FERRIS, C. D. 1973. A revision of the of the Drumheller Area, Alberta. Colias alexandra complex (Pieridae) Geological Survey of Canada Memoir aided by ultraviolet reflectance 370. 122 pp. photography with designation of a new subspecies. Journal of the 23TAVERNER, P. A. 1919. The birds of the Lepidopterists’ Society 27:57-73. Red Deer River, Alberta. Auk 36:1-21, 248-265. "FISHER, M. S. 1977. The taxonomy and identity of Papillo nitra W. H. Edwards 24THORMIN, T. W. 1977. The butterflies of in Colorado (Papilionidae). Bulletin of Beaverhill Lake. Edmonton Naturalist the Allyn Museum 47:1-8. 5:160-163. J

12 Blue Jay