ALBERTA SWALLOWTAILS AND PARNASSIANS: NATURAL HISTORY, KEYS, AND DISTRIBUTION

FELIX A.H. SPERLING, Section of Ecology and Systematics, Corson Hall, Cornell University, Ithaca, New York, 14853-2701, and NORBERT G. KONDLA, 6415-95 Avenue, Edmonton, Alberta. T6B 1A6

In this article we provide pictorial 1. Phoebus Parnassian - Parnassius keys, distribution maps, and natural his¬ phoebus (F.) tory information for Swallowtails and This species is characterized by: 1) Parnassians in Alberta. Except for intro¬ narrow white rings around each segment ductory sections, only information that of the antennae; 2) black wing markings, pertains to Alberta or is new is reported. particularly on the dorsal forewing disk; More general treatments for s^>ecies can and 3) a small, dark brown spragus on be found in existing literature.7 Common the end of the abdomens of females. names of host plants follow Budd's Both sexes tend to be smaller and darker Flora.6 In collaboration with C.D. Bird, at higher latitudes and elevations. All G.J. Hilchie, and E.M. Pike, we are populations in Alberta belong to P. preparing a book on the of Al¬ phoebus s mint he us Dldy. berta, but it will be a few more years before this project is complete. By sum¬ At Frank, Alberta, we have found lar¬ marizing what we know now, we hope vae on Lance-leaved Stonecrop (S. lan- to encourage further work on these but¬ ceolatum). Larvae are black, with dense terflies, particularly on the natural his¬ short hairs and small yellow spots on tory of more poorly known species. Fig¬ each segment. There is one generation of ure 1 shows Papilionidae for Alberta. adults. They have been reported from 2 June to 10 October. The main flight may Parnassians begin as early as mid-June in the Distinguished by hairy bodies and southern Alberta Rockies, but is not until wings with a white ground colour and August in the north. Males patrol slowly semi-transparent border, Parnassians are over open areas where the food plant also easily separated from Whites by grows, flying a metre above the ground, their moderately large size (50-65 mm searching for females. Adults with wingspan). Females are noticeably darker wing bases appear to be able to darker than males and have a spragus, a warm up better in the sun. Adults nectar waxy structure attached by males to the on a variety of Composites. end of the abdomen of females to prevent them from mating again with The Phoebus Parnassian is found other males. Parnassians are found in throughout the Rockies of Alberta in mountainous temperate regions and are subalpine meadows and dry, south¬ most diverse in Asia. Isolated popula¬ facing mountain slopes. The species is tions within the same species often show common in valley bottoms in the minor differences. southern mountains. There is an outlying population on the valley sides of the

49(4). December 1991 183 (D=dorsal; V=ventral; FW= lorewing; HW=hindwing) la. Tailed hindwing. 3. lb. No tails.Parnassius 2. Papilio Parnassius 2a. White-ringed antenna; black spot in DFW disk...P. phoebus 2b. All-black antenna; grey bar across DFW disk.P. clodius

3a. Striped center of FIW.4. 3b. Black gr yellow FIW center..6. P. phoebus P. clodius 4a. 1-tailed FIW.5. 4b. 2-tailed HW..P. multicaudatus Tb I 4a 4b I 5a. Ground color yellow or cream; stripes narrow...P. canadensis 5b. Ground color white; stripes wide.P. eurymedon

6a. FIW eyespot round and centered.P. zelicaon 6b. FIW eyespot connected to margin or low and oval...7.

7a. VFW disk mainly yellow and yellow hair under thorax...8 7b. VFW disk mainly black or > P. canadensis P. eurymedon black hair under thorax....9.

8a. On grassland or dry riverbanks..P. m. pikei 8b. Boreal/montane forest .P. m. hudsonianus

9a. On grassland or dry riverbanks..P. m. dodi 9b. Forested habitat. P. zelicaon X macbaon

Figure 1. Alberta Papilionidae.

184 Blue Jay Milk River north of Del Bonita. This eggs are spherical and green or yel¬ may be the only strictly prairie-dwelling lowish. Pupae are held upright against population in North America. It has been twigs by a silk girdle, resembling leaves found on the Saskatchewan side of the or broken branches. Pupae are either Cypress Hills, but not yet on the Alberta green or brown, depending on the back¬ side, though Stonecrop grows there.5 It is ground, and are the overwintering stage. less frequently collected in the northern They are often parasitized by large black Rockies, where it occurs on alpine and Ichneumon wasps ( spp.). upper subalpine rocky slopes. Old World Swallowtail Species Group 2. Clodius Parnassian - Parnassius Members of this species complex, the clodius Menetries group, are found This species is unlikely to be en¬ throughout most of the northern hemi¬ countered in Alberta except in Waterton sphere. There are about six species in the National Park. The species is distin¬ group. Late instar larvae have black and guished from the Phoebus Parnassian by green bands on each segment, inter¬ its: 1) completely black antennae; 2) spersed by small yellow or orange spots. grey rather than black markings on the The ratio of yellow to orange spots wing surfaces, especially in the dorsal varies geographically and can help to forewing disk; and 3) a large, white distinguish populations. Larvae feed on spragus on the female. There are only numerous species of Umbellifers, as well two confirmed records for P. clodius in as a few Composites, both of which are Alberta, from 1 and 12 August. They unusual food plant families for ¬ were collected by D. G. Wales at Bertha tails. Lake (5800’) and Goat Lake (6500’) in Waterton National Park. Early stages are The most common adult form in Al¬ unknown in Alberta. The Clodius Par¬ berta is predominantly yellow with black nassian is generally found in lush forest markings along the veins and outer bor¬ openings. ders. A dark morph, present in varying proportions, is characterized by a Swallowtails predominantly black pattern on the wings and yellow spots rather than a yel¬ Swallowtail butterflies of the low band on the abdomen. Males of the Papilio L. are found worldwide. There Old World Swallowtail group usually are more than 200 species in 42 species patrol grassy or sparsely treed hill tops groups; two of these groups have species with good nectar sources and a in Alberta. Research on these large and panoramic view. Encounters by patroll¬ beautiful butterflies has provided a great ing males lead to a short chase, while deal of support for conservation and in¬ females are pursued until they land in a sight into the process of evolution.1 In grassy spot and mate. Although species Alberta, Swallowtails are distinguished generally maintain their integrity when from other butterflies by their large size they meet, Swallowtails of the Old (wingspan more than 75 mm), hindwing World group sometimes form inter¬ tails, and a bold black and (usually) yel¬ specific hybrid populations.8'9 low colour pattern. Like Parnassians, the larvae have osmeteria, which are fleshy, 3. Old World and Artemisia Swallow¬ smelly tubes that larvae evert from just tails - Papilio machaon L. behind their head to discourage predators. Early instar larvae are black The Old World and Artemisia Swal¬ with a white patch in the middle and lowtails are best distinguished by the have spines with fleshy bases; they look shape of the hindwing eyespot on upper like unappetizing bird droppings. The side of the inner margin of the hindwing. 49(4). December 1991 185 Papilio machaort dodi, Drumheller, Alberta. Felix Sperling

3rd instar of Papilio zelicaon on 5 th instar of Papilio zelicaon on Angelica lanatum, Buck. leaf. Mountain, Alberta. Felix Sperling Felix Sperling

186 Blue Jay This eyespot is club-shaped and con¬ given year will emerge during the fol¬ nected to the margin, or at least oval and lowing year. A small proportion will low in the area of red scales. Three sub¬ wait an additional year or more to species occur in Alberta and, to reduce emerge, avoiding the kind of drought confusion, we refer to them by their that scorches host plants such as the one Latin names. P. m. dodi McDunnough is that occurred in southern Alberta in found in southern Alberta and was 1985. The larval food plant of P. m. hud- named after F. H. Wolly Dod, an early sonianus has still not been established. lepidopterist in Alberta. P. m. pikei Judging from a photo of a late instar Sperling was recently discovered in the larva taken by Gary Anweiler in Sas¬ Peace River region and named after katchewan, the food plant may be Pal- another Albertan, E.M. (Ted) Pike. P. m. mate-leaved Coltsfoot (Petasites pal- hudsordanus Clarke occurs, as its name matus). It is possible that larvae of P. m. suggests, in the Hudsonian region of hudsonianus also feed on Umbellifers. Canada. As Hooper later reported. Swal¬ lowtails originally identified as P. bair- P. m. dodi has been collected from 3 dii brucei Edwards in The Butterflies of May to 23 August. The peak for the first Saskatchewan actually are the summer generation is the second half of May and generation of P. m. dodi,4,5 Butterflies the first half of June, while for the that Hooper referred to as Badlands Old second generation it is late July and early World Swallowtails (“F\ machaon ssp.”) August. P. m. pikei has been recorded are the spring generation of P. m. dodi.A from 9 June to 9 July. P. m. hudsonianus Butterflies that Hooper referred to as has been collected in Alberta from 31 Cypress Hills Old World Swallowtails May to 28 July, but most were collected (“P. m. dodi") appear to be a hybrid in the second half of June. Males of P. population between P. zelicaon (the m. dodi and P. m. pikei tend to fly along Zelicaon Swallowtail) and one or more the upper edge of deep river valleys and subspecies of P. machaonA accumulate at prominences. Males also patrol slopes well below valley edges, Spring generation adults of P. m. dodi looking for females near larval food look much like P. zelicaon. Summer plant patches. Nectar sources include generation adults generally have more Groundsel (Senecio spp.). Sweet Broom yellow scales and hairs. Dark morph (.Hedysarum spp.) and Alfalfa adults comprise less than 2% of any {Medicago sativa). P. m. pikei and P. m. population in Alberta. P. m. dodi also dodi occur in dry grassland habitats tends to have more pointed forewings along eroding river banks, badlands, and and narrower hindwing tails than P. road cuts near the larval food plant. zelicaon. Both P. m. pikei and P. m. hud- Populations are patchily distributed sonianus are distinguished by yellow along major river valleys and badlands. hair extending around the underside of In contrast, P. m. hudsonianus is found the thorax and by yellow scales at the in the boreal forest region, especially base of the underside of the fore wing. openings along roads, bogs or sparsely The latter two subspecies are separated treed hill tops. It is infrequently collected by habitat and locality. in Alberta, but is distributed over a broad region. Larvae of P. m. dodi and P. m. pikei feed exclusively on Linear Leaved 4. Papilio zelicaon X machaon Wormwood {Artemisia dracunculus), as Hybrid individuals are difficult to do other subspecies of P. machaon recognize. By definition they are nor¬ found in the western United States. Not mally rare and, if back-crosses are pos¬ all pupae of P. m. dodi or P. m. pikei that sible, hybrids can exhibit the full range are produced in the late summer of a 49(4). December 1991 187 of character combinations between the easy to see how hybrids could occur. It parental species. P. zelicaon and P. is more difficult to understand why the machaon form hybrid populations in species don’t merge wherever they con¬ west-central Alberta and the Cypress tact each other. The frequency of hybrids Hills; Hooper refers to the Cypress Hills is higher in areas where contacting Old World Swallowtail hybrids.4,8 populations of the parental species have Hybrid adults look much like P. m. dodi, more similar habitat preferences. Thus and are best distinguished by molecular the relative viability of hybrid in¬ characters and by habitat and locality. dividuals, rather than different mating Most individuals from hybrid popula¬ behaviour, may determine whether tions have a club-shaped eyespot, but hybrid populations will form. However, have a rounded wing shape much more actual observations of natural matings like that of P. zelicaon. The parental are rare and the two species might also species seem to hybridize much less fre¬ have mating barriers in dry land regions, quently in the southern Alberta prairies in addition to staying out of each others’ and in the Peace River region, as well as way as an indirect consequence of dif¬ where their ranges overlap in the rest of ferent habitat preferences. western North America. Most hybrid populations occur in mixed forest, be¬ 5. Zelicaon Swallowtail - Papilio tween 1000 and 2000 m elevation. zelicaon Lucas

Adults of P. zelicaon are best identified Larvae of hybrid populations are by the round and centred anal eyespot on found on Heart-leaved Alexanders (Zizia the upper side of the hind-wing. They aptera) in the area west of Calgary and usually have more black coloration than P. on Cow Parsnip (Heracleum lanatum) in machaon, especially at the base of the the moister areas north and west of Red ventral forewing and on the thorax and ab¬ Deer. At Bragg Creek, immediately west domen. The black adult “nitra” morph may of Calgary, most larvae have yellow comprise up to 10% of some populations spots which distinguish them from the in southern Alberta. orange spotted larvae of P. m. dodi at Calgary and eastward. P. zelicaon X Larvae feed entirely on Umbellifers machaon hybrid Swallowtails have been and will not feed on Narrow-leaved collected in the Alberta foothills from 22 Wormwood. Eggs are laid on flower May to 2 August, with the main flight heads or leaves. In the Waterton area, period in the first half of June. Some¬ larvae tend to occur on Lyall’s Angelica times typical P. zelicaon adults are col¬ (Angelica arguta), while in northern Al¬ lected at the same localities later in the berta they tend to occur on Cow Parsnip. summer; these may have dispersed from Larvae have also been found in Alberta pure P. zelicaon populations in moun¬ on Mountain Parsnip (Angelica daw- tains to the west. Hybrid Swallowtails in soni), Kneeling Angelica (Angelica the Cypress Hills have mostly been col¬ genuflexa). Mountain Wild Parsnip lected in late June and early July. Where (Lomatium dissectum), Western Wild hybrids are rare, most specimens are col¬ Parsley (Lomatium triternatum). Water lected together with apparently pure P. Parsnip (,Sium suave), and Heart-leaved zelicaon or P. machaon. The latest Alexanders, as well as two cultivated records for any Swallowtail in the Umbellifers, Garden Celery (Apium southern Alberta prairies (28 September) graveolens) and Garden Parsnip {Pas- and the Peace River region (22 July) tinaca sativa). The main larval food were for hybrids. plant on the prairies of southern Alberta is unknown, though it seems likely to be The mating behaviour of P. zelicaon a Prairie Parsley {Lomatium spp.). and P. machaon is very similar, and it is 188 Blue Jay There is almost always a single 6. Canadian Tiger Swallowtail - generation per year for P. zelicaon. The Papilio canadensis Rothschild and Jor¬ main flight period is variable, with adults dan generally emerging later in cooler Previously considered a subspecies of habitats, especially in the mountains. the Eastern Tiger Swallowtail (Papilio Adults have been collected in Alberta glaucus), the Canadian Tiger Swallow¬ from 23 April to 19 August, though the tail has recently been given species main flight period varies from the status on the basis of a number of second half of May in the Wintering genetic differences and the relatively Hills south of Drumheller, to the first narrow hybrid zone it forms with P. half of July, in Waterton National Park. glaucus. It is the only member of the A few fresh adults have been collected species group that occurs in central and near Drumheller in early August. A com¬ northern Alberta. Tire ground colour of mon nectar source for hill-topping male males is always yellow when they are P. zelicaon is dandelion. P. zelicaon is fresh, but females may be either a lighter generally found in moist meadows in cream-yellow or a more orangy-yellow forested areas, but in Alberta it can also than the males. The wingspan is 8.5 to be found in habitats ranging from 10 cm in Alberta, and there is only one parkland and prairie hill tops to subal- tail per hindwing. The Western Tiger pine meadows. Adults occasionally hill¬ Swallowtail (Papilio rut ulus) sometimes top even on alpine peaks. Whenever hybridizes with the Canadian Tiger males of both colour morphs occupy the Swallowtail in southern British Colum¬ same hill top, it seems that the black bia. Apparently the only difference in ones occupy the choice spots on the very adult colour pattern is that Western Tiger peak of the hill. Very little data has been Swallowtail has no orange marginal collected on the mating behaviour of spots on the underside of the hind wing.7 these species, however, and this very in¬ teresting situation awaits someone with a Larvae have been collected on Aspen little time and patience to study it. Poplar (.Populus tremuloides), Willow (Salix sp.) and Domestic Crabapple The Tiger Swallowtail Species Group (Malus sp.) in Alberta. Eggs are laid on This species complex, the P. glaucus the middle of the upper side of small group, is the only species group of Swal¬ leaves, about 1 mm from the central lowtails that is restricted to North vein. The egg is bright green when laid, America and adjacent Central America. and has a deep yellow ring of cement at Like the P. machaon group, the P. the base, which dries clear in about 5 glaucus group is most diverse in western minutes. Larvae spin a silk mat on the North America. Six species are currently leaf on which to rest when they are not recognized, and most of these hybridize feeding. occasionally where their ranges meet or overlap. Late-instar larvae have a dis¬ Adults have been collected from 12 tinctive pattern, with large eyespots on May to 12 August, with the main flight the thoracic segments, followed by a in June. Adults are common along trails single band and a smooth green back¬ through mixed forest, or along the edges ground. They feed on the foliage of a of aspen parkland. Males tend to patrol wide variety of trees and shrubs. Pupae along streams, forest edges and trails, are similar to those of the P. machaon but also accumulate on hill tops or con¬ group, but are slightly narrower and less gregate on wet soil. Females fly in the bent. Adults have a distinctive pattern of same habitats, but have a faster and more black stripes on a yellow or white back¬ circuitous flight than males, as they ground - hence their common name. search for leaves on which to lay their

49(4). December 1991 189 eggs. Both sexes nectar at a variety of from British Columbia. In Alberta it has flowers, especially dandelions. The been collected from 23 June to 1 August. Canadian Tiger Swallowtail can be It is found in dry, montane forest and the found commonly throughout most of Al¬ species is our rarest Swallowtail in Al¬ berta except the southern grasslands, berta. where it is uncommon and local. Acknowledgements 7. Two-Tailed Swallowtail - Papilio We thank C.D. Bird, G.J. Hilchie, E.M. Pike, multicaudatus Kirby H. Pinel and many other collectors of Alberta This species looks much like the butterflies for natural history and Canadian Tiger Swallowtail, but has a distributional information. second, smaller tail on each hind wing. I. COLLINS, N.M. and M.G. MORRIS. Its wing stripes are also a bit narrower 1985. Threatened Swallowtail Butterflies and its wingspan is usually more than 10 of the world. International Union for Con¬ cm. Its background colour is the same servation of Nature and Natural Resour¬ ces, Gland, Switzerland. yellow shade as in male Canadian Tiger Swallowtails. There are no larval records 2. GUPPY, C.S. 1986. Geographic variation yet for Alberta. in wing melanism of the Parnas- sius phoebus F. (: Papilionidae). Can. J. Zool. 64: 956-962. Adults have been collected from 7 June to 2 September, and most are from 3. HAGEN, R.H., R.C. LEDERHOUSE, J.L. BOSSART and J.M. SCRIBER. 1991. late June. The Two-tailed Swallowtail is Papilio canadensis and P. glaucus are dis¬ found only on the U.S.-Alberta border, tinct species. J. Lepidop. Soc. 45: in press. at Waterton National Park, along the Milk River (especially at Police Coulee) 4. HOOPER, R.R. 1973. Butterflies of Sas¬ katchewan. Sask. Museum of Natural His¬ and rarely north to Lethbridge. It flies tory, Regina. along coulees and the edges of riverine forest. The large wingspan of this but¬ 5. -. 1986. Revised checklist of Sas¬ katchewan butterflies. Blue Jay 44: 154- terfly makes it a particularly fast flier 163. and well suited to soaring. It commonly nectars on thistles (Cirsium spp.). 6. LOOMAN, J. and K.F. BEST. 1979. Budd’s Flora. Research Branch, Ag. Can. Publ. 1662. 8. Pale Swallowtail - Papilio eurymedon Lucas 7. SCOTT, J.A. 1986. The butterflies of North America. Stanford Univ. Press, This species has the black wing stripes Stanford. 583 pp. shared by other species in the Tiger 8. SPERLING, F.A.H. 1987. Evolution of Swallowtail group, but its background the Papilio machaon species group in colour is pale white, without any yellow. western Canada. Quaestiones En- The wing bands are wider than the other tomologicae 23: 198-315. species and are usually substantially 9. -. 1990. Natural hybrids of Papilio (In- wider than the white areas between secta: Lepidoptera): poor or in¬ them. There is only one tail on each teresting evolutionary phenomenon? Can. J. Zool. 68: 1790-1799. hindwing, and the wingspan is 9 to 10 cm in Alberta.

Immatures of the Pale Swallowtail have not been recorded yet in Alberta. The species is found in the southwestern comer of Alberta, north to the Crowsnest Pass and east to Police Outpost. Many Waterton records may represent strays 190 Blue Jay 49(4). December 1991