Family Nymphalidae

Brush-footed Butterflies

This is the largest family of butterflies, with about 5,000 species worldwide. Traditionally the family Nymphalidae as now defined has been treated as a number of families, including the satyrs (Satyridae), monarchs (Danaidae), heliconians (Heliconiidae), and snouts (Libytheidae). More recent concepts of phylogenetic analysis and classification have resulted in these groups being treated as subfamilies of the Nymphalidae. By this revised definition, the Nymphalidae contains about 220 species in North America and 102 in Canada.

The forelegs of nymphalids are reduced in size, usually in both sexes (not in Libytheinae females), and covered with long hairs, so as to resemble a brush. They are so small as to be practically useless; nymphalids perch and walk using four legs only, with the front pair held up under the "face".

Unlike the previous families, nymphalids vary greatly in appearance in larval, pupal, and adult stages. Adults range from small to large, with most being of medium size. In North America, the most common colour is orange brown but there are many exceptions to this. Some species are powerful fliers (Polygonia) or migrants (Vanessa), while others (Euphydryas) are very weak fliers living in small, highly localized colonies.

Subfamily Libytheinae (Snouts)

This tiny subfamily contains only 12 species, with one found in North America. It is immediately recognizable by the extremely long palpi on the front of the head, the "snout". The forelegs of the male are reduced in size, but those of the female are not. The larva has a hump on the thorax but lacks the spines of most nymphalids, and the pupa is relatively unornamented.

Subfamily Heliconiinae (Heliconians)

This is a small subfamily, found mostly in tropical America; seven reach the southern U.S. and one migrates into the north and has twice been recorded in Canada. All species have long narrow wings, most (but not ours) are vividly coloured to advertise that they are unpalatable. Heliconian larvae have rows of branching spines, but lack a middorsal row, and there are more spines on the head. The pupae are irregular in shape with protruding wing-cases. All heliconians feed as larvae on various species of passion flowers that contain poisonous compounds.

The subfamily Heliconiinae is closely related to the next subfamily, the Argynninae, and the two have been combined in the Heliconiinae by some recent authors (e.g., Opler and Malikul, 1992).

Subfamily Argynninae (Fritillaries)

This mainly holarctic subfamily contains about 35 species in North America and 26 in Canada. They are divided into two groups, the greater fritillaries (Euptoieta and Speyeria) and the lesser fritillaries (Boloria). Both groups are orange brown above with mottled black patterning. The greater fritillaries are similar on the forewing underside, but on the hindwing underside they have a complex pattern of pale, usually metallic silver spots. The lesser fritillaries have complex underside patterns, but usually lack silver spots (except Boloria selene).

Eggs are laid singly, often not on the foodplant but in the leaf litter near the foodplant. Larvae have rows of branched spines, but lack both a mid-dorsal row of spines and head spines. All feed strictly at night and hide during the day away from the foodplant. Pupae are somewhat irregular in shape and have small dorsal bumps or cones.

The greater fritillaries offer a tremendous challenge to amateur and professional lepidopterists alike. Although experts are mostly in agreement on what constitutes a species, there is a confusing array of highly variable subspecies and forms. In some instances in western Canada, there is a greater difference among subspecies within a species than there is between species. There is also intergradation between forms in some areas, but no intergradation between the same forms in other areas. Species must be studied across their total geographical range. The authors have largely followed James Scott's classification (Scott, 1986). The ten Canadian species of Speyeria are all medium-sized and orange (bright to very pale) with black markings on the upper surface. Most have bright silver markings on the hindwing underside.

Two other species, the Coronis Fritillary (Speyeria coronis (Behr)) and the Great Basin Fritillary (Speyeria egleis (Behr)), recorded from Canada mostly on the basis of historical records, appear now to have been based on mistaken identifications; until recently there have been no specimens to substantiate the presence of either species in Canada. Both species, however, have been recorded regularly in Montana and Washington close to the Canadian border and could possibly be found in southern Alberta or British Columbia. Recently coronis has been taken in extreme southern Alberta, but egleis is still eluding capture north of the border. However, the possibility is still good and we include the diagnosis of egleis (under Speyeria zerene).

East of Manitoba, there are only three regularly occurring greater fritillary species and they can be determined reliably. It is in the west that great care must be taken and experts consulted for final determination. Habitat is important for determining species and subspecies, particularly between prairie, parkland, and boreal forms. The amount of solar radiation on larvae and pupae is believed to play a role in colour variation.

The larvae of all greater fritillaries feed on violets (Viola spp.). The first-instar larvae go into hibernation, without feeding, immediately after hatching from the egg. Adults usually appear in mid-summer.

The lesser fritillaries look like smaller versions of the greater fritillaries on the upper surface. Few have silver spots on the lower side. They are mostly boreal, tundra and alpine species in northern Canada and most are holarctic, being also found in Asia and Europe. Many have extensive black scaling to absorb as much solar radiation as possible in their cool habitat. They also fly low to the ground to prevent heat loss from the often constant winds in their arctic and alpine habitats.

The larvae of some species feed on violets, but most feed on a wide variety of other plants. Most overwinter in the fourth instar. Some northern species take two years to develop, overwintering in the first or second instar and again in the fourth instar.

Subfamily Melitaeinae (Checkerspots and Crescents)

This subfamily is restricted to the Northern Hemisphere and the American tropics. There are about 50 species in North America, of which 17 occur in Canada. They are mostly small to medium-sized orange-brown butterflies with black or dark brown markings; the best distinguishing marks are usually on the underside of the hindwings. Many are weak-flying butterflies that live in localized colonies and never stray far from the foodplant.

Most lay their eggs in large clusters of up to several hundred on the underside of the leaves of the foodplant, almost always herbaceous plants, usually in the families Asteraceae (Compositae) and Scrophulariaceae. Because of this habit, many checkerspots occur in concentrated colonies and this can lead to considerable local and geographical variation. The larvae have six rows of branching spines along the body but none on the head. They live together, often in a silken web, and hibernate as partly grown larvae; many species live singly after hibernation.

Because they are so similar, there has been a great deal of speculation concerning species status, with major revisions of genera, some quite recently. Distribution of species, particularly distributions based on old records unsubstantiated by specimens, has also been confusing, especially in the west where there have been major problems with identification and classification in all three genera for almost a century.

Subfamily Nymphalinae (Anglewings, Tortoiseshells, Thistle Butterflies, and Peacocks)

This worldwide subfamily of medium to large butterflies contains 26 species in North America, of which 16 occur in Canada. Larvae all have rows of branching spines, and pupae are irregular in shape, often ornamented with metallic silvery or golden spots. There are many distinct groups within this subfamily, of which four groups occur in Canada. The tortoiseshells (Nymphalis) lay their eggs in large groups and the larvae feed communally until about half-grown; the anglewings (Polygonia), thistle butterflies (Vanessa), and peacocks (Junonia) lay eggs singly or in very small clusters. The tortoiseshells and anglewings use both trees and herbs as foodplants, the thistle butterflies and peacocks use only herbs; species from the first three groups feed on nettles (Urticaceae).

Most anglewings and tortoiseshells are long-lived butterflies that hibernate as adults, becoming sexually mature only in their second year. The thistle butterflies are migrants, with none regularly overwintering in Canada; the Painted Lady (Vanessa cardui) is the most widespread butterfly in the world, having spread to all continents except Antarctica and South America.

The anglewing group (commas and the Question Mark) is made up of seven closely related species in Canada that cause many identification problems. Three species have two generations per year, in which the adults of each generation look quite different; there is considerable individual variation as well. All hibernate as adults, and specimens seen in the early spring look very different than those that are freshly emerged. There are various details of wing shape and the form of the silver "comma" mark on the hindwing underside that differ between the species, but even using these, there can still be difficulties.

The tortoiseshells are a small group with four North American species, all of which occur in Canada; some of these are also found in northern Eurasia. Like the anglewings, they all have cryptic undersides and irregular wing margins for camouflage. However, the upper sides have unique patterns and the species are easily distinguished. They feed as adults mainly on tree sap and animal droppings although one, the Milbert's Tortoiseshell, also feeds regularly on flowers.

The thistle butterflies get their name from the larval foodplant of the most widespread species, the Painted Lady. The Painted Lady is the most cosmopolitan of butterflies, rarely surviving the northern winters (only a few possible records in Canada), but populations are replenished in most years by specimens flying north into Canada from the south. The other three species overwinter in the U.S. and perhaps southernmost Canada, but expand their ranges farther into Canada during the summer.

The peacocks are a small group of mainly tropical species of which only one, the Common Buckeye, regularly migrates to Canada but cannot overwinter.

Genus Polygonia Hübner, 1819 (Anglewings)

The species in the genus Polygonia are collectively called anglewings; although the individual species are called either the Question Mark or various kinds of comma. There are seven species of anglewings in Canada and they can be a challenge to identify both as adults and larvae. As a group, adults can be recognized by the peculiar angled and lobed margins of the wings and the presence of a silver comma-like mark on the underside of the hindwing. Most species occur in two colour forms above and two below; these forms can be associated with season (spring versus summer forms) and with sex (females of the first four species are more unicoloured below than males).

The larvae, like those of many nymphalids, have series of branching spines along their back and sides. A feature that is exclusive to the genus Polygonia (and Nymphalis vaualbum) is that the branching spines include a pair on the top of the head. In Polygonia species, unlike Nymphalis vaualbum, the side branches on the spines along the back tend to cluster into one or two spots along the main spine, so they appear as whorls of side spines (like the branches of a pine tree) rather than scattered along the main spine. The larvae of Polygonia are extremely variable and the body can be pale with dark speckling and dashes to very dark with pale flecking; the spines may be mostly white, yellow, red, or black, or various combinations thereof. The dorsal spines tend to be white or yellow in most species, but are usually red or orange in Polygonia interrogationis. The larvae of Polygonia faunus, satyrus, gracilis, and oreas tend to be much paler on the back than the sides, whereas those of interrogationis, comma, and progne tend to be similarly patterned on the back and sides.

Subfamily Limenitidinae (Admirals)

This is a worldwide subfamily of medium-to-large butterflies, with 26 species in North America but only four are found in Canada.

The eggs of our species are laid singly on trees. The larvae resemble bird droppings when small and have two barbed spines on the top of the swollen thorax when fully grown. They feed on the leaves of a variety of trees, particularly willows (Salix spp.) and aspens (Populus spp.). They overwinter as partly grown larvae in a shelter made by rolling up a small part of a leaf tip.

The pupae are irregularly shaped. All of our species hybridize to some extent because they are so closely related. Some of these hybrid forms have been given varietal names.

Most admiral species are dark with a distinctive white band, but several, such as the Viceroy (Limenitis archippus) and the Red-spotted Purple (Limenitis arthemis astyanax) mimic other butterflies that are distasteful to birds.

All our Canadian species tend to be associated mostly with woodlands and favour rotting fruit or animal dung as the adult food source. Most are often seen sipping moisture from wet spots on the ground.

Subfamily Apaturinae (Emperors)

This small subfamily occurs worldwide in temperate and tropical regions. The emperors are a small group of medium-sized New World species, with 12 species occurring in North America and two in Canada.

The eggs are laid singly or in small groups on the leaves of the foodplant. The larvae feed in small groups; they lack branching spines but have two large barbed horns on the head. They are longitudinally striped and the last segment branches into two "tails". Hibernation occurs as partly grown larvae curled up in leaves of the foodplant. The pupa is somewhat flattened, with two headhorns; in both our species they are green and lie flat on a leaf, unlike most other nymphalid pupae, which hang.

The adults are tawny in colour with black and white dots, the females larger and lighter in colour than the males. In males the wings are slightly pointed; in females they are more rounded.

Mainly forest-dwelling butterflies, they occur where hackberry trees grow, sometimes in urban centres, such as Toronto and Montreal, where there are ornamental plantings of hackberry. The adults have a distinctive flap-and-glide flight and often alight on the trunks of trees or high up on the leaves. As with the admirals, they get their nourishment from animal dung, rotting fruit, or the sap of trees.

Subfamily Satyrinae (Satyrs and Wood-Nymphs)

This large worldwide subfamily contains about 50 species in North America and 34 in Canada. The adults are mainly medium-sized butterflies, almost always drab orange, brown, or greyish brown in colour, usually with at least a few eye-spots. All species have the base of some of the forewing veins visibly swollen; this is thought to be a hearing organ. Most have a weak, bobbing, dodging flight but some, for example species in the genus Oeneis, are powerful fliers.

The larvae are green or brown, well camouflaged, smooth, and somewhat tapered towards both ends, with the last segment usually forked into two "tails". All feed on monocotyledons (Canadian species on grasses and sedges). Pupae are smooth and unornamented, usually with two horns on the head.

Genus Erebia Dalman, 1816 (Alpines)

Erebia is a large genus of small and medium-sized satyrs that occur throughout Europe, Asia, and North America. There are more than a hundred species, mostly in the Old World. There are 14 species in North America, of which 13 are found in Canada. The species in the genus are known as alpines, which accurately describes the habitat of many, but not all, species.

Alpines have broad rounded wings and are always dark brown or blackish, usually with lighter brown, orange, or reddish patches and eye-spots on both wings. In many species the females are paler than the males. For many species the larvae are undescribed; known larvae are dull-coloured, plain, or faintly striped. They have large heads and their bodies taper towards the rear, ending in two short "tails". All feed on grasses, sedges, or rushes and pupate in a cell on the ground or even under rocks, sometimes in a rudimentary cocoon. Pupae are short and stout, with a rounded head, and lack hooks on the cremaster. There is never more than one generation per year, and in some northern species only one every two years.

Adults have a weak flight, low over the ground, and most are rarely seen on flowers.

Genus Oeneis Hübner, 1819 (Arctics)

The genus Oeneis includes 11 species in Canada, another in California, and many more in the Old World. About half of the species in Canada are also found in Eurasia. All are medium-sized butterflies with pointed forewings, rounded hindwings, short antennae, and hairy bodies. Most species are dark in colour, usually dark grey or brown, and this permits them to absorb heat while basking in the sun in their cold habitats; they often have a few small eye-spots. The undersides are cryptically coloured; the butterflies close their wings on landing and seem to disappear, whether the background is bare rock, lichens, tree trunks, or grass.

They are commonly known as arctics, and most are found in arctic, subarctic, and alpine areas. Not all of the larvae have been described, but known ones are hairy, cylindrical in shape, and tapering towards the rear, with two very short tails. They have a dark dorsal stripe and up to six alternating dark and light lateral stripes. Pupae are short and stout; pupation takes place on the ground or under rocks, often in a silk-lined cell. As far as is known all species feed on grasses and sedges, and have a two-year life history, passing the first winter in the first or second larval stage and the second winter in the fourth or fifth larval stage. In some areas, adults of some species are seen only every second year; other species fly every year, although they may show a considerable alternation in abundance.

Arctics usually have a powerful flight and even some of the weaker fliers can be very difficult to catch in their windy arctic and alpine habitats; when alarmed they lift up and allow themselves to be swept away by the wind. They are not often seen feeding at flowers or at mud.

Subfamily Danainae (Milkweed Butterflies)

This worldwide subfamily is found mainly in the tropics; four occur in North America and one, the Monarch, migrates every year into Canada. The adults are large, brightly coloured butterflies, distasteful to predators. The forelegs are reduced in size as in other Nymphalids.

The larvae are brightly coloured. They have no spines, but most have one or more pairs of fleshy filaments on the body. They feed on poisonous plants in the milkweed family (Asclepiadaceae) and dogbane family (Apocynaceae). The pupae are rounded, with no projections, but are ornamented with bright dots of gold and red.

© 2002. This material is reproduced with permission from The Butterflies of Canada by Ross A. Layberry, Peter W. Hall, and J. Donald Lafontaine. University of Toronto Press; 1998. Specimen photos courtesy of John T. Fowler.