The family Nymphalidae is the most speciose family of butterflies with about 6000 described species so far. The family contains many well-known species, such as the monarch, the Painted Lady, the buckeye, the fritillaries, checkerspots and the electric blue morphos. Indeed, nymphalids are in many places the most visible members of the local butterfly fauna. Due to their visibility and ease of study in the field and lab, many species of nymphalids have been used as model systems to understand the complexity of life on this planet.
This is a large family of small- to large-sized butterflies. The front feet are atrophied and used as sensory organs, while the remaining feet are used for locomotion. Brush-Footed butterflies are important visitors of various wildflowers, although a few species prefer tree sap, fermenting fruit, or dung. There are several subfamilies; only the more important ones will be described. Libytheinae (Snout Butterfly): Only a single migratory species occurs in the area, Libytheana carinenta bachmanii (American Snout). The wing undersides are brown-patterned (resembling a dried-out leaf), while the uppersides have patches of orange and brown with white dots. Projecting from the head is an elongated labial palp, or "snout," giving this butterfly a distinctive appearance. The caterpillars feed on leaves of hackberry. Limenitidinae (Viceroy et al.): These are large butterflies that often mimic other species. The species Limenitis archippus (Viceroy), is orange with black stripes and white dots. It mimics the Monarch butterfly. The caterpillars feed on willow, wild cherry, and other plants. The species Limenitis arthemis astyanax (Red-Spotted Purple) is black with rows of blue and orange dots. It is a mimic of the Pipevine Swallowtail, but the hindwings have no tails. Further to the north occurs another variety of this butterfly, Limenitis arthemis arthemis (White Admiral). It is black with conspicuous bands of white, along with rows of blue or orange dots. The caterpillars of Limenitis arthemis feed on plants that are similar to those that are described for the Viceroy. Nymphalinae (Painted Ladies et al.): This is a large subfamily, including the Red Admiral, Painted Ladies, Buckeye, Mourning Cloak, Comma, Checkerspots, Baltimore, and others. Most of these butterflies have brown patterns on the wing undersides (looking like a dead leaf), and orange/black patterns on the oversides with white dots. They are medium- to large-sized butterflies. The species Nymphalis antiopa (Mourning Cloak), however, is black with a band of yellow and blue dots along the margins of the wings. The caterpillars of these butterflies feed on a wide variety of plants, such as nettles, elm, willow, hops, plantains, turtlehead, and members of the Aster family. Heliconiinae (Fritillaries): These butterflies are checkered orange and black with white dots. Their larvae are nocturnal and feed primarily on violets, although some species feed on passion vines in southern Illinois. Danainae (Monarch): In our area, this consists of a single species, Danaus plexippes (Monarch), which migrates northward from Mexico, not arriving in central or northern Illinois until mid-summer. It is a large orange butterfly with black stripes and white dots. The caterpillars feed on milkweeds. Satyrinae (Wood Nymphs, Pearly Eyes): These are brown woodland butterflies with black/white eyes. The wing undersides are a lighter shade of brown than the oversides. They occasionally stray from woodlands to nectar at wildflowers in moist meadows. The caterpillars feed on various woodland grasses and sedges.
All species of Nymphalidae are united by a single morphological character, the tricarinate ridges found on the adult butterfly's antennae. Most also exhibit extreme reduction in the size of the forelegs, particularly in males (this feature is also exhibited by Riodinidae).
This is a large and diverse family of butterflies. Over 4,000 species of Brushfoots are found all around the world. In Michigan we have 37 species.
Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native )
This family of butterflies gets its name from its front legs. They are shorter than the other four legs, and they don't use them to walk or stand. These front legs don't have feet, just little brushes of hairs that the butterflies can use to smell and taste with. Sometimes the front legs are so small you can't see them.
This is a very diverse group of species. Some are brightly colored, some well camouflaged. Most have brown camouflage patterns on the underside of the wings, and brighter colors (often orange) on top, but there are lots of exceptions. Some have rounded wings, some have irregular edges with notches and little curves. Several species in this group are mimics, they look like other species that are toxic, and so avoid predators.
Many of the caterpillars have horns or spines or bumps to discourage predators. Some are dark colored, some are green or yellow, many have stripes or spots.
(See More Information on Butterflies and Moths for a general physical description of butterflies.)
Other Physical Features: ectothermic ; bilateral symmetry
Sexual Dimorphism: sexes shaped differently
Species in this family are so variable that it is hard to generalize. Brushfoot butterflies can be found in almost any habitat that has plants.
Habitat Regions: temperate ; terrestrial
Terrestrial Biomes: taiga ; desert or dune ; chaparral ; forest ; rainforest ; scrub forest ; mountains
Wetlands: marsh ; swamp ; bog
Caterpillars of different species of brushfoots eat many different kinds of plants. Many specialize on just a few species or one family of plants. Some specialize on thistles or nettles, some on willow trees, some on plants in the daisy family, some on violets.
Adults sometimes sip nectar, but many species in this group seem to prefer tree sap or rotting fruit, and some feed on dung or mud.
Some of these species are important herbivores, reducing the populations of their food plants.
Caterpillars hide and feed at night, or have spines and horns which may release toxic chemicals. In some species young caterpillars live together in groups, and they all thrash around and give off toxic chemicals if predators attack.
Adults may have camouflage colors on their wings, and can fly away from danger. Some species are toxic, and have warning colors to tell predators to leave them alone, and others mimic these colors but don't have the toxins.
- Soricidae (eat pupae)
- Sigmodontinae (eat pupae)
- Anura (eat adults)
- Araneae, especially crab spiders and orb-weavers (eat adults)
- Formicidae (eat caterpillars)
- Hymenoptera (eat caterpillars and adults)
- mantids (eat adults)
- Diptera (eat caterpillars)
- Coccinellidae (eat eggs)
- Chrysopidae (eat eggs)
- Acari (eat eggs)
Life History and Behavior
Communication and Perception
These butterflies communicate mainly with their scent and their colors. Males attract mates with scent and display, and females leave a scent mark on plants where they have laid eggs.
Like all Moths and Butterflies, this family has complete metamorphosis. See More Information on Butterflies and Moths for an explanation of this. Pupae do not make cocoons in this family, they are chrysalids. Usually it is the larvae that hibernate in this group, but a few species survive the winter as adults.
Development - Life Cycle: metamorphosis
In most species of Brushfoots, individuals only live for a year or a little more (not more than two winters). A few cold climate species may survive through two winters as larvae or pupae, but the adults only survive for a few weeks. Some temperate climate adults live the longest of any adult butterflies, surviving for 6 months or more.
Mating System: polygynous
After mating, females lay up to several hundred eggs. Some species lay their eggs one at time, others lay clusters together (this relates to the behavior of the caterpillars after they hatch).
Breeding season: Late Spring, Summer, and/or early Fall.
Key Reproductive Features: iteroparous ; seasonal breeding ; sexual ; fertilization (Internal ); oviparous
Once they have laid their eggs, there is no parental care in these species.
Parental Investment: no parental involvement
Evolution and Systematics
Discussion of Phylogenetic Relationships
The pages under Nymphalidae on the Tree of Life Web site are being worked on at this moment. Relationships are shown for the most part down to the level of genera, but there is still very little other information on the pages. The pages will be continuously updated, so please check back often.
The diversity in form and life style has meant that the phylogenetic relationships of nymphalids have been contentious. This in turn has meant that there has been no consensus on the classification of the group, with some authors splitting the family into up to 9 different families! The lack of a good phylogenetic hypothesis has also meant that the evolutionary history of the group has been shrouded in mystery. Recent molecular and morphological work is bringing light to the question of how different species and groups of species are related to each other. The tree shown above is the best hypothesis of subfamilial relationships based on as yet unpublished combined analyses of morphological and molecular data.
Many groups within Nymphalidae are currently under investigation, and we have endeavored to provide current hypotheses of relationships for each group. Where these are lacking, lists of taxa down to the species level are provided, in the hope that this will stimulate further research.
The digestive system of Monarch butterflies protects them from poisonous milkweed latex eaten to make themselves poisonous to predators.
"Milkweed gets its name from a poisonous latex that exudes from its broken stem. This is so toxic that it can give a small animal a heart attack. The monarch butterfly, however, has developed an immunity to it. Its caterpillars nibble away at the leaves with impunity. But they do not digest the poison. Instead, they appropriate it and use it for their own purposes. In some way they are able to separate the toxin in the latex and store it unaltered in their bodies. This not only prevents them from succumbing to it, but makes them poisonous to any predator that might swallow them." (Attenborough 1995:70-71)
Learn more about this functional adaptation.
- Attenborough, D. 1995. The Private Life of Plants: A Natural History of Plant Behavior. London: BBC Books. 320 p.
Molecular Biology and Genetics
Statistics of barcoding coverage
|Specimen Records:||41,600||Public Records:||15,573|
|Specimens with Sequences:||38,084||Public Species:||2,664|
|Specimens with Barcodes:||35,656||Public BINs:||2,173|
|Species With Barcodes:||4,527|
A few species in this family (but none in Michigan) are considered endangered or threatened, usually due to destruction of their habitat.
Relevance to Humans and Ecosystems
Economic Importance for Humans: Negative
These harmless butterflies don't directly affect humans much in positive or negative ways. A few species have caterpillars that can cause a rash if you touch them.
Negative Impacts: injures humans (bites or stings)
Nymphalidae is the largest family of butterflies with about 6,000 species distributed throughout most of the world. These are usually medium sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called brush-footed butterflies or four-footed butterflies. Many species are brightly colored and include popular species such as the emperors, admirals, tortoiseshells, and fritillaries. However, the underwings are in contrast often dull and in some species look remarkably like dead leaves, or are much paler, producing a cryptic effect that helps the butterfly disappear into its surroundings.
Rafinesque introduced the name Nymphalia as a subfamily name in diurnal Lepidoptera. Rafinesque did not include Nymphalis among the listed genera, but Nymphalis is unequivocally implied in the formation of the name (Code Article 126.96.36.199). The attribution of the Nymphalidae to Rafinesque has now been widely adopted (e.g., Vane-Wright & de Jong, 2003: 167; Pelham, 2008; Wahlberg, 2010).
For terms see External morphology of Lepidoptera.
In adult butterflies the first pair of legs are small or reduced, giving the family the other names of four-footed or brush-footed butterflies. The caterpillars are hairy or spiky with projections on the head, and the chrysalids have shiny spots.
The forewing has the submedial vein (vein 1) unbranched and in one subfamily forked near base; medial vein with three branches, veins 2, 3, and 4; veins 5 and 6 arising from the points of junction of the discocellulars; subcostal vein and its continuation beyond apex of cell, vein 7, with never more than four branches, veins 8–11; 8 and 9 always arising from vein 7, 10, and also 11 sometimes from vein 7 but more often free, i.e., given off by the subcostal vein before apex of cell.
The hindwing has internal (1a) and precostal veins. The cell in both wings closed or open, often closed in the fore, open in the hindwing. Dorsal margin of hind wing channelled to receive the abdomen in many of the forms.
Antennae always with two grooves on the underside; club variable in shape. Throughout the family the front pair of legs in the male, and with three exceptions (Libythea, Pseudergolis, and Calinaga) in the female also, is reduced in size and functionally impotent; in some the atrophy of the forelegs is considerable, e.g., Danainae and Satyrinae. In many of the forms of these subfamilies the fore legs are kept pressed against the underside of the thorax, and are in the male often very inconspicuous.
Systematics and phylogeny
The phylogeny of the Nymphalidae is complex. Several taxa are of unclear position, reflecting the fact that some subfamilies were formerly well-recognized as distinct families due to insufficient study.
The libytheine clade (basal)
The danaine clade (basal)
- Host plant families include Apocynaceae, Asclepiadoideae (subfamily of Apocynaceae), and Moraceae.
- Most species with long wings, some having transparent wings. Host plants in the families Apocynaceae, Gesneriaceae, and Solanaceae.
The satyrine clade
- Tropical canopy butterflies. Caterpillars often with head spines or projections. Mostly edible species with some Batesian mimics. Host plants in the families Annonaceae, Celastraceae, Convolvulaceae, Euphorbiaceae, Fabaceae, Flacourtiaceae, Lauraceae, Myrtaceae, Piperaceae, Poaceae, Rhamnaceae, Rutaceae, Santalaceae, and Sapindaceae.
- Include the spectacular neotropical Morphos. Food plants include the Arecaceae, Bignoniaceae, Fabaceae, Menispermaceae, Poaceae, and Sapindaceae.
- Brassolini (Owls. Neotropical with 70–80 species. Mostly crepuscular. Sometimes considered a subfamily Brassolinae.)
- Host plants in the families Arecaceae, Araceae, Cyperaceae, Heliconiaceae, Poaceae, and Selaginellaceae.
- Colourful tropical butterflies noted for Müllerian mimicry. All species use host plants in the family Passifloraceae.
- Apaturinae (Mostly tropical)
- Biblidinae (formerly in Limenitidinae)
- Cyrestinae (formerly in Limenitidinae)
- Nymphalinae (A large subfamily that sometimes includes the Limenitidinae and Biblidinae.)
- Some species migratory. Caterpillars sometimes covered in spines. Host plants include Acanthaceae, Caprifoliaceae, Convolvulaceae, Euphorbiaceae, Fagaceae, Flacourtiaceae, Lamiaceae, Loranthaceae, Moraceae, Plantaginaceae, Poaceae, Rubiaceae, Rutaceae, Salicaceae, Sapindaceae, Scrophulariaceae, Urticaceae, and Verbenaceae.
Example species from this family
- Archdukes, genus Lexias
- California Tortoiseshell, Nymphalis californica
- Comma, Polygonia c-album
- Common Buckeye, Junonia coenia
- Common Snout Butterfly, Libytheana carinenta
- Cracker butterflies, genus Hamadryas
- Crimson Patch, Chlosyne janais
- Edith's Checkerspot, Euphydryas editha
- Lorquin's Admiral, Limenitis lorquini
- Marsh Fritillary, Euphydryas aurinia
- Meadow Brown, Maniola jurtina
- Mourning Cloak, Nymphalis antiopa
- Monarch Butterfly, Danaus plexippus
- Blue Morpho, Morpho menelaus
- Painted Lady, Vanessa cardui
- Peacock, Inachis io
- Question Mark, Polygonia interrogationis
- Red Admiral, Vanessa atalanta
- Small Heath, Coenonympha pamphilus
- Small Tortoiseshell, Nymphalis urticae
- Gatekeeper, Pyronia tithonus
- Rafinesque, C.S. (1815) Analyse de la Nature, ou Tableau de l'Univers et des Corps Organisés. Jean Barravecceia: Palermo. 224 pages, p 127.
- Charles Thomas Bingham (1905). Butterflies, Volume 1. The Fauna of British India, Including Ceylon and Burma. London: Taylor and Francis.
- Niklas Wahlberg, Elisabet Weingartner & Sören Nylin (2003). Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea) (PDF). In Gisella Caccone & Giacomo Bernardi. "Papers presented at the Mammalian Phylogeny symposium during the 2002 Annual Meeting of the Society for Molecular Biology and Evolution, Sorrento, Italy, June 13–16, 2002". Molecular Phylogenetics and Evolution 28 (3): 473–484. doi:10.1016/S1055-7903(03)00052-6. PMID 12927132.
- Philip J. DeVries (2001). "Nymphalidae". In Simon A. Levin. Encyclopedia of Biodiversity. Academic Press. pp. 559–573. doi:10.1016/B0-12-226865-2/00039-0. ISBN 978-0-12-226865-6.
- Glassberg, Jeffrey Butterflies through Binoculars, The West (2001)
- Guppy, Crispin S. and Shepard, Jon H. Butterflies of British Columbia (2001)
- James, David G. and Nunnallee, David Life Histories of Cascadia Butterflies (2011)
- Pelham, Jonathan Catalogue of the Butterflies of the United States and Canada (2008)
- Pyle, Robert Michael The Butterflies of Cascadia (2002)