NYMPHALIDAE BRUSHFOOT BUTTERFLIES FAUNA PARAGUAY

NYMPHALIDAE - NYMPHALID BUTTERFLIES
An extremely variable group, the Nymphalids make up about 6000 species worldwide. A wide variety of wing shapes, sizes and colours are present.

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Characteristics
Nymphalids have elongated chaetosemata running parallel to the eye margin. Maxillary palpi are single segmented and labial palpi ascending. Most antennal segments have two antennal grooves. Epiphysis absent. Forelegs are reduced so that they may appear to be "four-legged insects" and are used only by female Libytheinae for walking. Males have a reduced number of tarsomeres and lack a pretarsus. Tibial spur formula 0-2-2 or 0-0-0. Humeral vein present in the hindwing. In some groups small tympanal organs are present in swollen bases of certain wing veins. (Scoble 1995).

Life Cycles
Life cycles are predictably diverse, but eggs are usually laid in clusters. Eggs may be spherical, barrel-shaped or conical. Early instar larvae are commonly gregarious and many construct silk shelters on their foodplants. Larvae are diverse, often possessing scoli, filaments and protruberances and secondary setae are abundant. Crochets are usually in triodrinal mesoseries, only occasionally uni- or biordinal. Pupae are usually suspended by the cremaster, but occasionally a weak cocoon is formed. (Scoble 1995).

Classification
Ten subfamilies of the family Nymphalidae are represented in Paraguay. We currently have images of 132 species in this family. Figures in brackets after the subfamilial names refer to the number of species for which we currently have images.

Superfamily Papilionoidea: Family Nymphalidae.

Subfamily Apaturinae (4) Emperors
Strikingly-coloured, robust Nymphalids with cryptic underwing, creating a flash effect to escape predators. Flight is fast and direct and they share many characteristics with the Charaxinae. One of the most obvious characters of the subfamily is the green proboscis which, despite the superficial resemblance of many species to the Limenitidini, immediately distinguishes the emperors. They feed on dung, carrion and rotting fruits and are often attracted to drink sweat from humans. Sexually dimorphic. Larvae with bifid "tails". (Canals 2003).

Subfamily Biblidinae (35) Biblids
The majority of the species in the Biblidinae are Neotropical. In older literature it was referred to as Eurytelinae.
Tribe Biblidini: Biblids
Approximately 340 species. Small to medium-sized, often colourful species, with rapid, bouncing flight style. Most species rest with wings folded, the large hindwing often obscuring much of the forewing.
Tribe Limenitidini: Sisters
Previously considered a separate subfamily, the Sisters have now been relegated to a tribe of the Biblidinae. The most distinctive characteristic of this group that is obvious in the field is the flap and glide flight style. Certain larval and pupal characteristics are unique to this group.
Tribe Cyrestidini: Daggerwings
Previously considered a separate subfamily, the daggerwings have now been relegated to a tribe of the Biblidinae. Daggerwings are fast-flying, boldly-coloured butterflies, with long, falcated forewings and cryptic underwing patterns. They frequently fly high and feed on rotting fruit, dung and carrion, coming to ground to "puddle" solitarily. Many species are "tailed" in the adult form. Globally there are only three genera and the tribe is most speciose in South East Asia, thought he genus Marpesia is entirely Neotropical.

Subfamily Charaxinae (6) Leafwings
Widespread in the tropics, but barely reaching temperate regions. They are robust forest dwellers, fast-flying and colourful dorsally, though the ventral surface is typically cryptic and leaf-like. They feed on carrion, dung and rotting fruit. Larvae feed on a variety of plants, though any one species is typically restricted to a single plant family or group of families. The larval head is distinctive and bears one or two pairs of horns. (Scoble 1995).

Subfamily Danainae (5) Monarchs
The Monarchs (or Milkweed Butterflies) are robust butterflies, at their most speciose in the Oriental region. Their monophyly is supported by paires, sheathed and eversible hairpencils on the male abdomen and strongly clubbed, spinose, four-segmented pretarsi in the female. Hairpencils are everted by the pressure of haemolymph and withdrawn with retractor muscles. Larval instars after the first stage have one to nine pairs of fleshy dorsal tubercles and aposematic colouration, typically with hoops or stripes. Foodplants are mainly Asclepiadaceae, Apocynaceae and Moraceae. Adults typically bear aposematic colouration and are impalatable to predators. Some species are migratory. (Scoble 1995).

Subfamily Heliconiinae (13) Longwings
Disputed taxonomic history and have at time been considered to be specialised members of the tribe Argynnini in the subfamily Nymphalinae. New World butterflies, the vast majority of which are Neotropical, with about 70 descibed species. Larvae feed on toxic Passifloraceae and the toxicity is carried into adulthood. Most species have aposematic colouration and a slow, fluttery flight designed to advertise their impalatability. Most adults feed on nectar, though some feed on pollen. Larvae have six rows of spines arranged longitudinally with a single pair of spines on the head. Many species are involved in mimic rings, especially with members of the Ithomiinae. (Scoble 1995).

Subfamily Ithomiinae (12) Clearwings and Tigerwings
Around 300 species restricted to the Neotropics. Many are involved in mimic rings with the toxic Heliconiinae, imitating both colouration and flight style, though they possess no toxins themselves and are perfectly palatable to would-be predators. The antennae are scaled. Tarsus of the female foreleg is four- or five-segmented but not strongly clubbed. Larvae feed largely on Solanaceae. In the past they have been considered part of the Danaiinae, but now considered to comprise a separate subfamily. (Scoble 1995).

Subfamily Libytheinae (1) Snouts
Labial palpi are greatly elongated, forming a distinctive "snout". They are widespread but only 10 species are recognised. Crochets of larvae form a pseudocircle of biordinal mesoseries with a few uniordinal crochets in lateroseries. They are smooth, lack head processes and do not have a bifid "tail". They feed on membes of the Ulmaceae. (Scoble 1995).

Subfamily Morphinae (8) Morphos and Owl Butterflies
Confined to the Neotropics, two distinct tribes are present, both formerly separated as subfamilies. The Morphos (Morphini) are arguably the most eyecatching of Paraguayan butterflies, being large and bright blue in colour. They feed largely on rotting fruit and excrement. Most larvae feed on dicotyledonous plants. The Owl Butterflies (Brassolini) are typically large and fast-flying, being characterised by the presence of eyespots on a cryptic underwing. Most take to the wing at dusk and inhabit shady areas. Larvae are smooth, often with a bifid "tail" and feed on monocotyledons. (Scoble 1995).

Subfamily Nymphalinae (20) Brushfoot Butterflies
Formerly much larger the Nymphalinae included several other groups that have now been separated into distinct subfamilies. Nymphalids have reduced forelegs in both sexes and the larvae often bear spines. Live cycles are diverse but many species feed on Acanthaceae or Urticaceae. (Scoble 1995).

Subfamily Satyrinae (28) Satyrs
Around 1500 species globally, the Satyrs are butterflies of shady areas and forest undergrowth. Typically brown in colour with cryptic underwing patterns often with lines and ocelli, and bases of wing veins are often swollen. Larvae usually bear cephalic horns and a bifid "tail". They are most closely related to the Charaxinae and it has been suggested that that group should be merged into the Satyrinae to create monophyly. Predominant foodplants are monocotyledons. (Scoble 1995).

References:
Canals GR 2000 - Mariposas Bonaerenses - LOLA, Buenos Aires.
Canals GR 2003 - Mariposas de Misiones - LOLA, Buenos Aires.
Scoble MJ 1995 - The Lepidoptera: Form, Function and Diversity - Oxford University Press, Oxford.
Smart P 1975 - The Illustrated Encyclopedia of the Butterfly World - Salamander Books, London
Turner D, Stanêk VJ 1992 - The Illustrated Encyclopedia of Butterflies and Moths - Select Editions, London.