Hedylidae
Wikipedia, the free encyclopedia - Cite This SourceHedylidae, the "American moth-butterflies", is a family of insects in the lepidopteran order, representing the superfamily Hedyloidea. They are an extant sister group of the butterfly superfamilies Papilionoidea and Hesperioidea. In 1986, Scoble combined all species into a single genus Macrosoma, comprising 35 currently recognized and entirely Neotropical species, as a novel concept of butterflies.
Taxonomy and systematics
Hedylidae were previously treated as a tribe of Geometridae: Oenochrominae, the "Hedylicae Prout considered they might even merit treatment as their own family. Scoble first considered them to be a hitherto unrecognised group of butterflies and also suggested Hedylidae might possibly constitute the sister group of the "true" butterflies (Papilionoidea, rather than of (Hesperioidea + Papilionoidea). Weintraub and Miller argued against this placement (but see). In 1995, Weller and Pashley found that molecular data did indeed place Hedylidae with the butterflies and a more comprehensive study in 2005 based on 57 exemplar taxa, three genes and 99 morphological characters, recovered the genus Macrosoma as sister to the ("true butterflies" + "skippers"). Since there are no obvious gaps between supposed species groups, according to basic morphological structure, Scoble (1986) synonymised the five pre-existing genera (33 of which had been described in Phellinodes) into just one genus. However, a phylogenetic analysis of all Macrosoma species is still needed.Nomenclatural notes
In Zoological nomenclature, there exist numerous junior homonyms of Macrosoma Hübner, 1818, (Macrosoma Leach 1819 (a reptile), Macrosoma de Haan 1826 (Odonata), Macrosoma Robineau-Desvoidy 1830 (Macrosoma multisulcata Berlese 1913 and M. floralis, Diptera: Muscidae), Macrosoma Brandt 1835 (Coelenterata), Macrosoma Hope 1837 (Coleoptera), Macrosoma Lioy 1864 or 1865 (Diptera: Sarcophagidae), Macrosoma Hammer 1979 (M.rugosa; Acarina: Oribatidae). To add to this potential confusion in lists of names, there exist two junior homonyms of Hedyle Guenée, 1857: Hedyle Bergh, 1895 (sea slugs in the order Opisthobranchia: superfamily "Acochlidioidea", family Hedylopsidae Odhner, 1952
that are currently placed in the genus Hedylopsis Thiele, 1931)
, and Hedyle Malmgren 1865 (a polychaete worm)
The sea slug family name "Hedylidae Bergh, 1895" (type species Hedyle weberi Bergh, 1895) is thus also invalid.
Morphology and identification
The eggs of hedylid moths have an upright configuration and are variable in shape: in Macrosoma inermis they are particularly narrow and spindle-shaped resembling those of some Pieridae and in the case of M. tipulata they are more barrel-shaped, like certain Nymphalidae. The larvae resemble probably convergently those of Apaturinae. Adult hedylids resemble geometer moths. They share many characteristics both (morphological and genetic) with the superfamilies Papilionoidea and Hesperioidea. The abdomen is very long and slim, like many Neotropical butterflies of the subfamilies Ithomiinae and Heliconiinae, hence the name of one Macrosoma species "heliconiaria". Unlike other butterflies however, the antennae are un-clubbed, but rather filiform or bipectinate. Unlike the family Geometridae, in which they had been placed by the geometer expert L.B.Prout, hedylids lack tympanic organs at the base of the abdomen, but have them on the wings (see under Behaviour). Unlike other butterflies, however, (except in the unique case of the remarkable Australian skipper butterfly Euschemon rafflesia whose males possess these structures), the single-spined frenulum, and retinaculum is not lost or reduced in males, except in three Macrosoma species where there is no functional wing coupling system; the retinaculum is always lost in females and the frenulum may be vestigial. The family have been fully catalogued and illustrated in an identification guide.Butterfly-like characteristics of Hedylidae
- "Mesoscutum" with "secondary line of weakness" near median "notal" wing process, as in some representatives of Papilionidea and Hesperioidea (potentially unique butterfly character;
- Mesothoracic aorta with horizontal chamber, as in other butterflies (not Papilionidae), but as also in Cossidae;
- Metathoracic "furca" resembling a blunt arrowhead; this a variable but potentially unique character in butterflies;
- Second median plate of forewing base lying partly under the base of vein "1A+2A", unlike the configuration in moths;
- "Postspiracular bar" on first abdominal segment;
- Female genitalic "anterior apophyses" reduced;
- Male genitalia relatively "deep" dorso-ventrally;
- Abdomen curved (especially in males), as in papilionoids;
- Abdominal first tergal segment is strongly "pouched" (Scoble 1986; as also in Thyatirinae moths;
- "Precoxal" sulcus joining "marginopleural" sulcus;
- Male Foreleg pretarsus lost,thus fused into two elements as in nymphalid butterflies, with the mid and hindlegs used for perching, but apparently redeveloped in hesperiids;
- Egg upright, spindle-shaped and ribbed as in some Pieridae (e.g. the Orange tip butterfly), some other butterflies, and as in some moth groups also;
- Larva with "anal comb", as in some Hesperioidea (not however Megathyminae) and Pieridae, but not in other Papilionoidea except one species (and also independently in Tortricidae), that is used for propulsion of frass away from the caterpillar;
- Caterpillar with horn-like processes and a "bifid" tail as in many Nymphalidae;
- Caterpillar with "secondary setae", as in Pieridae;
- Ventral larval proleg "crochet" hooks not forming a complete circle, unlike configuration in hesperiids and papilionoids; Image of 1st abdominal segment like in Pieridae (as also in some Geometridae, especially the subfamily Sterrhinae (in which the girdle is around the abdomen), but lost in many Papilionoidea);
- Pupal cocoon lost, as in papilionoids, and a few other groups of Lepidoptera;
- "Temporal cleavage line" lost in the pupa (as in papilionoids).
Distribution
Hedylidae range in Central America south from central Mexico and in South America through the Amazon from southern Peru (where there are a full 26 species, up to 12 at a single site: to central Bolivia and southwestern Brazil). In the Caribbean, they also occur in Cuba, Jamaica and Trinidad.Behaviour
Hedylids are attracted to articifial lights, but occasionally some species can be found flying by day. Thus, they may be involved in some mimicry complexes with Ithomiinae (e.g. the female only of Macrosoma lucivittata. A few species are white and resemble pierid butterflies (e.g. Macrosoma napiaria). Based on a study of Macrosoma heliconiaria , it has been found that hedylids have tympanic organs on their forewings for hearing apparently homologous to the "Vogel's organ" in some Papilionoidea that would help them evade bats at night. They have been shown to exhibit typical moth evasive behaviour towards bats such as erratic spiralling movements and dives. The resting posture is often at a curious angle
, with the thorax tilted and the posterior edge of the hindwings nearly touching the substrate (Scoble, 1986). The larvae which lack the prominent horns in the first instar tend to rest on the midrib of the leaf and often skeletonise leaves or at either side produce an untidy patchwork of holes
The elegant pupa is attached by a cremaster and silken girdle
and sometimes resembles a bird dropping
List of species
This list of species is largely based on pheneticcharacters.- Macrosoma tipulata Hübner, 1818Type Locality: Brazil
Range: Costa Rica as far as SE BrazilTYPE: type probably lostHostplants: Theobroma grandiflorum
, T.angustifolium, T. bicolor)
Card Pupa Larva Adult Adult - Macrosoma hyacinthina (Warren, 1905) (originally in Lasiopates)Type locality: Cayapas, N.W. Ecuador 1°5'0 N, 79°3'0 WRange: widespread from Mexico through Central America into South America (Colombia, Ecuador, Venezuela, Guiana, and Goiás, west Brasilia in central BrazilTYPE: BMNH, Holotype male Card Live Photo
- Macrosoma heliconiaria (Guenée, 1857)Type locality: Cayenne, French Guiana 4°56'0 N, 52°20'0 WRange: Widespread through tropical South America (Colombia, Venezuela, French Guiana, Brazil (to Belém) and to midwestern PeruTYPE: BMNH, Lectotype female Card Image 06-SRNP-100259
- =Hedyle heliconaria Walker, 1862 [Incorrect subsequent spelling of heliconiaria Guenée, 1857] Card
- Macrosoma semiermis (Prout, 1932) (originally in in Hedyle]Type locality: PanamaRange: Widespread from Mexico through Central America into Colombia, Ecuador, Venezuela and Guiana and Goiás, west BrasiliaTYPE: BMNH, Holotype male Card Adult Adult AdultHostplants: Byttneria aculeata
and Luehea sp.
) - =Hedyle inermis Prout, 1932Type locality: Sta. Cruz, Bolivia many possible locationsTYPE: BMNH, Holotype male Card
- Macrosoma albipannosa (Prout, 1916) (originally in Hedyle)Type locality: Cerro de Pasco, Huancabamba, E. Peru 10°21'0 S, 75°32'0 WRange: Andean Peru and Ecuador (Intas), to 10000 feet [3050 m.]TYPE: BMNH, Holotype male Card
- Macrosoma pectinogyna Scoble, 1990Type locality: Loja [Prov], Celica, 2000 m., Ecuador style="font-style : italic;"> S, 79°57'0 WRange: Celica, Loja in EcuadorTYPE: AMNH, Holotype female Card
- Macrosoma hedylaria (Warren, 1894) (originally in Phellinodes)Type locality: (Nord Brasilien?), S.AmericaRange: Widespread from Eastern Colombia, amazon of Venezuela, NW to NE and S Brazil beyond Sao Paulo and either side of the Peruvian Andes and west to LimaTYPE: BMNH, Holotype male Card Image
- Macrosoma conifera (Warren, 1897) (originally in Phellinodes)Type locality: Paramaribo, Surinam 5°50'0 N, 55°10'0 WRange: Widespread in Central America from Guatemala south to South American Paraguay (Villarica), Gorgon Island in Colombia, Cuzco in the Peruvian Andes to eastern Brazil, Amazonian Venezuela, Trinidad, Surinam, French Guiana, to Amazonian NE Brazil (Belém)TYPE: BMNH, Lectotype female, slide# 12341 Card Image Larva Larva Pupa AdultHostplants: Ochroma pyramidale, Croton schiedeanus, Hampea appendiculata, Conostegia xalapensis, Miconia argentea, Ossaea

- =Phellidones conifera gorgonensis (Prout, 1932)Type locality: Gorgona I., W. Colombia, 200' 3°24'0 N, 76°23'0 WTYPE: BMNH, Lectotype female Card
- =Phellidones latiplex Dognin, 1911Type locality: St-Laurent du Maroni, French Guiana 5°30'0 N, 54°2'0 WTYPE: NMNH, Holotype male USNM# 30855 Card
- =Phellinodes zikani (Prout, 1932)Type locality: Itatiaya, Brazil 19°20'0 S, 41°26'0 WTYPE: Senckenberg Museum Frankfurt, Holotype male Card
- Macrosoma intermedia (Dognin, 1911) (originally in Phellinodes)Type locality: ColombienRange: sparse records in Costa Rica, Colombia (St. Antonio), and Rio Ucayali, Peru TYPE: NMNH, Holotype male USNM# 30850 Card
- =Phellinodes biapicata Prout, 1917Type locality: Rio Ucayali, Contamamana, Peru 4°30'0 S, 73°27'0 WTYPE: BMNH, Holotype male Card
- =Phellinodes gratiosa Schaus, 1912Type locality: Tuis, Costa Rica 9°51'0 N, 83°35'0 WTYPE: NMNH, USNM# 17773 Card
- Macrosoma cascaria (Schaus, 1901) (originally in Hyphedyle)Type locality: Jalapa, Mexico style="font-style : italic;"> N, 92°40'0 W, or other locationsRange: Mexico (Tabasco, Misantla), south to northern VenezuelaTYPE: NHMN, Lectotype female USNM# 11906 Card Larva Larva Pupa Pupa Adult Adult AdultHostplants: Croton schiedeanus, C.megistocarpus, Conostegia xalapensis

- Macrosoma paularia (Schaus, 1901) (originally in Hephedyle)Type locality: São Paulo, S. Brazil] many locationsRange: Sparse records from Brazil (Mato Grosso, Planaltina, south to Sao Paulo and from northern Bolivia (Province del Sara)TYPE: NMNH, Lectotype female, USNM# 11907 Card
- Macrosoma albifascia (Warren, 1904) (originally in Phellinodes)Type locality: Carabaya, S[an] Domingo, SE PeruRange: eastern Andes of South America (Ecuador, Peru, Bolivia) TYPE: BMNH, Holotype male Card
- =Phellinodes albifascia expedita (Prout, 1932)Type locality: Macas, EcuadorTYPE: Senckenberg Museum Frankfurt, Holotype male Card
- Macrosoma stabilinota (Prout, 1932) (originally in Phellinodes)Type locality: Fonte Boa, Brazil, Amazonas 1°8'0 N, 67°12'0 W or 2°32'0 S, 66°1'0 WRange: northern Honduras, northern Peru, northwestern Brazil, French Guiana and JamaicaTYPE: BMNH, Holotype male Card Photo
- Macrosoma nigrimacula (Warren, 1897) (originally in Phellinodes)Type locality: 10 miles above Mapiri, Bolivien 10°31'0 S, 66°52'0 W or 15°15'0 S, 68°10'0 WRange: central Mexico south to C-S Peru and western Bolivia, and Amazonian Brazil from Fonte Boa to BelémTYPE: BMNH, Holotype female Card Adult
- =Phellinodes cellulata Dognin, 1911Type locality: Near Loja, El Monje, EcuadorTYPE: NMNH, USNM# 31774, Holotype male Card
- =Phellinodes interrupta Warren, 1904Type locality: Rio Cachabí, S[an] Javier, Ecuador, Holotype female 1°3'18 N, 78°49'5 W or 1°3'58 N, 78°46'38 W,or other locationsTYPE: BMNH, Holotype female Card
- =Phellinodes megalophysa Warren, 1908Type locality: St Jean, Maroni, French Guiana style="font-style : italic;"> N, 54°2'0 WTYPE: NMNH, USNM# 11377, Lectotype female Card
- =Phellinodes obstructa Warren, 1904Type locality: Pambilar, EcuadorTYPE: BMNH, Holotype male Card
- =Phellinodes parornata Dognin, 1911Type locality: Nouveau Chantier, French GuianaTYPE: NMNH, Holotype male, USNM# 30853 Card
- Macrosoma klagesi (Prout, 1916) (originally in Phellinodes)Type locality: Fonte Boa, Amazonas, Brazil 1°8'0 N, 67°12'0 W or 2°32'0 S, 66°1'0 WRange: Fonte Boa, Amazonas, Brazil (known only from holotype)TYPE: BMNH, Holotype male Card
- Macrosoma costilunata (Prout, 1916) (originally in Phellinodes)Type locality: Paramba, Ecuador 0°49'0 N, 78°21'0 WRange: sparsely recorded from northwestern Costa Rica and N and C EcuadorTYPE: BMNH, Holotype female Card
- Macrosoma muscerdata (Felder & Rogenhofer, 1875) (originally in Phellinodes)Type locality: AmazonasRange: central Costa Rica, eastern Colombia to French Guiana, through Ecuador and Peru south to Carabaya range, and in Amazonian Brazil from São Paulo de Olivencia to BelémTYPE: BMNH, Holotype female Card
- Macrosoma lucivittata (Walker, 1863) (originally in Hedyle)Type locality: Ega, [=Tefé], Brazil 3°22'0 S, 64°42'0 WRange: Honduras south to SE Peru, and Belém in Amazonian BrazilTYPE: BMNH, Holotype female Card
- =Phellinodes absentimacula (Warren, 1904)Type locality: Salidero, N.W. Ecuador
TYPE: BMNH, Holotype male Card - Macrosoma coscoja (Dognin, 1900) (originally in Phellinodes)Type locality: Environs de Loja, Equateur, Ecuador 4°0'0 S, 79°13'0 WRange: eastern Colombia, E-C and S Ecuador, C to SE Peru and eastern BoliviaTYPE: NMNH, Lectotype female, USNM# 30834 Card
- Macrosoma albistria (Prout, 1916) (originally in Phellinodes)Type locality: Carabaya, Rio Huacamayo, La Union, Peru 13°30'0 S, 69°40'0 W 13°32'0 S, 69°38'0 WRange: C and S Peru and southern BoliviaTYPE: BMNH, Holotype male Card
- Macrosoma bahiata (Felder & Rogenhofer, 1875) (originally in Phellinodes)Type locality: Amazonas, BahiaRange: Eastern Mexico (Quintana Roo Territory) through Central America to Lima in Peru and east to Belém in Amazonian BrazilTYPE: BMNH, Holotype male Card Larva larva Larva Adult Adult Adult
- Macrosoma uniformis (Warren, 1904) (originally in Phellinodes)Type locality: Rio Napo, E. Ecuador [Peru on specimen label]Range: C Colombia to S Peru and C Bolivia southeast to Brazil (Rio de JaneiroTYPE: BMNH, Holotype male ? Rio Napa 0°20'0 S, 78°54'0 W Card
- Macrosoma amaculata Scoble, 1990Type locality: Rincon National Park, 4 km. E. Cas[e]tilla, Guanacaste Prov., Costa Rica style="font-style : italic;"> N, 85°44'0 WRange: C-W and NW Costa RicaTYPE: BMNH, Holotype male Card
- Macrosoma leucophasiata (Thierry-Mieg, 1904) (originally in Phellinodes)Type locality: Rio Colorado, Peru, 2500' 13°34'0 S, 72°33'0 W; other locationsRange: SE Ecuador to S Peru and BoliviaTYPE: NMNH, Lectotype male, # 30852 Card
- =Hyphedyle divisa Warren, 1905Type locality: Chanchamayo, Peru 11°3'0 S, 75°19'0 W or 13°42'0 S, 75°48'0 W or 14°8'36 S, 75°41'14 WTYPE: BMNH, Holotype male Card
- Macrosoma albimacula (Warren, 1900) (originally in Hephedyle)Type locality: Paramba, W. Ecuador 0°49'0 N, 78°21'0 WRange: W and N Ecuador and C Peru (1630 m.)TYPE: BMNH, Holotype male Card
- Macrosoma leucoplethes (Prout, 1917) (originally in Phellinodes)Type locality: Alpayacu, Rio Pastaza, E. Ecuador 1°35'0 S, 77°45'0 WRange: E EcuadorTYPE: BMNH, Holotype male Card
- Macrosoma satellitiata (Guenée, 1857) (originally in Phellinodes)Type locality: BrasilienTYPE: BMNH Card Photo
- =Phellinodes praecostalis Dognin, 1911Type locality: Yuntas near Cali, Colombia 3°46'27 N, 76°44'40 W 3°25'0 N, 76°33'0 W, or other locationTYPE: NMNH, Holotype male, USNM# 30857 Card
- =Phellinodes zapotensis Prout, 1932Type locality: Zapote, Guatemala style="font-style : italic;"> N, 89°56'0 W, or other locationsTYPE: BMNH, Lectotype female Card
- Macrosoma subornata (Warren, 1904) (originally in Hyphedyle)Type locality: Carabayo, S. Domingo, Peru 13°50'0 S, 70°15'0 WRange: N Colombia and Ecuador to SE PeruTYPE: BMNH, Holotype male Card Adult photo identified as nigrimacula
- =Phellinodes desueta Prout, 1932Type locality: Pueblo Rico, San Juan, Choco, slopes of Colombia, 5200 ft. [1585 m.]TYPE: BMNH, Lectotype male Card
- Macrosoma lamellifera (Prout, 1916) (originally in Phellinodes)Type locality: Fonte Boa, Amazonas] 1°8'0 N, 67°12'0 W or 2°32'0 S, 66°1'0 WRange: sparse records fromupper Amazon of Brazil and SurinamTYPE: BMNH, Holotype male Card
- Macrosoma minutipuncta (Prout, 1916) (originally in Phellinodes)Type locality: La Oroya, Rio Inambari, SE Peru, 3100' 11°32'0 S, 75°54'0 W