Generic revision of the superfamily Pterophoroidea (Lepidoptera) C. Gielis Gielis, C. Generic revision of the superfamily Pterophoroidea (Lepidoptera). Zool. Verh. Leiden 290, l.xii.1993:1-139, figs. 1-241.— ISSN 0024-1652/ISBN 90-73239-22-2. Key words: Pterophoroidea; Pterophoridae; Macropiratidae; revision; phylogenetics; biogeography; checklist. The genera of the superfamily Pterophoroidea are redescribed. Three new genera are described: Dejongia, Stockophorus and Shafferia. The characters of the genera and their states are translated into a data matrix. A majority rule consensus tree composed with PAUP serves as a basis for a phylogenetic analysis. A cladogram, representing the views of the author, based on the data acquired, is presented with argumentation. The evaluation of the characters is discussed. A new subfamily is erected, the Deuterocopinae. The Pterophorinae are redefined. The Platyptiliinae are synonymized with the Pterophorinae. The Macropiratidae, considered by Minet (1991) a subfamily or a genus of the Pterophoridae, has been raised to family rank, and is placed as the sister group of the Pterophoridae in the superfamily Pterophoroidea. A discussion is given on the biogeographical data. The only known Pterophorid fossil is discussed and placed in a time scale. A checklist is produced of all known species of Pterophoroidea arranged according to the definition of the genera as (re)described in this publication. C. Gielis, c/o Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie), Postbus 9517, 2300 RA Leiden, The Netherlands. Contents I. Introduction 4 Α. General remarks 4 Β. Historical notes 6 C. Facing a problem 8 D. Terminology of wing venation 8 II. The genera of the Pterophoroidea 8 A. Pterophoridae 8 B. Macropiratidae 45 III. The sister group of the Pterophoroidea 45 IV. Phylogenetic analysis 46 A. Methods 46 B. Characters and character states 47 C. Results 51 D. Character evaluation based on the consensus tree 51 V. Taxonomic conclusions 62 VI. Biogeographical notes 63 VII. Acknowledgements 65 VIII. Checklist of taxa of the superfamily Pterophoroidea 65 IX. References 84 X. Illustrations 88 XI. Index Pterophoroidea 126 4 ZOOLOGISCHE VERHANDELINGEN 290 (1993) I. Introduction Α. General remarks* Within the Lepidoptera, the family of the Pterophoridae is well­defined by the cleft in the fore wings and the double cleft in the hind wings in most species. These characters differentiate them from nearly all other Lepidoptera. Only the families Alucitidae (formerly Orneonidae) with six lobes per wing and Oxychirotidae with a single cleft in both the fore and hind wing, have their wings cleft too. Cleft wings, with a single cleft in the fore wing and a double cleft in the hind wing, are not charac­ teristic for the entire family The character unique to the family Pterophoridae is the presence of venous scales at the underside of the hind wing, along veins M3 and Cu2. Currently, the family is divided into four subfamilies. Of these the subfamilies Agdistinae and Ochyrotinae have uncleft wings and the Pterophorinae and Platyptiliinae have cleft wings. The feather­like appearance of the cleft wing, is reflect­ ed in vernacular names of this family: Plume moths (English), Federmotten (German), Fjädermott (Swedish), Fjermöl (Danish) and Vedermot (Dutch). These "feathers", or lobes, as they are called in this publication are fringed by pronounced "hairs". The fringe hairs are differentiated in paler and darker parts. In the fringes pronounced scales lined up in a row, scale groups or scale teeth may be present. Minet (1991) in reconstructing the phylogeny of the ditrysian moths considers the presence of a fifth subfamily: Macropiratinae. The characters mentioned by him, supporting the relationship between the Macropiratinae and the other subfamilies of the Pterophoridae are: (a) ocelli absent; (b) no spinarea (a wing locking device occa­ sionally present in the Alucitoidea); (c) wing position in rest; (d) fenestra media (defined by Minet in 1982) absent or reduced; (e) tergal rim ventrally very narrow in posterior view; among others. The present author is of the opinion that the following characters, which are present in the Pterophoridae but absent in the Macropiratinae are sufficient to give the Macropiratinae family rank: (a) maxillary palpi absent; (b) metathorax with very narrow subalar sclerites; and (c) underside of hind wing with venous scales along M3 and Cul. The families Pterophoridae and Macropiratidae comprise the superfamily Ptero­ phoroidea. The wings are narrow. In rest the wings are placed rectangularly to the body, giv­ ing it a "T"­like appearance, and folded in such a way that only a minor surface remains exposed, making the specimens hard to detect by day. The way the wings are folded is variable and seems to vary per subfamily (Wasserthal, 1974). The wing venation is rather uniform. In the subfamily Pterophorinae some varia­ tion is found in the radial veins, and the differences are used in the generic grouping of the family. The presence of one or two veins in the third lobe of the hind wing has been used to differentiate the subfamilies of the Platyptiliinae and Pterophorinae. In the Agdistinae the markings on the fore wings are reduced to a small number of dots at the costa, at the margin of the naked field and faint spots at the dorsum. (The naked field is a terminal triangular field on the fore wing of Agdistis species, Note.— Wherever the names Platyptiliinae and Pterophorinae are mentioned in this chapter, they are used in their traditional meaning, as current before the results of the present study were known. GIELIS: GENERIC REVISION OF THE PTEROPHOROIDEA 5 containing fewer cover scales. It plays a role in wing folding.) Throughout the family spots at or before the base of the cleft are present. The remaining patterns are more or less restricted to a subfamily or/and generic grouping. In the Platyptiliinae, the Platyptilia group of genera have a remarkable costal triangle, and in the Oxyptilus group the transverse brown markings bordered by narrow white lines are a general phenomenon. In the generic group of Stenoptilia dots and lines before the cleft and in the fore wing lobes are present, combined with small scale groups in the terminal fringes of the fore wing. In the subfamily Platyptiliinae a group or row of pro• nounced scales may be present in the fringes of the termen or, more commonly, on the dorsum. In addition there is a scale tooth at the dorsum of the third hind wing lobe. In the Pterophorinae the markings are limited to a more or less pronounced number of spots at the costa, the dorsum, near the base of the cleft and in the sepa• rate lobes. The pronounced scale teeth, as seen in the Platyptiliinae, are not present in this subfamily. A character present in all species belonging to the family, and found in both males and females, are the rows of specialized scales on the underside of the hindwing. These scales have been referred to as androconial scales (German: Duftschuppen), but their presence in the females makes the function of these scales doubtful. It seems bet• ter to refer to these scales as venous scales. In general the head is broad and appressedly scaled, with a variable number of erect scales in the collar region. The frons may show a conical extension of the head. The eyes are ball-shaped, in lateral view. Ocelli and maxillary palpi are absent. The labial palpi have three segments with appressed scaling, sometimes with small "hair" brushes along the segments, and are protuberant or upcurved. Between the palpi a rather well-developed tongue is present. The antennae are filiform, appress• edly scaled with short ciliation; the first segment is pronounced. The antennae vary in length from half as long to as long as the fore wing. The thorax has a compact, cubical structure. The thoracical segments are appress• edly scaled. The moths are slender, with a long abdomen and long and slender legs. The fore legs are the shortest, the hind legs by far the longest. The tibial segments are longer than the coxal segments. The hind legs have a double pair of spurs on the tibiae. The spurs of each pair may be of equal or unequal length, and the length of the proximal and distal spur pair may be different as well. At the base of the spur pairs a scale brush may be present, as found in the genus Oidaematophorus. The five tarsal seg• ments show considerable differences in size, the basal segment being the longer. At the top of the fifth tarsal segment a pair of small claws is positioned. On the margins of the abdominal segments scale brushes may be present, often paired and placed dorsally. This feature is found in some species of the genus Stenoptilia and Pterophorus. In the Oxyptilus genus complex a lateral scale brush may be found along the ninth segment. On the dorsal and ventral sides of the abdomen linear markings or patterns may be present. The Pterophoridae are very widespread, and are found from the arctic and anti- boreal to the tropical zones. In mountainous areas considerable altitudes may be reached. As a consequence the species occur in very wide range of ecological condi• tions. Examples of extreme conditions can be found in the Agdistis species living in salt marshes along the coast, and under desert-like circumstances in southern Spain, 6 ZOOLOGISCHE VERHANDELINGEN 290 (1993) and in the species of Stenoptilia from the high altitudes in the Alps, and from Iceland and Greenland under arctic conditions. Other species live in shady woods among the herbaceous vegetation, or in clearings of woodlands. Also extreme wet environments are inhabitated, such as moorlands and wet meadows, and banks of rivers, brooks and ditches; drier areas like heaths and herbaceous road sides complete the variety of habitats.