Systematics and Zoogeography of the Family Neopseustidae with the Proposal of a New Superfamily (Lepidoptera: Neopseustoidea) DONALD R. DAVIS SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 210 SERIAL P U B L I C A T I O N S OF T H E S M I T H S O N I A N I N S T I T U T I O N The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These pub- lications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available. S. DILLON RIPLEY Secretary Smithsonian Institution S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 2 1 0 Systematics and Zoogeography of the Family Neopseustidae with the Proposal of a New Superfamily (Lepidoptera: Neopseustoidea) Donald R. Davis ISSuEO SEP 2 1975 SMITHSONIAN INSTITUTION PRESS City o? Washington 1975 ABSTRACT Davis, Donald R. Systematics and Zoogeography of the Family Neopseustidae with the Proposal of a New Superfamily (Lepidoptera: Neopseustoidea). Smith- sonian Contributions to Zoology, number 210, 45 pages, 98 figures, 1 table, 1975.? The general morphology, zoogeography, and classification are reviewed for the three genera and seven species of Neopseustidae. One new genus (Apoplania) and three new species (Apoplania chilensis, Neopseustis bicornuta, and N. sinen- sis) are described. The discovery of Apoplania chilensis marks the first record of this family from the Western Hemisphere. Distribution maps and keys are pro- vided for all species. A new superfamily, Neopseustoidea, is proposed for the Neopseustidae which is tentatively placed in the suborder Dacnonypha. OFFICIAL PUBLICATION DATF. is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5390. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging in Publication Data Davis, Donald Ray Systematics and zoogeography of the family Neopseustidae with the proposal of a new super- family (Lepidoptera: Neupseustoidea) (Smithsonian contributions to zoology, no. 210) Bibliography: p. Supt. of Docs, no.: SI 1.27:210 1. Neopseustidae. 2. Insects?Classification. 3. Insects?Geographical distribution. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 210. QL1.S54 no. 210 [QL561.N38] 591'.08s [595.7'8] 75-619049 Contents Page Introduction 1 Zoogeography 2 Review of Morphology and Classification 5 Abbreviations 13 Neopseustidae 13 Key to the Genera of Neopseustidae 15 Archepiolus Mutuura 15 A. schmidi Mutuura 16 Neopseustis Meyrick 16 Key to the Species of Neopseustis 18 N. calliglauca Meyrick 18 N. bicornuta, new species 19 N. archiphenax Meyrick 20 N. meyricki Hering 21 N. sinensis, new species 22 Apoplania, new genus 23 A. chilensis, new species 24 Literature Cited 26 Systematics and Zoogeography of the Family Neopseustidae with the Proposal of a New Superfamily (Lepidoptera: Neopseustoidea) Donald R. Davis Introduction Few families of Lepidoptera are as little known as the Neopseustidae, and even though this primi- tive group has been recognized as a valid and very distinct family for nearly fifty years, it is probable that the majority of lepidopterists today are little aware of its very existence. The reason for this obscurity is primarily one of rarity, apparently in nature as well as in collections, of both species and specimens. Until recently, for example, only one genus, comprising three rare species, had been described over the past sixty years. Furthermore, the family was known to occur only in the Indian region where probably fewer than eight collectors had ever encountered these insects in their natural habitats. Presently, only 26 specimens are known to exist of the three genera and seven species now recognized; all of these have been examined in the course of this study. Unfortunately, essentially nothing is known re- garding the life history of any member of this family. The adults may be primarily crepuscular or nocturnal in that they are readily attracted to lights and have not been noted active during the day. Their means of oviposition probably involves a rasping or tearing action perhaps similar to that described for the prodoxine moth, Parategeticula Donald R. Davis, Department of Entomology, National Mu- seum of Natural History, Smithsonian Institution, Washing- ton, DC. 20560. (Davis, 1967). The eggs likely are inserted into some part of a host plant possessing relatively firm composition, such as that of most young fruits or stems. The larvae are thus suspected of being internal borers, perhaps either in fruits or stems. The eggs are probably not inserted very deeply but may only be partially imbedded in shallow excavations similar to that formed by the females of Parategeticula. The principal aim of this report is to inform entomologists and collectors of this almost totally neglected group, and in particular, to review the zoogeography, general morphology, and classifica- tion of its components. Similar studies are cur- rently being pursued by the author on the almost equally primitive and poorly known families of Prototheoridae and Paleosetidae in an attempt to learn more of the antiquity and evolution of the Lepidoptera. Several individuals have aided me during the course of this review by providing critical infor- mation or special assistance. In this regard I wish to express my appreciation to Dr. Ian Common, CSIRO, Canberra City, Australia; Dr. Thomas Donnelly, State University of New York, Bing- hamton, N.Y.; Dr. John Dugdale, Dept. of Scien- tific and Industrial Research, Nelson, New Zealand; Prof. Syuti Issiki, formerly of the University of Osaka, Osaka, Japan; Dr. N. P. Kristensen, Uni- versitetets Zoologiske Museum, Copenhagen, Den- mark; Dr. Tosio Kumata, Hokkaido University, SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Sopporo, Japan; Dr. M. S. Mani, St. John's Col- lege, Agra, India; and Dr. Akira Mutuura, Ento- mology Research Institute, Ottawa, Canada. I am especially grateful to Mr. Luis Pena G., Universi- dad de Chile, Santiago, Chile, for the habitat photographs and specimens of Apoplania chilensis, new species. For assistance with illustrations I wish to thank the following staff artists of the Depart- ment of Entomology, Smithsonian Institution: Mr. Lawrence Druckenbrod, Ms. Elsie Froeschner (re- tired), Mr. Andre Pizzini (recently deceased), and Mr. George Venable. I am further indebted to Mr. Victor Kranz of the Smithsonian Photographic Laboratory and to Mr. Walter Brown and Miss Mary Mann of the Smithsonian Scanning Electron Microscope Laboratory for their much appreciated photographic assistance. In addition, I wish to thank my colleagues in the institutions and collec- tions listed below for allowing me to examine specimens under their care. BMNH British Museum (Natural History), London, Eng- land CNC Canadian National Collections, Entomology Re- search Institute, Ottawa, Canada HU Hokkaido University, Sopporo, Japan LEP Collection of Luis E. Pena, Universidad de Chile, Santiago, Chile USNM National Museum of Natural History (under cat- alog numbers of former United States National Museum), Smithsonian Institution, Washington, DC. ZMHU Zoologisches Museum der Humboldt-Universitat, Berlin, DDR-East Germany Zoogeography Previous to this report, members of the family Neopseustidae were known only from the Assam region of northeastern India, Burma, and Taiwan. Presented herein are the first records of neopseu- stids from the Western Hemisphere (the northern Valdivian forest region of Chile) and mainland China. Five of the seven known species occur at various points along the Himalayan Chain (Map I), thus suggesting this general region as an important refugium for the family. Undoubtedly, other members of Neopseustidae remain to be discovered from this area. The endemic Formosan species is now somewhat geographically isolated from the Himalayan species; however, it is closely related to two species of Neopseustis from Burma and Szechuan, China. The present distribution of this family corre- sponds to that of several other relict insect groups in being restricted to relatively cool, mountainous habitats. In this regard, it is interesting that the relict family Paleosetidae also shows an Assam- Taiwan distributional pattern, in addition to hav- ing endemic representatives in Australia. The limited capture data thus far indicate the Neo- pseutidae prefer forested habitats between 600 and 2300 meters. A consideration of the Himalayan Chain as an ancient faunal refugium may at first appear in strong contradiction to the observation that these mountains are relatively recent, beginning to form no earlier than the middle or late Miocene. The Himalayas, of course, have long been recognized as a Pleistocene refugium, with the present fauna being predominantly derived from the north (Asia). The lower or intermediate elevations, how- ever, apparently harbor much older, Gondwanian elements, which originally migrated from the ancient southern Indian Peninsula during the formation of the Himalayas. Migrations from the Peninsula to the north were perhaps most frequent following a retreat of the Glaciers (post-Pleistocene) and before the Indo-Gangetic Plain achieved its present condition as a type of xerophytic barrier. As pointed out by Mani (1974), an exchange of Gondwanian and Asiatic faunas probably occurred during preglacial as well as postglacial times, par- ticularly through the Assam gateway. Understanding the present distribution of not only the Neopseustidae but of all families of primitive Lepidoptera (i.e., Zeugloptera, Dacno- nypha, and Monotrysia) may prove to be a valuable means of establishing a better concept as to the age of the Order and the emergence of its major components. The inadequacy of fossil evidence in deducting phylogenies involving the Lepidoptera has been emphasized frequently. Although the Order is believed to have arisen over 200 million years ago, only one actual fossil remain?a frag- mented larval head capsule embedded in Creta- ceous amber?is known earlier than the Tertiary period (MacKay, 1970). Several authors (Tillyard, 1935; Forbes, 1932; Hinton, 1946) have proposed early Mesozoic or late Paleozoic origins for the Order, basing their beliefs more on conjecture or NUMBER 210 indirect comparisons with fossil evidence from other orders of insects. Renewed interest within the last decade and a half in the phenomenon of continental drift, in conjunction with certain principles proposed in the recent upsurge of phylogenetic systematics (Hennig, 1966a, 1966b; Brundin, 1966), have facilitated means for dating extant groups for which fossil evidence is largely lacking. Thus, as the approxi- mate occurrence of certain continental movements or land-bridge connections become better docu- mented, minimum age determinations, or the terminus post quern non of Hennig (1966a), can be proposed for groups whose present distributions can best be explained by those phenomena. Unfortunately, too little is known concerning the present distribution of the Neopseustidae to decide beyond a reasonable doubt on the origin and prin- cipal dispersal routes for this group. Although the recent discovery of the Andean genus Apoplania suggests a transantarctic dispersal route, a more northern, Laurasian origin cannot be excluded with the evidence available. As discussed recently by Ashlock (1974), under what he termed the "drift sequence rule," at least three continents must be involved in considerations of this nature before direct biotic connections can be confirmed. It is, of course, possible that additional members of this group might yet be discovered in other remnants of Gondwanaland, thereby indicating more strongly a Gondwanian origin. Climatic changes, however, during the late Tertiary and Cenozoic, as witnessed by the warming and drying trends in South Africa and Australia, may have exterminated any forms previously existing there. There seems little doubt that the Neopseustidae, 4 Archepioius schmidi Mutuura A Neopseustis caliiglauca Meyrick + Neopseustis archiphenax Meyrick ? Neopseustis sinensis, new species it Neopseustis bicornuta, new species ? Neopseustis meyricki Hering MAP 1.?Distribution of Old World Neopseustidae. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY as well as several other groups of aculeate Lepi- doptera, are ancient enough to have had their present distributions seriously affected by conti- nental drift, which, conversely, has been shown to have occurred late enough to be a major factor in the zoogeography of several extant groups of organisms. For example, evidence, based on mor- phology as well as biogeography, is beginning to appear which suggests that at least two major groups of Lepidoptera, represented by the present- day Micropterygidae and Hepialidae, arose early in the history of the Order. Both of these groups are considered among the most generalized Lepidop- tera existing today, and both demonstrate a broad, bipolar distribution suggesting a widespread dis- persal prior to the separation of Pangaea. If the present, widespread distributions of these two groups are the result of such a phenomenon, then this would suggest a minimum date for the exist- ence of the Micropterygidae and Hepialidae of approximately 170 million years ago, or during the Jurassic. Also interesting in this regard are the Incurvariidae which likewise show a broad, bipolar distribution with sister-group complexes existing at similar latitudes between both the Nearctic- Palearctic and the southern Ethiopian-Neotropical regions. The Micropterygidae have generally been con- sidered the most primitive surviving group of all the Lepidoptera. For example, Hinton (1946), emphasizing several primitive features in the adult mouthparts, believed the Micropterygidae were even more primitive than the Trichoptera and retained the family in a separate order, Zeugloptera, as earlier proposed by Chapman (1917). Recently (Friese, 1969), the Hepialidae have been consid- ered the most primitive and, therefore, the earliest of the Lepidoptera. Friese primarily arrived at his conclusions by deemphasizing characters of the mouthparts and instead utilized a partially differ- ent set of characters such as the relative develop- ment of the prothorax and the abdominal ganglia. Several primitive families of Lepidoptera show a Gondwanian origin; thus, those groups may have appeared later than the Micropterygidae and Hepialidae and after a separation of the northern and southern continents. Families falling into this category are the Agathiphagidae, Lophocoronidae, Mnesarchaeidae, Paleosetidae, Prototheoridae, and possibly Neopseustidae. The Eriocraniidae probably appeared during the same general period but arose, instead, in Laurasia. Thus, the Eriocraniidae should have some indication of a sister group in the Southern Hemisphere. Such a group may be repre- sented by the Lophocoronidae, a family only recently discovered (Common, 1973). All present indications suggest an Old World origin, now concentrated in the Himalayan Chain for the Neopseustidae. Archepiolus, in possessing the greatest number of plesiomorphic characters, particularly in its wing structure, clearly presents the most generalized member of the group. Pos- sessing a number of apomorphic characters (i.e., reduced epiphysis, reduced anal pocket in the fore- wing, and heavily sclerotized aedeagus), Apoplania chilensis, new species, represents an ancient deriva- tive from a common stock shared with Neopseustis. Just how ancient was this divergence is largely con- jectural at present, as pointed out previously. How- ever, for many of the same reasons summarized by Hennig (1966a) for certain Diptera, the mini- mum period for the separation of the common Neopseustis-Apoplania stem would appear to be somewhere in the mid-Cretaceous. The Atlantic Ocean was forming during that period (Smith et al., 1973), and the separation between North and South America was well established (Kossmatt, 1936). Furthermore, considering that South Amer- ica was isolated from North America for most of the Tertiary (Darlington, 1957), one would believe it improbable for the Neopseustidae to have mi- grated over such vast distances from Asia, or even from North America, during the late Tertiary and Cenozoic without leaving some evidence of their former existence. Certainly the Microlepidoptera fauna is far better known for the northern conti- nents than for the southern ones. Before general hypotheses such as those just dis- cussed can be confirmed, further investigations, based on phylogenetic principles, are needed on the general morphology and zoogeography of the primitive Lepidoptera. Additional field work in critical areas is particularly necessary. As an exam- ple of the latter, recent field discoveries as reported in this paper and by Dumbleton (1952), Mutuura (1971), and Common (1973) have added signifi- cantly to our present meager knowledge of the aculeate Lepidoptera. NUMBER 210 Review of Morphology and Classification The first example of a member of this family was discovered in the Khasi Hills of Assam, India, by a native collector. The specimen was sent to Edward Meyrick, who described it under the name Neopseustis calliglauca. Originally, Meyrick (1909, 1912) included the species in his heterogeneous concept of the Micropterygidae, subfamily Erio- cranianae [sic]. Martin Hering (1925), in describ- ing a second species of Neopseustis from Taiwan, recognized the uniqueness of the moths and pro- posed the family Neopseustidae. More recently, the Neopseustidae have been con- sidered as members of the suborder Dacnonypha and have been usually associated with the Erio- craniidae in the Eriocranioidea (Bourgogne, 1949; Common, 1970). The Neopseustidae and Mnes- archaeidae were considered by Kristensen (1968) to be monophyletic sister groups and to represent the most advanced dacnonyphous families. Mutuura (1972) stressed the relationship of Neopseustidae to the New Zealand Mnesarchaeidae (largely on the basis of the male genitalia) and to the South African Prototheoridae (on the basis of venation and male genitalia). Dugdale (1974), basing his interpretation on data provided by Mutuura (1972), tentatively placed the Neopseustidae with the Hepialioidea in the exoporian Ditrysia. Results of the present study have revealed the relative uniqueness of this family to such an extent that it becomes necessary to propose the new superfamily, Neopseustoidea. Because of the group's unusual morphology, coincident with the apparent extinction of annectant forms, it is now difficult to determine its relative position within the lepidopteran hierarchy. To date, numerous attempts have been made to subdivide the Lepi- doptera into various suborders, and most of these have presented almost as many queries as they have solved. As our basic knowledge of the aculeate Lepidoptera increases, various refinements have been added to the current subordinal system; how- ever, there is still a great need for additional infor- mation and improvements. Under the current sys- tem (as most recently defined by Dugdale, 1974), the Neopseustoidea most appropriately fall within the Dacnonypha. They are, however, only dis- tantly related to the Eriocranioidea, and, with some character systems, show as close a relationship to members of other suborders (e.g., Nepticu- loidea). Thus, in some features, the Neopseustoidea appear to occupy an intermediate position between the Dacnonypha and Monotrysia and, like the Nep- ticuloidea, as pointed out by Dugdale (1974:138), obscure the separation of these two suborders. Pre- viously postulated close affinities of the Neopseu- stoidea with the Hepialoidea and Mnesarchaeoidea are not supported by the comparative morphology of the female abdomen and reproductive system. Similarities in wing structure, however, do suggest some relationship between these groups. The uniqueness of the female reproductive system of the Neopseustoidea actually suggests a new sub- ordinal status for the group; but such a decision should await further studies of the other aculeate families, utilizing as many character systems as possible. Hopefully, it will eventually be possible to consider the morphology of the immature stages more in such a classification, as soon as those forms have been better collected and studied. The general morphology of the head, as reviewed by Kristensen (1968), displays a mixture of primi- tive and specialized features. Among the more primitive are the elongate, five segmented maxillary palpi, a relatively large (though fused) labrum, and well developed frontal chaetosemata. The lat- ter structures are usually present in the Neopseus- tidae as a pair of swollen areas on the frons beneath the antennal sockets, with another, often smaller pair situated above the antennae across the vertex. The chaetosemata of Lepidoptera were first described by Jordan (1923) and later studied fur- ther by Eltringham (1925), who demonstrated their nerve supply. Most likely these sensory structures are homologous to the "warts" of Trichoptera. In the Neopseustidae, the chaetosemata have been referred to as Eltringham's organs by Kristensen (1968) and Mutuura (1971). Among the characters of the head that may be considered as derived or modified are the loss of ocelli, the loss of the sensory pits (organs of vom Rath) from the apical segment of the labial palpi, the reduction of the mandibles, and the speciali- zation of the galeae. The head is hypognathous and broader than long. The compound eyes are unusually enlarged for the Dacnonypha, possessing an interocular index of 1.3-1.5 (see Davis, 1975). Similarly, the antennae are unusually long and many segmented (69-96 segments). A small inter- SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY calary sclerite (Figure 29) is present in Neopseu- stis and Apoplania between the scape and pedicel but absent in Archepiolus. The flagellar segments are usually submoniliform irv outline and bear scattered rows of prominent sockets from which arise slender scales. The length and arrangement of these rows are generally similar to those scat- tered over the legs (Figure 48). The general chae- totaxy and structure of the antennal segments are quite similar among all three genera. In addition to scales, relatively large setal sensillae (sensilla trichodea) also arise from prominent sockets in the antennae. The sockets of the latter are larger and more scattered and not arranged in rows. The sur- faces of these sensillae are densely covered by a mealy, waxlike substance (Figure 22). The func- tion of these sensillae may possibly be chemo- receptive in nature. The mandibles are enlarged but only weakly sclerotized and are nonfunctional. The galeae are very short, their length usually not exceeding the length of the labial palpi. According to Kristensen (1968), the structure of the food groove consists of transverse bars, possibly sub- divided into densely packed lamellae similar to the condition in higher Lepidoptera. The dorsal arms of the tentorium (Figure 26) are well devel- SLM2 PWP3 V- -H SE3 PLS3 - - - - EN3 AC3 AM2 Fl SA CS3 PRM2 K2 FICURE 1.?Neopseustis archiphenax Meyrick, lateral view of thorax. (Scale=0.5 mm.) NUMBER 210 oped in this family and nearly reach to the vertex. These arms are also well developed in the Mnes- archaeidae, Eriocraniidae, and Lophocoronidae but are reduced or absent in the Micropterygidae, Agathiphagidae, and nearly all other families of Lepidoptera. Ehrlich (1958), however, shows these arms secondarily developed in Graphium and Lamproptera of the Papilionidae. The general structure of the thoracic sclerites as viewed laterally (Figure 1) superficially resembles that of the trichopteran Rhyacophila as illustrated by Malicky (1973). The prothorax is reduced and free from the mesothorax. The pronotum is greatly reduced in size, very narrow, and is largely ob- scured dorsally by the mesonotum. The protho- racic furca and spina are well developed and elongate. The tegula is moderately large and nearly oval in full outline (Figure 7). The subtegular sclerite is relatively large and crescent shape. One of the most unusual features of the mesothorax is the fused condition of the furcal apophyses. The mesothoracic furcae of Apoplania and Neopseustis are very similar in structure with their apophyses extending medially and fusing, thus forming an internal bridge (Figures 3, 9). In several other families of primitive Lepidoptera examined (e.g., Micropterygidae, Eriocraniidae, Hepialidae, Pa- laeosetidae and Nepticulidae), these apophyses are either not well developed, or if developed, then widely separated. The metathoracic furcae of Neopseustis and Apoplania differ significantly in structure (Figures 5-6). In Apoplania the base of the furca is second- arily joined ventrally to the internal fold of the basisternum, thereby creating a lumen (Figure 6) immediately ventrad to the furca. In Neopseustis, the base of the furca is relatively reduced in length and does not extend to connect secondarily with the basisternum. The coxae of all legs are relatively long and slender in the Neopseustidae. An epiphysis is typically present in all species, but this becomes extremely reduced in Apoplania (Figure 48). The mesothoracic and metathoracic tibiae possess one and two pairs, respectively, of elongate, well- developed spurs. The terminal tarsal segment (Fig- ure 52) is typically lepidopterous with a median, setiform empodium; a broad, triangular arolium; and a pair of densely setose pulvilli. The tarsal claws are simple, with no secondary specializations. The wings are broad and usually thinly covered by slender scales (Figure 16) except for a sparse scattering of broad silvery scales on the forewings. Most Neopseustidae, thus, superficially resemble certain Neuroptera. Archepiolus is unusual in hav- ing the forewings clothed in predominantly dark, broad scales. The ultrastructure of the wing scales in this family is unusual. In Neopseustis calliglauca (Figures 38-39), the intervenal transverse striae are frequently interrupted by relatively large and often elongate lacunae. The lacunae are unique in being lined with a very distinct, thickened border. The wing scales of Apoplania are similar in general structure, except the lacunae are smaller (at least in the scales of the hindwing) and not as well defined. The scale structure of Neopseustidae in being lacuneate, thus differs significantly from the nonperforated type found in the Zeugloptera, Dacnonypha (Figure 46), and most Monotrysia. Instead, it bears more resemblance to the multi- perforated (lacuneate) type of the Hepialoidea (Figure 43), Mnesarchaeoidea, and many endopo- rian Ditrysia. The microtrichiae (Figure 40) are dense and are randomly scattered over both sur- faces of all wings. Although the wing venation of the Neopseustidae is of a very archaic pattern, certain specializations are to be noted. Archepiolus possesses the most generalized wing structure in the family. In this regard it resembles the Hepialoidea, particularly the Paleosetidae, in possessing rather broad wings with a branched subcostal vein in the forewing and a five branched radial system in both wings. Although Hering (1925) believed a jugum to be absent, this structure is present in all three genera. The venations of Apoplania and Neopseustis are quite similar, and both show such specializations as the fusion of R2 + 3 and the development of an anal "pocket" in the forewing. The latter is best developed in Neopseustis, where it is formed by the dorsoanterior bending of the base of the first anal vein over the base of the posterior cubital vein (CuP). This has caused the base of the wing to be pushed up and forward (Figure 36), resulting in a corresponding pocket to form underneath. Hering (1925) was greatly impressed with the sig- nificance of the "axillar-Wurzelschlinge" and largely on this basis, as well as on the supposed "absence" of a jugum, removed Neopseustis from the Micropterygidae and recognized its current tri?AM: - - B H 10 J_ FIGURES 2-10.?Thoracic morphology: 2, Apoplania chilensis, new species, caudal view of pro- thorax; 3, A. chilensis, new species, caudal view of mesothorax; 4, A. chilensis, new species, cephalic view of metathorax; 5, Neopseustis archiphenax Meyrick, lateral view of metathoracic furca; 6, A. chilensis, new species, lateral view of metathoracic furca; 7, N. archiphenax Meyrick, lateral view of tcgula; 8, N. archiphenax Meyrick, caudal view of prothorax; 9, AT. archiphenax Meyrick, caudal view of mesothorax; 10, N. archiphenax Meyrick, cephalic view of metathorax. (Scale=0.5 mm.) NUMBER 210 status as a distinct family. Hering proposed the function of the anal pocket as a different type of wing coupling mechanism that had developed to replace the function of the "missing" jugum. How- ever, examination of spread or partially spread specimens (i.e., in a condition somewhat similar to flight) shows the costal border of the hindwing rather remote from the pocket. It appears more likely that the anal pocket and the extended anal groove formed by the elevation of A l + 2 would be of more use when the insect is resting. At least in unspread specimens, the costal margin of the hindwing lies closely approximate to the anal pocket and groove of the forewing. Consequently, the groove and pocket may be a modification to assist Neopsenstis in closing its wings at rest and perhaps evolved in conjunction with a noticeable broadening of the wing base. In connection with a discussion involving wing venation, an important fact regarding the correct terminology of the cubital system in Lepidoptera perhaps should be emphasized. American entomol- ogists in particular have persisted in following Comstock's (1918) terminology by referring to the second primary branch of the cubitus (CuP) in Lepidoptera as the first anal vein (1A). The cor- rect terminology for the cubital system has been most recently reviewed by Martynova (1960) and has been followed in the present paper. Marty- nova's belief that no more than two anal veins exist in Lepidoptera, however, should be critically examined as several genera appear to exhibit a small third anal vein (e.g., in the jugal fold of Neopsenstis, and in several genera illustrated by Common, 1970). The ventral articulation between the metathorax and abdomen is generally similar to that of other primitive families in that no sternal processes pro- trude from the second abdominal sternite (Figure 54). Remnants of the first sternite are believed present in the form of a narrow, transverse, median sclerite weakly connected to the anterior margin of the second sternite by a median, furcate bridge. The first sternite extends laterally as a very slender sclerite which joins the anterolateral angle of the second sternite. The lateral margin of the second sternite is thickened somewhat to form an internal rod, which, however, is entirely fused to the ster- nite. This thickening may represent the precursor of the sternal processes so prominent in the Tin- eoidea (Brock, 1971). Within certain genera of the aculeate Lepidop- tera, variously modified, paired organs appear on the fourth and/or fifth sternites of the male and female abdomen. These structures were first studied in the Lepidoptera by Philpott (1925), who regarded them questionably as scent organs. In Table 1, I have summarized their occurrence among several representative genera. In addition to the Lepidop- tera, apparently homologous structures also occur in both sexes on the fifth sternite of certain primi- tive Trichoptera (Malicky, 1973). Philpott (1925) specifically mentions a well-developed pair of fifth sternal glands for a New Zealand philopotamid, Hydrobiosella stenocera Tillyard. Within the Neopseustidae no such organs appear on the fifth sternite of either sex; however, paired, circular, membranous areas are present on the fourth sternite of the female (Figure 57). As noted in Table 1, only two other families, the Eriocra- niidae and Nepticulidae, are known to possess similar structures on the fourth sternite. These structures are apparently synplesiomorphic char- acters that demonstrate yet another distant genet- ical link between these three otherwise very different groups. It should be noted that the erio- cranioid genus Acanthopteroctetes lacks any sternal organs, which may be another justification for removing this genus from the Eriocraniidae; how- ever, Kristensen has observed (pers. comm.) that the glands opening on the fifth sternum are sometimes lost in one or both sexes of certain Eriocrania. The sternal organs of the fourth segment in both the Neopseustidae and Eriocraniidae appear as thin, membranous "windows" in the otherwise pig- mented integument. They are usually spaced rather widely apart and near the caudal margin of the segment. In the Nepticulidae these structures are situated close together, reduced in size and possibly vestigial, and may be absent in some species. In contrast to the obviously glandular nature (i.e., with ducted glands) of those organs arising from the fifth sternite of certain genera, these structures of the fourth sternite exhibit no external openings or internal ducts. Their function may possibly be chemosensory in nature as compared to the appar- ent chemical production and secretory function of 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY the fifth sternal glands. Kristensen (pers. comm.), however, has recently found that the unpigmented patches of the fourth sternum in Eriocrania merely demarcate the areas to which the globular reser- voirs of the fifth sternal glands are attached. Kristensen (1972) has further suggested that the latter organs may serve a defensive purpose. Table 1.?Occurrence of sternal abdominal glands among aculeate Lepidoptera Species examined Micropterygidae Micropteryx aruncella Scopoli Epimartyria auricrinella Walsinqham Sabatinca inconqruella Walker Agathiphagidae Agathiphaga vitiensis Oumbleton Eriocraniidae Acanthopteroctetes bimaculata Davis Dyseriocrania auricyanea (Walsinqham) Eriocraniella aurosparsella (Walsinqham) Lophocoronidae Lophocorona pediasia Common Neopseustidae Archepiolus schmidi Mutuura Neopseustis meyricki Herinq Apoplania chilensis, new species Hepialidae Hepialus sequoiolus Behrens Paleosetidae Oqyqioses eurata Issiki & Strinqer Mnesarchaeidae Mnesarchaea loxoscia Meyrick Nepticulidae Nepticula variella Braun Opostegidae Opostega alboqaleriella Clemens Incurvariidae Incurvaria russatella Clemens Abdominal segments 4th 5th 5th 9 9 urn; 46, Eriocraniella aurosparsella (Walsingham), Eriocraniidae, detail of a ventral scale from discal cell of forewing, X 7000, scale = 1.7 u,m. Leg structure: 47, Xeopseustis meyricki Hering, epiphysis; 48, Apopla7iia chilensis, new species, epiphysis. NUMBER 210 35 49 50 TC E M - J_ FIGURES 49-52.?Leg structure: 49, Archepioltis schmidi Mutuura, scale = 0.5 mm; 30, Meopseu- stis meyricki Hering, scalc = 0.5 mm; 51, Apoplania chilensis, new species, scalc = 0.5 mm; 52, Neopseustis meyricki Hering, terminal tarsal segment of metathoracic leg, scale = 0.1 mm. 36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FIGURES 53-58.?Abdominal structure: 53, Neopseustis archiphenax Meyrick, first and second sternites; 54, Apoplania chilensis, new species, first and second sternites; 55, N. archiphenax Meyrick, sternal process of seventh (7) and eighth (8) segments of male; 56, Archepiolus schmidi Mutuura, sternal process of seventh segment of male; 57, Apoplania chilensis, new species, sensory organs of fourth sternite of female; 58, Eriocraniella aurosparsella (Walsingham), sen- sory organs of fourth (4) and fifth (5) sternites. NUMBER 210 37 61 FIGURES 59-63.?Archepiolus schmidi Mutuura, male genitalia, scale = 0.5 mm: 59, ventral view; 60, lateral view; 61, aedeagus, ventral view; 62, anellus-juxta, ventral view; 63, aedeagus, lateral view. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY PE 64 T a 65 68 FIGURES 64-68.?Neopseustis cdlliglauca Meyrick, male genitalia, scale = 0.5 mm: 64, ventral view; 65, lateral view; 66, anellus-juxta-parameres, lateral view; 67, ventral view of Figure 66; 68, ductus ejaculatorius-parameres, lateral view. NUMBER 210 39 71 69 T _L 70 73 FICURES 69-73.?Neopseustis bicortnita, new species, male genitalia, scale = 0.5 mm: 69, ventral view; 70, lateral view; 71, anellus-juxta-parameres, lateral view; 72, ventral view of Figure 71; 73, ductus ejaculatorius-parameres, lateral view. 40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 74 ' i_ 77 75 78 FIGURES 74-78.?Neopseustis meyricki Hering, male genitalia, scale = 05 mm: 74, ventral view; 75, lateral view; 76, anellus-juxta-parameres, lateral view; 77, ventral view of Figure 76; 78, ductus ejaculatorius-parameres, lateral view. NUMBER 210 81 T 79 LPA EHF 82 FIGURES 79-83.?Neopseustis sinensis, new species, male genitalia, scale = 0.5 mm: 79, ventral view; 80, lateral view; 81, anellus-juxta-parameres, lateral view; 82, ventral view of Figure 81; 83, ductus cjaculatorius-parameres, lateral view. 84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY - - - A - - LPA T 85 86 FIGURES 84-87.?Apoplania chilensis, new species, male genitalia, scale = 0.5 mm: 84, ventral view; 85, caudal view of aedeagus; 86, lateral view; 87, aedeagus and ductus ejaculatorius, lateral view. NUMBER 210 43 8S 8S 90 - - - P A CO V 91 FIGURES 88-91.?Neopseustis meyricki Hering, female reproductive system, scale = 0.5 mm: 88, external gcnitalia, lateral view; 89, ventral view of Figure 88; 90, median sagittal view of repro- ductive system; 91, ventral view of Figure 90. 44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY FIGURES 92-95.?Female genitalia: 92, Neopseustis meyricki Hering, ventral view; 93, N. mey- ricki Hering, apex of ovipositor, ventral view, X 200; 94, Apoplania chilensis, new species, ventral view; 95, A. chilensis, new species, apex of ovipositor, ventral view, X 210. NUMBER 210 45 T EC FIGURES 96-98.?Apoplania chilensis, new species, female genitalia: 96, ventral view, scale = 0.5 mm; 97, spermatheca, scale = 0.2 mm; 98, seventh abdominal sternite, ventral view. 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