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
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 21 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). Smithsonian 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. sinensis) are described. The discovery of Apoplania chilensis marks the first record of

this family from the Western Hemisphere. Distribution maps and keys are provided 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 superfamily (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 primitive 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 regarding 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 Museum of Natural History, Smithsonian Institution, Washington, 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 classification of its components. Similar studies are currently 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 information 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, Binghamton, N.Y.; Dr. John Dugdale, Dept. of Scientific and Industrial Research, Nelson, New Zealand;

Prof. Syuti Issiki, formerly of the University of

Osaka, Osaka, Japan; Dr. N. P. Kristensen, Universitetets Zoologiske Museum, Copenhagen, Denmark; Dr. Tosio Kumata, Hokkaido University,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sopporo, Japan; Dr. M. S. Mani, St. John's College, Agra, India; and Dr. Akira Mutuura, Entomology Research Institute, Ottawa, Canada. I am

especially grateful to Mr. Luis Pena G., Universidad 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 Department of Entomology, Smithsonian Institution: Mr.

Lawrence Druckenbrod, Ms. Elsie Froeschner (retired), 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 collections listed below for allowing me to examine

specimens under their care.

BMNH British Museum (Natural History), London, England

CNC Canadian National Collections, Entomology Research 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 catalog 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 neopseustids 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 corresponds 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 AssamTaiwan distributional pattern, in addition to having endemic representatives in Australia. The

limited capture data thus far indicate the Neopseutidae 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, however, 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, particularly through the Assam gateway.

Understanding the present distribution of not

only the Neopseustidae but of all families of

primitive Lepidoptera (i.e., Zeugloptera, Dacnonypha, 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 fragmented larval head capsule embedded in Cretaceous 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 approximate occurrence of certain continental movements

or land-bridge connections become better documented, 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 principal 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 Lepidoptera, are ancient enough to have had their

present distributions seriously affected by continental 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 morphology as well as biogeography, is beginning to

appear which suggests that at least two major

groups of Lepidoptera, represented by the presentday Micropterygidae and Hepialidae, arose early in

the history of the Order. Both of these groups are

considered among the most generalized Lepidoptera existing today, and both demonstrate a broad,

bipolar distribution suggesting a widespread dispersal 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 existence 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 NearcticPalearctic and the southern Ethiopian-Neotropical

regions.

The Micropterygidae have generally been considered 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 considered 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 different set of characters such as the relative development 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 represented 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. Possessing a number of apomorphic characters (i.e.,

reduced epiphysis, reduced anal pocket in the forewing, and heavily sclerotized aedeagus), Apoplania

chilensis, new species, represents an ancient derivative from a common stock shared with Neopseustis.

Just how ancient was this divergence is largely conjectural at present, as pointed out previously. However, for many of the same reasons summarized by

Hennig (1966a) for certain Diptera, the minimum 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 America was isolated from North America for most of

the Tertiary (Darlington, 1957), one would believe

it improbable for the Neopseustidae to have migrated 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 continents than for the southern ones.

Before general hypotheses such as those just discussed 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 example of the latter, recent field discoveries as reported

in this paper and by Dumbleton (1952), Mutuura

(1971), and Common (1973) have added significantly 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 Eriocranianae [sic]. Martin Hering (1925), in describing a second species of Neopseustis from Taiwan,

recognized the uniqueness of the moths and proposed the family Neopseustidae.

More recently, the Neopseustidae have been considered as members of the suborder Dacnonypha

and have been usually associated with the Eriocraniidae in the Eriocranioidea (Bourgogne, 1949;

Common, 1970). The Neopseustidae and Mnesarchaeidae 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 Lepidoptera 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; however, there is still a great need for additional information and improvements. Under the current system (as most recently defined by Dugdale, 1974),

the Neopseustoidea most appropriately fall within

the Dacnonypha. They are, however, only distantly related to the Eriocranioidea, and, with

some character systems, show as close a relationship

to members of other suborders (e.g., Nepticuloidea). Thus, in some features, the Neopseustoidea

appear to occupy an intermediate position between

the Dacnonypha and Monotrysia and, like the Nepticuloidea, as pointed out by Dugdale (1974:138),

obscure the separation of these two suborders. Previously postulated close affinities of the Neopseustoidea 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 subordinal 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 primitive 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 latter structures are usually present in the Neopseustidae 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 further 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 specialization 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 Neopseustis 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 scattered over the legs (Figure 48). The general chaetotaxy 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 surfaces of these sensillae are densely covered by a

mealy, waxlike substance (Figure 22). The function of these sensillae may possibly be chemoreceptive 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 subdivided into densely packed lamellae similar to

the condition in higher Lepidoptera. The dorsal

arms of the tentorium (Figure 26) are well develSLM2

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 Mnesarchaeidae, 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 obscured dorsally by the mesonotum. The prothoracic 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, Palaeosetidae 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 secondarily 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, welldeveloped spurs. The terminal tarsal segment (Figure 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 having 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 multiperforated (lacuneate) type of the Hepialoidea

(Figure 43), Mnesarchaeoidea, and many endoporian Ditrysia. The microtrichiae (Figure 40) are

dense and are randomly scattered over both surfaces 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 significance 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 prothorax; 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. However, 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 Al+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 entomologists 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 correct terminology for the cubital system has been

most recently reviewed by Martynova (1960) and

has been followed in the present paper. Martynova'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 protrude 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 sternite. This thickening may represent the precursor

of the sternal processes so prominent in the Tineoidea (Brock, 1971).

Within certain genera of the aculeate Lepidoptera, 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 Lepidoptera, apparently homologous structures also occur

in both sexes on the fifth sternite of certain primitive 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 Eriocraniidae and Nepticulidae, are known to possess

similar structures on the fourth sternite. These

structures are apparently synplesiomorphic characters that demonstrate yet another distant genetical link between these three otherwise very

different groups. It should be noted that the eriocranioid genus Acanthopteroctetes lacks any sternal

organs, which may be another justification for

removing this genus from the Eriocraniidae; however, 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 pigmented 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 apparent 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 reservoirs 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 <s

. . .

- + +

+ +

+

_

+ + +

+ + +

? ? +

? ?

+

+

-

-

+

-

-

The seventh and eighth abdominal sternites of

most species of Neopseustidae also display some

degree of specialization. For example, the males of

Neopseustis and Archepiolus possess a small, median, sternal lobe or process directed caudally from

the seventh sternite (Figure 55). This peculiar

process is but one of several synapomorphic characters that dictates the inclusion of Archepiolus

within the Neopseustidae. The sternal process is

absent in Apoplania, although a shallow, lightly

sclerotized, midventral depression is visible. In

addition, the males of most Neopseustis possess a

much broader median lobe near the anterior margin of the eighth sternite (Figure 55). This lobe is

very low in profile and is densely covered with

short, peglike spines. Both sternal processes are

represented by similar, homologous structures in at

least the females of Neopseustis meyricki. In the

latter, however, the eighth sternal lobe (and

spines) has moved even further anteriorly and is

contiguous with the caudal margin of the seventh

sternite. A similar, heavily spinose lobe exists in

the female of Apoplania chilensis. The sternal

spine of the seventh segment must serve a similar

function in both males and females (e.g., assisting

to support the body while resting in an upright

vertical position). The spines of the eighth segment

in the female, however, are believed to be used

primarily during ovipositioning.

The eighth abdominal spiracle is present and

well developed, in contrast to the Zeugloptera and

some Monotrysia where it is lacking (Dugdale,

1974).

The male genitalia of Neopseustidae is complex

to the extent that its corresponding structures are

difficult to homologize with those of other families.

In the terminology I have followed, I have

attempted to use standard terms familiar to lepidopterists even though in some cases their homologies are uncertain. My selection of these terms was

based primarily on their relative position as defined

by Klots (1970). In the only other comparable

study of the male neopseustid genitalia, Mutuura

(1971) partially followed terminology sometimes

used by trichopterists (Nielsen, 1970). Consequently, that author uses the term dorsal processes

for what I refer to as the uncus, and lateroventral

processes for what I collectively term the valvae.

The alimentary canal terminates near the center

of a largely membranous pad of tissue, immediately
NUMBER 210 11

ventrad and slightly caudad of the usually bilobed

uncus (ninth tergite). This typically inflated pouch

probably represents the tenth, or anal, segment.

The entire structure may be somewhat swollen

and protrusive, or it may be partially retracted

into the body beneath the tegumen. On each side

of the anal segment is situated a pair of densely

setose pads referred to as socii. The setae of the

socii are interesting in arising from prominent,

tubercular follicles. A median, unpaired, and frequently uncinate gnathos lies immediately ventrad

to the anal tube. Usually present at the lateroposterior margins of the tegumen is a pair of

broad, lobelike processes, or tegumenal lobes (TL).

Immediately ventrad to these lobes are the valvae

which are usually rigidly fused to the tegumen and

vinculum. The vinculum is superficially similar to

that found in a few other primitive families (e.g.,

Eriocraniidae and Nepticulidae) in possessing a

deeply excavate or concave cephalic margin.

Located in or arising from the diaphragma are

a number of highly modified structures of uncertain homologies. In general, these structures are

arranged into two more or less distinct areas: one

group situated primarily dorsad and laterad to the

aedeagus or gonopore; and a second, usually single

structure entirely ventrad. The entire assemblage

is rigidly joined to the center of the vinculum by

means of a transverse bar (TB). Mutuura has

referred to the dorsolateral plate and its appendages as the paramera. In contrast, I have termed

the dorsolateral sclerotizations as the anellus. The

most prominent features of this complex, the bilateral, armlike processes (LPA), are prominent in all

three genera and are very useful in distinguishing

species. In Neopseustis the anellus entirely surrounds the gonopore and is fused midventrally to

the vinculum by means of the transverse bar. The

ventral sclerotization in the diaphragma has been

termed the juxta. It may be relatively broad

and flat (e.g., in Apoplania and Archepiolus)

or very compressed and spatulate as observed in

Neopseustis.

An aedeagus is present in Apoplania and Archepiolus but absent in Neopseustis. In Apoplania

the aedeagus is highly modified with prominent

exophallic spines arranged in a rigid spiral around

the apex. The vesica and possibly part of the

ejaculatory duct are also modified and sclerotized.

The comparable situation in Neopseustis is a curious one with the ejaculatory duct terminating in

a broad cavity surrounded by the anellus. An

eversible vesica as well as any clear indication of

an aedeagus appear absent. Instead, from the ventral margin of the ejaculatory opening, or gonopore, arises a pair of long, free, digitate processes

(Figure 82) which, because of their relative position, have been termed parameres (Pa). Unfortunately, my usage of the term paramere does not

agree with that of Snodgrass (1957), nor of

Mutuura (1971). Usage of the term by Snodgrass

corresponds to what lepidopterists refer (probably

incorrectly so) as the valve, or valva. It is possible,

of course, that what is referred to in Neopseustis

as parameres may actually represent the adeagus or

structures derived from the aedeagus. If such is

true, however, then its present form and function

has diverged greatly from that of the typical

aedeagus present in nearly all Lepidoptera. The

parameres in Neopseustis are usually completely

separated and contain no external opening (phallotreme) for sperm emission.

In Neopseustis calliglauca and N. bicornuta the

parameres are poorly developed, membranous, and

devoid of setae. In N. archiphenax, N. meyricki,

and N. sinensis they are greatly lengthened with

the basal half encased in a sclerotized sheath and

the apical half densely pubescent on their ventral

surfaces. Because similar structures do not appear

in the two genera possessing a well-developed

aedeagus, the parameres may somehow assist in

sperm or spermatophore transfer in Neopseustis.

The female reproductive system also demonstrates several unusual features, which, in some

instances, were misinterpreted recently by Mutuura

(1972). Externally, one of the most conspicuous

modifications is in the ventral invagination of

the intersegmental region between the seventh and

eighth sternites (Figure 88). The anterior rim of

this pocket (formed largely by the seventh sternite)

is strongly sclerotized and equipped with a median

lobe bearing numerous short blunt spines (discussed previously). The pocket has no internal

opening or ducts leading to it. The development

of this pocket, in conjunction with a piercing type

of ovipositor, suggests that it may have evolved to

better anchor the apex of the abdomen during

oviposition.

The tenth segment is represented by a pair of

short but heavily sclerotized apophyses, which are
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fused at their caudal apex to form a blunt, rasping,

ovipositor. Thus, the method of oviposition may

be remotely similar to that performed by some

members of the Eriocranioidea and Incurvarioidea.

The extreme apex of the ovipositor is coarsely

serrated and is partially defined by a narrow

suture, which may represent the eleventh segment

as suggested by Mutuura (1972).

As shown in Figures 90, 91, the anal aperture is

a narrow slit immediately anterior to the apex of

the ovipositor. The rectum is large and inflated.

Arising from the anterior margin of the anus is

the duct of the undivided colleterial gland. In

most families of Lepidoptera these glands are

paired or occasionally lacking. Partially separating

the apertures of the colleterial gland and spermatheca is a pair of small sclerites representing

the ninth sternite. The spermathecal duct is sinuate

but not coiled as in the Nepticuloidea and most

higher families of Lepidoptera. A spermathecal

vesicle is present as an annular thickening at the

termination of the duct. An internal canal is present within the spermathecal duct, but no evidence

of a spermathecal papilla has been observed. The

spermatheca of Apoplania (Figure 97) differs significantly in structure from that of Neopsenstis. In

Apoplania, the spermathecal utriculus is more

reduced in size, and it possesses a filamentous,

anterior extension (receptacular gland), which,is

apparently absent in Neopseustis. Slightly anterior

to the spermathecal vesicle in Neopseustis arises a

slender duct representing the lateral lagena, as

figured by Dugdale (1974:137) for Nepticula. This

tube actually resembles more closely a homologous

structure shown by Unzicker (1968, therein referred

to as the prespermathecal diverticulum) to occur

in most Trichoptera. Although a lateral lagena is

present in Neopseustis, this structure does not

occur in Apoplania.

Anterior to the spermathecal aperture lies a

very broad vulva which immediately divides to

form a dorsal ductus bursae and ventral common

oviduct. All of the body openings just referred to,

including the anus, open into an abbreviated,

incomplete cloaca similar to that described for

certain Opostega by Dugdale (1974, fig. 21). Laterally the cloaca is partially exposed due to the

flaplike structure of the eighth sternite.

As pointed out by Dugdale (1974) and others,

the relative position of these various terminal apertures and their internal organs in the female is

extremely important toward understanding the

higher classification of the Lepidoptera. For example, the ventral position of the common oviduct

(versus dorsal as figured by Mutuura, 1972) in the

Neopseustidae amply distinguishes this family from

the Hepialoidea and Mnesarchaeoidea. In the latter two groups, the oviporous is separate from and

dorsal to the ostium bursae, thus typifying the

exoporian ditrysian condition first described for

the Hepialidae by Oiticica (1948). The ventral

position of the common oviduct is clearly the

primitive condition as universally demonstrated

by all panorpoid orders (Unzicker, 1968; Dugdale,

1974) except the Lepidoptera. Within the Lepidoptera, an oviduct situated dorsal to the bursa copulatrix exists only in the exoporian and endoporian

Ditrysia. Consequently, utilizing this character system, the Hepialoidea appear more specialized than

the Micropterygidae.

The retention of a vestige of the ninth sternite

separating the anus from the spermathecal aperturevulva may prove significant not only to the higher

classification of the Neopseustidae but to other

families as well. Certainly no such separations are

known to exist in any other members of the Dacnonypha. The presence of this sclerite (with its

resulting demarcations) in Neopseustis somewhat

obscures the definition of a true cloaca. Largely for

this reason, a cloaca is not believed to be present

in this family except in a rather incomplete or

intermediate stage.

In summary then, the Neopseustidae, with their

retention of a jugum, complete absence of a frenulum, rudimentary cloaca, sinuate spermathecal duct,

and ventral oviduct, appear most related to the

suborder Dacnonypha. They are, lvowever, amply

differentiated from the other members of the Dacnonypha and demonstrate certain affinities with

other aculeate groups. For example, except for the

rasping ovipositor, the female reproductive system

more closely resembles that of certain Nepticuloidea, particularly in the presence of a vestigial ninth

sternite. Similarly, the relatively broad wings and

basic pattern of venation and scale structure is

more typical of the Hepialoidea. If nothing else,

the Neopseustidae demonstrate some basic problems in clearly defining the Dacnonypha and

Monotrysia.
NUMBER 210

A

AC3

Am2

Ans

Ar

B2

Bm

Ca

CB

Cdo

CG

CO

CS3

DA

DB

Ea2

EC

Em

En2

Fl

FA

G

IC

IS

J

K2

LCS

LL

LPA

Me

Mn2

O

PA

Pam

Pe

PI S3

Pnl

Prm2

PrS

Ps

PWP3

RG

7S

Sa

Se2

Si

Slm2

Sm2

SP7

Spa

Ss

St

Sua

SV

8T

Ta

Abbreviations

anus

anapleural cleft of metathorax

anepisternum of mesothorax

anellus

arolium

basalare of mesothorax

basisternum

chaetosema

corpus bursae

cardo •

colleterial gland

common oviduct

coxal suture of metathoracic coxa

dorsal arm of tentorium

ductus bursae

eucoxa of mesothoracic coxa

external canal

empodium

epimeron of mesothorax

furca of prothorax

furcal apophysis

gnathos

internal canal

intercalary sclerite

juxta

katepisternum of mesothorax

lateral cervical sclerite (pre-episternum)

lateral lagena

lateroposterior process of anellus

mandible

meron of mesothoracic coxa

ovipositor (tenth abdominal segment)

posterior apophysis

patagium

paramere

pleural suture of metathorax

pleuron of prothorax

pre-episternum of mesothorax

prothoracic spina

pulvillus

pleural wing process of metathorax

receptacular gland

seventh abdominal sternite

spina

subalare of mesothorax

socii

scutellum of mesothorax

scutum of mesothorax

sternal process of seventh abdominal segment

spermatheca

stipes

sensilla trichodea

subtegula

spermathecal vesicle

eighth abdominal tergite

tegula

13

TB

TC

T L

T n

U

Us

V

Va

Ve

Vm

transverse bar

tarsal claw

tegumenal lobes

tegumen

uncus

utriculus

vulva

valva

vesicle

vinculum

NEOPSEUSTIDAE

ADULT.—Small to medium size moths with relatively broad and usually thinly scaled wings; coloration usually of a cryptic or somber pattern,

ranging in shades of pale gray to fuscous, variously

marked with white. Wing expanse ranging from

14 to 27 mm.

Head: Vestiture relatively sparse, hairlike, mostly

rough; occipital scales loosely appressed, directed

forward toward vertex and converging between

antennal sockets; a pair of usually large chaetosemata situated along front rim of eye below antennal sockets, each bearing an erect tuft of sensory

scales; a second, usually smaller and contiguous

pair of chaetosemata present slightly above and

between the sockets; ventral margin of gena extended into an angulate, subgenal process. Antennae

long, approximately equaling length of forewing,

with 69 to 96 segments; segments submoniliform

to subserrate, cupuliform, bearing two major types

of sensory setae in definite sockets (Figures 20-23);

entire shaft (except for a narrow, naked, midventral tract) thinly covered with narrow scales and

setae; scale rows somewhat indefinite due to scattered sockets but appearing under low magnification as one scale row per segment; scape large,

swollen, an intercalary sclerite usually present between scape and pedical (absent in Archepiolns).

Ocelli absent. Compound eyes rounded, enlarged,

interocular index approximately 1.3-1.5; eyes

naked. Labrum triangular. Mandibles present, but

reduced, weakly sclerotized, and nonarticulated.

Maxillary palpi long, folded against head in repose,

5-segmented; the penultimate (fourth) and apical

(fifth) segments the longest and of equal lengths;

galeae relatively short, approximately 0.5 the length

of maxillary palpi. Hypopharynx elongate, tapering to a triangular lobe ventrally. Labial palpi

moderately long, 3-segmented; apical (third) seg-
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ment elongate, equaling length of second. Dorsal

tentorial arms (Figure 26) completely sclerotized,

well developed, nearly reaching to vertex of cranium immediately posterior to antennal insertion.

Thorax: Mesothoracic furca with apophyses connected by their apices at midline (Figure 3). Legs

of moderate length (Figures 49-51); prothoracic

leg with epiphysis relatively long (0.3-0.5 the

length of tibia) or minute (length less than width

of tibia); epiphysis simple, not pectinate; tarsus

equaling length of tibia; mesothoracic leg with one

pair of apical spurs, tarsus equaling tibia; metathoracic leg with a pair of apical spurs and a pair

of spurs arising from distal third or fourth of

tibia; tarsus distinctly shorter than tibia; all paired

spurs of unequal lengths. Forewings relatively

broad, outer margin rounded, either 13 or 14

veined; Sc divided; R2 and 3 usually fused and

stalked or connate with R4+5, but stalked over

half their length and separate from R4+5 in

Archepiolus; R4 and 5 stalked; Ml and 2 usually

stalked (separate in Archepiolus); M3 separate;

base of medius well preserved; m-cu crossvein

present; CuAl and 2 stalked over one-third their

length; CuP weak; 1A and 2A separate at extreme

base but united for most of their length to margin;

3A present at jugal fold; jugum present; a prominent pocket formed ventrally at base of anal veins

in Neopseustis (Figure 36), which is poorly developed in Apoplania and absent in Archepiolus.

Hindwings 11 or 12 veined; Sc simple, undivided;

subcostal crossvein present at base of wing or

absent; R2 and 3 either fused or strongly stalked;

R4 and 5 stalked half their length or more; Ml

and 2 stalked; M3 separate; m-cu crossvein absent,

thus lower half of discal cell open; CuAl and 2

stalked; CuP weak, connected at base to 1A in

Neopseustis and Apoplania but not in Archepiolus.

Both wings with microtrichiae dense and evenly

distributed (Figure 37).

Abdomen: Eighth abdominal spiracle well developed, functional. Seventh sternite of male usually

with a median, connate, spinose process or with a

median, slightly depressed area; eighth sternite of

male sometimes with a median, spinose ridge,

though reduced or absent in some species. Female

with a bilateral pair of oval, membranous pads

near caudal margin of fourth sternite; entire

seventh sternite heavily sclerotized, either with or

without a median, connate process similar to male;

caudal margin of seventh sternite produced into a

low, median lobe bearing numerous short, blunt

spines. Intersegmental area between seventh and

eighth sternites deeply excavated.

Male genitalia: Uncus present, bifurcate, prominently developed in Archepiolus and Apoplania,

very reduced and largely fused to tegumen in

Neopseustis. Socii present, densely setose; setae

arising from prominent, tubercular follicles. Anus

terminating in middle of a usually inflated, membranous pouch which may either be extended or

withdrawn slightly. Gnathos present; typically a

median plate poorly connected to tegumen, and

usually with a median process arising ventrally.

Tegumen a narrow ring dorsally, expanding

slightly laterally and fusing with valvae and vinculum; a bilateral pair of lobelike processes arising

from lateroposterior margin of tegumen, reaching

their maximum development in Neopseustis. Vinculum broadened laterally with a pair of lobes

directed anteriorly; anterior margin of vinculum

usually deeply concave. Valvae short, usually simple but divided in Archepiolus; articulation absent,

with valvae fused to vinculum and tegumen. Diaphragma with two principal, rather complex,

sclerotized areas around ejaculatory opening or

aedeagus; plate (i.e., anellus) dorsal to gonopore

usually bearing a bilateral pair of elongate processes; ventral plate (i.e., juxta) usually broad with

a strongly compressed, median process extending

cephalad in Neopseustis. Aedeagus either absent

(in Neopseustis) and with ejaculatory duct terminating at base of a pair of elongate parameres, or

present (in Archepiolus and Apoplania) and with

parameres absent; endophallic cornuti absent.

Female genitalia: Tenth segment consisting primarily of a large pair of apophyses rigidly united

at caudal apex by a stout bridge; extreme apex

subacute and serrated, modified for rasping. Ninth

tergite at least partially subdivided into two lateral plates or setose pads. Ninth sternite reduced

to two lightly sclerotized plates embedded in dorsal

wall of cloacal chamber, anterior to anus. Eighth

tergite enlarged, hoodlike, covering most of ninth

and tenth segments. Eighth sternite typically a

relatively short but broad plate, possessing either

a truncate or deeply concave caudal margin. All

openings of the reproductive tract and alimentary

canal emptying into a very short, incomplete

cloacal chamber formed dorsally primarily by the
NUMBER 210 15

ninth segment and ventrally by the eighth sternite.

Colleterial gland present, undivided. Spermatheca

relatively simple and undivided; spermathecal duct

slightly sinuate, not coiled. Bursa copulatrix well

developed; ductus bursae usually thickened,

strongly sinuate in Neopseustis; corpus bursae

membranous, signum absent. Common oviduct arising ventrad to bursa copulatrix.

Key to the Genera of Neopseustidae

1. R2 and 3 forked nearly one-half their length in both fore- and hindwings. Wings fully

clothed with relatively broad scales (Figure 12) Archepiolus

R2 and 3 fused throughout their length in both wings. Wings thinly clothed with slender,

hairlike scales (Figures 13-19) 2

2. Epiphysis elongate, over 0.3 the length of foretibia; forewing with fused portion of CuA

approximately one-fifth the length of CuAl (Figure 35); distal end of 1A+2A abruptly

curving to hind margin Neopseustis

Epiphysis minute (Figure 48), less than the diameter of the foretibia in length; forewing

with fused portion of CuA over one-third the length of CuAl (Figure 34); distal end of

1A+2A gradually curving to hind margin Apoplania, new genus

Archepiolus Mutuura

Archepiolus Mutuura, 1971:1129.

TYPE-SPECIES.—Archepiolus schmidi Mutuura,

original designation.

ADULT.—Small, slender-bodied moth, with fully

scaled wings.

Head (Figure 24): Antennae without intercalary

sclerite between scape and pedicel; segments submoniliform along basal and apical one-fifth of

shaft, asymmetrically subserrate along middle

(Figure 25); sensory scales arranged in rows of

usually six sockets. Chaetosemata present on frons

and at vertex but not as swollen as in Neopseustis

and Apoplania. Galeae short, subequal in length

to labial palpus. Maxillary palpi moderately long,

slightly surpassing labial palpi in length, fourth

and fifth segments the longest and approximately

equal in size.

Thorax: Epiphysis 0.5 the length of foretibia

(Figure 49) and reaching to base of tarsus; metathoracic tibia with basal spurs arising from outer

fourth of segment. Forewings (Figure 33) 14-veined;

R2 and 3 stalked approximately 0.5 their total

length, arising separate from R4+5; Ml and 2

separate; discal cell relatively long, extending more

than half the length of wing; base of anal area not

raised, ventral axillary pocket absent; R2 and 3

stalked more than half their total length; crossvein

r-m slanted obliquely outward toward Ml; Ml and

2 separate; M3 arising relatively remote from crossvein r-m.

Abdomen: Ventral lobe of seventh sternite of

male rounded, short, not extending beyond margin

of seventh segment, densely covered by short setae.

Male genitalia: Uncus well developed, deeply

forked. Socii padlike, densely covered with slender

setae, situated on either side of bulbous anal tube.

Tegumenal lobes broad but short. Gnathos well

developed, not uncinate, consisting of an inverted

V-shaped plate from whose apex arises a prominent,

median, sclerotized lobe. Vinculum with anterior

margin deeply concave. Valvae bilobed, deeply

divided to form a relatively narrow, dorsolateral

cucullar lobe and a much broader and more ventral, saccular lobe; ventral margins of sacculi fused

partially together along midline. Juxta broad, platelike. Anellus consisting primarily of a bilateral pair

of rodlike processes. Aedeagus mostly sclerotized.

DISCUSSION.—The recent discovery of this interesting genus has reaffirmed my earlier belief that

the foothills and lower forested elevations of the

Himalayan Chain may harbor an array of primitive Lepidoptera yet to be collected. In general

appearance, Archepiolus does not resemble the

other members of Neopseustidae, particularly in

the darker, broader scaling of its wings. Its relationship to that family, however, is strongly suggested by morphological similarities of the head

and male genitalia. In this regard, the discovery

of Apoplania, an unquestionable member of the

Neopseustidae from Chile, has been significant in

demonstrating the range of morphological extremes

in this family. It should prove very enlightening
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

to examine the female reproductive system of

Archepiolus, whenever that sex is collected. Certainly, any further meaningful discussion on the

the family affinities of this genus must await the

discovery of the female.

As pointed out by Mutuura (1971), the venation

of Archepiolus is among the most primitive of the

Lepidoptera. The radial system of both wings is

very similar with all five branches of R distinct,

thus differing from the 4-branched condition in

Neopseustis and Apoplania. In addition, Archepiolus lacks the basal modification of the forewing

(i.e., the anal pocket) developed in varying degrees

in the two above-mentioned genera.

Archepiolus schmidi Mutuura

FIGURES 12, 24-25, 33, 49, 56, 59-63

Archepiolus schmidi Mutuura, 1971:1129.

ADULT (Figure 12).—Wing expanse: $ , 16 mm.

Head: Rough, scales apparently dark, perhaps

fuscous (not described by Mutuura prior to dissection). Antennae thinly covered with narrow, whitish scales; 94 segmented; segments slightly depressed; sensory scales arranged primarily in one

band around segment, consisting typically of a

longitudinal row of 6 raised sockets. Eyes large;

interocular index approximately 1.4.

Thorax: Dorsum of thorax fuscous. Forewings

rather heavily covered by broad scales with mostly

bidentate apices, fuscous mottled with paler shades

of gray and silvery white; a large, somewhat obscure area of dark fuscous situated midway along

costa and extending approximately halfway across

wing; relatively large, silvery white spots present

near costa at Sc 1 and 2; a series of 3 or 4 irregular

whitish spots continuing around outer margin to

M3; cilia fuscous. Hindwing uniformly pale gray,

thinly scaled; scales mostly narrow with acute

apices, except for broader, bidentate scales near

apex of wing.

Abdomen: Coloration unknown (previously

mounted in balsam), probably fuscous.

Male genitalia (Figures 59-63): Uncus deeply

bifurcate; lobes widely separated, elongate, nearly

equaling valvae in length. Caudal margin of

gnathos serrulate near base on either side, becoming strongly but bluntly serrate on median claviform process. Tegumenal lobes broadly rounded,

reduced in length, less than 0.5 the length of

valvae. Valvae deeply divided; outer (costa and

cucullus) lobe more narrow with evenly rounded

apex; inner lobe (principally sacculus) with ventral margins fused at midline for nearly half their

length, apex irregular, bidentate. Vinculum a narrow ring ventrally, anterior margin deeply concave, expanding laterally and anteriorly to form

elongate and widely separate arms. Caudal half of

juxta a broad, thin plate with caudal margin

excavate; anterior half compressed to a slender,

keellike process similar to Neopseustis. Anellus

with a bilateral pair of elongate, rodlike processes

equaling valve in length. Aedeagus with base

bilobed, slightly flared and sharply set off at an

angle from main axis; main shaft straight, partially

membranous, without cornuti.

TYPE.—Holotype, $ , no. 11629; in the Canadian

National Collection.

TYPE-LOCALITY.—Jhum La, Kameng, Frontier

Division, Assam, India.

DISTRIBUTION (Map 1).—Known only from the

type-locality located southwest of the town of Rupa

near the border of Bhutan.

DISCUSSION.—Unfortunately, only a single male

specimen of this interesting insect is known at

present. Its collector, Dr. F. S. Schmid of Ottawa,

Canada, has personally informed me that the specimen was collected May 13, 1961, at light (gasoline

pressure lantern) near a mountain stream. The site

was reportedly heavily forested with an estimated

elevation of approximately 7000 feet [2154 m].

Neopseustis Meyrick

Neopseustis Meyrick, 1909:436; 1912:3.—Hering, 1925:143-

145.—Fletcher, 1929:147; 1933:83.—Viette, 1946:25.—Dumbleton, 1952:26.—Kristensen, 1968:148.—Mutuura, 1971:

1129—Common, 1973:11, 13, 16.

Neopseustie [sic] Meyrick, 1909:436 [misspelling of Neopseustis].

Nepseustis [sic] Kristensen, 1968:141 [misspelling of Neopseustis].

Formopseustis Matsumura, 1931:1115, new synonym.

TYPE-SPECIES.—Neopseustis calliglauca Meyrick,

1909, by monotypy.

ADULT.—Small to medium size moths with thinly

scaled, semitransparent wings. Wing expanse ap

proximately 15-27 mm.

Head (Figures 26-27): Antennal segments submoniliform, slightly depressed and asymmetrical;
NUMBER 210 17

intercalary sclerite between scape and pedicel present; sensory scales arranged mostly in longitudinal

rows of 4 raised sockets (Figures 21-23) in a somewhat staggered band around distal half of each

segment; microtrichia evenly scattered over enlarged part of segment. Eyes large; interocular

index approximately 1.3-1.4. Chaetosemata well

developed on frons and at vertex of head. Galeae

reduced, extending to base of terminal segment of

maxillary palpus. Maxillary palpi very long, nearly

doubling the length of labial palpi; fourth and

fifth segments the longest and of equal size. Labial

palpi elongate, slender, slightly surpassing length

of galeae.

Thorax: Epiphysis well developed (Figure 47),

approximately 0.3 the length of tibia; metathoracic

tibiae with basal spurs arising from distal fourth

of segment. Forewings (Figure 35) 13 veined; R2

and 3 completely fused, shortly stalked to base of

R4 + 5; Ml and 2 stalked nearly half their total

length; discal cell relatively short, .closed basally

by junction of radial and medial veins, extending

less than 0.5 the length of wing; base of anal area

raised to form a large pocket ventrally; fused portion of CuA approximately 0.2 the length of CuAl;

distal end of Al+2 abruptly curving to hind margin. Hindwings 12 veined; crossvein present at

base of costa; R2 and 3 completely fused; crossvein

r-m slanted obliquely inward toward Ml+2; Ml

and 2 stalked over 0.3 their length; M3 arising

approximate to crossvein r-m.

Abdomen: Relatively short and broad. Caudal

margin of seventh sternite of male with an acute,

setigerous, median lobe (Figure 55); eighth sternite

sometimes with a small median, slightly bilobed

process at caudal margin. Female with a bilateral

pair of membranous disk-shaped pads present near

caudal margin of fourth sternite; seventh sternite

strongly sclerotized, caudal margin curved toward

center producing a median, heavily spinose lobe;

a slender, spinose sternal process arising ventrally

and anterior to median lobe.

Male genitalia: Uncus greatly reduced, apparently fused with tegumen and indicated merely by

a small, bilobed structure at the apex of the ninth

tergite. Tegumen also reduced, largely membranous, represented principally by a thin, dorsal,

lightly sclerotized ring weakly joined at caudal

apex. Tegumenal lobes extended to form a bilateral pair of slender, slightly curved arms projecting

beyond apex of uncus. Gnathos consisting of a thin

median plate with either one or two heavily sclerotized processes projecting ventrally. Socii densely

setose, widely separated on either side of a bulbous

anal tube. Vinculum with anterior margin deeply

concave, produced laterally into a pair of slender,

armlike anterior lobes; length of vincular lobes

approximately 0.8 that of tegumenal lobes. Valvae

broad, relatively short, rigidly fused to vinculum,

narrowly separated ventrally. Anellus consisting of

a sclerotized sheath partially surrounding opening

of ejaculatory duct and slender parameres; typically

with a large, bilateral pair of armlike processes

arising laterally and directed caudally. Juxta elongate, strongly compressed, forming a paddlelike

extension anteriorly from a usually broad, transverse bar rigidly fused ventrally to vinculum.

Aedeagus undeveloped, with ejaculatory duct terminating at base of paired, variously developed

parameres.

Female genitalia: Posterior apophysis relatively

stout, forming a heavily sclerotized, tapering ovipositor; apex of ovipositer coarsely serrate, relatively broad. Ninth tergite partially subdivided into

two lateral plates. Ninth sternite reduced, consisting

of two, lightly sclerotized plates embedded in dorsal wall of cloacal chamber. Eighth tergite enlarged,

hoodlike. Eighth sternite broad, with caudal margin deeply concave. Ductus bursae elongate, extending approximately 0.5 the length of entire bursa

copulatrix, walls heavily thickened, irregular in

outline; corpus bursae membranous. Spermatheca

well developed, extending anteriorly almost to fifth

segment; lateral lagena present, filamentous; receptacular gland absent.

DISCUSSION.—The genus Neopseustis, for many

years represented by only three poorly known

species, is now believed to consist of at least five

species. Additional members of this interesting

genus will surely be discovered after the Microlepidoptera fauna of southeastern Asia has been better

sampled. Unfortunately, the holotype of Neopseustis archiphenax is lacking an abdomen, thereby

prohibiting a positive diagnosis of that species;

however, an additional male specimen, believed

to be conspecific with N. archiphenax, has been

studied.

On the basis of the available material, the genus

appears to fall into two groups. One, characterized

principally by well developed, densely pubescent
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

parameres (Figures 77-82), includes the species

N. archiphenax, N. meyricki, and N. sinensis, new

species. The second group contains the smaller

species, JV. calliglauca and N. bicornutus, which

possess male genitalia with poorly developed, naked

parameres. In both groups the best means for

species distinction is provided by the widely divergent forms of the anellar lobes in the male genitalia. Because the females of only one species (AT.

meyricki) have been collected, it is not known how

great the female genitalia vary between species in

this genus.

Key to the Species of Neopseustis

(based largely on males)

1. Wing expanse exceeding 25 mm Neopseustis archiphenax

Wing expanse less than 25 mm 2

2. Parameres greatly lengthened, curved, extending to apices of tegumenal lobes ( Figure 82) 3

Parameres reduced, indistinct, relatively straight, not surpassing apices of lateral processes of

anellus (Figure 66) 4

3. Lateral processes of anellus thickened; spinose projections, irregular, stout (Figure 82)

Neopseustis sinensis, new species

Lateral processes of anellus slender, each process immediately subdividing to form two, slender, nearly symmetrical branches (Figure 77) Neopseustis meyricki

4. Lateral processes of anellus simple, acute, resembling a pair of inwardly curving horns

(Figure 72) Neopseustis bicomuta, new species

Lateral processes not corneate, broader with rounded apices, divergent (Figure 66)

Neopseustis calliglauca

Neopseustis calliglauca Meyrick

FIGURES 13, 37-39, 64-68

Neopseustis calliglauca Meyrick, 1909:436; 1912:3, 4.—Hering,

1925:146.—Meyrick, 1928:404.—Fletcher, 1929:147; 1933:

83.—Mutuura, 1971:1129.

ADULT (Figure 13).—Wing expanse: $ 15-17.2

mm.

Head: Vertex and frons brownish fuscous. Antennae with 69-72 segments; dorsum of scape and

pedicel brownish fuscous, flagellum light brown;

venter of scape and pedicel stramineous. Maxillary

palpi light brown. Labial palpi brownish fuscous.

Thorax; Dorsum brownish fuscous; venter stramineous. Legs with coxal and femoral segments

stramineous, distal segments mostly brownish fuscous. Forewings with basal two-thirds light brown,

mottled with irregular spots and transverse streaks

of fuscous; a series of dark, transverse bars especially concentrated between bases of medial veins;

distal third of wing whitish, heavily bordered with

light grayish brown along apex and termen; an

oblique, triangular patch of brown at outer third

of costa at Sc2. Hindwings with basal two-thirds

light brownish fuscous; distal third whitish with

broad, terminal band of light grayish brown.

Abdomen: Sparsely clothed with slender, lightbrown scales. Seventh sternite with rather acute,

triangular median lobe; median lobe of eighth

sternite absent.

Male genitalia (Figures 64-68): Gnathos a broad

cordate plate with a pair of compressed, bilateral

processes arising ventrally; distal ends of processes

flared, broadly bifurcate. Posteroventral margin of

valvae extended to form a short, rounded lobe.

Anellus a broad, thin plate dorsally with median

caudal margin folded ventrally; lateral lobes extending to bases of socii, broadly compressed with

rounded apices and a small, serrated ridge situated

laterally near middle. Juxta with a broadly compressed median lobe extending anteriorly slightly

beyond lateral arms of vinculum. Parameres consisting of a pair of slender, unsclerotized, relatively

inconspicuous, digitate processes extending caudally about halfway to apex of anellar lobes.

TYPE.—Holotype, $ ; in the British Museum

(Natural History).

TYPE-LOCALITY.—Khasi Hills, Assam, India.

DISTRIBUTION (Map 1).—Known only from the

Khasi Hills of northeastern India.

MATERIAL EXAMINED.—4 males.

INDIA. Assam: Khasi Hills: 1 $ (holotype), Oct. 1906

(BMNH). Shillong: 1 $, 20 May 1928 (BMNH); Shillong,

5000 ft.: 1^,19 May 1928 (BMNH); 1 $ , May 1964 (USNM).
NUMBER 210 19

DISCUSSION.—This species appears rather closely

related to the following, N. bicornuta, in possessing

relatively reduced, unsclerotized parameres that are

in sharp contrast to those developed in N. sinensis,

new species, and N. meyricki. The two species,

however, can be easily separated by their very different gnathosi, N. calliglauca is the only member

of the genus with a bifid gnathos.

Meyrick determined the sex of the holotype as

a male, although this cannot be confirmed now

due to the loss of the abdomen. He also gave the

wing expanse as 20 mm, although it is actually

closer to 17 mm. There is, however, no serious

question concerning the identity of the holotype,

and it agrees in maculation with the material later

collected by Fletcher. Fletcher's note (1933) relating

to the capture of this species is important in that

it provides us with details of the source of his

Shillong material, "a very restricted area just above

the stream at the top of the Fruit Garden." This

author further reported that the moths were found

during the day resting on leaves and closely resembled birds' droppings.

Neopseustis bicornuta, new species

FIGURES 14, 23, 69-73

ADULT (Figure 14).—Wing expanse: $ , 19 mm.

Head: Vertex and frons brownish fuscous. Antennae brownish, darkest along basal fourth of

shaft; underside of scape stramineous. Maxillary

palpi stramineous. Labial palpi brownish above,

stramineous below.

Thorax: Dorsum brownish fuscous; venter stramineous. Legs uniformly light brown to stramineous. Forewings with basal half predominantly

light brown, mottled with irregular patches of

brownish to dark fuscous and with a sparse scattering of broad, silvery white scales; distal half

primarily whitish, mottled with relatively large,

pale, grayish-brown spots especially along wing

margin from Scl to CuAl; a triangular spot present on costa between Scl and 2; an elongate costal

spot situated between Sc2 and apex, and a third

large spot along termen between Ml and 3. Hindwings with basal half pale brown; outer half whitish with an irregular series of light-brown spots

along outer margin; apical spot between Sc and R4

the largest.

Abdomen: Sparsely clothed with slender, brownish scales. Median process of seventh sternite in

male triangular, acute; median process of eighth

sternite absent.

Male genitalia (Figures 69-73): Gnathos a large,

stout, uncinate process projecting ventrally from a

triangular base between socii. Valvae with ventral

margin extended caudally to form a short, spinelike

process. Lateroposterior lobes of anellus corneate

heavily sclerotized and curved inward toward each

other, extending to base of gnathos. Juxta with a

bilateral pair of curved, basal processes extending

laterally around opening of ejaculatory duct to

anellus and an elongate, broadly compressed median process projecting anteriorally beyond lateral

arms of vinculum. Parameres similar to N. calliglauca, consisting of a pair of slender, unsclerotized,

relatively inconspicuous, digitate processes extending caudally about halfway to apex of anellar lobes.

HOLOTYPE.— $, Mt. Omei, 4400 feet [1351 m],

Szechuan, China, D.C. Graham coll., USNM 72100;

in the National Museum of Natural History, Smithsonian Institution.

DISTRIBUTION (Map 1).—Known only from the

type-locality, Mt. Omei, located in the southwestern

area of Szechuan Province, China.

DISCUSSION.—The discovery of this species, along

with Neoposeustis sinensis, new species, represents

the first report of Neopseustidae from mainland

China. Their occurrence in Szechuan Province is

not surprising, however, since this area is largely

an extension of the main Himalayan Plateau and

general habitat of the genus Neopseustis. The type

locality of N. bicornuta is located approximately

80 miles [50 km] south of Chengtu, near the village

of Omei. The area is quite mountainous, with the

maximum elevation of Mt. Omei listed at 9957 ft.

[3064 m]. All specimens of N. bicornuta and N.

sinensis were collected by the Rev. David C.

Graham, an American missionary who lived in this

area of Western China from 1910 to 1948.

Neopseustis bicornuta may be distinguished most

easily from its supposed nearest relative, N. calliglauca, on the basis of the male genitalia. Whereas

the gnathos of N. bicornuta is a single, uncinate

process, that of N. calliglauca is bifid with flared

apices. The lateral processes of the anellus also

differ with those of N. bicornuta, being more slender and corneate (hence the specific epithet,

bicornuta).
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Neopseustis archiphenax Meyrick

FIGURES 1, 5, 7-10, 15, 21-22, 35, 53, 55

Neopseustis archiphenax Meyrick, 1928:404.—Fletcher, 1933:

83.—Mutuura, 1971:1129.

ADULT (Figure 15).—Wing expanse: $ , 26-27

mm.

Head: Largely devoid of scales (badly rubbed)

exposing dark fuscous integument. Maxillary and

labial palpi similarly sparsely scaled, fuscous.

Thorax: Dorsum mostly naked, dark fuscous,

sparsely covered with slender brownish scales.

Forewings light brown to stramineous, heavily

mottled with various shades of fuscous; basal twothirds of wing dull white to stramineous, with 3

or 4 rather indistinct fuscous spots along costal

cell; a large, slightly curved fuscous spot extending

between M2 and CuAl; two triangular fuscous

spots situated along hind margin at CuAl and

CuA2; a series of smaller, transverse spots scattered over posterior half of wing; a few broad,

silvery white scales scattered somewhat randomly

over basal two-thirds though concentrated along

major veins; outer third of wing whitish, with numerous irregular spots of light brown; a series of

mostly confluent spots bordering apex and termen.

Hindwings considerably paler; basal two-thirds

light grayish brown; outer third whitish with a

nearly continuous, broad row of light grayish

brown spots bordering outer margin from apex to

CuAl.

Abdomen: Thinly covered with slender, hairlike,

brownish scales. Seventh sternite of male with a

prominent median, conical process bearing 6

minute, closely appressed spines; eighth sternite

with low median ridgelike lobe bearing approximately 23 irregular, rather elongate, blunt spines

along its caudal margin.

Male genitalia: Gnathos a slender, uncinate

process arising between socii and projecting ventrally. Valvae with a small, slender, bilateral pair

of clavate lobes arising from base of sacculus and

nearly contiguous along midline. Lateroposterior

lobes of anellus bilaterally symmetrical, deeply

forked near base, giving rise to two elongate, asymmetrical processes; inner branch more stout,

roughly serrated along entire length of outer (lateral) edge. Parameres greatly lengthened, surpassing apex of uncus, densely pubescent over ventral

half.

TYPE.—Holotype, £ ; in the British Museum

(Natural History).

TYPE-LOCALITY.—Kangtang [?], upper Burma.

DISTRIBUTION (Map 1).—Presently known from

upper Burma and Szechuan Province in southwestern China.

MATERIAL EXAMINED.—2 males.

BURMA. Kangtang: 1 $ (holotype), coll E. A. Swann

(BMNH).

CHINA. Szechuan Province: Sungpan: 1 $, July, D. C.

Graham coll. (USNM).

DISCUSSION.—An examination of the holotype of

this species unfortunately revealed that the specimen at one time had been partially damaged and

the abdomen lost. A search in the drawer containing the holotype did not reveal the missing abdomen. Consequently, it is now difficult to establish

beyond question the identity of the species. Figure

15 was developed from an old negative in the British Museum's files, obviously taken prior to the

loss of the abdomen.

A male specimen in the collection of the Smithsonian Institution from Sungpan, China, closely

agrees with the holotype in size and maculation.

Because the outer marginal band of the hindwings

is not as heavily pigmented as the holotype, it

could represent yet another undescribed species of

Neopseustis. However, in lieu of better preserved

material from the type locality and because of the

above similarities, this specimen has been considered conspecific with N. archiphenax. Consequently, all remarks concerning the genitalic

characters of N. archiphenax are based upon the

specimen from China.

The species most resembling N. archiphenax

appears to be N. meyricki. In the male genitalia of

both species the lateroposterior lobes of the anellus

are deeply divided to form two slender branches;

however, in N. archiphenax the two branches are

heavier with the inner branch coarsely serrated

along one edge, as compared to the completely

smooth condition in the Formosan species. The

difference in wing expanse between the two is

probably significant, although little is known regarding the size variation in N. archiphenax.

Meyrick (1928) lists the type locality for the

species as Kangtang, in upper Burma. A search for

this locality, including a query to the Burmese

Embassy in Washington, D.C., has thus far proved
NUMBER 210 21

futile. However, the names of two towns whose

spellings approximate Meyrick's rendering have

been located in the southern Shan State near the

Laotian border. The towns, Keng Tung and Keng

Tawng (or Kengtawng) are situated approximately

125 air miles [78 km] apart, and one may actually

represent the type locality of N. archiphenax. Thus,

in Map 1, the occurrence of this species has been

provisionally indicated in the Shan State.

Neopseustis meyricki Hering

FIGURES 16, 36, 47, 50, 52, 74-78, 88-91, 92-93

Neopseustis meyricki Hering, 1925:146-147.—Kristensen, 1968:

137, 141-147.—Mutuura, 1971:1129, 1133; 1972:1065.

Formopseustis takamukui Matsumura, 1931:1115, new synonym.

ADULT (Figure 6).—Wing expanse: $ , 20 mm;

9, 20-22 mm.

Head: Vertex and frons brownish fuscous. Antennae brownish fuscous; underside of scape pale

brown. Maxillary palpi pale brown to stramineous.

Labial palpi brownish fuscous, becoming lighter,

more stramineous on underside toward base (first

segment).

Thorax: Dorsum brownish fuscous with a small,

bilateral tuft of whitish scales arising anteriorly

near margins of tegulae; venter stramineous, without conspicuous banding. Forewings with basal

two-thirds predominantly light brown, mottled with

irregular patches of darker brown to dark fuscous

and with a sparse scattering of broad, silvery white

scales, particularly along veins; distal third primarily whitish, mottled with rather large spots of

brownish fuscous bordering outer margin; largest

and most conspicuous spot extending along costa

from Sc2 almost to apex. Hindwings with basal

two-thirds pale grayish brown; distal third primarily whitish with series of partially contiguous

spots of pale grayish brown bordering wing margin

from Rl to CuAl.

Abdomen: Sparsely clothed with slender, brownish scales. Male with a relatively long (3 times its

width in length) stout, median spinelike process

arising from caudal margin of seventh sternite;

apex of process minutely bidentate; caudal margin

of eighth sternite with a median, low, broad, indistinctly bilobed process bearing a single transverse

row of coarse, irregular teeth. Female with median

spine of seventh sternite stout, acute, and serrulate

along its ventral margin; caudal margin of seventh

sternite with a median, rugose patch of 40-60

blunt teeth; bilateral pair of pads on fourth sternite nearly 0.5 the length of segment in diameter.

Male genitalia (Figures 74-78): Gnathos a slender, uncinate process arising between socii and projecting ventrally. Valvae with a small, slender,

bilateral pair of clavate lobes arising from sacculus

and nearly contiguous along midline. Lateroposterior lobes of anellus rather symmetrical in outline, deeply forked near base, giving rise to two

slender processes (Figure 77); a relatively short,

bilateral pair of processes arising nearly contiguous

to one another from median posterior margin of

anellus; apices of these processes not acute, instead,

irregularly serrate. Parameres greatly lengthened,

surpassing apex of uncus, densely pubescent over

ventral half, slightly enlarged near acute apices.

Female genitalia (Figures 88-91): Apex of ovipositor broad, heavily sclerotized, with a triangular,

tricuspidate, and bluntly rounded median lobe

and a bilateral pair of more acute lobes. Lateral

margin of ninth tergite produced into a bilateral

pair of bilobed, somewhat scalloped, ridgelike

processes. Caudal margin of eighth sternite deeply

excavated, with lateral margins extending posteriorly as rather broad, triangular lobes. Ductus

bursae relatively long, strongly curved in a sagittal

plane so as to appear sigmoid in outline viewed

laterally; walls heavily thickened. Corpus bursae

completely membranous, spherical in outline.

TYPES.—Holotype, 9 (Neopseustis meyricki); in

the Zoologisches Museum of Humboldt-Universitat,

Berlin. Lectotype, 9 (Formopseustis takamukui,

present designation), bearing the following labels:

Horisha, Formosa, 1917; Formopseustis takamukui

M.; type, Matsumura; 9 genitalia on slide 2940,

D. R. Davis; lectotype 9 , Formopseustis takamukui

Mats., by Davis 1975.

TYPE-LOCALITIES.—Kosempo, Formosa (Neopseustis meyricki); Horisha, Formosa (Formopseustis

takamukui).

DISTRIBUTION (Map 1).—This species has been

collected only from the central highlands of Taiwan (Formosa), where it appears to occur rather

widely at elevations exceeding 1000 meters.

MATERIAL EXAMINED.—2 males and 6 females.

TAIWAN. Horisha [Pu-Ii]: 1 9 (lectotype, F. takamukui),

1 £ (paralectotype, F. takamukui), 1917 (HU): 1 9 (paralec-
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

totype, F. takumukui), 1917 (USNM). Kanshirei [Kuan-tzuling]: 2$, 3 April 1909 and 7 July 1908 (BMNH). Kosempo

[Chia-hsien]: 1 $ (holotype, N. meyricki), 1 $ (paratype, N.

meyricki), Sept. 1909 (ZMHU). Raisya [Lai-i-ts'un]: 1$ , 23

Nov. 1934 (USNM).

DISCUSSION.—As many specimens have been collected of this species as of any other member of the

family. With the exception of a single female of

Apoplania chilensis, new species, this small series

of eight specimens contains the only known females

collected in this group. As is true for the Neopseustidae in general, very little is known concerning the biology or behavior of this species. Through

recent discussions with Prof. Syuti Issiki, I was

able to learn that the specimen he found at Raisya

in southern Taiwan was collected in a horticultural garden for medicinal plants. Prof. Issiki further informed me that the three specimens collected

by Matsumura at Horisha were found near floodlights of a hydroelectric station. It is possible that

the species may be bivoltine as adults have been

collected in April and July as well as later in

September and November.

Although N. meyricki has never been found on

the Chinese mainland, its closest relatives, N. archiphenax and N. sinensis, new species, do occur there.

The male genitalia of these species bear striking

similarities, particularly in the peculiar development of the pubescent parameres and in the clavate

saccular lobe that arises from the base of the valve.

The three may be easily distinguished by the different form of their anellar lobes.

Neopseustis meyricki was described from two

specimens, supposedly both males as stated originally by Hering (1925). An examination of both

of these specimens, however, has revealed some

discrepancies in Hering's description. The holotype,

for example, is not a male but instead a female.

More significantly, it was discovered that the male

genitalia was damaged in Hering's slide preparation of the paratype, with the gnathos, juxta, and

lateral processes of the anellus, obviously broken

and missing. This discrepancy can be noted in

Hering's rather crude sketches (Figures 3, 4) of

the male genitalia.

A review of the literature has revealed a second

species of Neopseustidae, Formopsenstis takamukni,

described from Taiwan which western entomologists have apparently overlooked. An examination

of the two females and one male comprising the

syntypic series of this moth has resulted in the

synonymizing of both the generic and specific

names. The "rediscovery" of this series in the collections of Hokkaido University has enabled me

to correct the faulty description of the male genital

structure by Hering.

Neopseustis sinensis, new species

FIGURES 17, 26-27, 79-83

ADULT (Figure 17.)—Wing expanse: $, 23-24

mm.

Head: Vertex and frons brownish fuscous with

a slight suffusion of whitish scales near lateral

margin of occiput. Antennae brownish fuscous,

underside of scape light brown to stramineous.

Maxillary palpi whitish to pale stramineous. Labial

palpi brownish fuscous below, pale brown to stramineous above.

Thorax: Dorsum light brown; venter pale stramineous. Legs light brown to stramineous, without

distinct banding on tarsi. Forewings primarily

whitish to stramineous, heavily mottled with various shades of fuscous; basal two-thirds of wing

dull white to stramineous, with 3 or 4 rather large,

indefinite spots of dark fuscous along costal cell,

two or more very irregular areas of dark fuscous

transversing wing from near apex of discal cell to

middle of cell M3; numerous scattered patches of

broad, silvery white scales concentrated mainly

along costal cell and some of the major veins;

outer third of wing thinly covered with narrow,

whitish scales except for infusion of light brown,

partially confluent spots along apex and termen,

as well as an irregular series of brownish striae

scattered across outer third of wing. Hindwings

whitish to pale grayish brown over basal two-thirds

and whitish over outer third except for a series of

light grayish brown spots along outer margin at

termination of principal veins.

Abdomen: Sparsely clothed with slender, pale

grayish brown scales. Male with median process of

seventh sternite sharply acute. A broader, median,

sometimes bilobed process present near caudal

margin of eighth tergite; lobes shallow, each bearing 3 or 4 minute peglike spines. •

Male genitalia (Figures 79-83): Gnathos a slender uncinate process arising between socii and

projecting ventrally. Valvae with a bilateral pair
NUMBER 210 23

of clavate lobes arising from sacculus and nearly

contiguous along midline. Lateroposterior lobes of

anellus irregular in outline, spicate in appearance

with numerous, acute, secondary projections of

various sizes; an elongate, bilateral pair of sharp

processes arising posteriorly and closely parallel

from median area of anellus. Parameres greatly

lengthened, surpassing apex of uncus, densely pubescent over ventral half, gradually tapering to

acute apices.

HOLOTYPE.—•$ , Beh Luh Din, 30 mi [18:5 km]

north of Chengtu, Szechuan Prov., China, D. C.

Graham coll., USNM 72099; in the National Museum of Natural History, Smithsonian Institution.

PARATYPE.—CHINA: Szechuan Prov.: Same data

as holotype, 1 $ (USNM). Described from two males.

DISTRIBUTION (Map 1).—Known only from Szechuan Province of southwestern China.

DISCUSSION.—This species approximates Neopseustis archiphenax in size, and it may eventually be

shown, whenever a large series of both are obtained, that their wing spans may actually overlap

to some extent. The wing maculation of the two

differ, however, and this should provide a ready

means of identification. The principal distinction

can be observed in the hindwing with the marginal,

pale grayish brown markings of N. sinensis being

more reduced and separated than those of N.

archiphenax. A more reliable means of separating

these two species, as well as N. meyricki, is in the

form of the lateroposterior lobes of the anellus. In

N. sinensis these lobes are very stout and not deeply

divided; in N. archiphenax they are more slender

and are divided for most of their length with one

branch coarsely serrated on one side. As pointed

out under the discussion for the latter species, however, the genital descriptions of N. archiphenax are

based on a questionably identified specimen.

Apoplania, new genus

TYPE-SPECIES.—Apoplania chilensis, new species.

ADULT.—Relatively small, moderately heavy bodied moths with very thinly scaled wings.

Head (Figures 28-32): Antennae with minute

intercalary sclerite present between scape and pedicel (Figure 29); segments submoniliform, of similar form over most of flagellum; sensory scales

arranged mostly in longitudinal rows of 4 or 5 contiguous sockets each (Figure 20). Chaetosemata

well developed on frons and vertex of head. Galeae

short, subequal to entire labial palpus and slightly

surpassing third segment of maxillary palpus. Maxillary palpi very long, nearly doubling the length

of labial palpi; fourth and fifth segments the greatest in length and of equal size.

Thorax: Epiphysis present but extremely reduced in size, approximately twice its width in

length (Figure 48); metathoracic tibia with basal

spurs arising from distal fourth of segment. Forewings (Figure 34) 13-veined; R2 and 3 completely

fused, usually connate with R4+5, but occasionally

stalked for a short distance; Ml and 2 stalked

approximately 0.3 their total length; discal cell

relatively long, extending more than half the

length of the wing; base of anal area raised to

form a prominent depression underneath wing at

base of discal cell and a poorly developed pocket

at base of anal veins; fused portion of CuA over

0.3 the length of CuAl; distal end of Al+2 gradually curving to hind margin. Hindwings 12-veined;

crossvein present at base of costa; R2 and 3 completely fused; crossvein r-m slanted obliquely inward toward Ml+2; Ml and 2 stalked over

one-third their length; M3 arising approximate to

r-m crossvein.

Abdomen: Relatively short and broad. Seventh

sternite of male without median lobe. Female with

a bilateral pair of setigerous dark-shaped pads

present near caudal margin of fourth sternite;

seventh sternite strongly sclerotized, with caudal

margin curved posteriorly at middle and bearing

a small spinose lobe.

Male genitalia: Uncus well developed, elongate,

deeply clefted. Tegumenal lobes broadly rounded.

Gnathos a relatively simple, median plate without

elongate processes. Valvae simple, bases widely

separated. Vinculum with anterior margin moderately convex, lateral arms reduced. Juxta broad,

platelike. Anellus primarily consisting of a bilateral

pair of long, slender processes. Aedeagus heavily

sclerotized, with a prominent array of stout and

rigid exophallic spines.

Female genitalia: Posterior apophyses stout, relatively short, extending anteriorly to broad, internal vulva; apex of ovipositor strongly tapering,

relatively narrow. Ninth tergite with caudal margin deeply concave; lateral lobes without secondary

ridges. Ductus spermathecae nearly straight, only
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

slightly sinuate; annular vesicle prominent; lateral lagena absent; utriculus slender, reduced in

size; receptacular gland present, filamentous. Ductus bursae very short, walls heavily thickened with

bilaterally paired, internal pockets or folds.

Corpus bursae elongate, completely membranous.

Anterolateral angles of eighth tergite with short

processes, perhaps representing rudimentary anterior apophyses.

DISCUSSION.—Although the two taxa superficially

resemble one another closely, several major features separate Apoplania from its closest known

Old World ally, Neopseustis. Prominent among

these are: the greatly reduced epiphysis, heavily

sclerotized aedeagus, and strongly developed uncus

in Apoplania; the differential development of the

saccus between the two genera; and the presence

of paired parameres and the more extended, armlike tegumenal lobes in Neopseustis. The two

resemble each other rather closely in venation,

although the peculiar anal pocket at the base of

the forewing in this group is much better developed in Neopseustis.

The generic name Apoplania is treated as feminine and is derived from the Greek, apoplanias,

meaning wanderer or fugitive.

Apoplania chilensis, new species

FIGURES 2-4, 6, 18-19, 20, 28-32, 34, 40-42, 48, 51, 54, 57,

84-87, 94-95, 96-98

ADULT (Figure 18).—Wing expanse: $ 15-19

mm.; $ , 20 mm.

Head (Figures 28-32): Vestiture stramineous to

pale ochreous. Eyes large, interocular index approximately 1.35. Antennae whitish below, light

tan above, 94-96 segmented; sensory scales arranged

mostly in two bands (distal and medial) of alternating longitudinal rows; each row usually with

4 or 5 raised sockets (Figure 20); interspersed

between distal rows at edge of segment are solitary sockets containing large sensory setae; microtrichiae largely restricted to basal third of each

segment. Maxillary palpi and galeae whitish. Labial palpi whitish ventrally, pale tan dorsally.

Thorax: Dorsum pale tan, venter whitish. Legs

pale tan to whitish; tibial and tarsal segments

ringed with brown; usually two rings present on

each tibia; distal ring the broadest; tarsal segments

predominantly brown. Forewings translucent, thinly scaled, mottled with irregular patches of brown

and white; scales slender with acute apices; a conspicuous brownish patch at wing apex and a slender, oblique streak of brown from outer third of

costa across radial veins to Ml. Hindwings similarly clothed, less mottled with brown, with two

conspicuous brownish patches corresponding to

those mentioned in forewing; namely, a slender,

triangular patch from outer third of costa to Ml,

an apical patch extending from Rl to Ml.

Abdomen: Sparsely clothed with slender scales;

pale tan above, more whitish beneath. Female with

caudal margin of seventh sternite bearing approximately 30 stout, short, spinules on and around

median lobe.

Male genitalia (Figures 84-87): Uncus broad

and ventrally concave at base; apex narrowly

divided nearly 0.5 its length. Socii reduced, narMAP 2.—Distribution of Apoplania chilensis, new species.
NUMBER 210 25

rowly separated, and situated on a bulbous, membranous anal pad. Tegumenal lobes evenly

rounded, approximately 0.3 the length of valves.

Gnathos a relatively flat plate, without any median process projecting ventrally, but with a

median extension of plate caudad bearing numerous short, stout spines. Valvae relatively narrow,

without lobes or processes; apex subacute. Median

excavation of vinculum shallow, lateral arms short,

broadly rounded. Juxta peltate, with a bilateral

pair of compressed arms extending caudad. Anellus

a narrow ring, developed dorsally (of aedeagus)

into a bilateral pair of sinuate processes projecting

ventrally, surpassing length of valvae. Apex of

aedeagus with an irregular series of approximately

five large spines radiating outward in a somewhat

spiral manner from main shaft of aedeagus; ejaculatory duct with heavy, sclerotic thickenings for a

distance of 1.5 times the length of the adeagus.

Female genitalia (Figures 96-98): Apex of ovipositor with a bluntly rounded median lobe and a

bilateral pair of minute, acute lobes. Lateral margins of ninth tergite produced into a bilateral pair

of rounded, setigerous lobes on either side of ovipositor apex. Caudal margins of eighth sternite

truncate. Ductus bursae short, very broad, heavily

thickened internally by three bilaterally symmetrical pockets or folds. Corpus bursae long and rather

narrow, over 3.0 the length of posterior apophysis.

HOLOTYPE.—Chile, Nuble Province, Recinto, elevation approximately 1000 m, $, 4-6 Mar. 1968,

coll. Flint and Pena, USNM 72098; in the National

Museum of Natural History, Smithsonian Institution.

PARATYPES.—CHILE: Linares Province: Fundo

Malcho [50-60 km east of Parral], Cordillera

FICURE 11.—Habitat of Apoplania chilensis, new species, Linares Province Chile.

(Photograph courtesy of Mr. L. Pena.)
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Parral, elevation approximately 600-1000 m, 3 $ ,

11-20 Nov. 1964 (USNM); 2$, same data (LEP);

1 <$ , same data (BMNH). Maule Province: Costa

Maule, Pelluhue, 600 m, 1 $ , 2 Dec. 1953 (USNM).

Described from a total of seven males and one

female.

DISTRIBUTION (Map 2).—Known only from heavily forested areas of central Chile between latitudes

of 35°S to 37°S at elevations of 600 to 1000 m.

DISCUSSION.—This species occurs in forest habitats dominated by such genera as Nothofagits and

Chusquea. Most of its known range lies within

the faunal region characterized by Peiia (1966) as

the Northern Valdivian Forest, such as shown in

Figure 11. The holotype of Apoplania chilensis

was collected by O. S. Flint during the day by

sweeping bushes bordering a mountain stream near

Recinto. All paratypes were collected by L. Pefla

at night from a white sheet suspended before a

gasoline pressure lantern.

Literature Cited

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1974. The Uses of Cladistics. Annual Review of Ecology

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1949. Note sur la systematique des lepidopteres et creation de deux superfamilles. Revue Francaise d'Entomologie, 16 (2):74-77.

Brock, J. P.

1971. A Contribution towards an Understanding of the

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Brundin, L.

1966. Transantarctic Relationships and Their Significance, as Evidenced by Chironomid Midges. Kungl,

Svenska Velenskapsakademiens Handlingar, Fjarde

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Chapman, T. A.

1917. Micropteryx Entitled to Ordinal Rank; Order Zeugloptera. Transactions of the Royal Entomological

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1970. Lepidoptera. Pages 765-866 in CSIRO, The Insects

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Comstock, J. H.

1918. The Wings of Insects. 430 pages. Ithaca, New York.

Darlington, P. F., Jr.

1957. Zoogeography: The Geographical Distribution of

Animals, xi + 675 pages. New York.

Davis, D. R.

1967. A Revision of the Moths of the Subfamily Prodoxinae (Lepidoptera: Incurvariidae). United

States National Museum Bulletin, 255:170 pages.

1975. West Indian Moths of the Family Psychidae with

Descriptions of New Taxa and Immature Stages.

Smithsonian Contributions to Zoology, 188:66 pages.

Dugdale, J. S.

1974. Female Genital Configuration in the Classification

of Lepidoptera. Netc Zealand Journal of Zoology,

1 (2): 127-146.

Dumbleton, L. J.

1952. A New Genus of Seed-Infesting Micropterygid

Moths. Pacific Science, 6(l):17-29.

Ehrlich, P. R.

1958. The Comparative Morphology, Phylogeny and

Higher Classification of the Butterflies (Lepidoptera: Papilionoidea). University of Kansas Science

Bulletin, 39 (8):305-370.

Eltringham, H.

1925. On a New Organ in Certain Lepidoptera. Transactions of the Entomological Society of London,

1,2:7-9, plate 2.

Fletcher, T. B.

1929. A List of the Generic Names Used for Microlepidopteia. Memoirs of the Department of Agriculture

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1933. Life-Histories of Indian Microlepidoptera (Second

Series): Cosmopterygidae to Neopseustidae. The

Imperial Council of Agricultural Research, Scientic Monograph, 4: 85 pages, 77 plates.

Forbes, W. T. M.

1932. How Old Are the Lepidoptera? The American

Naturalist, 56:452-460.

Friese, G.

1969. Zur Phylogenie der altern Teilgruppen der Lepidopteren. Bericht tiber die 10 Wanderversammlung

Deutscher Entomologen Tagungsberichte 80, 2:203-

222.

Hennig, W.

1966a. The Diptera Fauna of New Zealand as a Problem

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1966b. Phylogenetic Systematics. 263 pages. Urbana.

Hering, M.

1925. Ueber die Gattung Neopseustis Meyrick (Lep.)-

Mitteilungen aus dem Zoologischen Museum, Berlin, 12:141-147.

Hinton, H. E.

1946. On the Homology and Nomenclature of the Setae

of Lepidopterous Larvae with Some Notes on the

Phylogeny of the Lepidoptera. Transactions of the

Royal Entomological Society of London, 97:1-37.
NUMBER 210 27

Jordan, K.

1923. On a Sensory Organ Found on the Head of Many

Lepidoptera. Novitates Zoologicae, 30:155-158, plate

2.

Klots, A. B.

1970. 20, Lepidoptera. Pages 115-130 in S. L. Tuxen,

Taxo7iomist's Glossary of Genitalia in Insects. 2nd

edition, 359 pages. Darien, Conn.

Kossmatt, F.

1936. Palaegeographie und Tektonik. Berlin.

Kristensen, N. P.

1968. The Skeletal Anatomy of the Heads of Adult

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(3917):379-380.

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1973. 29, Ordnung Trichoptera (Kocherfliegen). Handbuck der Zoologie, 4(2) 2/29:1-114.

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1974. Biogeographical Evolution in India. Pages 698-724

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1960. Wing Venation in Lepidoptera. Entomological Review, 39 (2): 190-191. [English translation of Entomologicheskoe Obozrenia, 39 (2):296-299.]

Matsuda, R.

1970. Morphology and Evolution of the Insect Thorax.

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Matsumura, S.

1931. 6000 Illustrated Insects of Japan-Empire. 1687

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Meyrick, E.

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1971. A New Genus of Homoneurous Moth and the

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1970. 19, Trichoptera. Pages 104-115 in S. L. Tuxen,

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1948. Sobre a Genitalia das Femeas de Hepialidae. Summa

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1966. A Preliminary Attempt to Divide Chile into Entomofaunal Regions, Based on the Tenebrionidae

(Coleoptera). Postilla, 97:1-17.

Philpott, A.

1925. On an Abdominal Scent-organ (?) in Sabatinca

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Smith, A. G., J. C. Briden, and G. E. Drewry

1973. Phanerozoic World Maps. Pages 1-42 in N. F.

Hughes, Organisms and Continents through Time.

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Palaeontology, 12: 334 pages. London.

Snodgrass, R. E.

1957. A Revised Interpretation of the External Reproductive Organs of Male Insects. Smithsonian Miscellaneous Collections, 135(6):l-60.

Tillyard, R. J.

1935. The Evolution of the Scorpionflies and Their

Derivatives. Annals of the Entomological Society

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Unzicker, J. D.

1968. The Comparative Morphology and Evolution of

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1946. Revision du Catalogue des Lepidopteres Homoneures, Ire Note: Famille des Micropterygidae.

Revue Francaise d'Entomologie, 14(1)24-31.
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

12 13

14 15

16

FIGURES 12-17.—12, Archepiolus schmidi Mutuura, $ holotype, wing expanse 16 mm; 13, Neopsettstis calliglauca Meyrick, $, wing expanse 16 mm; 14, N. bicornuta, new species, $ holotype,

wing expanse 19 mm; 15, N. archiphenax Meyrick, $ holotype, wing expanse 27 mm; 16, N.

meyricki Hering, $, wing expanse 22 mm; 17, N. sinensis, new species, $ holotype, wing

expanse 23 mm.
NUMBER 210 29

FIGURES 18-23.—Adults: 18-19, Apoplania chilensis, new species, $ holotype, wing expanse 18

mm. Antennae: 20, A. chilensis, new species, antennal segments 40-43, X 200; 21, Neopseustis

archiphenax Meyrick, antennal segments 27-29, X 230; 22, AT. archiphenax Meyrick, surface

detail of sensillum trichodea (see Figure 21, ST), X 12,000; 23, N. bicornuta, new species,

antennal segment 21, X 600.
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

25

h- H

FIGURES 24-27.—Head structure: 24, Archepiolus schmidi Mutuura, anterior view of head

(scale = 0.5 mm); 25, A. schmidi Mutuura, antennal segments 15-18; 26, Neopsenstis sinensis, new

species, lateral view of tentorium (scale = 0.2 mm); 27, N. sinensis, new species, anterior view of

head (scale = 0.5 mm).
NUMBER 210

-— IS

32

i CDO

; Ss

83!

30

T

L

is- -a

31

FIGURES 28-32.—Apoplania chilensis, new species, head structure: 28, anterior view of head

(scale = 0.5 mm); 29, intercalary sclerite (IS) between scape and pedicel; 30, maxilla (scale =

0.25); 31, caudal view of head; 32, antcnnal segments 25-28 (scale = 0.25 mm).
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

R l R2

Scl

FIGURES 33-36.—Wing venation: 33, Archepiolus schmidi Mutuura; 34, Apoplania chilensis, new

species; 35, Neopseustis archiphenax Meyrick; 36, N. meyricki Hering, detail of anal pocket

(shaded area).
NUMBER 210

FIGURES 37-42.—Scale structure: 37, Neopsenstis calliglauca Meyrick, dorsal scales of discal

cell, forewing, X 500; 38, detail of Figure 37 (x), X 10,000; 39, detail of Figure 38, X 30,000,

scale=1.3 um; 40, Apoplania chilensis, new species, microtrichium (aculea) from discal cell of

hindwing, X 12,500; 41, A. chilensis, new species, dorsal scales of discal cell, hindwing, X 500;

42, detail of Figure 41 (x), X 15,000, scale = 2 urn.
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

'TT^ • - •

•• mi i- it Mi.fJ

mm

FIGURES 43-48.—Scale structure: 43, Hepialus sequoilus Behrens, Hepialidae, detail of a dorsal

scale from discal cell of forewing, X 5000; 44, detail of Figure 43, X 20,000, scale = 2 \im; 45,

Mnesarchea loxoscia Meyrick, Mnesarchaeidae, detail of a dorsal scale from discal cell of forewing, X 10,000, scale=l.r> 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, Meopseustis 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), sensory 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 reproductive 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. meyricki 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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