A Revision of the

North American Moths of the

Superfamily Eriocranioidea

with the Proposal of a New Family,

Acanthopteroctetidae (Lepidoptera]

DONALD R. DAVIS

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 251
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SMITHSONIA N CONTRIBUTION S TO ZOOLOG Y • NUMBE R 251

A Revision of the

North American Moths of the

Superfamily Eriocranioidea

with the Proposal of a New Family,

Acanthopteroctetidae (Lepidoptera)

Donald R. Davis

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978
ABSTRAC T

Davis, Donald R. A Revision of the North American Moths of the Superfamily Eriocranioidea with the Proposal of a New Family, Acanthopteroctetidae

(Lepidoptera). Smithsonian Contributions to Zoology, number 251, 131 pages,

344 figures, 3 tables, 6 maps, 1978.—The general biology, biogeography, morphology, and classification are reviewed for the five genera and 16 species

(including one subspecies) of North American Eriocranioidea. One new genus,

Neocrania; one new subgenus, Disfurcida, of Eriocraniella; nine new species,

Ei'iocrania breviapex, Eriocraniella xanthocara, E. longifurcula, E. platyptera,

E. variegata, E. trigona, E. falcata, Neocrania bifasciata, and Acanthopteroctetes

unifascia; and one new subspecies, Eriocrania semipurpurella pacifica, are

described. A new family, Acanthopteroctetidae, is proposed for the endemic

North American genus, Acanthopteroctetes. The Australian-Fijian family

Agathiphagidae is removed from the Eriocranioidea and placed in the new

superfamily Agathiphagoidea. Distribution maps and keys are provided for all

species. The larval stages for nearly half of the known Nearctic eriocraniid

species are described for the first time. All stages are fully illustrated by line

drawings and photographs.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Davis, Donald Ray.

A revision of the superfamily Eriocranioidea with the proposal of a new family, Acanthopteroctetidae (Lepidoptera)

(Smithsonian contributions to zoology ; no. 251)

Bibliography: p.

1. Eriocraniidae. 2. Acanthopteroctetidae. 3. Insects—Classification. 4. Insects—North America. I. Title: A revision of the superfamily Eriocranioidea .. . II. Series: Smithsonian

Institution. Smithsonian contributions to zoology ; no. 251.

QL1.S54 no. 251 [QL561.E8] 591'.08s [595.7'8] 77-24967
Contents

Page

Introduction 1

Biology 2

Distribution 2

Life History 4

Parasitism 5

Hosts 6

Morphology 7

Adult Morphology 10

Pupal Morphology 29

Larval Morphology 32

Abbreviations 34

Systematics and Evolution 35

Subordinal Classification 35

Family Classification 36

Generic Classification 38

Early History 41

Checklist of Nearctic Eriocranioidea 42

ERIOCRANIOIDEA 42

Key to the Nearctic Families of Eriocranioidea 44

ERIOCRANIIDAE 44

Key to the Genera of Nearctic Eriocraniidae 46

Key to the Larvae of Nearctic Eriocraniidae 47

Dyseriocrania Spuler 47

Key to the Species of Nearctic Dyseriocrania 48

D. griseocapitella (Walsingham) 48

D. auricyanea (Walsingham) 56

Eriocrania Zeller 60

Key to the Species and Subspecies of Nearctic Eriocrania 61

E. semipurpurella semipurpurella (Stephens) 61

E. s. pacifica, new subspecies 63

E. breviapex, new species 67

Eriocraniella Viette 68

Key to the Subgenera of Eriocraniella 70

Subgenus Eriocraniella Viette 70

Key to the Species of the Subgenus Eriocraniella 70

E. (£.) aurosparsella (Walsingham) 71

E. (E.) xanthocara, new species 76

E. (E.) longifurarfa, new species 81

E. (E.) platyptera, new species 85

Disfiircula, new subgenus 87

Key to the Species of the Subgenus Disfurcirfa 87

E. (D.) variegata, new species 87
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page

E. (D.) trigona, new species 88

E. (D.) falcata, new species 90

Neocranidj new genus 92

N. bifasciata, new species 94

ACANTHOPTEROCTETIDAE, new family 95

Acanthopteroctetes Braun 96

Key to the Species of Acanthopteroctetes 96

A. tripunctata Braun 97

A. bimaculata Davis 97

A. unifascia, new species 98

Literature Cited 100

Figures 166-344 104
A Revision of the

North American Moths of the

Superfamily Eriocranioidea

with the Proposal of a New Family,

Acanthopteroctetidae (Lepidoptera)

Donald R. Davis

Introduction

The superfamily Eriocranioidea contains some of

the most primitive elements within the order

Lepidoptera; consequently, a thorough study of its

members should be of considerable importance to

anyone interested in the phylogeny and systematics

of the order. The superfamily is further interesting

in that it is believed to represent one of the first

successful phyletic lines within the Lepidoptera to

explore and fully utilize the leaf-mining habitat.

Largely for these reasons, it has become imperative

that a revision of our North American species be

undertaken. From the relative number of new taxa

and information presented on the following pages,

it should be apparent that such an investigation

has been long overdue.

The present study represents the first attempt to

revise the Nearctic Eriocranioidea. Although the

completion of this revision has been somewhat curtailed by the absence of a similar treatment for the

Palearctic species, much information concerning

the Old World fauna has been derived from an

examination of most of the known species as well

as from critical data generously supplied me by sevDonald R. Davis, Department of Entomology, National

Museum of Natural History, Smithsonian Institution, Washington, DC. 20560.

eral colleagues. The European Eriocraniidae have

been superficially reviewed by such early workers

as Spuler (1910), or reviewed more recently for

specific countries by Heath (1958, 1976) and Razowski (1975).

This report marks the first in a series of investigations by the author on select families of North

American leaf-mining moths. The research has been

partially supported by grants from the Smithsonian

Research Foundation. In addition to this aid, I

would like to acknowledge several individuals who

have helped me during the course of this study by

providing critical information or special assistance.

In this regard I wish to thank Dr. Robert Carlson,

United States Department of Agriculture, Washington, D.C.; Mr. John Heath, Monks Wood Experimental Station, Huntingdon, England; Prof.

Syuti Issiki, Ito City, Japan; Dr. Niels Kristensen,

Zoologisk Museum, Copenhagen, Denmark; Dr.

Tosio Kumata, Hokkaido University, Sapporo,

Japan; Dr. Akira Mutuura, Entomology Research

Institute, Ottawa, Ontario; Dr. Paul Opler of the

Office of Endangered Species, United States Department of the Interior, Washington, D.C.; and Dr.

A. K. Zagulajev, Zoological Institute, Academy of

Sciences, Lenningrad, USSR. I wish especially to express my appreciation to Mr. Heath and Dr. Kristensen for assisting me with numerous inquiries re-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lating to this problem. Most of the line illustrations

in this paper were completed under contract by

Miss Ann Szymkowicz, variously assisted by me

and by the following staff artists of the Department

of Entomology, Smithsonian Institution: Miss Biruta Akerbergs, Mr. Lawrence Druckenbrod, Mr.

Ambrose Nelson, and Mr. George Venable. I am

indebted to Mr. Victor Kranz of the Smithsonian

Photographic Laboratory and to Miss Mary Mann

and Mr. Walter Brown of the Scanning Electron

Microscope Laboratory for their much appreciated photographic assistance. Finally, I wish to

thank my colleagues at the institutions and collections listed below for allowing me to examine

specimens under their care or for other assistance.

AMNH American Museum of Natural History, New York,

New York

BMNH British Museum (Natural History), London,

England

BrM Collection of Bryant Mather, Clinton, Mississippi

CAS California Academy of Sciences, San Francisco

California

CNC Canadian National Collection, Entomology

Research Institute, Ottawa, Canada

CPK Collection of Charles P. Kimball, Barnstable,

Massachusetts

DCF Collection of Donald C. Frack, La Puente,

California

JBH Collection of John B. Heppner, Gainesville, Florida

LACM Los Angeles County Museum, Los Angeles,

California

NSM Nova Scotia Museum, Halifax, Nova Scotia

RHL Collection of Ronald L. Leuschner, Gardena,

California

UCB University of California, Berkeley, California

UCR University of California, Riverside, California

USNM United States National Museum collections (in the

National Museum of Natural History, Smithsonian

Institution, Washingon, D.C.)

Biology

DISTRIBUTION

MAPS 1-6

All members of the Acanthopteroctetidae and

Eriocraniidae are confined to the Holarctic region.

Prior to this study, the Eriocraniidae was known

principally from the Palearctic region, but recent

collecting in North America has shown the family

to be even more diversified in the New World.

Meyrick (1921b) lists 12 species of Eriocraniidae

as occurring in Europe; one or two of these names

have since been synonymized, and at least two additional names were proposed subsequently. The

great majority of the Palearctic species are known

primarily from western Europe, but at least two

species have been reported from northern Japan.

The distribution of this family in Asia is very

poorly known because of insufficient collecting in

several critical areas. From the scant records presently available, it appears that the northern and

southern distributional limits of the family through

this area may roughly approximate that of one of

the common host species, Betula pubescens Ehrhart

as outlined by Schmucker (1942, map 69) and

partially reproduced herein on Map 1.

Previously, only three genera including seven

species of Eriocranioidea were known from North

America. One of these species, Micropteryx [sic]

luteiceps Walker, has been found to be a new (junior) synonym of Paraclemensia acerifoliella (Fitch),

a member of the monotrysian family Incurvariidae.

Another species, Dyseriocrania cyanosparsella (Williams) is treated as a junior synonym of D. auricyanea (Walsingham) in this paper. Despite these

new synonymies, the North American fauna has

been found to include at least 15 species and one

subspecies distributed in two families and five

genera. Of these, one family and three genera appear to be indigenous to the Nearctic region.

Briefly mentioned, but not named in the present

study, are representatives of five additional, endemic species. One of these, known only from a

unique male in imperfect condition from Ontario,

Canada, may also represent an additional, undescribed genus. The general distribution of the superfamily in North America is intimately associated

with the two principal montane features of our

continent, the western North American Cordillera

and the eastern Appalachian Highlands. No members of Eriocranioidea are known to occur from the

vast Interior Plains.

The genus Dyseriocrania is presently represented

by two Nearctic species and at least one western

Palearctic species. The two North American species

are amply separated by the Interior Plains of the

continent, with the eastern D. griseocapitella (Walsingham) ranging widely through the Appalachian

Highlands and the adjacent Atlantic Coastal Plain;

the western D. auricyanea (Walsingham) is confined to the southern ranges and coastal valleys of

the Pacific Mountain system.
NUMBER 251

MAP 1.—General distribution of Palearctic Eriocraniidae (shaded area) in relation to Betula

pendula Roth {dotted line).

Although the genus Eriocrania contains several

Palearctic species, only two distinct members of this

group have been discovered thus far in North

America; however, some evidence exists for yet another species. One of our North American species,

E. semipurpurella (Stephens), also ranges widely

through the Palearctic region, reportedly from

Great Britain to Japan. In North America, the

typical form of this species is restricted primarily

to the northern Appalachian Highlands and St.

Lawrence Valley region, with a widely disjunct,

western form, E. s. pacifica, new subspecies, occurring at low elevations along the northern Pacific

Coast Ranges. One new species, E. breviapex, from

the Appalachian Plateau of New York is apparently

endemic to North America.

The endemic genus Eriocraniella has its main

center of distribution and probable origin in the

western United States where two subgenera and

seven species are known to occur. Evidence for an

eighth species exists. Only one member of this group

is known from the entire eastern United States,

this being E. platyptera, new species, from the Appalachian Plateau of New York. In this regard, it

is of interest to note that the latter shares closest

affinities with E. longifurcula, new species, the most

eastern ranging of the western species and the only

member of that assemblage not restricted to cismon-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tane California.

The new genus Neocrania presently contains

only one species endemic to the coastal ranges of

southern California.

The family Acanthopteroctetidae is represented

by a single genus Acanthopteroctetes, which is indigenous to western North America. Although only

three species are currently recognized for our fauna,

evidence for two others is available. Most of the

collection records to date indicate that the members

of this genus prefer relatively cool, wet, boreal

habitats; however, the only known habitat of A.

unijascia, new species, is that of a relatively dry,

coastal chaparral biota.

LIFE HISTORY

AH records to date indicate the members of Eriocranioidea to be consistently univoltine with the

adult moths emerging relatively early in the spring

or late winter, usually during February to late May

depending upon the latitude or elevation. Some

adults of the genus Acanthopteroctetes have been

collected as late as mid-July. The early seasonal

emergence of the adults has undoubtedly contributed to the usual sparseness of specimens in collections and the frequent lack of adequate study

material and distributional records.

The Eriocranioidea are of minor economic importance, with only a single reference to date alluding to larval damage by a member of this group.

Keifer (1927) reported a heavy infestation of Dyseriocrania auricyanea (Walsingham) in the leaves

of live oak, Qitercus agrifolia Nee, in San Francisco, California.

The adults may often be found resting on foliage

or tree trunks in the early morning. Their flight,

according to Williams (1905), tends to be in a slow,

direct manner. Williams also noted that the moths,

when captured by net, would sometimes feign death

and lie motionless in the bottom of the bag. The

adults are typically diurnal and are capable of

flight anytime during the daylight or evening hours.

Opler (1974) observed the adults of Dyseriocrania

auricyanea to be most active in the hours just before sunset.

The highly modified, piercing ovipositor enables

the female to insert the egg under the leaf epidermis. The eggs are usually laid singly into young

leaves in the process of opening and expanding.

Wood (1890) presents evidence to suggest that the

females of Eriocrania hazuorthi Bradley ( = purpurella Haworth) deposit more than one egg per

site and often three. Because of the partially unopened condition of the developing leaves, a female

eriocraniid frequently oviposits on the underside of

a leaf. As summarized by Needham et al. (1928),

the tim.e required for the female to cut an incision

and deposit an egg may vary from 2.5 to 15 minutes, largely depending upon the relative temperature. After the eggs have been deposited, they may

swell to more than twice their original size.

As a direct result of the damage done to the leaf

tissue during oviposition, an oval hole, or insertion

scar (Figure 134), typically develops soon after an

egg has been deposited. The scar may be obliterated

later by the developing mine, but often it persists

as a small hole> usually measuring 1.5-2.0 mm in

length and 1.0-1.2 mm in width. Normally evident

along one side of the insertion scar is a small, reddish brown, cylindrical ovate egg pouch (Figure

164) often measuring 0.7-0.9 mm by 0.5-0.4 mm.

Inside each egg pouch can normally be found the

remains of a single egg. The formation of the egg

pouch is not completely understood, but it may

result from the relative adhesiveness of the egg

chorion to the leaf epidermis with the eventual drying and collapsing of the leaf epidermis around the

eggWithin 7 to 15 days the first instar larva emerges

and initiates a narrow, rather serpentine mine

directed either toward or along the outer margin

of the developing leaf. Again, this stage of the mine

may sometimes be obliterated by the later feeding

of the larva. Frequently, in Dyseriocrania griseocapitella, the linear mine forms a line of weakening

across the leaf that later may rupture forming a

narrow fissure. The length of the linear mine varies

according to the species, usually ranging from 5 to

40 mm in length.

The termination of the linear, serpentine mine

probably signals the initiation of the second larval

instar. After this the mine either gradually or

abruptly enlarges to form an irregular blotch that

often is longer than wide. In some species the

blotch tends to be very elongate and follows the

margin of the leaf; in others an entire section (e.g.,

the apical third) of a leaf may be mined resulting

in a much broader blotch. All of the parenchyma is

devoured within the blotch, thus producing a full
NUMBER 251

depth, transluscent mine. As noted by Kearfott

(1902), the larva while feeding normally sweeps in

small circles of 5 mm or more radius with its anal

end functioning as a pivot. The final pattern of the

mine usually results from a series of these semicircular movements marked by a series of fine, concentrically curved lines. The frass produced by

eriocraniid larvae is characteristic in consisting of

relatively long, slender segments or threads that

are loosely scattered and intertwined within the

mine. As noted by previous authors (Wood, 1890;

Opler, 1974), a total of four larval stages are believed to exist, with the last three being responsible

for the blotch mine. At maturity, only a single

larva normally occupies each mine. In the case of

Eriocrania haworthi ( = purpureIla), however, Wood

(1890) has reported as many as six larvae per mine.

Larval development proceeds rather rapidly and

within 7 to 10 days all feeding ceases.

The immature stages of Eriocraniidae are not

restricted to a leaf-mining habitat in that pupation

occurs in the soil. In emerging from the leaf, the

larva cuts a small semicircular slit in the upper

epidermis and drops to the ground. The larva then

digs down into the soil to a depth of 2 to 10 inches,

probably depending in part upon the relative compactness of the soil. There the larva creates a small

earthern cell within which it spins a thin but firm,

oval cocoon (Figure 102), frequently measuring 2

mm by 3-4 mm. The cocoon is composed largely of

whitish silk with small grains of earth and sand

firmly incorporated into the matrix.

The entire life cycle of these moths is synchronized with the production of new, tender leaves containing a relatively low tannin content (Feeney and

Bostock, 1968; Morris 1974). Throughout the summer and winter months, the larva remains inactive

within the cocoon. A total of nearly 11 months may

be spent underground with actual pupation perhaps not occurring until the final month, or a few

weeks prior to adult emergence.

Pupal emergence from the cocoon was first described by Chapman (1893) and is accomplished

primarily with the aid of the greatly enlarged and

completely functional mandibles. Probably assisting

the mandibles in this activity is a well-developed

and elongate frontal ridge. The pharate adult first

cuts a rather large opening in one end of the cocoon. Utilizing vigorous motions of the mandibles

and head, this stage is able to work its way gradually out of the cocoon to the soil surface. The legs

are essentially useless during these activities and

are held relatively motionless against the body as

the pharate adult forces its way to the surface. In

contrast, the mandibles are fully capable of being

open and closed as well as being raised and lowered

in conjunction with similar motions of the rather

freely movable head. After emerging from the

ground, the pharate adult remains quiescent for an

indefinite period until final ecdysis can be accomplished.

Chapman (1893) reports that the adult of Eriocrania haworthi emerges in the morning about 6 to

7 A.M. The imaginal wings already are nearly fully

expanded within the wing cases prior to emergence.

After leaving the pupal shell, the moth normally

climbs to some support for final wing expansion

and drying. Within a relatively short interval, the

adult is capable of flight.

PARASITISM

Very few examples of parasitism have been reported for the Eriocraniidae. As summarized in

Table 1, these records involve at least four different

wasp parasites of the family Ichneumonidae attacking a possible total of seven species of Eriocraniidae. Most of the wasps are not closely related but, instead, represent three different subfamilies of Ichneumonidae. It is of interest to note

that no other families of Lepidoptera are known to

be parasitized by any member of the tribes that

these wasps represent.

The Ctenopelmatinae (=Scolobatinae) and Tryphonini almost exclusively attack sawflies. According to Townes (1971), the Tersilochinae are parasitic on beetle larvae, particularly those of Curculionidae. An important biological correlation

among all the known hosts is that their larvae are

typically feeders in plant tissue with pupation occurring in the soil. Thus, the sporadic distribution

of host-parasite records to date indicates a situation

involving certain wasps that have evolved primarily

as parasites of other insect orders but which have

later adapted to the Eriocraniidae largely because

of similarities in the general life history of the

hosts.

It is difficult to comment to any extent concerning relative host specificity because of the limited

number of host-parasite relationships that have
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Host-parasite relationships of Eriocraniidae

Host

Dyseriocrania griseocapitella (Walsingham)

Dyseriocrania sobpurpwcella (Haworth)

Eriocrania haworthi Bradley (-rubroaurella Haworth)

Briocrania salopiella Stainton

Eriocrania sangii Wood

Eriocrania semipurpurella semipurpurella (Stephens)

Heringocrania unimaculella (Zetterstedt)

Subfamily

of

Ichne umonidae

Ctenopelmatinae

Tersilochinae

undetermined

Ctenopelmatinae

Tryphoninae

Ctenopelmatinae

C tenopelmat i nae

Ctenopelmatinae

Tryphoninae

Ctenopelmatinae

Reference

Rohwer, 1914

new record

Heath,

Heath,

Heath,

Heath,

Heath,

Seyrig

Heath,

Heath,

1961

1961

1961

1961

1961

, 1928

1961

1961

Lathrolestes mnemonicae Rohwer

?Tersilochus

undetermined

Lathrolestes clypeatus (Zetterstedt)

Grypocentrus basalis Ruthe

Lathrolestes clypeatus (Zetterstedt)

Lathrolestes clypeatus (Zetterstedt)

Lathrolestes clypeator [sicl eriocraniae Seyrig

Grypocentrus basalis Ruthe

Lathrolestes clypeatus (Zetterstedt)

been reported. It may be possible that the wasp

parasites are largely linked to plant host-moth complexes and will attack any eriocraniid feeding within a particular plant group. Heath (1961), for

example, concluded that the two ichneumonid species he reared probably parasitized all birch-feeding Eriocraniidae.

HOSTS

As summarized in Table 2, nearly all host records for the family Eriocraniidae are included in

two plant families, the Betulaceae and Fagaceae

of the order Fagales. According to Cronquist (1968)

and nearly all other authorities, these two families

are closely related and are derived by floral reduction from ancestors similar to the Hamamelidaceae.

Only two possible host records have been reported

from non-Fagales genera. Eriocrania argyrolepidella

of Europe is suspected of feeding on Populus tremnla in that adults have been observed flying

around this plant. No eriocraniid larvae, however,

have yet been collected from any member of the

Salicales. Eriocraniid larvae, possibly representing

the immature stages of the North American Eriocrania semipurpurella pacifica, new subspecies, have

been found mining the leaves of a member of

the Rosales, Holodiscus discolor. Because no adults

have been definitely associated with these larvae

or reared from Holodiscus the generic and specific

identification of the larvae remains questionable.

With the possible exceptions of Eriocrania alpinella, E. argyrolepidella, and E. semipurpurella pacifica, all members of the genus Eriocrania appear

to feed exclusively on Betula. Only one other eriocraniid, Heringocrania unimaculella, is reportedly

monophagous on Betula. The generic placement of

this particular species is questionable, and it may

only represent a subgenus of Eriocrania (BirketSmith and Kristensen, 1974; Razowski, 1975).

Heath (1958) has noted that the birch-feeding

species show a marked preference for young trees

under 15 feet (4.6 m) in height.

The members of the predominantly North

American genera Dyseriocrania, Eriocraniella, and

possibly Neocrania, new genus, are almost entirely

restricted to plants of the family Fagacae. Only D.

subpurpurella of Europe has been reported feeding

on a nonfagaceous host (Carpinus). Dyseriocrania

griseocapitella appears to be the most polyphagous

of the North American eriocraniids in having been

reared from at least three species of chestnuts and

from representatives in both the red and white oak

groups.

Unfortunately, nothing is known concerning the

host associations of the family Acanthopteroctetidae.

Many facts relating to the distribution of the

Eriocraniidae can be explained by reviewing the

foregoing host data. For example, the restrictions of

nearly all larvae to members of the Fagaceae and

Betulaceae account for the general occurrence of

the Nearctic Eriocraniidae over much of the broadleaf forests of eastern and western North America

as well as for their observed absence from the Interior Plains. Although some members of the Fagaceae range far south into Central America {Qiiercus) or are entirely restricted to the southern Hemisphere (Nothofagus), no examples of Eriocraniidae

have yet been found south of the 25°N parallel.
NUMBER 251

TABLE 2.—General distribution and the known host plants of Eriocraniidae

Species Distribution Host plants

Dyseriocrania

auricyanea (Walsingham).

griseocapitella (Walsingham).

subpurpurella (Haworth)

Briocrania

alpinella Bunnann

argyrolepidella Fuchs

havorthi Bradley

(-purpurella Haworth;

(Tubroaurella Haworth;

saloplella Stainton

sangii Wood

?semipurpurella pacifica Davis

semipurpurella semipurpurella (Stephens)

sparmannella Bosc

Eriocraniella

aorosparsella (Walsingham)

?longifurcula Davis

xanthocara Davis

species +

falcata Davis

Heringocrania Kuznetsov

chrysolepidella Zeller

unimaculella (Zetterstedt)

Neocrania Davis

bifasciata Davis

+ fossil record

eastern North America

western Europe

western Europe

western Europe

Queicus agrifolia Nee*

Q. douglasii Hooke £ Arnott

Q. dumosa Nuttall

0. lobata Nee

Q. wislizenii Alphonse de Candolle

Castanea dentata (Marshall) Borkhausen

C. mollissima Blume

C. pumiia (L.) P. Miller

Quercus alba L.

0. ?borealis F. Michaux

Q. Tfalcata A. Michaux

0. velutina Lamarck

Carpinus species

Castanea species

Quercus petraea (Mattuschka) Lieblein

Q. robuz L.

Alnus viridis Aug. de Candolle

Betula species

TPopulus tremila L.

Betula pendula Roth

western Europe Betula species

western Europe Betula species

northwestern North America Holodiscus discolor Maximowicz (Pursh)

Europe Betula pendula Roth

Japan B. platyphylla var. japonlca (Miquel) Hara

northeastern North America B. pubescens Ehrhart

western Europe

California, Oregon

Arizona

Idaho

California

western Europe

western Europe

B. pendula Roth

B. platyphylla var. japonica (Miquel) Hara

Quezcus kelloggii Newberry

?Quercus gambelli Nuttall

X arizonica Sargent

Quercus agrifolia Nee

Q. wislizenii Alphonse de Candolle

Quercus simulata Knowlton

Quercus chrysolepis Liebmann

Alnus species

Carpinus betulus L.

Corylus avellana L.

Ostrya species

Betula species

?Quercus chrysolepis Liebmann

Qpler, 1974

Hering, 1957

Lhomme, 1949

Burmann, 1958

Hering, 1957

Lhoune, 1949

Lhoome, 1949

Hering, 1957

Kumata (in litt.)

Lhomme, 1949

Toll, 1959

Kumata (in litt.)

Lhomme, 1949

Opler, 1973

Hering, 1957

Hering, 1957

Morphology

The following discussion is not intended to be

an exhaustive treatment on the morphology of the

various stages for the members of the Eriocranioidea. Instead, it is a review of those anatomical

features investigated during the course of this and

related studies that have been considered significant
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 1-4.—Dyseriocrania griseocapitella (Walsingham), adult head structure: 1, lateral view

of head, X 55. 2, compound eye and ocellus, X 170; 3, inner surface of galea, X 1300; 4,

scales of labrum, X 620. (Photographs reproduced at 100%.)
NUMBER 251

FIGURES 5-10.—Eye structure: 5, Dyseriocrania griseocapitella (Walsingham), surface detail of

ocellus, X 1200; 6. facets and microtrichia of compound eye, X 1700; 7, corneal nipples with

globular, secondary divisions, X 30,000; 8, Rhyacophila fuscula (Walker), facets and microtrichia, X 1300; 9, detail of microtrichia and corneal nipple array, X 7000; 10, corneal nipples

with smooth surfaces, x 30,000. (Photographs reduced to 83%.)
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in understanding the systematics of this group. The

publications of Kristensen (1968b) and Birket-Smith

and Kristensen (1974) may be consulted for further

details on the anatomy of the head, alimentary

canal and male genitalia.

ADULT MORPHOLOGY

HEAD.—The head (Figures 1, 11, 17) is hypognathous and sparsely covered with elongate, hairlike

scales, except for the lower part of the frons and

labrum; here the scales tend to be broader and triangular in form (Figure 4). The head capsule differs

markedly in length between the two families. In the

Eriocraniidae the postocular region is more extended and possesses a pair of prominent ocelli.

One result of the lengthened head in the members

of this group is that the compound eyes appear to

be situated farther forward. The postocular region

is more shortened in Acanthopteroctetidae, and the

ocelli are absent. The size of the compound eye is

slightly smaller in the Eriocraniidae, with most

species possessing an interocular index (see Davis,

1975a:5) of approximately 0.70 to 0.75.

The surface of the eriocraniid compound eye appears naked under low magnification; actually,

however, it is sparsely covered with minute microtrichiae (Figures 2, 6) arising between certain facets.

Their length is less than the diameter of a single

facet. The surface of each facet is composed of

myriads of submicroscopic divisions (Figure 7), referred to as corneal nipples by Bernhard and Miller, 1962, and M0ller, 1965. According to these

authors, the function of the corneal nipple array is

that of a natural impedance transformer whereby

light reflection from the corneal surface may be reduced and transmission increased. They may also

function as interference filters as concluded by

Bernard and Miller (1968) for certain Tabanidae

and operate as a contrast filtering device for vision.

The corneal nipples vary considerably in structure

and general arrangement among different insect

groups, with some orders (e.g., Coleoptera and

Orthoptera) lacking them (Miller et al., 1966).

Under increased magnification (i.e., X20,000 and

above), each corneal nipple in eriocraniids can be

observed to be subdivided into more than 15 globular units. Similar secondary subdivisions, or subnipples, have also been observed by the author in

moths of the genera Greya and Acrolophus but

absent in the butterflies Eurema and Pyrgus and

the caddisfly Rhyacophila (Figures 8-10) Consequently, the ultrastructure of the corneal nipples

may have some significance in the higher systematics of the Lepidoptera.

The best review to date on the corneal nipple

array in insects is that by Bernhard et al. (1970).

These authors reported well-developed corneal nipples in several families of Lepidoptera, including

the Eriocraniidae. Within most of the families examined, both nippled and nonnippled corneas were

found represented. No mention was made of the

more minute, secondary units comprising each nipple, and it is possible that these have never been

noted previously.

The cranial sulci generally are better developed

in the Eriocraniidae than in the Acanthopteroctetidae. For example, the laterofacial sulci (LfS) are

reduced in the latter family but extend to the anterior tentorial pits in the Eriocraniidae. The postfrontal sulci (PfS) terminate differently in the two

groups. In the Eriocraniidae they curve anteriorly

to the dorsal rim of the eye; in the Acanthopteroctetidae the postfrontal sulci continue in an almost

straight line from the midcranial sulcus (MS), joining the eye posteriorly. The occipital sulci (OS),

although evident in the Eriocraniidae, are obsolete

in Acanthopteroctetidae. Internally, the dorsal tentorial arms are slender and elongate in both families, extending nearly to the vertex immediately

behind the antennal insertions. Posterior to the

origin of the dorsal arms, the anterior tentorial

arms are variously developed in the Eriocraniidae

and may either be strongly developed (in Heringocrania) or barely discernible and threadlike (in

Dyseriocrania and Eriocrania). In Acanthopteroctetes the posterior half of the anterior arms is relatively well developed.

A pair of large, elongate chaetosemata are situated transversely across the occipital ridge of the

head. Similarly raised areas are present elsewhere

on the head (e.g., on the postmental area), which

also may represent chaetosemata.

The antennae are relatively short and simple. A

small, sometimes poorly defined intercalary sclerite

is present between the scape and pedicel. The external vestiture of the entire antenna consists

primarily of slender, elongate scales. Immediately

beneath these scales are scattered, highly modified

chemosensory sensilla, of the general type referred
NUMBER 251 11

FIGURES 11-16.—Dyseriocrania griseocapitella (Walsingham), adult head structure: 11, anterior

view (scale = 0.5 mm); 12, lateral view; 13, maxilla (scale = 0.5 mm); 14, lateral view of tentorium; 15, posterior view; 16, dorsal view.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 17-22.—Acanthopteroctetes bimaculata Davis, adult head structure: 17, anterior view

(scale = 0.5 mm); 18, lateral view; 19, maxilla (scale = 0.5 mm); 20, lateral view of tentorium;

21, dorsal view; 22, posterior view.
NUMBER 251 13

to by Jefferson et al. (1970) as sensilla auricillica.

Callahan (1975) discusses this and other forms of

antennal sensilla and explains their general function with analogies to dielectric antennae. Sensilla

auricillica probably occur in a wide variety of

moths and have been reported by the above authors

in the Noctuidae. I have also observed them present

in the Nepticulidae but found them lacking in the

Micropterigidae, Lophocoronidae, and Neopseustidae. The sensilla auricillica of Eriocraniidae are

relatively small with longitudinally grooved surfaces and multiple apertures, thus, similar in general structure to that figured by Callahan (1975) for

Heliothis. In Acanthopteroctetidae, these sensilla

are considerably larger and lack grooves (Figure

26) and are densely covered with minute pores

(Figure 34).

The eriocranioid labrum is relatively elongate

and somewhat ovate in outline with a subaccuminiate apex. It superficially resembles that of Micropterigidae and certain Agathiphagidae, except that

in these families the labrum is broader and of a

more quadrate outline. The pilifers with their

usual array of prominent setae are not pronounced.

The clypeolabral suture is usually well defined, although in Acanthopteroctetes this suture is barely

discernible. In the latter group, the labrum is

slightly reduced in size and more slender.

Although the adult mandibles have lost their

normal function in the Eriocraniidae and are partially membranous, their relative size is still

considerable in comparison to that of all other Lepidoptera, except the Micropterigidae and Agathiphagidae. More significantly, the internal apodemes

of the mandibles, particularly the adductor, are

relatively well developed (Figure 218). Studies by

Kristensen (1968b) suggest a glandular function for

the adductor apodeme in Eriocrania. The mandibles in Acanthopteroctetidae are only questionably

present. An examination of the genal (or mandibular) cavity that the mandibles normally occupy reveals only a slightly thickened ridge that appears to

be the sole vestige of mandibles in that family. The

mandibular apodemes are similarly reduced and

barely discernible. The mandibular condyles are

essentially absent in the Eriocraniidae although

slight indications of these still persist in some

species (Figure 219).

Although the maxillae in the Eriocraniidae exhibit a very primitive condition, they are decidedly

more specialized than those of the Agathiphagidae

and especially Micropterigidae in possessing a functional haustellum and in having largely lost the

lacinia. Kristensen has reported the lacinia as

vestigial, being represented perhaps by a membranous protuberance arising from the inner surface of

the stipes.

The maxillary palpi are elongate and consistently

five segmented with the fourth being the longest;

often it measures 2.0 to 2.5 times the length of the

fifth. The fourth segment characteristically possesses

a subapical, unpigmented zone. The apex of the

fifth segment is variously modified depending on

the species. The remainder of the segment is

densely covered by primary (i.e., the longest) and

secondary setae, numerous microtrichiae, and an

occasional scale. In Acanthopteroctetidae the apex

(Figure 217) is relatively unspecialized and rounded

with several, relatively elongate, divergent setae. In

the Eriocraniidae the apex is usually naked except

for typically eight, stout, sensory setae arising from

one to as many as five separate lobes. In the Nearctic eriocraniids the relative development of the

setae and their supporting lobes are closely correlated with other characteristics (e.g., in venation

and often male genitalia) and are helpful in delimiting genera. In the Palearctic species, however, a

few exceptions to these correlations have been

noted. For example, the apex of the fifth segment

in the genera Dyseriocrania and Eriocrania are similar in that there is a single apex (i.e., not divided)

usually bearing a crown of eight, short, stout setae

(Figures 185, 191). The apex is slightly more attenuated in Dyseriocrania with two or three sensory

setae situated more subapically. But in the European species Eriocrania sangii Wood and E.

haworthi, the apex is divided into two, nearly symmetrical lobes, each bearing four short setae (Figure 195). Superficially, the apex of E. sangii resembles that of Neocrania, new genus, except in the

latter the lobes are slightly subequal in length, and

the apical setae are more elongate with striated

sides (Figure 210). Under high magnification it can

be determined that the apical sensory setae of Eriocraniella and Neocrania possess longitudinal ridges

or striae (Figures 200, 214). The apical setae of

Dyseriocrania, Eriocrania, and Heringocrania lack

these striations and appear smooth.

Often correlated with these features of the apical
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 23-28.—Antennae (near middle of flagellum): 23. Dyseriocrania griseocapitella (Walsingham), fully scaled, X 35; 24-25, partially denuded, X 315 and X790; 26, detail of sensory

scale (sensillum auricillicum) X 7900; 27, Eriocrania breviapex, new species, fully scaled, X 600;

28, partialy denuded, X 825. (Photographs reduced to 83%.)
NUMBER 851 15

FIGURES 29-34.—Antennae {near middle of flagellum): 29, Eriocraniella variegata, new species,

partially denuded, X 390; 30, detail of sensory structure, X 3900; 31, E. aurosparsella (Walsingham), partially denuded, X 825; 32, sensory scales, X 5000; 33, Acanthopteroctetes bimaculata

Davis, partially denuded, X 1025; 34, sensory scale {sensillum auricillicum) X 5400. (Photographs

reduced to 83%.)
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lobes and setae is the barbed versus smooth condition of the primary setae of the fifth palpal segment. In those species with a divided apex (except

Eriocraniella platyptera, Eriocrania sangii, and E.

haworthi) and striated apical setae, the primary

setae are barbed along their outer edges (Figure

212). Conversely, in those species with an undivided

apex and unstriated sensory setae, the primary

setae are simple and smooth (Figure 213). Barbed,

primary setae exist in Eriocraniella and Neocrania,

whereas Dyseriocrania, Eriocrania, and Her in gocrania possess smooth primary setae.

The apex of the fifth palpal segment of Eriocraniella (Figures 198, 200) has been found to be

the most complex of the types examined. A single,

large apical lobe is present that bears typically one

long and three short striated setae. The remaining

four elongate setae all arise from four separate,

slender lobes arranged in two pairs. The apical

most pair of lobes is intermediate in size and the

most subapical pair is the smallest.

The eriocranioid haustellum, although relatively

short, is functional and is probably used to inbibe

liquid nourishment such as water and honeydew

secreted by certain Homoptera. Kristensen (1968b:

259) has noted, in living specimens drinking from

water drops on a petridish, that, although the

haustellum is normally united, it tends to split

apart when pressed against a surface. The haustellum is composed of two galeae, each bearing a

broad mesal groove, which when joined, form the

typical sucking tube or food canal in Lepidoptera.

The inner sides of the groove are completely lined

with relatively large plates (Figure 3), each distally

overlapping the basal edge of the succeeding plate.

The base of the labium is relatively simple and

uniform among the species in consisting of a small,

oval or triangular, slightly elongate median sclerite

or prementum. In Acanthopteroctetidae this sclerite

is somewhat more narrow. The anterior margin of

the prementum is usually slightly cleft. The labial

palpi differ greatly in structure between the two

families. In the Eriocraniidae they are elongate and

three segmented with all segments of approximately

equal lengths. The apex of the third segment bears

a noticeable sensory pit (organ of von Rath) that

often contains from 15 to 26 short, peglike, basiconic sensillae (Figures 215-216). These structures

are assumed to possess a chemosensory function. The

palpi in Acanthopteroctetidae are reduced to two,

very short segments that barely surpass the first

maxillary palpal segment in length. The second

segment is usually very short and less than one-half

the length of the first. The third segment has been

lost along with any evidence of an organ of von

Rath.

THORAX.—The general structure of the thorax

appears uniform throughout the superfamily. The

pronotum is quite small and mostly obscured dorsally by the mesoscutellum. The prosternum is extended anteriorly as a relatively slender sclerite

free from the pleuron. Its basal area is sometimes

slightly expanded laterally. The lateral cervical

sderites (LCS) are well developed, with broadly

triangular bases (viewed anteriorly) that taper to

slender rods anteriorly. The prothoracic spina and

furcal apophyses are moderately developed and

similar in form to those of Neopseustidae (Davis,

1975b).

The mesothoracic furcasternum possesses extremely short, almost obsolete apophyses (Figure

51). The katepisternum is well defined by a distinct paracoxal suture. The mesomeron is not

greatly reduced and approaches the meseucoxa in

size. The furcal apophyses of the metathorax are

well developed, arising subparallel and diverging

at their apices. Inside the metathorax, a broadly

compressed, anteromedial process (APM) extends

from the stem of the metafurcasternum toward the

intercoxal lamellae of the basisternum (Figure 48).

At this point the anterior end of the anteromedial

process is shortly divided with elongate, threadlike tendons extending into the coxae on either side

of the intercoxal lamellae. The anteromedial furcal

process in Acanthopteroctetidae is very similar in

structure except its anterior end is more distinctly

separated from the intercoxal lamellae (Figure 49).

It is possible that this structure (i.e., the anteromedial process) may be homologous with the mesal

lamella of Brock (1971, fig. 31b, etc.). The metakatepisternum is largely unsclerotized and reduced

in width. The metacoxae of the Eriocraniidae

differ from those of the Acanthopteroctetidae in

having the eucoxa more well developed and the

meron proportionately largely unsclerotized and

reduced in width. The metacoxae of the Eriocraniidae differ from those of the Acanthopteroctetidae

in having the eucoxa more well developed and the

meron proportionately smaller. The metamera of

both families are reduced and in that feature
NUMBER 251 17

resemble the reduced metamera of the Micropterigidae and certain ditrysian genera (Shepard,

1930).

The legs of both families are similar in retaining

a single apical spur on the mesothoracic tibiae. The

pro- and mesothoracic tibiae of the Eriocraniidae

are somewhat more reduced in length. The tarsi

(Figure 182) are of typical structure in possessing

a pair of simple claws and a slender empodium, a

broad, padlike arolium, and a lateral pair of relatively large, spinose pulvilli. Although an epiphysis is present in the genera Eriocraniella and

Neocrania, it has been lost in Dyseriocrania and

Eriocrania as well as in the Acanthopteroctetidae.

The wings of Eriocranioidea are moderately

slender with subacute apices. The fore and hind

wings are approximately equal in size but differ

in venation according to genus. The venation is

considered primitive for the Lepidoptera, particularly in the hind wings. The hind wings of all Eriocraniodea possess a four-branched radius with R3

absent (probably through fusion with R2). In

the forewings the radius may be either four or five

branched with Rl frequently subdivided. A relatively well-developed jugum is present. The subcosta is sometimes subdivided in the forewings but

normally not in the hind wings. Another primitive

venational feature in this superfamily is the preservation of the base of the medial vein within the

discal cell of both wings, a plesiomorphic trait

that persists into certain primitive ditrysian families (e.g., many Tineoidea). As is true for nearly

all members of Zeugloptera, Dacnonypha, and

Monotrysia, all surfaces of both wings are uniformly and densely covered with microtrichiae. Besides being somewhat more slender, the wings of

Acanthopteroctetidae differ significantly from those

of the Eriocraniidae. In the former, vein Ml is

stalked with the stem of R4+5 in both wings, and

the anal veins are completely separate in the forewings. In the Eriocraniidae, all three anal veins

fuse distally in the forewing, and Ml arises separate. A pterostigma is also typically evident in

particularly the forewings of Eriocraniidae but is

absent in Acanthopteroctetidae.

The broad discal scales of the forewings differ

significantly in structure between the two North

American families of Eriocranioidea. In the Eriocraniidae, most of the scales over the dorsal wing

surface (i.e., the broader scales) possess minutely

serrated apices and are of a nonperforated type

(Figure 36-37) with a dense array of transverse

striae interconnecting the rather prominently

raised, longitudinal ribs. In cross section, as viewed

through the Transmission Electron Microscope,

these scales resemble a very thin, approximately

88 nm thick (Figure 45) plate, resulting from the

fusion of the dorsal and ventral cuticle. In the

ditrysian Lepidoptera, these surfaces are well separated in nearly all wing scales, thereby creating a

relatively large internal lumen that normally is

in direct contact with the surrounding air through

the dorsal perforations, or lacunae. As Kristensen

(1970) first noted, some variation in scale structure exists over the wing with the marginal, more

hairlike scales frequently possessing a well-defined

lumen (Figure 46). Even in these scales, however,

no dorsal lacunae are present in the Eriocraniidae.

Also evident in Figure 46 and characteristic for the

slender marginal or fringe scales of most Lepidoptera are the presence of prominent ventral ribs. The

latter are lacking in the typically broad lepidopteran

scale, with the ventral surface being instead quite

flat.

Another prominent feature of the typical eriocraniid scale is that the longitudinal ribs extend

slightly beyond the terminal margin of the scale

(Figure 37). In this respect they agree with the

families with which they are most closely allied

(Agathiphagidae, Neopseustidae, and Lophocoronidae), as well as with the Micropterigidae

(Figures 38-40).

In contrast to the above, the discal scales in Acanthopteroctetidae possess prominently dentate apices

in which the ribs do not extend beyond the apex

of the scale (Figure 42). In addition, all scales

(including the more hairlike ones) are of the perforated type with a well-developed internal lumen

(Figure 44).

In both families the scales of the ventral surfaces

of the wings and of the dorsal surface of the hind

wings are typically more slender, and in the case

of the latter, frequently hairlike. The dorsal scales

in the hind wings of Dyseriocrania, Eriocraniella,

and Neocrania are relatively broad.

ABDOMEN.—The first abdominal segment in this

superfamily, as is true in varying degrees in all

Lepidoptera, has undergone reduction. The tergite

is largely membranous, with sclerotization largely

restricted to the lateral margins. The first sternite
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 35—40.—Wing scales {from dorsal surface of discal cell of forewing): 35, Dyseriocrania

griseocapitella (Walsingham), X 500; 36, detail of scale in Figure 35, X 7500; 37, apex of scale,

X 7500; 38, Lophocorona pediasia Common, apex of scale, X 10,000; 39, Agathiphaga queenlandensis Dumbleton, apex of scale, X 7500; 40, Epimartyria auricrinella Walsingham, apex

of scale, X 7500. (Photographs reduced to 83%.)
NUMBER 251 19

45 46

FIGURES 41-46.—Wing scales: 41, Acanthopteroctetes bimaculata Davis, dorsal scales from discal

cell of forewing, X 500; 42, apex of scale in Figure 41, X 5000; 43, detail of scale in Figure

41, X 15,000; 44, cross-sectional view of scale from Figure 41, X 10,200; 45, Dyseriocrania

griseocapitella (Walsingham), cross-sectional view of scale from Figure 35; 46. cross-sectional

view of slender scale from marginal fringe of forewing, X 10,200. (Photographs reduced to 83%.)
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SLM2

SE3

SM2

TA

PLS3

47

- A MN3

— V EA3

49

FIGURES 47-49.—Thoracic structure: 47, Acanthopleroctetes bimaculata Davis, lateral view

of thorax; 48, Dyseriocrania griseocapitella (Walsingham), lateral view of metafurcasternum;

49, A. bimaculata Davis, lateral view of metafurcasternum.
NUMBER 251 21

52

FA

55

FIGURES 50-55.—Caudal view of thoracic segments: 50, Dyseriocrania griseocapitella (Walsingham), prothorax; 51, mesothorax; 52, metathorax; 53, Acanthopteroctetes bimaculata Davis,

prothorax; 54, mesothorax; 55, metathorax. (Scale = 0.5 mm.)
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

is extremely reduced, resulting in fragmented remnants that are difficult to homologize. In Dyseriocrania (Figure 56) the first sternite appears to be

represented by a small, strongly bifid median plate

with slender arms extending laterally. Immediately

posterior to this is another, extremely narrow, crescentic sclerite that is, however, more elongate and

interrupted at the midline. Each arm of the latter

sclerite is superficially connected medially to the

second sternite and fused laterally to the apices of

the anterolateral processes of the second sternite.

The medial connection in most species is by means

of a small but well-defined furcate plate. The second, narrow sclerite, thus, may either represent a

fragmentation of the first sternite or, more likely,

the anterior edge of the second sternite. Consequently, as interpreted in this paper, the second

sternite consists of a narrow, anterior margin separated from the main body of the sternite by a

broad intrasegmental membranous area. The main

body of the second sternite is rather broad and

possesses a bilateral pair of slender, relatively

elongate anterolateral processes referred to above.

These processes, together with the fused apices of

the anterior margin of the second sternite, are

weakly articulated laterally to a bilateral pair of

posterolateral processes of the first tergite that are

situated well posterior of the first abdominal spiracles. There is no evidence of a prespiracular connection.

In Acanthopteroctetes the first abdominal tergite

is similar in shape and degree of sclerotization to

that of Dyseriocrania. The first sternite (Figure 57),

however, is less modified and consists of a simple,

narrow band. Immediately posterior to this is a

second more narrow band extending completely

across the venter. Unlike its homologue in Dyseriocrania this sclerite is entirely free from the main

body of the second sternite. Also, unlike Dyseriocrania, the anterolateral angles of the second sternite are not produced, resulting in an even weaker

connection with the posterolateral process of the

first tergite. In general, the posterolateral processes

of the first tergite are also less developed in this

genus.

The third abdominal segment is usually unmodified in the Eriocranioidea other than the presence

of paired hyaline areas on the sternite of certain

species. The male of Acanthopteroctetes unifascia,

new species, however, possesses a bilateral pair of

unusual, pedunculate organs arising laterally near

the third abdominal spiracles. These structures

(Figure 165) consist of a triad of clavate filaments

originating from a small, oval sclerotized plate. The

enlarged terminal end superficially appears glandular in composition, although the exact function of

these peculiar structures remains unknown. Similar

organs are absent in the males and females of other

species of Acanthopteroctetes. A determination of

their existence in the opposite sex of A. unifascia

will have to await the discovery of a female specimen.

The fourth and fifth abdominal sternites of most

Eriocaniidae are variously modified in association

with the presence of specialized, paired abdominal

glands. These organs also occur in certain Trichoptera, and their general distribution among Lepidoptera has been reviewed by Philpott (1925),

Razowski (1975a), and Davis (1975b). The sternal

modifications of the fourth segment in the female

typically resemble oval to circular, membranous

windows, or fenestrae (Figures 59, 60) in the otherwise darkly sclerotized sternite. In the North American Eriocraniidae, fenestrae are absent from the

fourth sternite in the males but are normally present in all females except Neocrania bifasciata, new

species. Internally, the fenestrae are intimately associated with the small, globular reservoirs of the fifth

sternal glands (Figure 59). In dissections of female

abdomens of Dyseriocrania griseocapitella fixed

in KAAD, the reservoirs were observed closely

pressed against the inner wall of the fenestrae,

held in place by strands of connective tissue and

overlaid by fatty deposits. Thus, the fenestrae have

no independent secretory function but merely appear to demarcate the presence of internal glands

opening on the following sternite. The reservoirs

in male D. griseocapitella (as well as in other male

eriocraniines) appear to be as closely associated

with the inner wall of the fourth sternite as in the

female, but for some reason are rarely delimited

by fenestrae externally. The same is probably true

for females of Neocrania bifasciata. The external

surface of the fenestrae (Figure 61) differs markedly

from the rest of the sternite in lacking microtrichiae.

The relative density of the body scales, however,

is the same as on the surrounding area.

Similar, typically paired, hyaline areas can also

be found on other abdominal segments of various

Lepidoptera. For example, such a pair may be
NUMBER 251 23

58

FIGURES 56-59.—Abdominal structure: 56, Dyseriocrania grheocapitella (Walsingham), ventral

view of first and second abdominal sternites. 57, Acanthopteroctetes bimaculata Davis, ventral

view of first and second abdominal sternites; 58, D. griseocapitella {Walsingham), lateral view of

first and second abdominal segments; 59. internal view of fourth and fifth abdominal sternites

and fifth abdominal gland.

59

observed in various stages of development in the

males and females of some species of Eriocraniidae

near the anterior margin of the third sternite.

These are particularly evident in Neocrania bifasciata (Figure 243).

The sternal glands are completely separated internally and discharge through a pair of variously

modified tubercules located near the anterior margin of the fifth sternite. Two distinct forms of

tubercules are known in this family. The sternal

tubercules of Dyseriocrania are unique in resembling greatly enlarged, flattened pads (Figure 62).
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 60-65.—Abdominal structure: 60, Dyseriocrania griseocapitella (Walsingham), fenestra

and tubercule of fourth and fifth sternites respectively, X 170; 61, surface detail of fenestra,

X 1075; 62, reniform sternal tubercule, X 420; 63, detail of Figure 62 showing aperture of

fifth sternal gland, X 2100; 64, Eriocraniella falcata, new species, fenestra and tubercule (see

arrow) of fourth and fifth sternites respectively, X 280; 65, papillose sternal tubercule, X 100.

(Photographs reduced to 84%.)
NUMBER 251 25

In all other eriocraniid genera, the tubercules are

reduced in size and are papiliform. In both types

the external surface of the tubercules is covered

with a dense reticulated pattern of interconnecting

grooves radiating outward from a centrally located

pore (Figure 63). The likely function of such a

finely divided surface is to provide a greater overall

surface area and a more efficient means of dispersing the secretions of the glands. Because the glands

are usually present in both sexes (if present at all),

it would seem most likely that they serve a defensive purpose. At least one Japanese eriocraniid

species examined and all Acanthopteroctetidae lack

any evidence of sternal tubercules or fenestrae.

The eighth segment in the male is unspecialized

and is composed of a well-defined tergite and sternite. In the female, these two sclerites are largely

united to form a relatively large, cylindrical cone

that normally encloses the ninth and 10th segments

(i.e., the ovipositor). The eighth segment in the

females of Acanthopteroctetidae is very similar to

that of the Eriocraniidae and differs only in the

possession of a single row of elongate setae from

the caudal margin of the former. A prominent,

apparently functional spiracle is present in both

groups immediately dorsal to the basal insertions

of the anterior apophyses.

MALE GENITALIA.—Several structures in the male

genitalia differ greatly between the two families.

Among the small number of species within each

family, however, the genital morphology is remarkably similar. In the following discussions and descriptions I have attempted to use terms familiar

to lepidopterists (as defined by Klots, 1970) even

though the exact homologies of some structures are

questionable.

The ninth segment in the Eriocranioidea resembles that of a few other primitive groups in being

cylindrical and undifferentiated into a distinct,

dorsal tegumen or ventral vinculum. Although the

latter sclerites are not well defined in this superfamily, for the sake of brevity, these terms have

been used in the following taxonomic descriptions

to designate the general dorsal and ventral areas

of the ninth segment. The anterolateral angles of

the vinculum in the Eriocranioidea are either

merely sharply defined (as in Acanthopteroctetidae)

or greatly elongated into a prominent pair of

apophyses. Somewhat similar developments in the

vinculum are also evident in varying degrees in

such families as the Neopseustidae and certain species of Nepticulidae. In one subgenus of Eriocraniella, a prominent furcate process, or furcula (Fra),

arises midventrally near the caudal margin of the

vinculum. Its function is unknown; possibly, it

plays some role during copulation and may be

brought into contact with the spinose pocket on

the ninth segment of the female, as discussed later.

The 10th tergite, or uncus, is well developed in

all species. It differs in form between the two families with that of the Eriocraniidae being more

prominently divided or lobed. Any evidence of a

sternite (gnathos, etc.) is lacking.

Often arising near the caudal margin of the

ninth tergite and from the intersegmental region

between the valvae and the base of the uncus is a

bilateral pair of small digitate lobes, each bearing

a single, elongate seta. These structures are typically present in Eriocrania, Eriocraniella, and Neocrania but absent in Dyseriocrania and Acanthopteroctetes. They have also been secondarily lost in

Eriocrania sparmannella, (Birket-Smith and Kristensen, 1974). Largely because of their position (i.e.,

ventrad to the base of the uncus), they have been

regarded as socii in the present paper. Because of

the general absence of socii throughout the homoneurous families (with the possible exception of

the Neopseustidae), however, the true homology of

the socii in the Eriocraniidae is questionable. In

their study of the male genitalia of Eriocrania,

Birket-Smith and Kristensen (1974) have considered

these structures to be basal processes of the valvae.

The valvae also differ greatly in form between

families but are very uniform within each group.

Those of the Eriocraniidae are greatly reduced and

probably incapable of appreciable movement. Birket-Smith and Kristensen (1974), however, have

found the valvae of Dyseriocrania subpurpurella

to be movable by muscles inserted on their bases.

The valvae of Acanthopteroctetidae are considerably more elongate, freely movable, and possess a

peculiar membranous fringe along portions of the

ventral margin.

One of the most complex areas of the male genitalia involves various sclerotizations in the diaphragma surrounding the aedeagus. In the Acanthopteroctetidae, these consist of two distinct sclerites that I have referred to as the transtilla (dorsal

to aedeagus) and juxta (ventral to aedeagus). The

transtilla (Figure 289) in this genus is in the form
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of a heavily sclerotized transverse arch extending

between, but separate from, the bases of the valvae.

An elongate, similarly sclerotized process extends

caudally from the center of the arch to the apex

of the uncus. The apex of this median process typically curves dorsally to form an uncinate lobe in each

of the three known species. The main purpose of

this process is probably to serve as a supporting

guide for the aedeagus. The juxta in this subfamily

lies relatively free along the ventral side of the

aedeagus. It tends to be long and spatulate but

differs in form and the relative amount of sclerotization among the known species.

A transtilla is entirely absent in the Eriocraniidae.

The juxta is present but consists of a much more

complex morphology. Its structure is described in

detail by Birket-Smith and Kristensen (1974)

wherein it is referred to as the median plate. As

shown by these authors, the lateral edges of the

juxta are continuous with the membranous walls

of the genital cavity, thus separating this cavity

into a dorsal genital chamber and a smaller, ventral

subgenital crypt.

The 11th segment is largely represented by the

anal tube. Usually the tube is entirely membranous,

but occasionally it may contain numerous minute

spinules within its walls (in Dyseriocrania) or small

bilateral sclerites basally (in Eriocrania). The latter may represent vestiges of the 11th sternite, as

suggested by Birket-Smith and Kristensen (1974).

The aedeagus in the Eriocraniidae is prominently

divided into a primary dorsal branch (containing

the vesica) and a secondary ventral branch. The

vesica lacks well-developed cornuti, although a few

scattered minute spines do sometimes appear. According to Birket-Smith and Kristensen (1974) both

phallic branches are inserted during copulation.

The base of the aedeagus is typically inflated and

bulbous with the ventral branch often loosely

articulated. The ventral branch tends to be more

varied in form between species and may be variously lobed or even provided with apical external

spines or cilia.

Internally, the male reproductive system in

Dyseriocrania continues as an elongate, partially

coiled ejaculatory duct (ED) extending anteriorly

from the base of the aedeagus (Figure 66). Near its

termination this duct divides but remains contiguous until the paired ducts completely separate near

the junction of the inflated, slightly curved region

As

FIGURE 66.—Male internal reproductive system of Dyseriocrania griseocapitella (Walsingham).
NUMBER 251 27

of the vas deferens, or vesicula seminales (VeS).

Beyond the latter the vasa deferentia constricts to

a very slender duct leading to a reduced, relatively

indefinite mass of tissue, the testis, which is largely

enclosed by fat body cells. The interpretation of

the testis as being present in such a vestigial condition in the Eriocraniidae has been suggested by

Kristensen (pers. comtn.), who will discuss his findings more fully in a forthcoming publication. One

of the most revealing discoveries by Kristensen to

support the above interpretation was the presence

of spermatozoa in the vesicula seminales and testes

and their pronounced absence elsewhere in the reproductive system.

Anterior to the junction of the paired vesicula

seminales continue two elongate, somewhat convoluted ducts that lead to a firm, bilobed mass of

tissue resembling the more typical lepidopteran

testes but interpreted herein as the accessory glands.

Gross dissections of this organ failed to reveal any

indication of testicular follicles, thus supporting

Kristensen's belief that the organ does not represent the true testes.

In the Acanthopteroctetidae the aedeagus is not

divided and appears more simple except for a prominent array of endophallic cornuti. The particular

arrangement of the cornuti varies according to

species. The base of the aedeagus is relatively simple and not swollen.

FEMALE GENITALIA.—The principal features of

systematic importance in the female reproductive

system have been recently reviewed by Stekol'nikov

(1967), Mutuura (1972), Dugdale (1974), and Davis

(1975b). In the Eriocranioidea, I have interpreted

the ovipositor as consisting of structures derived

from the ninth and 10th abdominal segments, normally enclosed largely by the eighth segment. In this

regard I thus agree with the interpretations of Chapman (1917) and not with Philpott (1927). As already stated, the eighth segment is enlarged, conical

in form, and normally (at rest) contains the retractible ninth and 10th segments. Two sets of

elongate, rodlike apophyses are present that enable

the female to insert her eggs under the leaf epidermis. The anterior apophyses arise from the eighth

segment immediately below the eighth spiracle. The

posterior pair originate in the 10th segment and

are fused distally to form the actual piercing organ.

The fused apex is strongly depressed and equipped

with a lateral, subapical series of usually seven to

12 minute teeth. Closely associated with the apex of

the posterior apophyses in Eriocraniidae is an accessory pair of short, tertiary apophyses (TAp) that

are ventrally situated within the largely membranous body wall of the 10th segment. As early suggested by Wood (1891:178), these flattened rods may

represent remnants of the 10th sternite. The tertiary apophyses lie parallel to one another and more

or less laterad of the cloacal aperture. Whereas

their caudal ends appear free, anteriorly the tertiary

apophyses are weakly connected to a minute digitate process or articularly process (AP), arising

ventrally and well anterior of the apex of the posterior apophyses. Although articulary processes are

absent in Acanthopteroctetidae, tertiary apophyses

are present but even more poorly defined.

The ninth segment in female Eriocraniidae is

poorly defined and largely membranous. In one

subgenus of Eriocraniella, however, a specialized

midventral invagination has developed near the

caudal end of the ninth segment. The lateral margins of this shallow depression, herein referred to

generally as the spinose pocket (SPt) are typically

densely covered with minute spines (Figures 310,

317). Its function is not known, but it may be involved in copulation as hypothesized under the

discussion for the genus Eriocraniella.

Only one posterior body opening is present in

the females of this superfamily and that is situated

ventrally on the 10th segment (and not the eighth

as stated by Mutuura, 1972), slightly anterior to the

extreme apex of the ovipositor. Through this common opening, or cloacal aperture, all reproductive

and alimentary functions pass. A rather elongate,

well-defined cloaca is present, formed by the junction of the lower alimentary canal, or hindgut,

with the vagina.

The typically enlarged portion of the vagina is

sometimes referred to as the vestibulum. In the

Eriocraniidae, a highly specialized, variously modified vaginal sclerite (VS) encircles the vaginal passage. Normally the vaginal passage is constricted at

the point where it passes through the aperture of

the sclerite. Because of its unusual position, the

vaginal sclerite was simply termed the "valve" by

Philpott (1927). Stekel'nikov (1967) used the same

terminology but somewhat incorrectly stated that it

was located at the base of the common oviduct. As

noted by Stekel'nikov, large muscles occupying much

of the eighth segment are attached to the vaginal
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

CG

SPE8

T8

FIGURES 67-70.—Female reproductive system: 67, Dyseriocrania griseocapitella (Walsingham);

68, apex of ovipositor; 69, Acanlhnpterocteles bimaculata Davis, detail of spermathecal papilla;

70, A. bimaculata Davis. (Scale = 0.5 mm.)
NUMBER 251 29

sclerite, particularly to the variously modified anteroventral half. In some eriocraniids this end is

broadly depressed; in others, a large, compressed

keel-like process develops ventrally. Although variable within a few species, the form of the ventral

keel usually differs considerably between species

and, consequently, can be most useful in species

identification and, in some instances, generic delimitation. For example, the vaginal sclerite in

Eriocrania characteristically possesses a sclerotized

median process (MP) extending caudally from the

base of the midventral keel; this process apparently

does not occur in other genera.

Dorsal to the vaginal sclerite arises the slender

duct from the usually bifid colleterial glands (CG).

The anterior end of the vagina is marked by the

junction of both the ventral common oviduct (CO)

and the dorsal ductus spermathacae to the caudal

end of the ductus bursae. In the Eriocraniidae, the

ductus spermathacae is sinuate and possesses a

rather prominent, elongate internal thickening.

The spermathecal papilla (SP) is mostly membranous and cupuliform.

In the Acanthopteroctetidae, the vagina shows no

indication of a specialized sclerite and tends to be

much larger in diameter. The walls are considerably thicker and heavily folded, with one or more

secondary lobes projecting in an irregular manner.

In Acanthopteroctetes bimaculata, the common oviduct appears to emerge laterally from such a lobe

(Figure 330). Probably what has occurred is that the

normal, ventral junction of the common oviduct

has migrated to a lateral lobe of the vagina as this

lobe developed. As summarized by Dugdale (1974)

and Davis (1975b), a ventral origin of the common

oviduct is clearly the primitive condition within

the panorpoid orders, and in no known instance

does this passage arise laterally from the reproductive tract. In another but unnamed species of

Acanthopteroctetes, the vestibulum (i.e., vagina) is

even more irregularly lobed (Figure 332) and possesses a rather slender tube, assumed to be the common oviduct, arising from a more typical midventral position.

The spermathecal papilla is variously modified

in the Acanthopteroctetidae and may be either

membranous, as in the Eriocraniidae, or darkly

sclerotized (Figure 69). The general structure of the

spermatheca is similar to that of the Eriocraniidae

in possessing a relatively small, elongate utriculus

(Us) with a terminating and very slender, receptacular gland (RG). The spermathecal duct, however,

does not possess any internal thickening.

Although no signum appears within the corpus

bursae of any eriocranioid species, the walls of at

least one species of Dyseriocrania are finely covered

with a scattering of minute spinules (Figure 296).

PUPAL MORPHOLOGY

Ever since Stainton described and illustrated a

pupa of Eriocrania }purpnrella Haworth in 1863,

the unusual morphology of the eriocraniid pupa

has attracted wide interest. The completely functional, rather grotesque mandibles (Figures 71-78)

are the most spectacular feature of this stage. Superficially the eriocraniid pupae resemble the decticous

pupae of many Trichoptera, but, in the case of the

latter, the mandibles are normally smaller and function in a more opposable manner. The broad, truncate apices of the eriocraniid mandibles are particularly striking and bear closest resemblance to those

found in agathiphagid pupae. The unusual form

of the mandibles probably assists the pharate adult

in pushing its way from its underground cocoon to

the soil surface. Except for the heavily sclerotized

mandibles and certain structures that serve to

strengthen the head, the eriocraniine pupal cuticle

tends to be very thin and flexible. All appendages

of the body are free and usually capable of varying

degrees of movement; even the head is capable of

some motion.

Another prominent feature of the head is the

elongate frontal ridge that protrudes free of the

head as far as the labrum. This ridge, or cocoon

cutter, undoubtedly plays some role in assisting the

mandibles to tear through the wall of the subterranean cocoon. The labrum is well defined and

bears at least four radiating pair of elongate setae.

The longest of these setae extend almost to the

apices of the mandibles. Another pair or two of

elongate setae arise from the vertex of the head.

From the little comparative data available thus far,

the chaetotaxy of the head seems very uniform.

Likewise, the mandibles appear very similar in

structure, judging from the only two species available for study (Dyseriocrania griseocapitella and

Eriocrania species, perhaps scmipurpurella). For

example, a comparison of the highly modified

mandibular apices of both species (Figures 76, 78)
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 71-74.—Pupa of F.riocrania 'fsemipurpurclla seniipurpurella (Stephens) from England:

71, ventral view, X 34; 72, lateral view, X 34; 73, ventral view, X 60; 74, lateral view, X 60.

(Photographs reproduced at 100%.)
NUMBER 251 31

FIGURES 75-78.—Pupal structure: 75, Eriocrania hemipurpurella seinipiirpurella (Stephens),

detail of mouthparts of Figure 71, X 130; 76, apex of mandible, X 320; 77, Dyseriocrania

griseocapilella (Walsingham), lateral view of mandible, X 120; 78, apex of mandible, X 240.

(Photographs reduced to 94%.)
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

reveals essentially the same arrangement of four

progressively larger spines and a slightly raised

marginal ridge surrounding a relatively broad, flattened distal end. The only difference detected was

in the minutely serrated ridge of Eriocrania species

compared to the entire or smooth condition of the

ridge in D. griseocapitella. The inner surface of

the mandibular shafts of both species appears similarly serrated.

As pointed out by Hinton (1946a), the decticous

pupa has no muscles of its own. Thus, the muscles

that move the pupal mandibles belong to the adult.

These are attached to the mandibular apodemes of

the adult, which in turn encase (except at their

juncture with the pupal mandibles) the pupal

apodemes that actually project into the head of the

adult. Consequently, the situation in the genus

Acanthopteroctetes, wherein both the adult mandibles and mandibular apodemes appear extremely

reduced, may indicate a reduction and even loss of

function of the pupal mandibles. Certainly, it will

be most interesting to examine the pupal stage of

this genus, whenever it is discovered, for possible

reduction or loss of articulation of the mandibles.

The cases enclosing the developing imaginal

wings are elongate and frequently extend to the

caudal end of the abdomen. The tarsal segmentation, as is true for the antennae, is clearly defined.

Often even the scales of the wings and body are

visible through the thin, membranous cuticle. As

shown by Mosher (1916), the chaetotaxy of the

abdomen is relatively simple, typically consisting of

three pairs of dorsal setae over the first eight abdominal segments. All of the abdominal segments

are capable of independent movement except for

segments VIII-X, which are fused.

LARVAL MORPHOLOGY

Most of the larvae examined during the course of

this study and described in the text are believed to

represent the final, or fourth, instar. In the case of

Eriocrania }semipurpurella pacifica, new subspecies,

however, it is possible that only third instar larvae

were studied. From the few species examined, it was

apparent that the chaetotaxy of the last two, and

possibly three, instars was essentially identical;

however, it was noted for some species that younger

larvae tended to be more heavily pigmented and

may even possess darkly sclerotized tergal plates

over the abdomen. Because the immature stages of

the Acanthopteroctetidae are unknown, the following remarks pertain only to the Eriocraniidae.

Although the larvae of Eriocraniidae are leafminers throughout their entire feeding cycle and,

consequently, do demonstrate several specializations

for this behavior, they are not as highly specialized

as many leaf-mining Lepidoptera. For example, in

general form they superficially resemble the last

instar (i.e., the spinning larva) of many Gracillariidae and Tischeriidae in that the head and body

are cylindrical and not strongly depressed. They

further resemble the Tischeriidae in possessing a

well-developed spinneret throughout most of their

larval life and in the possession of spinose tufts or

brushes on the labrum and mandibles. The relatively few (i.e., four) instars exhibit no evidence of

hypermetamorphism, although the first instar is yet

to be studied.

HEAD.—The head capsule is partially retracted

into the thorax, resulting in the extreme reduction

of most of the posterior cranial setae. Several of the

cranial setae are so reduced that it is difficult to

examine them without resorting to either interference or electron microscopy. Frequently, for example, distinguishing between setal pores and setae

becomes a serious problem. The vertex is deeply

divided with the coronal notch extending to the

frontal sclerite, or frons. The adfrontal sutures are

prominent and externally mark very broad and

thickened, inflected ridges internally. One of the

diagnostic features of the head is in the development of the ecdysial lines, or frontal sutures, sometimes incorrectly referred to as as the adfrontal sutures (Hinton, 1948). In the Eriocraniidae these

sutures are very distinct and extend anteriorly from

the coronal notch around the antennae, terminating

on the posterior rim of the antennal socket. As a

result, the adfrontal sclerite is sufficiently broad

anteriorly to include seta Al. The latter is occasionally absent, thereby providing a useful character in

species determination. The ecdysial suture itself

forms a very irregular and often rather widely

separated fissure. In some specimens, the fissure is

broad enough to include certain marginal setae as

AF2 and PI. The latter is often greatly reduced in

size comparable to those of the V series. The North

American species examined all agreed in having

lost A3 and the third ocellar seta. In contrast, Hinton (1946, figs. 10-11) shows these to be present in
NUMBER 251 33

at least one European species. Two very minute

gular setae are consistently present, although the

pores associated with these setae appear variable

in number as well as position.

As shown by Jayewickreme (1940), the internal

structure of the head is unusual in possessing an

extremely delicate tentorium. The anterior tentorial arms are further peculiar in being attached

to small, anteriorly directed spurs of the epistomal

ridge.

The ocelli (more properly termed, stemmata) are

reduced to a single, vestigial element immediately

beneath the antennae (Figure 158). Although it normally appears as a darkly pigmented spot, the lens

is usually not well defined by a distinct margin. The

cellular structure of the ocelli in Eriocrania semipurpurella was discussed in some detail by Hackman (1942). This author observed that the primary

nerve leading to the ocellus possessed a rudimentary branch, thus suggesting the existence of a

second ocellus at one time.

The antennae (Figure 91) are very short and

appear two-segmented due to the reduction of the

first segment. All the species examined possessed

essentially the same antennal chaetotaxy. The

second segment is reduced in length and bears two

prominent apical setae, one more than twice the

length of the other. Also arising from the second

segment near the base of the third are two short

basiconic sensillae and one minute spinelike seta.

The small, very abbreviated third segment bears

one basiconic sensilla and one minute spinelike seta

of approximately the same length but of a much

smaller diameter.

The labrum appears relatively normal in form

except for the very spinose condition of the anterior

margin (Figure 87) and ventral surfaces. Although

most of these accessory spines may originate from

the underlying "epipharynx," several appear to

arise from the anterior margin of the labrum itself.

Although, as Jayewickreme (1940) has observed,

three pairs of epipharyngeal setae and several

supernumerary ventral spines are typically present,

true epipharyngeal spines are absent. The latter,

when present in Lepidoptera, normally arise from

the dorsal wall of the buccal cavity immediately

posterior to the epipharyngeal setae. The primary

labral setae consist of six pairs with the lower ones

distinguishable from the adjacent spines by their

socketed bases. The lower three pairs of setae (L3,

M2, and M3) may either be simple (Figure 87) or

broadly compressed (Figure 155).

The mandibles are relatively unmodified and

function in an opposable manner. A total of four

cusps are consistently present with the fourth being

considerably broader and less pronounced than the

others. In addition to the normal pair of lateral

setae, some species also possess a specialized tuft

of barbed setae arising from the inner, mesal edges

(Figures 95-100). Similar setal tufts are also present

in the larvae of Tischeria.

The hypopharynx and portions of the maxillae

are also densely covered with filamentous spines

(Figure 93). A majority of the hypopharyngeal

spines are barbed. Most, if not all, of these spinose

structures, including those referred to above on the

labrum and mandibles, probably serve a collective

purpose in monitoring the passage of both solid

and liquid food into the buccal cavity. As suggested

by Tragardh (1913), Jayewickreme (1940), and

Gerasimov (1952), some of these structures may

also aid in the absorption of plant liquids. The

maxillary palpi are three segmented, with the apex

of the third either truncate or terminating in a

shallow apical depression containing usually seven

to eight rather uniformly short, basiconic sensilla.

The second segment bears usually three elongate,

rather broad spines that arise from the dorsolateral

rim at the apex. The palpigers possess a single,

moderately large, ventral apical spine and three to

four considerably smaller, lateral apical spines.

The apices of the galeae resemble those of the

maxillary palpi in being truncate with usually

three short, basiconic sensilla. The galeae, however, are more reduced in length, and the apical

sensilla tend to be more scattered. The relative

lengths of the sensilla occasionally differ between

species, with, for example, those of Eriocraniella

aurosparsella (Figure 127) being larger than those

of Dyseriocrania griseocapitella (Figure 90). Dorsally, the basal areas of the galeae are covered with

several, moderately large, fleshy spines. The stipes

bears one large basal seta and one much smaller

distal seta.

The labium is relatively well developed in all

instars studied. The submentum is rather unusual

in having lost all setae. In most lepidopterous

larvae, a long pair of midventral setae arises near

the lateral margins of the submentum. The labial

palpi typically consists of a single, elongate basal
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

segment and a minute apical segment with two,

elongate apical setae. Tragardh (1913) reports a

three-segmented labial palpus in Eriocrania species

(possibly Dyseriocrania subpurpurella). A rather

indefinite groove (Figure 89) is sometimes present

around the basal area of what is termed herein as

the first palpal segment, which may represent a

vestigial basal segment. From all indications, the

spinneret is present throughout the life of the

larva. Its function, however, may be restricted only

to the last (fourth) instar as indicated by Tragardh

(1913). In form it resembles an elongate cone, with

a simple, circular orifice.

THORAX.—The thorax of later instar larvae

normally exhibits brownish pronotal and prosternal

plates with the remainder of the thorax being

whitish in color. Although thoracic legs are absent,

paired callosities, or calli, that assist in locomotion

are present. These are particularly evident on both

the dorsal and ventral surfaces of the meso- and

meta-thorax. The prothoracic subventral group of

setae is of interest in that some North American

species (e.g., Dyseriocrania auricyanea and D.

griseocapitella) possess all three setae. This is in

contrast to Hinton's statement (1946b:20) that the

SV series is always bisetose in Lepidoptera. The

arrangement and number of the microscopic proprioreceptor setae of the thorax and abdomen are

constant in all the species examined and agree with

those listed by Hinton (1946b:21).

ABDOMEN.—The abdominal setae, similar to those

of the head and thorax, tend to be reduced in

length. The prolegs and crochets are completely

absent. Generally the abdomen is whitish in color,

at least in the later instars. The cuticle is densely

spinose (Figure 94) over large areas of both the

dorsal and ventral surfaces of the body. Minute

spines are particularly evident over the thoracic

calli and are a further aid to locomotion. Seta LI

is often diagnostic in either being situated unusually high on the segment well above the

spiracles and SD1, or totally absent.

The chaetotaxy of the ninth segment is somewhat

questionable, especially in Dyseriocrania griseocapitella. This species demonstrates the most primitive setal pattern examined in that the ninth segment shows no setal reduction from that of most of

the preceding abdominal segments. Normally in this

family, only seven primary setal pairs exist on the

penultimate segment, but in D. griseocapitella, 10

pairs remain. The most controversial nomenclature

involves the setae SD1, LI and SD2 (if present).

In this paper I have followed the more typical

AA

AC

AG

Am

AP

APM

As

AT

ATP

B

BaS

BS

CA

CB

Cdo

CG

CO

cs

Cta

DA

DB

Ea

ED

En

FA

Fm

Fna

Fra

IL

IS

J

K

LfS

LCS

Lpa

Me

MK

ABBREVIATIONS

Anterior Apophyses

Anapleural Cleft

Accessory Gland

Anepisternum

Articulary Process

Anteromedial Process

of Metafurcasternum

Aedeagus

Anal Tube

Anterior Tentorial

Pit

Basalare

Basal Sclerite of

Anal Tube

Barlike Spot (Bracerod)

Cloacal Aperture

Corpus Bursae

Cardo

Colleterial Gland

Common Oviduct

Coxal Suture

Chaetosema

Dorsal Arm of

Tentorium

Dorsal Branch of

Aedeagus

Eucoxa

Ejaculatory Duct

Epimeron

Furcal Apophysis

Furcasternum

Fenestra of Fourth

Abdominal

Segment

Furcula

Intercoxal Lamella of

Basisternum

Intercalary Sclerite of

Antenna

Juxta

Katepisternum

Laterofacial Sulcus

Lateral Cervical

Sclerite (Preepisternum)

Laterophragma

Mandible

Midventral Keel of

Vaginal Sclerite

Mn

MP

MS

Oc

OS

PA

Pam

PfS

Pha

PiS

PIS

Pin

Pn

PoS

Prm

PrS

Psm

PWP

R

RAG

RG

S

Sa

SAF

Se

Si

Sim

Sm

SP

Spa

Spe

SPt

Ss

T

Ta

T A p

T r

Ts

U

Us

Va

VD

Vm

VS

VeS

Meron

Median Process of

Vaginal Sclerite

Midcranial Sulcus

Ocellus

Occipital Sulcus

Posterior Apophysis

Patagium

Postfrontal Sulcus

Phragma (Postnotum)

Postinterocellar

Sulcus

Plcural Suture

Piemen turn

Pleuron

Postoccipital Sulcus

Pre-cpisternum

Prothoracic Spina

Pros tern um

Pleural Wing Process

Rectum

Reservoir of Abdominal Gland

Receptacular Gland

Sternite of Abdomen

Spina

Secondary Arms of

Furcasternum

Subalare

Socii

Scutellum

Scutum

Spermathecal Papilla

Spermatheca

Spiracle

Spinose Pocket

Stipes

Tergite of Abdomen

Tegula

Tertiary Apophyses

Transtilla

Testis

Uncus

Utriculus

Valva

Vas Deferens

Vinculum

Vaginal Sclerite

Vesicula Seminalcs
NUMBER 251 35

interpretation of considering SD1 as arising dorsal

to LI on the ninth segment, even though this is

not true on segments I to VIII of those species that

have retained LI.

The 10th anal segment in later instar larvae

seldom displays a distinct anal plate. The chaetotaxy is somewhat reduced in that only two pairs

of dorsal setae are present, with the most caudal

pair often being greatly reduced in size. Present in

apparently all eriocraniid larvae near the lateroposterior margins of the 10th tergite is a pair of

sclerotized, barlike spots (Figure 125). Gerasimov

(1952) has referred to these slender structures as

"brace-rods." In some species these rods are nearly

obsolete, whereas in others (e.g., Eriocraniella

aurosparsella) they are well defined and possess an

enlarged, punctate cephalic end. Similar sclerotized spots are also present on the anal segment of

certain Nepticulidae as shown by Klimesch (1972).

Systematics and Evolution

SUBORDINAL CLASSIFICATION

The heterogenous and probably paraphyletic suborder Dacnonypha previously included two superfamilies and four families (Davis, 1975b). The

superfamily Eriocranioidea (Bourgogne, 1949) included the Australian-Fijian family Agathiphagidae, the Australian Lophocoronidae, and the

Holarctic Eriocraniidae (Common, 1975). The

Chilean—southeast Asian family Neopseustidae is

also considered a possible member of this suborder,

although it has been further removed from the

Eriocraniidae and placed in a separate superfamily,

Neopseustoidea (Davis, 1975b).

The Agathiphagidae differs from the Eriocraniidae in so many major respects that it should

be removed from the Eriocranioidea. The Lophocoronidae are difficult to relate at present due to

our ignorance regarding the female (i.e., no females

have yet been collected). Based on the morphology

of the male, however, this family's closest affinities

appear to be with the Holarctic Eriocraniidae and

may represent a sister group of the latter.

The Mnesarchaeidae, although frequently considered a member of the Dacnonypha (Hinton,

1946b; Friese, 1969; Birket-Smith and Kristensen,

1974; etc.), have recently been shown to be more

closely allied to the Hepialoidea (Dugdale, 1974),

based on characters of the female reproductive

system.

In his attempt to improve upon an earlier phyletic scheme proposed by Hennig (1953), Friese

(1969) divided the Lepidoptera into two new

suborders, the Gymnocera (including only the

Hepialoidea) and the Angiocera (containing all

other Lepidoptera). The Dacnonypha (including

only the Eriocranioidea) are still retained as a

group but are relegated to Friese's rank of "subcohort" in opposition to the Heteroneura (sensu

Tillyard, 1918). In concluding that the Hepialoidea

constitute the most primitive extant group of Lepidoptera (i.e., more primitive than the Zeugloptera),

Friese has misinterpreted the phylogenetic significance of several morphological features, involving,

particularly, the adult mouthparts and female genital system. Furthermore, the significance of the

morphological diversity witnessed in the larval and

pupal stages of the Lepidoptera was largely ignored

or similarly misinterpreted. These oversights

largely invalidate most of Friese's conclusions.

Recently, Razowski (1974) has revived a subordinal classification popularized by Tillyard

(1918), but first proposed by Karsh (1898), in which

the Lepidoptera are partitioned into two major

suborders, the Homoneura and Heteroneura (Isoneuria and Anisoneuria, respectively, of Karsh).

Following this scheme, Razowski does not recognize

the taxon Dacnonypha and, instead, places the

Eriocranioidea within the Homoneura.

In this report I have essentially followed the

classification proposed by Hinton (1946b) and later

modified by Dugdale (1974), wherein four suborders are recognized. Within the Dacnonypha I

recognize the following three superfamilies: Agathiphagoidea, new superfamily, containing the single

family Agathiphagidae; Eriocranioidea, consisting

of the Eriocraniidae, Acanthopteroctetidae, and

Lophocoronidae; and questionably, the Neopseustoidea, with the single family Neopseustidae.

Problems effecting this subordinal classificational

scheme and involving particularly the Dacnonypha

still persist. For example, no synapomorphies have

yet been proposed for the groups currently associated within the Dacnonypha.

The Agathiphagoidea, as defined essentially by

Kristensen (1967) and Common (1973), may be

distinguished from the Eriocranioidea in possessing

functional and fully articulated mandibles, non-
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

haustellate galeae, and relatively well-developed

lobular laciniae. Some of the most divergent apomorphies in adult Agathiphaga may be observed

in the structure of the female genitalia. As illustrated by Common (1973) for A. queenslandensis

Dumbleton, the ovipositor in this group is extensible but nonpiercing. The anterior apophyses are

unique among the Lepidoptera in consisting of a

double pair, with the dorsal pair arising from the

anterior margin of the 8th tergite and the ventral

pair from the anterior margin of the 8th sternite.

The posterior apophyses are unusual in fusing

before the caudal apex to form a single, slender

rod. The ultrastructure of the wing scales of

Agathiphaga (Figure 39) is unusual but similar to

that of Micropterix and Epimartyria in possessing

a minute network of obliquely oriented striae between the longitudinal ribs, underlaid by a layer

of mostly transverse striae. After examining the

apodal larva of Agathiphaga, Hinton (1958) concluded that it was typically lepidopterous in form

and closely related to the Eriocraniidae.

It is becoming apparent that a cladistic approach

similar to that attempted by Hennig (1953), wherein

the major evolutionary steps within the Lepidoptera are emphasized, will provide the most logical

and useful scheme for subdividing the order.

Following such an approach, one may arrange the

major phyletic lines as currently recognized within

the Lepidoptera in the following sequence:

(1) Zeugloptera, (2) Agathiphagoidea, (3) Eriocranioidea, (4) Neopseustoidea, (5) Exoporia (including the Hepialoidea and Mnesarcheoidea),

(6) Monotrysia, and (7) Ditrysia. A few of these

hierarchical groupings will require new names,

although previously coined names are already

available for most. The relative position of the

Neopseustoidea and Exoporia are still questionable,

but the eventual accumulation of new data (e.g.,

on the immature stages of the Neopseustoidea) may

help to resolve most of this uncertainty.

FAMILY CLASSIFICATION

One of the major problems in analyzing the

systematic relationships of the most primitive Lepidoptera (i.e., generally those with homoneurous

venation) is that phenotypic as well as phyletic distances between the most related, extant taxa tend

to be greatly emphasized due to the extinction of

annectant forms. The preponderant absence of

adequate fossil evidence compounds this problem

considerably. The paucity of taxa throughout

these groups, with the possible exception of the

relatively large family Hepialidae, is often encountered at all levels. For example, most of the homoneurous families are now recognized as comprising

monotypic superfamilies. Similarly, familial concepts are frequently based on only one to a small

number of genera, each typically containing a

relatively small number of species. Consequently,

the definitions of most homoneurous families tend

to be rather restrictive, with the result that new or

"aberrant" taxa (including the discovery of fossil

forms) may be difficult to classify. Certain principles

of phylogenetic systematics, as expounded by

Hennig (1966) and Brundin (1966), can assist in

resolving many of these problems; however, using

this philosophy but placing emphasis upon different character systems can still result in questionable

schemes of classification (cf. Friese, 1969). A phyletic analysis of the Eriocranioidea should be conducted eventually, but only in conjunction with a

study of all the known taxa.

A reexamination of the aberrant genus Acanthopteroctetes has required the recognition of a

new family, Acanthopteroctetidae. For reasons discussed above, the decision regarding the proper

systematic position of this taxon was not decided

easily. Acanthopteroctetes differs from the more

homogenous members of Eriocraniidae to such an

extent that its inclusion within that family has

been questioned (Mutuura, 1971; Common, 1973).

These differences involve major features of general

head structure, mouthparts, wing venation and

male genitalia. The significance of many of these

differences is debatable, however, as similar magnitudes of variation are known to occur within other

generally accepted moth families as well. For example, in some groups of Lepidoptera (e.g., the

Noctuoidea) relatively minor, but consistent, differences in venation are relied upon in distinguishing

families; in others (e.g., Tineoidea), however, rather

major venational divergences may occur within

presently recognized families. In this regard, the

differences between the forewing pattern in the anal

veins of Acanthopteroctetidae and Eriocraniidae

appear somewhat comparable to those observed

among major groups within the Psychidae. Similarly, the presence or absence of ocelli is a variable
NUMBER 251 37

character within several families, particularly the

Noctuidae. A partial reduction of the labial palpi

comparable to that which occurs in Acanthopteroctetes may also be witnessed within other groups

(e.g., Psychidae and Prodoxinae).

As pointed out earlier, the extreme reduction of

the adult mandibles in Acanthopteroctetes may possibly indicate a similar reduction in the prominent

pupal mandibles usually considered so characteristic for the family. Even if the pupal mandibles

are found to be reduced and nonfunctional in

Acanthopteroctetes, however, this alone may not be

justification for family separation, as similar extremes in variation are also known for other families with "decticous" pupae. Wiggins (1960) has

shown the pupal mandibles in the caddisfly family

Phryganeidae to be either normally well developed

and functional or degenerate. He further found

that in those species possessing vestigial pupal mandibles, the anterior sieve membrane closing the

larval (and pupal) case had been similarly lost.

Thus, significant modification in the primary organ

used in escaping from the pupal case more than

likely indicates some modification in this phase of

an insect's biology. It is intriguing to hypothesize

that the same may be true for Acanthopteroctetes.

Although the morphology of the male genitalia

differs greatly between the two families recognized

herein, the overall differences appear no greater

than those present in certain other families, such as

the Micropterigidae. One of the unusual features

of the male in Eriocraniidae is the presence of a

divided aedeagus, as compared to the more typical,

entire form in Acanthopteroctetidae. A divided

aedeagus also occurs within other families, such as

the Micropterigidae (Epimartyria) and Tineidae

(Diataga and Praecedes).

Of greater significance than most of the morphological differences discussed above between the Eriocraniidae and Acanthopteroctetidae are the pronounced differences in the structure of their wing

scales. As described previously, the broad, discal

scales of the forewings of Acanthopteroctetes differ

markedly in basic structure from those of Eriocraniidae and resemble, instead, those of nearly all

more recently evolved heteroneurous families. The

wing scales of Neopseustidae and the exoporian

Ditrysia also possess a basic structure most similar

to that of Acanthopteroctetes. Thus, the major

transition from the thin, platelike, nonperforated

wing scale so typical of the most primitive Lepidoptera to the more advanced, perforated scale possessing a large internal lumen is evident within the

Eriocranioidea. I believe that this apomorphy, associated with the previously debated characters, such

as the loss of ocelli, the mandibular reduction, the

origin of Ml from R4+5, the distal separation of

all three anal veins in the forewings, the less modified first abdominal sternite, and the several markedly different features of the male genitalia, sufficiently distinguish the Acanthopteroctetidae as a

family separate from the Eriocraniidae.

Of at least equal importance in determining the

relationship of Acanthopteroctetes to the Eriocraniidae is the evaluation of numerous morphological similarities that exist between the two groups.

These involve both symplesiomorphic and, more

importantly, synapomorphic character states. Symplesiomorphic similarities of the head involve such

notable features as the prominent genal processes,

the well-developed dorsal arms of the tentorium,

the five segmented maxillary palpi, the prominent

chaetosemata, and, possibly, the secondary subdivisions of the corneal nipples. The most prominent

symplesiomorphic similarities in wing structure are

the uniform distribution of microtrichia, the presence of a jugum in the forewings, and a fourbranched radius in the hind wings. One of the most

primitive features of the male genitalia is the undifferentiated, cylindrical form of the ninth segment, associated with a bisymmetrical pair of anteroventral processes or apophyses. In the female

reproductive system, the ventral origin of the common oviduct, the sinuate ductus spermathecae, and

well-developed cloaca must be considered simplesiomorphic. The presence of functional spiracles on

the 8th abdominal segment is another primitive

character shared by a few other homoneurous

families.

Several synapomorphies exist that strongly indicate a close relationship between the Eriocraniidae

and Acanthopteroctetidae. Some of the more notable synapomorphies include the subapical depigmented zone on the fourth maxillary palpal segment; the unpaired apical spur on the mesotibiae;

the largely fused, ringlike 8th abdominal segment

in the female; the short tertiary apophyses of the

10th segment, as well as the elongate, piercing, posterior apophyses. The latter superficially resemble

those found in the Incurvarioidea and even in a
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

few Lyonetiidae; but in each group the piercing

ovipositor is believed to have evolved independently. In addition, the reduced metameron in both

families, as well as the development of the anteromedial process of the metafurcasternum, may represent synapomorphies. Although the meron is greatly

reduced in the metathorax of Micropterigidae, in

this family there is no anteromedial process extending anteriorly from the furcasternum.

The general structure of the metafurcasternum in

Eriocraniidae and Acanthopteroctetidae is rather

similar to that found in most Monotrysia in possessing a well-developed anteromedial process. In

the latter group, however, the anterior pair of tendons (i.e., anterior to those arising from the furcal

apophyses) typically extends dorsally instead of ventrally and evidently connects with different muscles

than do the anterior tendons in Eriocraniidae and

Acanthopteroctetidae, which are directed ventrally

into the coxae.

GENERIC CLASSIFICATION

The generic concepts within the subfamily

Eriocraniidae have recently been questioned by

Birket-Smith and Kristensen (1974), resulting in the

recognition of most previously accepted genera

(Dyseriocrania, Eriocraniella, and Heringocrania) as

subgenera of a single taxon, Eriocrania. This treatment has been followed by Razowski (1975) and

Heath (1976), but it is not in agreement with the

findings of the present study. As a result of this investigation, it is apparent that the Nearctic Eriocraniidae have evolved into a minimum of four

distinct groups. Most of these groups (Dyseriocrania,

Eriocrania, and Eriocraniella) are characterized by a

number of correlated characters, involving such

diverse character systems as maxillary palpi, wing

venation, scales of the hind wing, and male genitalia. In addition, the members of Dyseriocrania

are further unique in possessing very broad, padlike tubercules on the fifth abdominal segment.

Furthermore, it is evident that the endemic North

American genus Eriocraniella has, at one time, subdivided to form two subgroups or subgenera, each

containing more than a single species.

It is important to note that at least two of these

genera are Holarctic in distribution, each possessing

species endemic to both the Nearctic and Palearctic

regions. In other words, the evidence to date

strongly suggests that both genera were not only

in existence but were widely distributed prior to

the ultimate separation of the two biotas. Although

the known distribution of extant Eriocrania does

not preclude a more recent trans-Beringian faunal

connection, the geographical vicariance of Dyseriocrania appears to have resulted, in part, from the

disruption of the North Atlantic faunal connection,

an occurrence that McKenna (1975) fixes during the

early Eocene, or approximately 49 million years

ago. It seems, therefore, that the Nearctic-Palearctic

counterparts of Dyseriocrania have evolved surprisingly little over the last 50 million years.

Because the three taxa referred to above are distinguished by a number of unrelated characters

that more than likely originated as early as the

Paleogene, I feel that it is more logical to recognize

these taxa as genera. I also believe that such recognition is more equivalent to the generic concepts

currently followed in the great majority of other

moth families. The main problem remaining is to

equate the other taxa of Eriocraniidae with the

three genera above.

The genus Neocrania, as typified by N. bifasciata,

new species, demonstrates several apomorphic characters that easily distinguish it from all other eriocraniids. Among the most characteristic differences

are the bilobed apices of the labial palpi from

which arise eight moderately long, striated setae;

the relatively short, broadly bilobed uncus; the

very irregular form of the aedeagus and the subapical cilia of the ventral phallic branch; and the

asymmetry of the bases of the anterior apophyses

of the female genitalia. Neocrania exhibits its closest affinities with certain elements of Eriocraniella

and, thus, probably represents a derivative from

that partially sympatric genus. Neocrania is also

similar to Eriocraniella in retaining the epiphysis,

a plesiomorphic trait that has been independently

lost in Dyseriocrania, Eriocrania, and Acanthopteroctetes. Eriocraniella falcata shows some phenotypic intermediacy between the two genera, especially with regard to wing venation, but it shares

more in common with the subgenus Disfurcula of

Eriocraniella as partially characterized by the specialized chaetotaxy at the apex of the maxillary

palpi.

The European genus Heringocrania presents the

greatest difficulties in that, according to BirketSmith and Kristensen (1974), no synapomorphies
NUMBER 251 39

• •

if 'A'Subgenus Eriocraniella

* Subgenus Disfurcula

Apomorphic

O Plesiomorphic

FIGURE 79.—Hypothetical phylogenetic relationships of

Nearctic eriocranioid genera.

are known for the two presently included species.

The venation of the type-species, H. unimaculella,

evidently varies somewhat as evidenced by the connate condition of R4 and 5 in the hind wings as

figured by Common (1973) and the stalked condition as shown by Razowski (1975b). Razowski has

also drawn Sc as being undivided in the forewing.

Except for the latter variation, Razowski's rendition closely agrees in venation with Dyseriocrania

in having R4 and 5 stalked in both wings. The

fifth maxillary palpal segment is also similar to

that of Dyseriocrania and especially Eriocrania in

consisting of a single lobe crowned by a cluster of

eight short, unstriated setae (Figure 196.)

As summarized in Figure 79, Dyseriocrania is believed to exhibit more plesiomorphic features than

any other genus in the superfamily. This conclusion

is based upon what is presently known concerning

the biogeography of the Eriocranioidea as well as

adult and larval morphology. An evaluation of

some of the more significant adult characters is

summarized in Table 3. The evaluation of at least

one of these (i.e., the fifth abdominal tubercules)

is rather questionable, although a consideration of

the enlarged, flattened form of the tubercules in

Dyseriocrania as plesiomorphic seems the most logical, especially in light of other correlated morphological and biogeographical data. Perhaps of some

significance in this regard is the fact that at least

some Trichoptera also possess broad, flattened sternal tubercules. As shown by Philpott (1925), the

New Zealand philopotamid, Hydrobiosella sternocera Tillyard, possesses well-developed, reticulated

tubercules very similar in form and relative size to

those of Dyseriocrania. The preservation of the

epiphysis in Eriocraniella and Neocrania and its

absence in the supposed earlier evolved genera

Dyseriocrania and Eriocrania is also problematic.

If the scheme outlined in Figure 79 is correct, then

the most logical assumption would be that the

epiphysis has been independently lost in both

Dyseriocrania and Eriocrania as well as in Acanthopteroctetes.

Although it is still somewhat premature to evaluate larval characters in this superfamily due to the

lack of sufficient data for a majority of the species,

some observations can be presented at this time. Of

the known larvae, that of Dyseriocrania griseocapitella possesses the most primitive chaetotaxy in retaining SV3 on the prothorax, LI on nearly all segments, and a full complement of 10 primary setal

pairs on the ninth segment. The presence of the

mandibular setal tuft may possibly represent a

plesiomorphic condition, with the loss of this struc-
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 3.—Subjective analysis of selected character states in Nearctic genera of Eriocranioidea

Character

1. Maxillary palpi,

4th segment

2. Apical spurs

of mesotibiae

3. Metafurcasternum

4. Eighth abdominal segment

(»)

5. Ovipositor

6. Tertiary apophyses

of forewing

8. Ocelli

9. Mandibles

10. Labial palpi

11. Vein Ml (both wings)

12. Abdominal glands

(5th stemite)

13. Anal veins (forewings)

14. First abdominal sternite

15. Aedeagus

16. Abdominal tubercules

(5th sternite)

17. Lobes of uncus (tl)

18. Apex of maxillary palpi.

19. Vein R3 (forewings)

20. Lobes of uncus ((?*)

21.. Anterior apophyses (9)

22. Apex of maxillary palpi.

23. Vinculum (<S)

24. Ninth abdominal segment

()

Character state

Plesiomorphic Apomorphic

entirely pigraented

ithout anteromedial

process (MA)

tergite and sternite separate

unspecialized, not piercing

undeveloped

solid, without a lumen

present

relatively large

3 segments

separate from R5

present

separate

relatively unmodified, ligulate

simple

large, reniform

short, poorly differentiated

simple

present

elongate

symmetrical

bilobed

unmodified, smooth

unmodified, membranous

subapically depigmented

anteromedial process

developed

tergite and sternite partially fused

specialized for piercing

developed

with an internal lumen

absent

extremely ^educed

2 segments

stalked to RS

absent

partially fused

reduced, furcate

divided

reduced, papiliform

elongate, well defined

lobed

absent

reduced

asymmetrical

5-lobed

with median furcula (Fra)

with ventral spinose pocket

'Regressive apomorphic trend.

ture secondarily occurring in at least three North

American species. The complete absence of true

legs and prolegs is a synapomorphic condition evidently present throughout the Eriocraniidae.

As is often the case in most groups of holometabolous insects, the larval stage in the Eriocraniidae

demonstrates a general lack of congruency with the

adult. From the limited number of species investigated in this study (i.e., six), it has been observed

that larvae belonging even to closely allied species

may differ significantly. For example, the larvae of

Eriocraniella aurosparsella and E. xanthocara,

whose adults are barely distinguishable morphologically, disagree with regard to major setal placements (e.g., the presence or absence of Al and LI)

as well as other important features (the presence

or absence of the mandibular setal tuft). Thus, the

application of larval characters to possible superspecific groupings of the adults does not appear

feasible. Too little is presently known concerning

the pupa to comment on the relative systematic

congruency of this stage.
NUMBER 251 41

EARLY HISTORY

The earliest record of this superfamily, according

to Martynova (1962), is from Paleogene deposits of

Baltic amber. Although Kusnetzov (1941:70) lists

two species of extinct, fossilized Eriocraniidae, it is

possible that neither one actually represents a

member of this family. Micropterix pervetus Cockerell, which Kusnetzov placed in Dyseriocrania,

agrees in all important aspects of venation with the

extant Australian-New Zealand genus Sabatinca

and probably does represent a true member of the

Micropterigidae.

The second species, Electrocrania immensipalpa,

which Kusnetzov (1941) described from Baltic amber, is shown with a venation so aberrant (Figures

251-252) that it is difficult to relate to any known

moth family. To make matters even more confusing, Kusnetzov presents two different venational

interpretations of the type specimen and, most

likely, neither is correct. Unfortunately, it is probably no longer possible to reexamine the type, because, as Kusnetzov notes on page 20, footnote 2,

the amber sample containing the specimen crumbled after having been soaked for five days in

toluene.

Although most of the deposits in which Baltic

amber now occurs are typically stated to be of

Oligocene age, it is possible, as pointed out by

Kusnetzov (1941:91) and others, that the actual

amber may have originated in even earlier preEocene deposits. Consequently, the age of Electrocrania immensipalpa is most likely somewhere between 30 and 40 million years or well after the

suspected minimum date for the existence of the

genus Dyseriocrania and, perhaps, Eriocrania and

Eriocraniella. The source of Micropterix prevetus

was Burmese amber, which generally occurs in even

later deposits of Miocene age.

Opler (1973) has identified a fossilized mine of

Eriocraniella species on Quercus simulata Knowlton from the middle to upper Miocene of Thorn

Creek, Idaho. This record, in addition to those of

several other extant genera of oak-mining Lepidoptera listed by Opler, clearly indicates the antiquity

of the association of these particular leaf miners

with the Fagaceae. Opler (1974:74) also refers to the

Eriocraniidae a Miocene leaf mine record originally

believed by Lewis (1969) to have been caused by a

nepticulid larva. Earlier, Opler (1973) considered

this mine to have been produced by a species of

Caloptilia (Gracillariidae). Thus, the identity of this

particular mine is questionable, but, if Lewis' description is correct, "The mine .. . is of a linear

serpentine type coursing into a blotch-like area,

and terminates as a linear-type mine," then most

likely it was not caused by an eriocraniid larva.

As hypothesized previously (Davis, 1975b), the

Eriocraniidae appear to have a Laurasian origin.

Unfortunately, the prime reasons for this view, as

well as many of the following opinions, are partially founded on negative evidence; however, the

absence to date of any records of this family from

the southern continents suggests the strong possibility that the Eriocraniidae arose in the Northern

Hemisphere at sometime following the separation

of Laurasia from Gondwana. As summarized by

Takhtajan (1969) and Schuster (1976), the plant

family Fagaceae is also presumed to possess a Laurasian origin, perhaps at sometime prior to the

middle Cretaceous. Although Takhtajan records

the remains of Fagns from the mid-Cretaceous,

Wolfe (1973) indicates that the earliest unquestionable evidence of this genus may only be from the

Eocene. As far as the principal host plants for the

Eriocraniidae are concerned. Betula demonstrates

the earliest fossil history as having been reported

from the late Cretaceous (Muller, 1970). Wolfe

(1973) states that Quercus has been positively identified from the early Eocene.

There is at this time no reason to assume that the

present host association of the Eriocraniidae with

the Amentiferae (particularly Betulaceae and Fagaceae) has not existed since the early origins of both

components. Consequently, it is possible that the

Eriocranifdae first arose sometime prior to the middle Cretaceous within the vast Boreal-Cretaceous

region in association with the first Amentiferae. As

characterized by Takhtajan, this region was dominated by an entirely temperate flora consisting

mostly of mesophilous, broad-leaved, mainly deciduous trees and shrubs, such as Acer, Alnus, Betulus,

Cocculus, Corylus, Credneria, Fagus, Grewiopsis,

Lindera, Magnolia, Platanus, Quercus, Sassafras, Viburnum, and Ziziphus, as well as Ginkgo, various

conifers, and ferns. The Eriocraniidae may have

early undergone a radiation comparable to that

experienced by these early angiosperms. Since the

late Eocene, however, the Eriocraniidae have remained remarkably conservative and, except for the
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

possible extinction of many forms, may have

evolved relatively little within the last 50 million

years. Although present evidence does not support

a pre-Cretaceous origin for the Eriocraniidae, future discoveries (e.g., from the Southern Hemisphere) may modify this situation. Certainly, several

plesiomorphic features observed in the overall

morphology of this group, particularly in the decticous pupa, strongly suggest an earlier origin for

the family.

Judging from the typical form of the Miocene

mine of Eriocraniella (Opler, 1973), it also appears

that the morphology of the mine and feeding behavior of the larvae has changed little since the

Tertiary. If ontogeny bears any relationship to phylogeny with regard to larval leaf-mining behavior,

then there may be some evidence to indicate that

the earliest lepidopterous miners constructed linear,

serpentine galleries. The first instar larvae of Eriocraniidae construct such mines as do most of the

Nepticulidae, a successful, predominantly leaf-mining family that, like the Eriocraniidae, are perhaps

the most primitive survivors of their phyletic stock.

Among the more advanced, ditrysian leafminers,

however, so many later specializations have occurred, even among those that initiate serpentine

galleries, that it becomes almost meaningless to

refer generally to primitive or derived types of

mine morphology. In reference to this discussion,

it may be of some significance to note that one of

the earliest fossil leaf mines reported thus far is

of a serpentine type from the early Eocene of Wyoming (Hickey and Hodges, 1975). The very extended, central frass line, clearly evident within

the mine of that specimen, strongly sugests that it

was produced by a species of Phyllocnistis, an extant genus of Gracillariidae.

CHECKLIST OF NEARCTIC ERIOCRANIOIDEA

(synonyms only in italics)

Eriocraniidae

Dyseriocrania Spuler, 1910

Mnemonica Meyrick, 1912

griseocapitella (Walsingham, 1898)

auricyanea Busck and Boving, 1914 [misidentification]

auricrinella (Fracker, 1930) [misidentification]

auricyanea (Walsingham, 1882)

cyanosparsella (Williams, 1908)

Eriocrania Zeller, 1851

semipurpurella semipurpurella (Stephens, 1834)

amentella (Zeller, 1850)

inconspicuella (Wood, 1890)

semipurpurella pneifica, new subspecies

semipurpurella Clarke, 1942 [misidentification]

breviapex, new species

Eriocraniella Viette, 1949

Eriocraniella Viette, 1949, subgenus

aurosparsella (Walsingham, 1880)

xanthocara, new species

aurosparsella Opler, 1974 [misidentification]

longifurcula, new species

platyptera, new species

Disfurcula, new subgenus

variegata, new species

trigona, new species

falcata, new species

Neocrania, new genus

bifasciata, new species

Acanthopteroctctidae

Acanthopteroctctes Rraun, 1921

tripunctata Braun, 1921

tripunclella Davis, 1969 [misspelling]

bimaculata Davis, 1969

unifascia, new species

ERIOCRANIOIDEA

ADULT.—Small moths with moderately slender,

fully scaled wings. Wing expanse ranging from

6 to 16 mm.

Head: Hypognathous, often broader than long,

slightly more extended in Eriocraniidae; cranial

sulci usually well developed; sometimes absent;

dorsal arms of tentorium slender, elongate, nearly

extending to vertex. Vestiture relatively sparse,

hairlike, mostly erect and divergent except for a

small tuft of nearly appressed, relatively broad

scales often covering basal half of labrum. Chaetosemata present, consisting of a large, elongate pair

of raised areas extending transversely across occiput and nearly meeting near vertex; a single, circular chaetosema may also be present ventrally on

postmentum. Ocelli either present or absent. Antennae simple, of moderate length, usually 0.5-

0.75 the length of forewings, typically covered with

narrow, relatively long, appressed scales; each segment also bearing numerous and usually scattered,

small, often oval and highly modified, sensilla

auricillica (Figures 26, 34) immediately beneath

outer scale covering; sensilla auricillica absent in

Lophocoronidae. A minute intercalary sclerite present between scape and pedicel. Compound eyes

relatively large in Acanthopteroctetidae and Lophocoronidae, more reduced in diameter and posi-
NUMBER 251 43

tioned farther forward in Eriocraniidae; cornea

sparsely covered with minute microtrichiae with a

length less than diameter of a single facet. Labrum

relatively well developed, elongate, constricted at

base, more reduced in Lophocoronidae; pilifers

obsolete. Ventral margin of gena extended into an

angulate subgenal process. Mandibles vestigial,

nearly absent in Acanthopteroctetidae, much more

developed in Eriocraniidae, possessing a large adductor and very small abductor apodemes but with

condyles greatly reduced or absent. Maxillae consisting of a distinct cardo and stipes, a relatively

short, functional haustellum and an elongate,

5-segmented palpus (4-segmented in Lophocoronidae); each galea with a series of large, slightly overlapping plates lining the food canal; maxillary

palpi with the 4th segment the largest and at least

twice the length of 5th; a small depigmented zone

usually situated subapically on penultimate segment; apex of 5th segment either relatively simple

or with minute, secondary lobes; primary setae

of 5th segment either simple or barbed; laciniae

absent or vestigial. Labium with relatively narrow,

elongate prementum and either 2- or 3-segmented

palpi; sensory organs of von Rath either present

(at apex of 3rd palpal segment) or absent in those

species with 2-segmented palpi (in genus Acanthopteroctetes).

Thorax: Meso- and metascuta well developed,

relatively elongate. Furcal apophyses of mesothorax

greatly reduced, barely evident; furcal apophyses

of metathorax well developed, prominently divided, arms relatively slender and not strongly

divergent, subparallel; length of metafurcal apophyses either equaling or slightly exceeding the

distance between their terminal apices. Metathoracic coxae with meron reduced, approximately

0.5-0.8 the length of eucoxa. Prothoracic tibiae

either with or without epiphyses, often with 2-3

rather large apical setae; tarsal segments of all

legs usually with 3-4 apical setae; mesothoracic

tibiae with a single large apical spur (spurs paired

in Lophocoronidae); metathoracic tibiae with 2

pair of long spurs, 1 pair at distal apex and another arising from distal two-thirds; all spurs of

approximately equal lengths; tarsae of typical form,

with simple claws and empodium, a broad padlike

arolium, and a lateral pair of large, spinose pulvillae. Forewings relatively narrow, subacute with

a rather prominent jugum at extreme base of hind

margin; Sc either simple or divided; radius with

either 4 or 5 main branches; Rl either simple or

divided, often with a moderately developed pterostigma over outer third of Rl; R3 sometimes absent; R5 terminating near apex, either slightly

above or below, or remote and on termen (in

Lophocoronidae); medius well preserved within cell,

3-branched; Ml usually separated but stalked with

R4 and R5 in Acanthopteroctetidae; anal veins

either fused beyond basal half or completely separate. Hind wings with Sc simple; radius usually

with 4 primary branches, sometimes 5; Rl simple;

area around apex of Rl sometimes slightly pigmented; R3 present (in Lophocoronidae) or absent; R4 and 5 either stalked or separate. Ml

usually arising separate but stalked to R4 and R5

in Acanthopteroctetidae. Both wings with microtrichiae dense and evenly distributed.

Abdomen: Pleural area of 3rd segment usually

unmodified, rarely with a cluster of small, filamentous, clavate appendages. Fourth sternite either

with or without a bilateral pair of circular, membranous fenestrae. Fifth sternite usually with a

bilateral pair of reticulated, glandular tubercules

usually arising near anterolateral margins; tubercules either large and reniform in outline or small

and papiliform, with a central exit pore leading

to a large ducted gland internally; reservoir of

gland often closely associated with fenestrae of 4th

sternite; glands and tubercules occasionally absent.

Eighth segment with spiracles well developed and

apparently functional; entire segment greatly enlarged in female and normally enclosing 9th and

10th segments or their derivatives.

Male Genitalia: Uncus either well developed,

bifid, usually separated from 9th segment or absent

(Lophocoronidae). Ninth segment in the form of

a broad sclerotized cylinder, undifferentiated into

a dorsal tegumen or ventral vinculum; cephalic

margin of ventral area (vinculum) usually concave,

with ventrolateral angles slightly to greatly extended anteriorly. Anal tube mostly membranous,

sometimes with minute, scattered spinules or with

a bilateral pair of small, disarticulated sclerites

near base. Socii either present or absent; gnathos

absent. Valvae either very short and relatively

broad (Eriocraniidae) or more elongate and movable (Acanthopteroctetidae and Lophocoronidae).

Anellus variously modified, either consisting of a

single but complex plate, or juxta, ventral to
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

aedeagus and often fused laterally to 9th segment;

or consisting of 2, very distinct, usually well-sclerotized structures, 1 dorsal (transtilla) to aedeagus

and 1 ventral (juxta); transtilla heavily sclerotized

with medial area greatly extended caudally. Aedeagus either divided for most of its length into a

dorsal and ventral phallic branch, or undivided

and with prominent cornuti.

Female Genitalia: Ovipositor of the piercing

type with a depressed, relatively broad, triangular

(viewed dorsally) apex possessing minutely serrated

edges. Two pairs of very elongate, rodlike apophyses present, the anterior pair arising from the

8th abdominal tergite and often extending as far

forward as the 5th segment; the posterior apophyses

approximately twice the length of anterior pair

and arising from the 10th segment, where they fuse

to form the piercing ovipositor; a small, slightly

elongate pair of articulary processes usually present

near junction of 9th and 10th segments; a pair of

very short tertiary apophyses present within the

10th segment extending from the small articulary

processes previously described to the cloacal aperture. Ninth segment largely membranous, roughened externally with numerous rows of minute

spines dorsally and ventrally. Cloaca well developed, consisting of a relatively elongate tube opening on 10th segment immediately anterior and

ventral to caudal apex of ovipositor; only 1 terminal body opening present, serving as common exit

for both alimentary and reproductive systems. Corpus bursae variable in size; usually large with completely membranous walls; signa absent. Spermatheca typically arising from either a sclerotized

or mostly membranous funnel-shaped papilla; spermathecal duct slightly sinuate, not coiled, often

with an internal, rodlike thickening (in Eriocraniidae). Common oviduct typically joining ductus

bursae ventrally opposite spermathecal papilla; oviduct rarely arising laterally. Colleterial gland arising dorsally from vagina, immediately anterior to

junction of alimentary tract with cloaca; glands

usually bifid. Vagina either membranous and partially encircled by a well-defined sclerite or without vaginal sclerite but with walls thickened and

heavily folded.

Key to the Nearctic Families of Eriocranioidea

Ocelli present. Ml arising separate from R4 and 5 in both wings; forewings with all three

anal veins joining distally. Abdomen with a pair of glandular, reticulated tubercules frequently present on 5th sternite. Male genitalia with uncus prominently bilobed; valvae

extremely reduced, considerably less than 0.5 the uncus in length; anellus completely

membranous dorsal to aedeagus; aedeagus divided for most of its length. Eighth segment of

female without ring of elongate setae ERIOCRANIIDAE

Ocelli absent. Ml stalked with R4 and 5 in both wings; forewings with all 3 anal veins

separate. Abdomen without sternal tubercules on 5th segment. Male genitalia with uncus

only slightly cleft, not prominently bilobed; valvae slender, elongate, surpassing total length

of uncus; anellus with a strongly sclerotized, elongate transtilla dorsal to aedeagus; aedeagus

undivided. Female with a prominent row of elongate setae encircling caudal margin of

8th abdominal segment ACANTHOPTEROCTETIDAE, new family

ERIOCRANIIDAE

ERIOCRANIIDES Tutt, 1899:104, 109, 110, 114, 131.

CHAP.MAMIDAE Borner, 1925:370.

TYPE GENUS.—Eriocrania Zeller, 1851.

ADULT.—Moths of very small size, with a wing

expanse of 6-13 mm; wings moderately slender.

Head: Relatively elongate with vertex well developed and extended; laterofacial sulci prominent.

Lateral ocelli present. Compound eyes relatively

small, interocular index approximately 0.75; eyes

positioned more anteriorly on head capsule. Mandibles vestigial but relatively large with well-developed

adductor apodemes; condyles usually evident but

very reduced in size. Apex of terminal (5th) segment of maxillary palpi variously modified, frequently with minute, secondary lobes. Labial palpi

elongate, 3-segmented; apex of distal segment with

a well-developed organ of von Rath containing

several minute, basiconic sensillae.

Thorax: Prothoracic legs relatively reduced in
NUMBER 251 45

size, foretibiae approximately 0.3-0.33 the length

of hind tibiae; epiphysis either present or absent.

Forewings moderately slender, width approximately

0.30-0.34 the length; Sc either simple or shortly

divided near costa; radius usually with 5 primary

branches, sometimes 4-branched with loss of R3;

Rl usually secondarily divided nearly a third its

length; a faint pterostigma typically present near

outer third of Rl; R4 and 5 either separate or

stalked; Ml widely separated from base of R5;

all 3 anal veins arising separate at base of wing,

then abruptly fusing near distal half; scales over

dorsal surface of forewing mostly broad with longitudinal ribs projecting slightly beyond apical margin and without internal lumen or lacunae; scales

of marginal fringe of termen mostly slender and

with internal lumen. Hind wings with pterostigma

usually faintly present near apex of Rl but greatly

reduced; R3 absent; R4 and 5 usually separate or

connate, sometimes stalked about 0.5 their length;

Ml arising widely separate from R5; scales ranging from hairlike to relatively broad.

Abdomen: Pleural area of 3rd segment unmodified, without appendages. Fourth sternite usually

with a circular to elliptical pair of membranous

fenestrae in female; fenestrae absent in male. Fifth

sternite usually with a pair of reticulated, reniform

or papiliform tubercules in both sexes, rarely absent; a relatively large pair of globular glands

with separate, slender ducts opening ventrally

through a central pore in each tubercule present

in all species possessing tubercules. Eighth segment

of female without encirclement of elongate hairs.

Male Genitalia: Uncus prominently divided

into 2 frequently large lobes; lobes usually slightly

divergent and well separated with rather broadly

rounded apices. A pair of usually elongate apophyses extending anteriorly from anterolateral margin of vinculum (i.e., venter of 9th segment). Anal

tube membranous, occasionally with numerous,

scattered minute spines or with a basal pair of

small sclerites. Socii either present or absent. Valvae very reduced and relatively broad, total length

less than 0.4 that of uncus; apex of cucullus strongly

spined. Anellus not sclerotized dorsally; ventral

half consisting of a large, rather complex plate

(i.e., juxta) that fuses laterally to ventrolateral

walls of 9th segment, thereby partially dividing

cavity within 9th segment. Aedeagus divided most

of its length into a primary dorsal branch containing the vesica and a secondary ventral branch that

may be variously modified; ventral branch usually

loosely articulated to prominent bulbous base of

aedeagus.

Female Genitalia: Corpus bursae enlarged; ductus bursae membranous. Common oviduct arising

ventral to bursa copulatrix usually opposite spermathecal papilla. Ductus spermathecae typically

with an internal rodlike thickening; spermathecal

papilla usually consisting of a shallow cuplike

depression that may be either flush with vaginal

wall or raised on a membranous, tubular extension.

Vagina partially encircled by a usually well-defined,

variously modified sclerite. Dorsal colleterial gland

usually arising from vaginal wall approximately

midway along sclerite. A short, slightly elongate

articulary process arising ventrally from posterior

apophyses near juncture of 9th and 10 segments.

Caudal region of 9th segment usually unmodified

but occasionally with a midventral pocket densely

lined laterally with minute spines.

LARVA.—Body mostly whitish in color, cylindrical

and somewhat fusiform in outline, without thoracic legs or abdominal prolegs. Integument densely

spinose. Total length usually not exceeding 11 mm.

Head: Prognathous, partially retracted into

thorax, usually pale brown to stramineous in color.

Vertex deeply divided to frons. Adfrontal sclerite

broad with ecdysial line joining anterior margin

of head laterad to antennae; ecdysial line very

well defined, broad, irregular, sometimes enclosing

PI. A single, rudimentary ocellus present near anterior margin of head immediately ventral to antennae. Majority of cranial setae very reduced in

length; PI either greatly reduced or moderately

long; Al either present or absent; if present, then

arising from adfrontal sclerite; A3 either present

or absent; C2 considerably longer than Cl; O3

either present or absent. Labrum with anterior

margin bearing numerous spines (possibly derived

all or in part from underlying epipharynx); setae

L3, M2, and M3 either simple or broadly compressed. Mandibles opposable, with 4 cusps; 4th

cusp broad and relatively indistinct; inner, mesal

margins of mandibles either with or without a

dense tuft of elongate, barbed setae. Hypopharynx

densely covered dorsally with numerous, rather

elongate setae. Spinneret well developed, tubular,

with a circular orifice.

Thorax: Usually whitish in color except for
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

large, brownish pronotal and prosternal plates.

Prothorax with SV3 either present or absent; L3

present. Thoracic legs absent. Meso- and metathorax with paired, slightly raised dorsal and

ventral calli.

Abdomen: Whitish in color; prolegs and crochets absent. LI either present or absent, if present

then arising well above level of spiracles; SV2

usually present but variable and occasionally absent; 9th segment usually with 7 pair of primary

setae but ranging to as many as 10 pair in certain

species. Tenth segment usually with anal plate

poorly defined, only 2 pair of setae present dorsally and with a pair of small, elongate barlike

spots situated lateroposteriorly.

PUPA (Figures 71-78).—Decticous with elongate,

heavily sclerotized, functional mandibles. Pupal

cuticle very thin, mostly transparent, with all appendages and most body segments free and movable. Length approximately 3.0-3.5 mm; diameter

1.3-1.8 mm.

Head: Vertex with 1-2 pair of elongate setae

arising immediately above and posterior to antennal insertions. Frontal ridge greatly developed,

raised and extending free of head to labrum. Antennae free, with all segments easily discernible,

terminating just short of apices of wing pads.

Labrum well defined, reniform, with pilifers prominent and bearing 6 pair of elongate setae; length

of longest setae nearly 1.5 times the diameter of

eye. Mandibles very elongate, their length approximately 2.0 the diameter of eye, consisting of a

strong but relatively slender, elongate, stalklike

base that abruptly enlarges near apex; extreme

apex truncate, with a broad, circular, median depression typically bordered peripherally by a progressively enlarged series of 4 broad spines and by

a low, sometimes serrated ridge that encircles nearly

one half of apical depression; basal stalk of mandibles with 1-2 indefinite rows of numerous, short

spines extending along dorsal-inner surface; mandibles fully articulated and capable of strong movement.

Thorax: Appendages all free, segmentation

clearly evident. Mesothoracic spiracles situated in

the conjunctiva connecting the pro- and mesothorax. Forewing and hind wing pads separate,

sometimes reaching to end of abdomen. Distal ends

of metathoracic legs usually extending beyond last

abdominal segment and sometimes curling around

end of abdomen.

Abdomen: Pupal cuticle extremely thin and

membranous. Segments I-VII movable in both

sexes; VIII-X without definite sutures and probably incapable of independent movement; segment

X larger in female and curved more ventrad than

in male. Spiracles small, circular, not produced;

first 8 abdominal spiracles present and apparently

functional. Abdominal tergites typically with 2

longitudinal rows of elongate setae on either side,

1 row above and 1 below spiracular line, and a

paired row of very short setae on either side of

middorsal line; each setal row with 1 seta per

segment; tergites lacking transverse rows of spines.

Key to the Genera of Nearctic Eriocraniidae

(based on the adult)

1. Sternal tubercules of 5th abdominal sternite broad, diameter approximately 0.5 the length

of sternite, relatively flattened, reniform. Male genitalia without socii (Figure 225)

Dyseriocrania

Sternal tubercules much smaller in diameter, more raised, papiliform (Figure 224). Socii

present in male (Figure 267) 2

2. Forewings with Rl simple, undivided, R3 absent. Epiphysis absent. Terminal segment of

maxillary palpi relatively simple (Figure 191), not lobed Eriocrania

Forewings with Rl divided; R3 present. Epiphysis present. Terminal segment of maxillary

palpi variously lobed (Figures 198, 210), not simple 3

3. Forewings with Sc usually simple, though sometimes divided. Terminal segment of maxillary palpi with 1 relatively large apical lobe and 2 pair of smaller, subapical lobes (Figures 198, 202). Male with uncus divided for most of its length. Vaginal sclerite of female

with a prominent, midventral keel (Figure 311) Eriocraniella

Forewings with Sc typically divided. Terminal segment of maxillary palpi with 2 apical

lobes of nearly equal size (Figure 210). Male with uncus divided about 0.5 its length.

Vaginal sclerite without midventral keel Neocrania, new genus
NUMBER 251 47

Key to the Larvae of the Nearctic Eriocraniidae

1. Mandibles with a prominent mesal tuft of barbed setae (Figure 105) 2

Mandibles without mesal setal tuft 4

2. Seta Al present, arising from adfrontal sclerite (Figure 104) Dyseriocrania auricyanea

Seta Al absent 3

3. Ninth abdominal segment with 10 pair of primary setae. PI minute in size. Distribution:

eastern United States Dyseriocrania griseocapitella

Ninth segment with 7 pair of primary setae. PI moderately long. Distribution: western

United States Eriocraniella xanthocara, new species

4. Abdominal segments I to VIII with LI present. Prothorax with subventral setae usually

trisetose. Labrum with setae L3, M2, and M3 simple, not compressed

Eriocrania ?semipurpurella pacifica, new subspecies

Abdominal segments without LI. Subventral setae of prothorax bisetose. Labrum with L3,

M2, and M3 broadly compressed 5

5. Labrum with ventral margins of L3, M2, and M3 simple (Figure 155). not dentate

Eriocraniella Hongifurcula, new species

Labrum with ventral margins of L3, M2, and M3 variably dentate (Figure 129)

Eriocraniella aurosparsella

Dyseriocrania Spuler

Dyseriocrania Spuler, 1910:483.—Fletcher, 1929:72, 141 [synonym of Mnemonica].—Hering, 1932:2.—Kuznetsov, 1941:

19, 20.—Lhomme, 1949:1220.—Viette, 1949:30, 32; 1951:

43, 44.—Gerasimov, 1952:189 [synonym of Eriocrania].—

Toll, 1959:16, 21, 23.—Kristensen, 1968b:241, etc.—Common, 1973:16.—Birket-Smith and Kristensen, 1974:158, etc.

[subgenus of Eriocrania].—Razowski, 1975:42 [subgenus of

Eriocrania].—Heath, 1976:157 [synonym of Eriocrania].

Mnemonica Meyrick, 1912a:5; 1912b: 18.—Barnes and McDunnough, 1917:197.—Tillyard, 1919:97, etc.; 1923:181,

195.—Crampton, 1920:26, 27.—Forbes, 1923:65 [subgenus of

Eriocrania].—Philpott, 1924:361; 1927:320.—Meyrick, 1928:

870, 872.—Fletcher, 1929:72, 141.—McDunnough, 1939:110.

—Kuznetsov, 1941:19 [synonym of Dyseriocrania].—Hinton,

1946:11.—Viette, 1949:30 [synonym of Dyseriocrania]; 1951:

43, 44 [synonym of Dyseriocrania].—Gerasimov, 1952:189

[synonym of Eriocrania].—Heath, 1958:117, 120.—Kristensen, 1968:242 [synonym of Dyseriocrania].—Dugdale, 1974:

134.—Birket-Smith and Kristensen, 1974:159 [synonym of

Dyseriocrania].—Razowski, 1975:42 [synonym of Dyseriocrania].—Heath, 1976:157 [synonym of Eriocrania].

TYPE-SPECIES.—Tinea subpurpurella Haworth,

1829; original designation by Spuler, 1910.

ADULT.—Small, relatively broad winged moths;

wing expanse: 9-13 mm.

Head: Maxillary palpi with basal 0.9 of terminal segment densely setose; apical 0.1 largely

naked; primary setae simple, without lateral barbs;

apex of terminal segment subacute, simple, without

secondary lobes but with a scattered series of usually 8-10 minute, subacute setae with smooth,

unstriated sides; setae of rather uniform size, very

short and stout, usually less than 0.3 the diameter

of apex in length.

Thorax: Forewings with Sc divided near costa;

Rl divided approximately one-third its length; R2

and 3 separate; R4 and 5 stalked approximately

0.4 their length. Hindwings with Sc and Rl simple;

R2 and 3 entirely fused; R4 and 5 stalked approximately one-third their length; scales relatively

short, broad, not hairlike.

Abdomen: Fenestrae present on fourth sternite

of female, relatively large and oval in form, absent

in male. Sternal tubercules present on fifth sternite

of both sexes, similar in form, very large and reniform, forming a broad, padlike plate.

Male Genitalia: Uncus bilobed but with lobes

poorly developed and not deeply divided; lobes

separated by a relatively shallow concave depression. Socii absent. Anal tube without basal sclerites

but with numerous, minute spinules scattered over

walls of tube. Vinculum with median caudal margin variously produced into shallow lobes, not

smoothly convex; furcula absent; cephalic margin

evenly concave between apophyses, without median

process. Caudal apex of juxta variously modified

but typically symmetrical. Aedeagus with ventral

phallic branch extremely slender, often less than

0.3 the dorsal branch in diameter; apex of ventral

branch simple, acute, without lobes or processes.

Female Genitalia: Vaginal sclerite variable in

form, either with or without midventral keel. Ninth

segment without a midventral spinose pocket.

DISCUSSION.—Dyseriocrania is represented in
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

North America by two very distinct species, auricyanea and griseocapitella, and in Europe by the

type of the genus, D. subpurpurella (Haworth). The

European member of this group is readily distinguished from the American species on the basis of

male genital characters, particularly in the relative

development of the paired apophyses. The cephalic

margin of the vinculum is not as deeply divided

in D. subpurpurella with the result that the apophyses are much shorter (length of apophyses less

than 0.5 the length of undivided vinculum) than

in the American species.

Key to the Species of Nearctic Dyseriocrania

Forewings light golden bronze, heavily mottled with minute specks of darker scales. Male

genitalia (Figure 255) with lobes of uncus widely separated; vinculum with median caudal

margin shallowly bilobed. Vaginal sclerite (Figure 295) of female without midventral keel;

corpus bursae without spinules. Distribution eastern United States

Dyseriocrania griseocapitella

Forewings light golden brown, faintly crossed by 3 irregular, purplish fasciae. Male genitalia

(Figure 257) with lobes of uncus narrowly separated; vinculum with a pronounced, median,

caudal lobe. Vaginal sclerite (Figure 298) with a prominent ventral keel; walls of corpus

bursae with a scattering of numerous tiny spinules. Distribution cismontane California

Dyseriocrania auricyanea

Dyseriocrania griseocapitella (Walsingham)

FIGURES 1-7, 11-14, 23-26, 35-37, 45-46, 48, 50-52, 56, 58-63,

66-68, 77-78, 80-108, 166, 184-185, 213, 215, 220-221,

244, 255-256, 293-295, 333; MAP 2

Eriocrania griseocapitella Walsingham, 1898:162-—Kearfott

1902:129.—Forbes, 1923:66.

Eriocephala griseocapitella (Walsingham).—Dyar, 1902 [1903]:

581, no. 6619.—Kearfott, 1903:125, no. 7166.

Mnemonica griseocapitella (Walsingham).—Meyrick, 1912a:

19; 1912b:5.—Barnes and McDunnough, 1917:197, no.

8479.—McDunnough, 1939:110, no. 9854.

Mnemonica auricyanea Busck and Boving, 1914:151 [misidentification, not Walsingham, 1898].—Rohwer, 1914:141.

—Mosher, 1916:35.—Forbes, 1923:65—Crampton, 1920:33,

34.—Philpott, 1924:361; 1925:458; 1926:723.—Hin ton, 1946:

figures 10-12— Muesebeck et al., 1951:329.—Gerasimov,

1952:92.—Friese, 1969:209.—Razowski, 1975a:5.

Eriocrania auricyanea (Busck and Boving).—Forbes, 1923:66.

—Forbes, 1928:534, no. 8477.—Needham, Frost, and Tothill, 1928:72.—Shepard, 1930:243.

Dyseriocrania auricyanea (Busck and Boving).—Kristensen,

1968:242—Opler, 1974:17.—Davis, 1975:10.

Epimartyria auricrinella Fracker, 1930:57 [misidentification,

not Walsingham, 1898].

ADULT (Figure 166).—Wing Expanse: tf, 10-13

mm; 9, 9-12.5 mm.

Head: Vertex and frons covered with long, predominantly white hairs intermixed with fuscous.

Antennae approximately 0.53-0.57 the length of

forewing, 43- to 47-segmented; scape predominantly

white, irrorated with fuscous; flagellum white beneath, fuscous above. Maxillary and labial palpi

white, irrorated with fuscous. Maxillary palpi with

apex of 5th segment naked except for 5-6 apical

setae and 2-3 subapical setae; setae very short,

length not exceeding 2.0 their width.

Thorax: Dorsum almost equally intermixed

with elongate hairs and broader, appressed scales

of white and fuscous. Venter white. Legs white

ventrally, light fuscous dorsally, usually darker on

prothoracic legs. Forewings golden bronze, heavily

mottled with minute specks of darker scales which,

under low magnification may appear iridescent

bluish purple; forewings rather evenly mottled except for a small, pale subtornal spot usually present on hind margin; fringe bronzy brown. Hind

wings paler, grayish with a slight purplish luster,

sparsely covered with moderately broad scales.

Abdomen (Figures 56, 58-63, 220-221): Sparsely

covered with long brownish hairs dorsally, whitish

ventrally. Fourth sternite of female with fenestrae

nearly equaling sternal tubercules in size, diameter

approximately 0.5 the length of sternite; fenestrae

absent in male. Fifth sternite of both sexes with a

pair of large, reniform tubercules that are usually

wider than long, length approximately 0.5 the

length of sternite. Caudal margin of 8th sternite

and tergite of female relatively indistinct, subtruncate, not deeply clefted.

Male Genitalia (Figures 255-256): Uncus shallowly bilobed; lobes reduced, widely separated.

Anal tube membranous, with minute, scattered

spines; basal sclerites absent. Vinculum with caudal
NUMBER 251

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S03

F,CURES 80-8i.-Dyseriocrania griseocapitella (Walsingham), larval chae otaxy: 80, lateral view

of prothorax, mesothorax, and abdominal segments 1, 6, 8. and 9; 81. dorsal view of head

(scale = 0.3 mm); 82, left mandible (scale = 0.1 mm); 83, dorsal view of abdominal segments

8-10; 84, lateral view of head.
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4 86

FIGURES 85-90.—Dyseriocrania griseocapitella (Walsingham), larval head structure: 85, dorsal

view of anterior margin of head, X 220; 86, anterior view of labrum and mandibles, X 525;

87, chactotaxy of labrum, X 1250; 88, anterior view of mouthparts, X 360; 89, ventral view

of mouthparts, X 420; 90, details of maxilla and spinneret, X 800. (Photographs reduced to

84%.)
NUMBER 251 51

FIGURES 91-96.—Dyseriocrania griseocapitella (Walsingham), larval head structure: 91, antenna,

X 930; 92, surface detail of antennal basiconic sensilla, X 9425; 93, anterior view of buccal

cavity and chaetotaxy of hypopharynx, X 870; 94, integument of fifth abdominal tergite

between setae Dl; 95, ventral view of left mandible, X 500; 96, mesal view of left mandible,

X 500. {Photographs reduced to 84%.)
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 97-100.—Dyseriocrania griseocapitella (Walsingham), larval mandibles: 97, dorsal view

of setal tuft, X 950; 98, mesal view of setal tuft, X 1000; 99, base of setal tuft, X 3100; 100,

detail showing barbs of setal tuft, X 7650. (Photographs reduced to 84%.)

margin superficially bilobed; anterior margin with

apophyses elongate, approximately 0.75 the length

of undivided vinculum. Juxta broadest anteriorly,

tapering caudally to a narrow, minutely clefted

apex; greatest width approximately 0.6-0.65 its

length. Base of aedeagus prominently swollen; ventral branch of aedeagus very slender, simple, without any apical projections or basal lobes.

Female Genitalia (Figures 293-295, 333): Apex

of ovipositor acuminate; lateral edges serrulate,

with 10-12 minute teeth. Bursa copulatrix relatively elongate, extending noticeably beyond apices

of anterior apophyses; walls of corpus bursae entirely membranous. Vaginal sclerite moderately

broad, greatest width about 0.45-0.5 its length,

depressed, not darkly sclerotized, and without ventral keel; general structure as shown in Figure 294.

LARVA (Figures 80-100).—Length of largest larva

10 mm; diameter 1.8 mm.

Head: Mostly pale brown to light yellow in

color; adfrontal suture outlined in dark brown.

Greatest width 0.6 mm; length 0.85 mm. Va variable in position, usually either laterad or posterior

to VI. PI very reduced in size, approximately equal

to A2, usually separate from ecdysial line. A2 situated either slightly anterior or posterior to PI. Al

absent. Labrum with all setae simple, acute; anterior margins of pilifers and epipharynx densely

spinose; M3 arising much closer to M2 than to

L3. Mandibles with an inner, median tuft of 12-18
NUMBER 251 53

elongate setae; each seta finely branched, appearing somewhat serrate.

Thorax: Pronotal and prosternal plates well

developed, dark brown. Meso- and metathorax

without sclerotized plates, whitish in color. L2

relatively well developed on meso- and metathorax,

approximately equal to LI in size. Subventral setae

trisetose on prothorax.

Abdomen: Integument without pigmentation,

usually whitish except for a pair of minute, longitudinal, brownish spots near lateroposterior margin of anal segment; barlike spots without enlarged cephalic end. LI present on all segments,

situated relatively high above SD1 and spiracles

on I to VIII. SV2 absent in segment I, usually

present on II, VII, and IX but variable. Segment

IX with 10 pair of primary setae; both SD2 and

SV2 present and well developed.

TYPE.—Holotype, o"; in the British Museum

(Natural History).

TYPE-LOCALITY.—Washington, D.C.

HOSTS.—Fagaceae: "chestnut [Castanea dentata

(Marsh.) Borkh.]; chinquapin [Castanea pumila (L.)

Mill.]; oak [Quercus species]," Busck and Boving

(1914). Castanea mollissima Blume; Quercus alba

L.; Quercus }borealis (Michaux) Farw.; Q. }falcata

Michaux; Q. velutina Lamarck.

PARASITES.—Ichneumonidae: "Lathrolestes mnemonicae (Rohwer)," Rohwer (1914); near Tersilochus species.

FLIGHT PERIOD.—Late February to late May;

univoltine.

DISTRIBUTION (Map 2).—This species ranges

widely along the Atlantic states of North America

from Nova Scotia south to northern Florida. It is

known to occur as far westward as Michigan,

Illinois, and Mississippi.

MATERIAL EXAMINED.—97 males, 154 females, and 41 larvae.

CANADA. NOVA SCOTIA: Bridgewater: \$, \ Jun <NSM).

UNITED STATES. CONNECTICUT: New Haven Co: Bethany:

l £ , 19 May (USNM). Hamden: 19, 2 May (USNM). New

Haven: 1^, 5 May (USNM). Yale Preserve, New Haven:

4 £ . 5 $ , 18 Apr-9 May (USNM). West Rock Pk, New Haven:

2 $ , 24 May (USNM). DISTRICT OF COLUMBIA: Washington:

• DYSERIOCRANIA AURICYANEA (MALSINGHAM) "•. /" " \

» DYSERIOCRANIA GRISEOCAPITELLA (WALSINGHAM)

MAP 2.—Distributions of Nearctic Dyseriocrania in the United States.
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1 $ (holotype, E. griseocapitella) (BMNH); 1 3 , 1 $ (LACM);

1 ? 4 larvae, 20 May (USNM); Ft Totten: 3 larvae, 15 May

(USNM). FLORIDA: Alachua Co: San Felasco Hammock, 5

mi [8 km] SE Alachua: 1 9 , 22 Feb QBH). ILLINOIS: Palos

Pk: 1 $, 25 Apr (USNM). MARYLAND: Montgomery Co:

Forest Glen: \$, 2 May (USNM). Plummers Island: 4,$,

7 Apr (USNM). Prince Georges Co: Adelphi: 19, 2 May

(USNM). MASSACHUSETTS: Barnstable Co: Barnstable: 1 $,

15 May (CPK). Franklin Co: Leverett: \$ (USNM). Norfolk Co: Cohasset: 1^,1 1 May (USNM). Suffolk Co: Forest

Hills: 1£ , 19 , 12 May (USNM). MICHIGAN: Allegan Co:

T2N, R14W, Sec 7: 29 , 23 May (LACM); 19 , 23 May

(USNM). MISSISSIPPI: Oktibbeha Co: Mississippi State Univ:

1 $, 25 May (BrM). NEW HAMPSHIRE: Rockingham Co:

Hampton: 1$, ? May (LACM); 19 , 17 May (USNM). NEW

JERSEY: Essex Co: 1 $, 19 , 19 Apr (USNM). Essex Co Pk:

1 $, 19 , 21 Apr-^ May (USNM). Ocean Co: Lakehurst:

13, 6 May (USNM). NEW YORK: Bronx: Bronx Pk: 1 $,

29 Apr (AMNH)). Tompkins Co: Dryden, Ringwood Reserve: 39 . 8-15 May (USNM). Ithaca, Six Mile Ck: 15£ ,

58 9 , 7-15 May (USNM). Westchester Co: Pelham: 34 $,

68 9 , 21-30 Apr (AMNH); 1$, 19 , 21-30 Apr (USNM).

PENNSYLVANIA: Philadelphia Co: Mt. Airy (USNM). SOUTH

CAROLINA: Berkeley Co: McClellanville, The Wedge: 19 ,

15 Apr (RBD); 13 $ , 69 , 17-22 Mar (USNM). VIRGINIA:

Augusta Co: Staunton: 10 larvae, 22 May (USNM). Fairfax

Co: Falls Church: 19 , iss. 23 May (AMNH); 9$, 89 , iss.

9 Mar-3 Apr (USNM); 23 larvae, 9-18 May (USNM);

Minor Hills, E Falls Church: 1 larva, 15 May (USNM).

DISCUSSION.—Most of the references to this insect

in the literature, unfortunately, have been issued

under the name auricyanea. As discussed later in

this paper, under the latter species, an examination

of the type of auricyanea has revealed it to be the

senior synonym of D. cyanosparsella, a species

known only from California. Previous authors, evidently misled by Walsingham's (1882) statement of

the original type deposition (Peabody Academy of

Science, Salem, Massachusetts), assumed that auricyanea was an eastern insect. Actually, no mention

was made by Walsingham concerning the origin

of his specimen, and very likely he did not realize

himself that it was probably collected in cismontane California. Consequently, later authors, without closely examining the type specimen of auricyanea, preferred to use this earlier name in lieu

of D. griseocapitella, which was proposed 16 years

later. Dyseriocrania griseocapitella was never officially synonymized under auricyanea, although its

distinction had been questioned (Forbes, 1923).

Because of the rather poor condition of the

types of both D. auricyanea and D. griseocapitella,

casual comparison of the two would probably lead

one to conclude they were the same. Examination

of their genitalia, however, leaves no doubt regarding their true identity. In the male, the caudal margin of the vinculum in D. griseocapitella is superficially bilobed; the lobes of the uncus are widely

separated; and the base of the ventral branch of

the aedeagus is without lateral lobes. In D. auricyanea, the caudal margin of the vinculum is

deeply excavated on either side of a median lobelike extension, the apical lobes of the uncus are

not widely separated, and the ventral branch of

the aedeagus possesses a bilateral pair of small

lobes at its base. The females of these two, closely

related species are most easily separated by comparing the very different forms of their vaginal sclerites (Figures 294 and 297).

The first observation of the immature stages of

this species was by Kearfott (1902). This author

discovered larvae mining the leaves of Castanea

americana (=C. dentata) near Montclair, New Jersey. Because he was not able to rear any adults,

Kearfott was uncertain as to the identity of the

larvae in question; he did, however, suggest Dyseriocrania griseocapitella as a possibility.

The most complete account treating the life history of D. griseocapitella is that by Busck and

Boving (1914). The adults emerge in late winter

to early spring, which may be from late February

through March in the southern parts of its range

to as late as April and May farther north. According to Busck and Boving, the female abdomen

contains approximately 40 eggs. These are reported

to be relatively large, oblong (0.5 mm long and 0.2

mm in diameter), soft in texture with a whitish

color, and finely sculptured with minute dots. The

eggs are inserted singly into the young leaves of

oak and chestnut before the leaves have fully

opened. Usually the eggs are deposited near the

edge over the outer half of the leaf. Needham

et al. (1928) state that the eggs require 7 to 15 days

to hatch.

The mine (Figure 101) commences as a narrow,

linear passage extending toward the leaf margin.

This early stage of the mine is usually obliterated

as the mine is enlarged and often causes a characteristic fissure to form eventually in the leaf. Immediately following the serpentine stage, the mine

broadens to form a large, somewhat inflated blotch

suggestive of many coleopterous mines, so much

so that Kearfott (1902), after first discovering the

mines of this moth, believed them to be caused by
NUMBER 251 55

T

FIGURE 101.—Leaf mine of Dyseriocrania griseocapitella

(Walsingham) on Quercus alba L. (Scale = 10 mm.)

a beetle. The rather abrupt change from a linear

mine to a blotch mine probably signals the initiation of the second larval instar often characteristic

in certain other lepidopterous leafminers. Kearfott

noted that the larva usually sweeps in small circles

of 5 mm or more radius in feeding, with its anal

end functioning as a pivot. The final outline of

the mine often results from a series of these semicircles marked by a series of fine, concentric curved

lines. The blotch becomes full depth (i.e., with the

entire parenchyma within the mine being devoured), causing the mine to become semitransparent. Consequently, the larva and its dark frass,

usually lying loosely inside the mine, may be easily

observed through the leaf epidermis.

Larval feeding is rather rapid. Within 7 to 10

days the larva drops to the ground and burrows

into the soil as deep as 12 inches (30 cm). A relatively tough oval cocoon (Figure 102) measuring

approximately 3 mm long and 2 mm in diameter

is spun of silk and small particles of soil. The

larva diapauses inside this cocoon for several

months until sometime during the winter when it

finally pupates. Busck and Boving report the

FIGURE 102.—Pupal cocoons of Dyseriocrania griseocapitella

(Walsingham). Length of largest cocoon 3.1 mm.

greatly developed pupal mandibles to be heavily

relied upon by the pupa not only for emerging

from the cocoon but also for digging its way to the

surface of the soil. After emerging from the ground,

the pupa remains quiescent for an indefinite period

until final ecdysis can be performed. Rather soon

after emerging from the pupal shell, the adult is

capable of flight.

The larva of D. griseocapitella is easily distinguished from that of its nearest relative, auricyanea,

by several setal characters. Most significantly, seta

Al is lost in griseocapitella but present in auricyanea. The ninth abdominal segment of the former

demonstrates the most primitive chaetotaxy of the

family studied thus far in possessing 10 pair of

primary setae, including what is assumed to be

SD2 and SV2. In contrast, the ninth segment of

auricyanea possesses the more normal complement

of seven primary setal pairs.

As noted by Hinton (1946:11), Fracker's reference to the larva of Epimarlyria auricrinella Walsingham is incorrect. The description he provides

actually pertains to an eriocraniid, most probably

D. griseocapitella.

The larva and possibly pupa of this moth are

known to be parasitized by two genera of Ichneumonidae. Only one of these, Lathrolestes mnemoni-
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cae Rohwer, was previously reported. A second

parasite was discovered in the collections of the

National Museum of Natural History within a

cocoon collected and preserved by A. Busck. The

latter may represent an undescribed genus, near

Tersilochus, as determined by R. W. Carlson of

the U.S. Department of Agriculture. Both parasites

are believed to oviposit in the larva of D. griseocapitella during its mining phase but do not kill

the larva until it has completed its cocoon underground.

This particular host-parasite relationship appears

somewhat of an anomoly in that no other members in the respective subfamilies of these two

wasps are known to parasitize Lepidoptera. For

example, Townes (1969:13), evidently overlooking

previous literature references to the above host

record of Lathrolestes mnemonicae, stated that all

species of the subfamily Scolobatinae are internal

parasites of sawflies. Similarly, Townes (1971:33)

summarized that all members of the subfamily

Tersilochinae are parasitic on coleopterous larvae,

especially those of Curculionidae. Regarding the

latter, he further adds that the host species attacked

seem to be those with larvae in plant tissue and

pupation in the soil. Consequently, it appears that

at least some parasitic wasps that normally attack

other orders of insects have secondarily adapted to

D. griseocapitella largely due to certain similarities

in the general life history of the host. Further attempts to rear adult eriocraniids will undoubtedly

reveal additional associations of parasites.

Dyseriocrania auricyanea (Walsingham)

FIGURES 103-108, 167, 186-187, 222-223, 257-258,

296-298, 334; MAP 2

Micropteryx auricyanea Walsingham, 1882:204.—Riley, 1891:

111, no. 6016.

Eriocrania auricyanea (Walsingham).—Walsingham, 1898:162.

Eriocrania aurocyanea [sic] {Walsingham).—Walsingham,

1898:162.

Eriocephala auricyanea (Walsingham).—Dyar, 1902[1903]:581,

no. 6618.—Kearfott, 1903:125, no. 7165.

Mnemonica auricyanea (Walsingham).—Meyrick, 1912a: 19;

1912b:5— Barnes and McDunnough, 1917:197, no. 8477.—

McDunnough, 1939:110, no. 9852.

Eriocrania cyanosparsella Williams, 1908:14 [new synonym].

—Needham, Frost and Tothill, 1928:78.

Mnemonica cyanosparsella (Williams).—Meyrick, 1912a: 19;

1912b:5— Barnes and McDunnough, 1917:197, no. 8476.—

Keifer, 1927:138.—McDunnough, 1939:110, no. 9851 —

Armitage, 1951:111.

Dyserioaania cyanosparsella (Williams).—Opler, 1974:16.

ADULT (Figure 167).—Wing Expanse: d\ 10.5-13

mm; ?, 9.3-10.5 mm.

Head: Vertex and frons covered with long,

divergent white to predominantly white hairs intermixed with a scattering of pale brownish hairs.

Antennae approximately 0.65 the length of forewings, 44-48 segmented; scape whitish; flagellum

white ventrally and laterally, light fuscous dorsally.

Maxillary palpi pale brown; apex of terminal segment largely naked except for 5-6 apical setae and

2-3 subapical setae; setae very short, length less

than 2.0 their width; surface of apical region

rougher than in D. griseocapitella due to a scattered series of rudimentary, appressed spines. Labial

palpi whitish, particularly on ventral and inner

surfaces, with scattered, elongate brownish scales.

Thorax: Dorsum almost equally intermixed

with elongate hairs and broader, appressed scales

of white and light brown. Venter white. Legs

mostly white ventrally and bronzy brown dorsally.

Forewings pale golden brown, and rather distinctly

marked with several small patches of darker, purplish scales; heaviest patches of purplish scales

typically arranged into three narrow, irregular, and

approximately parallel bands obliquely slanted

from hind margin toward costa and apex; the

median fascia usually the most distinct with the

outer fascia the least well developed; purplish spots

also present along costal margin and outer margins

of wings; fringe uniformly light golden bown.

Hind wings slightly darker in color, more grayish

brown with a slight purplish luster.

Abdomen (Figures 222-223): Sparsely covered

pale grayish white hairs, usually more whitish ventrally. Fifth sternite of both sexes with a pair of

large, reniform tubercules situated toward lateral

margin of sternite; surface of tubercules swollen,

reticulated; tubercules wider than long; length approximately 0.5-0.6 the length of sternite. Fourth

sternite of female with a pair of moderately large

fenestral areas about half the diameter of 5th sternal tubercules; fenestrae situated more mesad than

glands; 4th sternite of male without fenestrae. Posterior margin of 8th segment of female not deeply

clefted, nearly simple.

Male Genitalia (Figures 257-258): Uncus shallowly bilobed, lobes reduced, rather narrowly sepa-
NUMBER 251 57

rated. Anal tube membranous, with several minute,

scattered spines; basal sclerites absent. Median lobe

from caudal margin of vinculum pronounced; margin deeply excavated on either side of median lobe;

anterior apophyses elongate, approximately 0.8 the

length of undivided vinculum. Juxta broadest anteriorly, then narrowing gradually to acute caudal

apex; greatest width approximately 0.7 its length.

Base of aedeagus moderately swollen, much less so

than in D. griseocapitella; ventral branch of aedeagus slender, with bilateral pair of small, rounded

lobes at basal sixth; apex of ventral branch simple,

acute.

Female Genitalia (Figures 296-298, 334): Apex

of ovipositor subacuminate to acute; lateral edges

serrulate, with 12-14 minute teeth. Bursa copulatrix of moderate length, not surpassing cephalic

apices of anterior apophyses when extended; walls

of corpus bursae with numerous minute spines

concentrated especially over caudal two-thirds. Vaginal sclerite moderately sclerotized; broad, greatest

width approximately 0.65 its length, not depressed,

with a prominent, triangular ventral keel; structure

of sclerite as in Figure 297.

LARVA (Figures 103-107).—Length of largest

larva 8.5 mm; diameter 1.2 mm.

Head: Relatively dark brown except for blackish outline of frontal sutures. Greatest width 0.6

mm; length 0.7 mm. Va situated approximately

midway between VI and V2. PI moderately elongate, arising either within or very approximate to

ecdysial line. Al present. Chaetotaxy of labrum

similar to that of D. griseocapitella; all setae simple, acute; anterior margin of pilifers and epipharynx densely spinose; M3 arising much closer

to M2 than L3. Mandibles with an inner median

tuft of 12-15 elongate, finely branched setae.

Thorax: Pronotal shield consisting of 2 pair

of irregular, brownish plates partially confluent

across midline. Prosternum with a large pair of

brownish plates partially confluent at midline;

plates roughly triangular in form with apex of

triangle directed anteriorly. Meso- and metathorax

without sclerotized plates. L2 relatively well developed on meso- and metathorax, approximately

equal to LI. Subventral setae trisetose on prothorax.

Abdomen: Integument without pigmentation,

appearing whitish except for a minute pair of longitudinal, brownish spots near lateroposterior margin of anal segment; barlike spots without enlarged cephalic end. LI present on all segments,

situated relatively high above SD1 and spiracles

on I to VIII. SV2 variable, but typically present

on segments I to VIII. Segment IX with 7 pair of

primary setae; SD2, L3, and SV2 absent.

TYPES.—Holotype, $, No. 1622 (Eriocrania auricyanea); in the Museum of Comparative Zoology,

Harvard University. Holotype cT, No. 3219 (Eriocrania cyanosparseila); in the California Academy

of Sciences.

TYPE-LOCALITIES.—Unknown [probably California] (Eriocrania auricyanea). At the base of Mt.

Tamalpais, Marin County, California, in a small

grove of oaks (Quercus agrifolia), (Eriocrania cyanosparsella).

HOST.—Fagaceae: "Quercus agrifolia Nee\"

Keifer, 1927; "Putative mines on Quercus douglasii

H. and A., Q. dumosa Nutt., Q. lobata Nee1 and

Q. wislizenii A.DC," Opler, 1974.

FLIGHT PERIOD.—Early March to mid-April; univoltine. The December through March emergence

dates of Keifer (see "Material Examined") have

not been considered here because they are suspected

to be laboratory rearings and, thus, probably do

not reflect natural emergence dates.

DISTRIBUTION (Map 2).—This species is known

only from California, where it occurs, according

to Opler (1974), from Shasta County southward in

the Coast ranges and western foothills of the Sierra

Nevadas to Santa Cruz Island and the San Gabriel

Mountains.

MATERIAL EXAMINED.—35 males, 17 females, and 1 larva.

UNITED STATES. CALIFORNIA: Alameda Co: Alameda:

1 $, 19 (CAS). Strawberry Canyon, Berkeley Hills: 1 9 ,

23 Mar, 1 larva, 11 Apr (UCB). Lake Co: Lakeport: 6$,

1-15 Apr (CAS). Lower Lake: 2$, 1$ , 15 Apr (CAS).

Marin Co: Mt Tamalpais: 1$ (holotype), 3 Mar (CAS);

1 $ (paratype), 3 Mar (CAS); 2 $ (paratypes), 3 Mar (USNM).

Novato: 19 , 6 Apr (CAS). San Luis Obispo Co: Black

Mts: 1 9 , 29 Mar (USNM). San Mateo Co: San Francisco:

2$, 14 Apr (LACM); 15 $ , 99 , 6 Dec-30 Mar (CAS);

3 3 , 39 , 7 Dec-29 Mar (USNM). Sonoma Co: Boyes Hot

Springs: 1 $, 31 Mar (CAS). Cazadero: 1 $, 12 Apr (CAS).

Other Records (not examined): CALIFORNIA: Alameda

Co: Berkeley, Univ. Calif, campus: 2 $, 26 Feb (UCB).

Contra Costa Co: Antioch: 29 , 26 Mar (UCB). 2 mi [3.2

km] E Antioch: 3$, 26 Mar (UCB). 2 mi [3.2 km] N

Orinda: 2$, 19 , 26 Feb (UCB). Russel Farm, 4 mi [6.4

km] NE Orinda: 1 $, 19 Apr (UCB). Richmond: 19 10 Apr

(UCB). El Dorado Co: Cool: 2 9 , 23 Mar (UCB). Marin

Co: Vicinity of Phoenix Lake: 1 £ , 2 Apr (UCB). Mendocino Co: 3 mi [4.8 km] N Branscomb, 1400 ft T427 ml:
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

• ••• ; ' •

^ —

• ^ • • ' - - • • •• • • ' :'•' •

^ y •

SV2-

>

n

•

^~-

"0

103

4DI

- t l

SVl

•

— -

FICURES 103-107.—Dyseriocrania auricyanea (Walsingham), larval chaetotaxy: 103, lateral view

of prothorax, mesothorax, and abdominal segments 1, 6, 8, and 9; 104, dorsal view of head

(scale = 0.3 mm); 105, left mandible (scale = 0.1 mm); 106, dorsal view of abdominal segments

8-10; 107, lateral view of head.
NUMBER 251

2$, 49 , 16-17 May (UCB). San Bernardino Co: 10 mi

[16 km] N Fontana, Lytle Creek Ranger Sta: 1$ , 17 Apr

(UCB). Santa Barbara Co: Prisoner's Harbor, Santa Cruz

Island: 4 9, 16 Mar (UCB). Santa Clara Co: Herbert Cr,

3 mi [4.8 km] W New Almaden: 1$, 18 Apr (UCB). Sierra

Co: Shenanigan Flat, 14 mi [22.4 km] W Donnieville: 1 $,

18 Apr (UCB). Ventura Co: Oxnard: 1$, 11 Apr (UCB).

DISCUSSION.—An examination of the holotype of

Dyseriocrania auricyanea has clearly shown that it

does not represent our common eastern eriocraniid

as once believed; instead it was discovered to be

the senior synonym of the California species

formerly known as D. cyanosparsella. This change

in names is an unfortunate but necessary action.

As summarized in the discussion of the preceding

species, D. griseocapitella, a number of morphological characters, particularly in the male and female

genitalia, easily distinguish these two closely related species. They can most quickly be identified

by their widely disjunct distributions and different

forewing maculation. The forewings of D. auricyanea are lighter in color with a definite golden

luster and are more distinctly marked with dark,

oblique but irregular fasciae.

The life history of D. auricyanea has recently

been reviewed (under the name cyanosparsella) by

Opler (1974). In general, its life history is very

similar to that of its eastern counterpart. The eggs

are inserted singly into opening leaves in early

spring. Normally, the usual oviposition site is on

the apical half of the leaf. A small egg pouch is

normally loosely attached to one side of the oviposition scar formed by the impression of the previously hatched egg. The eggs are believed to hatch

in approximately the same period (7-15 days) as in

D. griseocapitella.

Upon emerging, the larva develops rapidly,

passing through at least four instars. Opler encountered larvae in the field as early as 14 March

in the southern part of its range and as late as

19 May farther north. The first instar mine begins

as a linear or slightly curved passage to the leaf

margin. Following this stage, the mine is quickly

enlarged to an inflated, full length blotch. Larval

feeding during this period usually progresses in a

series of semicircular swaths. At maturity, the larva

leaves the mine through a slit in the upper leaf

epidermis and drops to the ground. It burrows into

the litter beneath the host tree and forms a whitish

silken cocoon covered with darker soil particles.

FIGURE 108.—Leaf mine of Dyseriocrania auricyanea (Walsingham) on Quercus agrifolia Nee. (Scale = 10 mm.)

The larva diapauses through the summer until

sometime during the winter when it pupates.

The adults emerge in the late winter or early

spring. Opler reports the moths most active in the

hours just before sunset, although they are capable

of flight at any time during the daylight hours.

The mature larva is easily distinguished from

that of its eastern sister species, D. griseocapitella,

by the presence of Al arising from the adfrontal

sclerite (absent in griseocapitella) and by the reduction of setae on the ninth abdominal segment.

In D. auricyanea only seven primary setae are present compared to 10 in griseocapitella. From Eriocraniella aurosparsella, the larvae of D. auricyanea

differ in retaining setae SV3 on the prothorax, LI

on abdominal segments I-VIII, as well as the man-
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dibular setal tuft (all absent in E. aurosparsella).

The larva of the only other California eriocraniid

studied, E. xanthocara, new species, differs from D.

auricyanea in having lost the cranial seta Al.

Eriocrania Zeller

Eriocrania Zeller, 1851:323.—Chapman, 1900:94, 96.—Rebel,

1901:246.—Meyrick, 1912a:4; 1912b: 17—Chapman, 1917:

310, 312.—Tillyard, 1919:97, etc; 1923:181, 195.—Forbes,

1923:65.—Philpott, 1926:371, 374; 1927:319.—Meyrick, 1928:

870.—Seyrig, 1928:261, 262.—Fletcher, 1929:84.—Hering,

1932:2—Kuznetsov, 1941:19, 20.—Hinton, 1946:6, etc.—

Lhomme, 1949:1221.—Viette, 1949:32; 1951:43, 44.—Hering,

1951:40, etc.—Gerasimov, 1952:189.—Heath, 1958:117.—

Toll, 1959:16, 20.—Kristensen, 1968a:70, 71; 1968b:241, etc.

—Mutuura, 1972:1055.—Common, 1973:13, 15, 16.—Dugdale, 1974:134.—Birket-Smith and Kristensen, 1974:157,

etc.—Razowski, 1975a:5, 7.—Heath, 1976:157.

Eriochania [sic].—Forbes, 1923:65, fig. 38.

Chapmania Spuler, 1910:483 [preoccupied, not Montic, 1893]

[type-species: Lampronia semipurpurella Stephens, 1835]—

Fletcher, 1929:45, 84 [synonym of Eriocrania].—Hering,

1932:2—Kuznetsov, 1941:19 [synonym of Eriocrania].—

Viette, 1951:43, 44 [synonym of Eriocrania].—Razowski,

1975:48 [synonym of Eriocrania].—Heath, 1976:157 [synonym of Eriocrania].

Allochapmania Strand, 1917:137 [type-species: Lampronia

semipurpurella Stephens, 1835].—Fletcher, 1929:10, 84

[synonym of Eriocrania].—Viette, 1949:32; 1951:43, 44

[synonym of Eriocrania].—Gerasimov, 1952:189 [synonym

of Eriocrania].—Toll, 1959:16, 20, 23.—Razowski, 1975:48

[synonym of Eriocrania].—Heath, 1976:157 [synonym of

Eriocrania].

TYPE-SPECIES.—Lampronia semipurpurella Stephens, 1835; designated by Meyrick, 1912a.

ADULT.—Small moths with relatively broad wings;

wing expanse: 7.8-13.5 mm.

Head: Maxillary palpi with basal 0.9 of terminal segment densely setose; apical 0.1 largely naked;

primary setae simple, without lateral barbs; apex

of terminal segment rounded to subacute, simple,

without secondary lobes but with 7-8 minute, unstriated, subacute setae concentrated around extreme apex; setae mostly of uniform length, very

short and stout, usually less than 0.5 the diameter

of apex in length.

Thorax: Venation somewhat reduced, with Sc

and Rl simple in both fore and hind wings; R2

and 3 completely fused in both wings; R4 and 5

stalked approximately half their length in both

wings. Hind wings with scales elongate, very slender

and hairlike.

Abdomen: Fenestrae usually present on sternite of female, either absent in male or rudimentary. Sternal tubercules present on 5th sternite of

both sexes, relatively small, papiliform.

Male Genitalia: Uncus deeply bilobed; lobes

divergent, lateral margins straight, tapering slightly

to apex. Socii present. Anal tube completely membranous except for a bilateral pair of minute, relalatively free sclerites near base. Vinculum with median caudal margin only smoothly convex or slightly

produced to form a short lobe, not deeply clefted;

cephalic margin with a median, triangular process

projecting anteriorly between lateral apophyses.

Caudal apex of juxta distinctly asymmetrical, subacute, with a short, median longitudinal fold and

an even shorter, transverse fold to the left of median fold. Aedeagus with ventral phallic branch

very slender, usually half the diameter of dorsal

branch, and simple, without prominent lobes or

processes.

Female Genitalia: Vaginal sclerite variable in

structure, usually in the form of a broad, depressed

plate with cephalic margin of plate broadly

rounded; cephalic 0.5-0.8 of plate sometimes curved

sharply ventrad forming a prominent ventral process; midventral keel relatively low and poorly developed; a sclerotized, median process typically extending caudally from basal area of midventral keel.

Ninth segment without midventral spinose pocket.

DISCUSSION.—The genus Eriocrania appears to be

a rather well-defined, monophyletic assemblage of

several Holarctic species. The group may be easily

distinguished by several synapomorphic features,

such as the loss of one of the radial veins (R3,

probably through fusion with R2) in both the fore

and hind wings and by a number of characters of

the male genitalia, principal among these being

the presence of paired, basal sclerites embedded in

the anal tube; the asymmetrical caudal apex of the

juxta with its peculiar folds; and, perhaps, the median projection of the cephalic margin of the vinculum. The male genitalia appear so uniform

among the members of this genus that frequently

it is difficult to identify species on this basis alone.

The female genitalia may be partially characterized by the presence of a slender, sclerotized process

(MP) extending medially and caudally from the

base of the relatively poorly developed midventral

keel of the vaginal sclerite.

Most of the previous references to the original

description of this genus (e.g., Meyrick, 1912a, b,
NUMBER 251

61

etc.) are in error in listing page 338 of Zeller's

1851 paper. The author and title are correct but

the pagination is not. The name Eriocrania actually is not mentioned on page 338 but is instead

first referred to on page 323. On pages 322 and 323

Zeller explains that he believes Micropterix auctorum to consist of two groups. The species he includes in his first group (i.e., Micropterix) comprises those species he numbers 1 through 10 in his

paper. The latter group consists of the species he

lists starting with number 11 (sparmannella Bosc)

through 16, and for these he proposes the name

Eriocrania on page 323. The pagination given by

Meyrick and others refers to the page on which

the included species of Eriocrania begin (no. 11)

and not the actual reference of the generic name.

The following treatment includes the description

of one new species and one new subspecies for this

genus. An additional and possibly undescribed species has not been named at this time but is briefly

discussed under E. semipurpurella pacifica, new

subspecies.

Key to the Species and Subspecies of Nearctic Eriocrania

1. Apex of ovipositor relatively short; extreme tip not strongly attenuated (Figure 335)

Eriocrania breviapex, new species

Apex of ovipositor more attenuated (Figure 336) 2

2. Ventral phallic branch with approximately 4-6 minute, subapical spines (Figure 262). Distribution northeastern United States and Canada

Eriocrania semipurpurella semipurpurella

Ventral phallic branch smooth, without subapical spines (Figure 259). Distribution northwestern United States Eriocrania semipurpurella pacifica, new subspecies

Eriocrania semipurpurella semipurpurella

(Stephens)

FIGURES 71-76, 168, 192-194, 224-225, 261-262, 305-309,

336; MAP 3

Lampronia semipurpurella Stephens, 1834, vol. 4:359.—Wood,

1845:230.

Micropteryx semipurpurella (Stephens).—Stainton, 1850:34.—

Zeller, 1851:322, 343 [division Eriocrania].—Stainton 1854:

45.—Herrich-Schaffer, 1855, vol. 5:394.—Wocke, 1877 [1876]:

779.—Snellen 1882:1068.—Wood, 1890:3,4.

Eriocrania semipurpurella (Stephens).—Rebel, 1901:247.—

Meyrick, 1912a:4; 1912b:2.—Chapman, 1917:314, pi. 86.—

Philpott, 1925:458, 459; 1926:723, 1927:320.—Meyrick, 1928:

870.—Needham, Frost, and Tothill, 1928:72—Seyrig, 1928:

259, 260.—Pierce and Metcalf, 1935:110.—Lhomme, 1949:

1221—Ford, 1949:206.—Kuznetsov, 1941:19.—Gerasimov,

1952:193.—Hering, 1957, vol. 1:178.—Heath, 1958:118.—

Kristensen, 1968:241, etc; 1970:404.—Common, 1973:15,

20.—Birket Smith and Kristensen, 1974:157, etc.—Razowski, 1975a:51; 1975b:4, 5, 6.—Heath, 1976:165.

Chapmania semipurpurella (Stephens).—Spuler, 1910:482,

483.

Allochapmania semipurpurella (Stephens).—Toll, 1959:17 etc.

—Issiki, 1971:7.

Micropteryx amentella Zeller, 1850:63.—Stainton, 1854:45

[synonym of Micropteryx semipurpurella].—Herrick-Schaffer, 1955, vol. 5:394.—Wocke, 1877[1876]:779.—Razowski,

1975:51 [synonym of Eriocrania semipurpurella].

Eriocrania amentella (Zeller).—Rebel, 1901:247 [synonym of

Eriocrania semipurpurella].—Meyrick, 1912a:4 [synonym

of Eriocrania semipurpurella]; 1912b:3 [synonym of Eriocrania semipurpurella].

Micropteryx inconspicuella Wood, 1890:3.—Razowski, 1975:

51 [synonym of Eriocrania semipurpurella].—Heath, 1976:

165 [synonym of Eriocrania semipurpurella].

Eriocrania inconspicuella (Wood).—Rebel, 1901:247 [synonym

of Eriocrania semipurpurella].—Meyrick, 1912a:4 [synonym

of Eriocrania semipurpurella]; 1912b:3 [synonym of Eriocrania semipurpurella]; 1928:870 [synonym of Eriocrania

semipurpurella].—Pierce and Metcalf, 1935:110 [synonym

of Eriocrania semipurpurella].—Ford, 1949:206 [synonym

of Eriocrania semipurpurella].—Gerasimov, 1952:193 [synonym of Eriocrania semipurpurella].—Heath, 1958:118

[synonym of Eriocrania semipurpurella].

ADULT (Figure 168).—Wing Expanse: tf, 10-

13.5 mm; $ , 10-12.5 mm.

Head: Predominantly covered with long whitish hairs but almost equally intermixed at vertex

with light fuscous hairs. Antennae relatively long,

approximately 0.65-0.70 the length of forewing,

34- to 36-segmented; scape brownish fuscous, irrorated with white; flagellum mostly whitish beneath,

brownish fuscous dorsally. Maxillary palpi predominantly brownish fuscous, with scattered white

scales ventrally; apex of terminal (5th) segment

relatively blunt, with approximately 8 short, apical

setae of equal length. Base of galeae sparsely covered with silvery white scales dorsally. Labial palpi
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

brownish fuscous above, silvery white below and

on inner margins.

Thorax: Dorsum predominantly brownish fuscous, irrorated with whitish hairs; tegulae equally

intermixed with long whitish and brownish fuscous hairs. Venter sparsely clothed with slender

whitish scales. Legs brownish fuscous above with

most of leg, particularly the femur, white ventrally.

Forewings mostly fuscous, with a distinct purplish

iridescence, heavily mottled with pale, whitish

spots; markings small and randomly scattered over

wing except for a large, narrowly triangular subtornal spot extending halfway across wing from

hind margin and extending into basal margin of

fringe immediately basad of tornus; fringe along

outer margin uniformly grayish fuscous. Hind

wings much paler, sparsely covered with very

slender scales, uniformly grayish.

Abdomen (Figures 224-225): Sparsely covered

with long grayish to pale brownish hairs above,

whitish ventrally. Fourth sternite of female with a

pair of moderately large, oval fenestrae approximately 0.35 the length of sternite in diameter;

fourth sternite of male without fenestrae. Sternal

tubercules present on 5th sternite of both sexes,

papiliform, diameter about 0.27-0.30 the length

of sternite. Caudal apex of 8th tergite of female

distinctly and deeply divided; caudal apex of 8th

sternite slightly clefted about 0.3 the distance of

tergal cleft.

Male Genitalia (Figures 261-262): Uncus deeply

bilobed; lobes prominent, widely separated at apex

for a distance equivalent to their length. Anal tube

membranous; a bilateral pair of minute, basal

sclerites present. Vinculum with median area of

caudal margin only slightly raised, smoothly curved

posteriorly; apophyses elongate, approximately 0.7

the length of undivided vinculum. Juxta broadest

anteriorly, gradually tapering posteriorly to a rather

broad, subacute apex; greatest width approximately equaling length. Base of aedeagus prominently swollen; ventral phallic branch slender without apical or basal lobes, about 0.5 the diameter

of dorsal branch; apex of ventral branch acute,

relatively simple except for 4-6 subapical exophallic spines.

Female Genitalia (Figures 305-309; 336): Apex of

ovipositor accuminate; lateral edge serrulate, with

12-14 minute teeth. Bursa copulatrix of moderate

length, usually not surpassing cephalic apices of

posterior apophyses when fully extended; walls of

corpus bursae completely membranous, without

spines. Vaginal sclerite darkly sclerotized, broad,

often relatively short but variable in length, greatest width approximately equaling length (viewed

dorsally); cephalic apex of sclerite extended ventrally into a broad, elongate, slightly curved process

as in Figures 306-309.

TYPES.—Syntype, o" (Lampronia semipurpurella); in the British Museum (Natural History).

Syntypes, 1 tf and 1 $ (Micropteryx [sic] amentella);

in the British Museum (Natural History). Syntype(s),

unmarked and unsegregated {Micropteryx [sic] inconspiniella); presumably in the J. H. Wood collection of the British Museum (Natural History).

TYPE-LOCALITIES.—Darenth Wood [50°25'40"N,

0°16'E], Kent, England (Lampronia semipurpurella). Not specifically stated (Micropteryx amentella). Not stated (Micropteryx inconspicuella); presumably in the vicinity of Tarrington, Ledbury,

England.

HOST.—Betulaceae: "Betula pendula Roth,"

Lhomme, 1949; "Betula pubescens Ehrh.," Toll,

1959; "Betula platyphylla var. japonica (Miq.)

Hara," Kumata (in Litt.).

FLIGHT PERIOD.—Late April to late May; univoltine.

DISTRIBUTION (Map 3).—In North America this

species is known to occur from eastern Ontario

east to Nova Scotia and south to southern New

York. According to Heath (1976), it occurs widespread in Great Britain, through northern and

central Europe to Japan.

MATERIAL EXAMINED (North American Records only).—

3 males and 5 females. CANADA. NOVA SCOTIA: Boulderwood, Halifax: \$, 11 May (NSM); 1^,1 4 May (USNM).

Kings Co, Coldbrook: 1 $ , 18 May (NSM). Ontario: Black

Sturgeon Lake [49°20'N, 88°52'W]: 19 , 31 May (CNC).

Ottawa: 19, 4 May (CNC). QUEBEC: Lac Mondor [46°37'N,

72°46'W], Ste. Flore: 1$, 4 May (CNC); 1$ , 15 May

(USNM). UNITED STATES. NEW YORK: Westchester Co:

Pelham: 1 9 , 21-30 Apr (AMNH).

DISCUSSION.—The vaginal sclerite of this species

appears to be quite variable, both in the Old and

New World populations. The rather broad, spatulate anterior process is the most variable feature

of the sclerite, both in relative length and in the

degree of ventral curvature. Figures 306 and 309

provide some idea as to the range of this variation

in North American specimens.
NUMBER 251 63

The aedeagus of the male provides the most

diagnostic character in recognizing this species and

in separating it from the following new subspecies

known only from the northwestern United States.

The ventral phallic branch of E. semipurpurella

semipurpurella possesses a row of approximately

four to six short, rather appressed subapical spines

that are lacking in E. semipurpurella pacifica.

Although fenestrae are typically absent from the

fourth sternite in all eriocraniid males, a few males

of E. semipurpurella semipurpurella exhibit paired,

faintly depigmented areas on this sternite. These

areas are even more evident in the males of E. semipurpurella pacifica.

Lectotypes have not been designated for this species in the present study because tentative selections, to be published eventually, have already been

made by others.

Eriocrania semipurpurella pacifica, new subspecies

FICURF.S 109-120, 169, 190-191, 218, 226-227, 259-260,

299-301, 337; MAP 3

Eriocrania semipurpurella.—Clarke, 1942:276 [misidentification; not Stephens, 1835].

Eriocrania species.—Heath, 1976:165.

ADULT (Figure 169).—Wing Expanse: tf, 10-11

mm; 9, 10-11 mm.

Head: Sparsely covered with brownish fuscous to

grayish hairs. Antennae moderately long, 0.55-0.6

the length of forewing, 35-37 segmented; scape and

flagellum uniformly clothed with brownish fuscous

scales. Maxillary palpi predominantly brownish fuscous with basal segments whitish beneath; apex of

terminal (5th) segment relatively simple, not lobed,

with an apical cluster of 6-7 minute, subacute setae

of equal lengths, length of setae approximately 0.3

the diameter of apex. Base of galeae largely naked,

pubescent, but with a small tuft of whitish scales

over extreme base arising from labrum. Labial

palpi predominently brownish fuscous, with inner

sides strongly suffused with white.

Thorax: Dorsum and tegulae brownish fuscous.

Venter sparsely clothed with brownish fuscous to

grayish hairs. Legs mostly pale fuscous to grayish

above, paler, more whitish beneath. Forewings

brownish fuscous with a slight golden brassy iridescence; a small, whitish tornal spot present along

margin near vein CuA2; fringe brownish fuscous

except for white at tornal spot. Hind wings paler,

more thinly scaled; grayish in color; scales slender,

hairlike.

Abdomen (Figures 226-227): Sparsely covered

with long brownish scales above, more grayish beneath. Fourth sternite of female with a pair of welldefined, transverse-elliptical fenestrae approximately

0.35-0.40 the length of segment in greatest diameter,

greatest (transverse) diameter about 1.8 the smallest

diameter; 4th sternite of male with fenestrae poorly

defined, consisting of a pair of irregular, unpigmented areas. Sternal tubercules present on 5th

sternite of both sexes, tubular in form, diameter

approximately 0.20-0.22 the length of segment.

Caudal apex of 8th tergite of female well defined,

deeply forked; apex of 8th sternite only slightly

divided, but with furcations widely separated.

Male Genitalia (Figures 259-260): Uncus deeply

bilobed; lobes prominent, widely separated at apex

for a distance equivalent to their length. Anal tube

membranous, with a bilateral pair of minute, basal

sclerites present. Vinculum with median area of

caudal margin slightly convex, smoothly curved

posteriorly; anterior apophyses elongate, approximately 0.72-0.75 the length of undivided vinculum.

Juxta broadest anteriorly, gradually tapering posteriorly to a relatively broad, subacute apex; greatest width approximately 0.6 its length. Base of

aedeagus moderately swollen; ventral phallic branch

very slender, about 0.5 the diameter of dorsal

branch, apex smooth without minute spines; length

of ventral branch slightly shorter than dorsal

branch.

Female Genitalia (Figures 299-301, 337): Apex

of ovipositor accuminate, lateral edges serrulate,

with 12-14 minute teeth. Bursa copulatrix relatively

short, terminating far short of cephalic apices of

posterior apophyses when extended; walls of corpus

bursae completely membranous, without spines.

Vaginal sclerite darkly sclerotized, relatively broad,

but with slender caudal extension, greatest width

approximately 0.4 its length; cephalic two-thirds of

sclerite a flattened, oval plate with a low, median

ventral keel as shown in Figure 301.

LARVA (Figures 109-119).—Length of largest larva

7 mm; diameter 1 mm.

Head: Uniformly pale brown except for dark

outline of adfrontal sutures and mandibles. Greatest width 0.4 mm; length 0.55 mm. Va located usually posterior to VI, sometimes laterad; PI minute,
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

o

/ •

•

FIGURES 109-113.—Eriocrania fsemipurpurella pacifica, new subspecies, larval chaetotaxy: 109,

lateral view of prothorax, mesothorax, and abdominal segments 1, 6, 8, and 9; 110, dorsal

view of head (scale = 0.3 mm); 111, left mandible (scale - 0.1 mm); 112, dorsal view of

abdominal segments 8-10; 113, lateral view of head.
NUMBER 251 65

FIGURES 114-119.—Eriocrania }semipurpurella pacifica, new subspecies, larval head structure:

114, dorsal view of anterior margin of head, X 245; 115, chaetotaxy of labrum, X 925; 116.

lateral view of head, X 285; 117, gular setae, X 540; 118, anteroventral view of mouthparts,

X 300; 119, ventral view of labrum, X 350. {Photographs reduced to 84%.)
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

arising closely adjacent to ecdysial line; Al present;

chaetotaxy of labrum very similar to that of Dyseriocrania, with all setae simple, not compressed;

anterior margin of pilifers and epipharynx densely

spinose; M2 and M3 closely approximate, nearly

contiguous, arising near median cleft remote from

L3. Mandibles without inner medial setal tuft.

Thorax: Pronotal plate consisting of 2 pairs of

usually confluent, irregular, light brownish areas.

Prosternum with a single, large brownish plate;

meso- and metathorax without pigmentation. L2

subequal to LI on meso- and metathorax. Subventral setae usually trisetose on prothorax.

Abdomen: Integument without pigmentation, appearing whitish (in alcohol) except for a pair ot

minute, longitudinal dark brownish spots near

lateroposterior margin of anal segment; barlike

spots without enlarged cephalic end. LI present on

all segments, situated relatively high above SD1 and

spiracles on I to VIII. Segment IX with 7 pair of

primary setae; SD2, L3, and SV2 absent.

HOLOTYPE.— cf, Seward, Alaska, 23 May 1951,

W. J. Brown; in the Canadian National collection.

PARATYPES.—UNITED STATES. ALASKA: Third Judicial

Division: Seward: 1 $, 23 May 1951, W. J. Brown, slides

USXM 18232, 18233 (USXM). WASHINGTON: Whatcom Co:

1 3 , 3$ , 3-8 Apr 1923, J. F. Clarke, slides USNM 18070,

18256, 18257, 18258 (USNM). Described from a total of 3

males and 3 females.

Other Material Examined: CANADA: British Colombia:

Vancouver Island: 6 larvae, coll. B. H. Tothill, USNM slides

18237, 18260-18262, 18273 (USNM).

HOST.—Questionable; possibly Rosaceae: "Holodiscus discolor Maxim," from specimen label.

FLIGHT PERIOD.—Early April to late May; univoltine.

DISTRIBUTION (Map 3).—Adults of this subspecies

are presently known from only two localities in

southern Alaska and northwestern Washington.

Larvae, suspected to represent this insect, have also

been collected on Vancouver Island. All localities

represent low coastal habitats bordering the northern Pacific Coast Mountain System.

DISCUSSION.—This subspecies, except for minor

morphological differences, is very similar to the

typical subspecies, Eriocrania semipurpurella semipurpurclla. It may be distinguished from the latter

by the entirely smooth apex of its ventral phallic

branch as compared to the typically spinose condition in E. s. semipurpurella (Figures 259, 262).

Their vaginal sclerites differ slightly, although individual variation obscures these differences somewhat. For example, the slender, caudal third of the

sclerite appears more strongly ridged dorsally in

E. s. pacifica than in the nominate subspecies. The

larval and host associations summarized below, if

proven correct, also indicate at least subspecific recognition for E. s. pacifica. The name pacifica is

intended to refer to the general distribution of this

subspecies, which presently is restricted to the

northwestern borders of the Pacific Mountain

System.

Six, probably immature, larvae collected from

Vancouver Island, British Columbia, have been tentatively identified as this subspecies. No adults have

been reared or otherwise definitely associated with

these larvae; consequently, the present determination will have to remain questionable until rearings

can be conducted. The specimens are of interest

because they reportedly were mining the leaves of

Holodiscus discolor Maxim (r: Spiraea discolor). If

the host identification is accurate, then these larvae

represent not only the first record of an eriocraniid

feeding on a member of the Rosaceae, but also a

host association that is rather remote from the more

typical Betulaceae-Fagaceae associations typical of

the other members of Eriocraniidae. Holodiscus discolor is a woody shrub attaining a maximum height

of approximately 5 to 6 meters. It occurs naturally

in western North America from British Columbia

south to California. The shrub is also utilized somewhat as an ornamental and, thus, may occur more

widely under cultivation.

In chaetotaxy the larvae resemble Dyseriocrania

auricyanea closely except they lack the prominent

mandibular setal tuft of that and a few other species. From Eriocraniella aurosparsella they differ in

typically possessing three subventral setae on the

prothorax and seta LI on the abdominal segments

I to VIII. The prothoracic seta SV3 may occasionally be lacking (as in specimen slide USNM 18273)

because of individual variation.

A closely related, but possibly distinct species,

may also occur within the general range of E. s.

pacifica. A single male specimen with identical male

genitalia (including smooth, ventral phallic branch)

was collected by Walsingham at Camp Watson in

Grant County, Oregon, during April 1872. This

specimen is now deposited in the British Museum

(Natural History). I am indebted to N. P. Kristen-
NUMBER 251 67

T

1

FIGURE 120.—Leaf mine of Eriocrania hemipurpurella pacifica,

new subspecies, on Holodiscus discolor Maxim. (Scale =

10 mm.)

sen for calling my attention to this specimen as well

as to the fact that the wings of the Oregon specimen, particularly the hind wings, are unusually

narrow. For example, the width of the hind wings

of the Oregon specimen is only 0.25 their length,

a ratio considerably less than any other eriocraniid

examined during the course of this study. By comparison, this same ratio measures 3.0 or more in the

specimens of E. s. pacifica. Because the Oregon

specimen is unique, it can not be decided now

whether the specimen is atypical with regard to

relative wing width. Any further consideration of

the taxonomic status of this specimen will have to

await the discovery of additional material.

Eriocrania breviapex, new species

FIGURES 27-28, 170, 188-189, 228, 245, 302-304, 335; MAP 3

ADULT (Figure 170).—Wing Expanse: 9, 7.5-9

mm.

Head: Sparsely covered with whitish hairs intermixed with light fuscous. Antennae relatively long,

0.6-0.7 the length of forewing, approximately 35-

segmented; scape whitish; flagellum mostly brownish fuscous and irrorated with white above, entirely

white beneath. Maxillary palpi light brownish fuscous; apex of terminal segment rather blunt,

crowned with a series of about 8 short spines of

equal lengths. Base of galeae clothed with whitish

scales. Labial palpi whitish, with a few scattered

fuscous hairs ventrally.

Thorax: Dorsum mostly light brownish fuscous

with a slight bronzy luster; tegulae with a tuft of

long white and brownish fuscous hairs equally intermixed. Venter white. Legs typically light brownish fuscous dorsally, white ventrally. Forewings

light fuscous with a slight bronze to purplish iridescence, irregularly marked with small pale golden

to brassy spots; a distinct and relatively large subtornal spot present, extending halfway across wing

to middle of discal cell; fringe uniformly pale

brownish fuscous except for whitish suffusion of

subtornal spot. Hind wings distinctly paler than

forewings, sparsely scaled with long, slender, hairlike brownish-fuscous scales.

Abdomen (Figure 228): Sparsely covered with

long grayish brown hairs above, whitish ventrally.

Fourth sternite with a pair of moderately large

oval fenestrae 0.52-0.54 the length of sternite in

greatest diameter. Sternal tubercules present on 5th

sternite, papiliform, diameter about 0.23-0.26 the

length of sternite. Caudal apex of 8th tergite distinct and deeply divided, branches divergent; caudal apex of 8th sternite likewise distinct and

divided approximately 0.5 the distance of tergal

cleft.

Female Genitalia (Figures 302-304, 335): Apex

of ovipositor relatively broad, abruptly narrowing

to a short, acute tip; lateral margins serrulate,

usually with 11-12 minute teeth. Bursa copulatrix

relatively long, surpassing ceplialic ends of posterior
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

apophyses when fully extended by nearly half its

length; walls of corpus bursae entirely membranous. Vaginal sclerite darkly sclerotized, broad,

greatest width approximately 0.5 its length; cephalic end broad, relatively flat, not extended

ventrally; a low, median keel present as shown in

Figure 304.

HOLOTYPE.— 9, Six Mile Creek, Ithaca, Tompkins County, New York, 12 May, 1961, R. W.

Hodges, photograph on file USNM, USNM Type

72102; in the National Museum of Natural History,

Smithsonian Institution.

PARATYPES.—UNITED STATES. New York: Tompkins

Co: Six Mile Cr, Ithaca: 1 9, 10 May 1959, R. W. Hodges,

9 genitalia slide USNM 17206, head slide USNM 18208

(USNM); 1$, 7 May 1961, R. W. Hodges, 9 genitalia

slide USNM 17199 (USNM); 1 $. 13 May 1961, R. W.

Hodges, wing slide USNM 16156, 9 genitalia slide USNM

17499 (USNM). Described from a total of 4 females.

HOST.—Unknown.

FLIGHT PERIOD.—Early to mid-May; univoltine,

DISTRIBUTION (Map 3).—This species is presently

known only from the type-locality, which is located

in the Cayuga Lake Basin of northwestern New

York.

DISCUSSION.—Although no males have been examined thus far, Eriocrania breviapex is easily distinguished from all other Eriocraniidae by the

rather abbreviated apex of its ovipositor. This

diagnostic feature has suggested its name, which

is derived from the Latin brevis (short) and apex

(top).

This species demonstrates greatest affinities to

E. semipurpurella pacifica, new subspecies, from

the northwestern United States, particularly with

regard to the similar form of their vaginal sclerites.

That species, however, as well as the northeastern

E. semipurpurella semipurpurella, can be readily

separated from E. breviapex on the basis of the

above-mentioned ovipositor character.

Eriocraniella Viette

Eriocraniella Viette, 1949:30.—Opler, 1973:1321— BirketSmith and Kristensen, 1974:159.

TYPE-SPECIES.—Micropteryx aurosparsella Walsingham, 1880; original designation and by monotypy.

ADULT.—Small moths with relatively broad wings;

wing expanse: 6-10 mm.

Head: Maxillary palpi with terminal segment

densely setose nearly to apex; primary setae barbed

laterally; apex of terminal segment complex, consisting of a single relatively large, apical lobe bearing 4 stout setae and 2 pair of smaller, subapical

lobes, each bearing a single large seta; the most

subapical pair of lobes the most reduced in size;

all setae arising from the above lobes longitudinally

striated.

Thorax: Epiphysis present. Forewings with Sc

usually simple, rarely divided; Rl divided approximately 0.4 its length; R2, 3, 4, and 5 all arising

separate from cell. Hind wings with Sc and Rl

simple; R2 and 3 entirely fused; R4 and 5 separate;

scales relatively short and broad, not hairlike.

Abdomen: Fourth sternite of females with a pair

of well-defined, relatively small, oval to circular

fenestrae; fenestrae absent in male. Sternal tubercules present on 5th sternite of both sexes, similar

in form, papiliform, relatively small.

Male Genitalia: Uncus well developed, deeply

divided to form 2 elongate lobes with rounded or

tapered apices; lobes usually distinctly curved from

middle outward. Anal tube entirely membranous,

without basal sclerites. Socii present. Caudal margin

of vinculum variously modified, either produced

into a slightly raised, median lobe (subgenus Disfurcula) or with a prominent, median furcate

process, or furcula, arising submarginally (subgenus

Eriocraniella); cephalic margin of vinculum

smoothly concave, without a median process.

Caudal apex of juxta broad, subtruncate to slightly

concave; caudal end with a series of minute, transverse ridges. Aedeagus with ventral phallic branch

stout, usually equaling dorsal branch in diameter;

apex of ventral branch with a nearly transparent,

spinose flabellate process in subgenus Eriocraniella; apex more simple and acute in subgenus Disfurcula but with a single, subapical conical or

spinose process.

Female Genitalia: Vaginal sclerite with cephalic

end typically broad, usually slightly flared, and

with a distinct midventral keel. Ninth segment

with a midventral, invaginated pocket laterally

bordered with numerous minute spines in subgenus Eriocraniella; spinose pocket absent in subgenus Disfurcula.

DISCUSSION.—As presently defined, the genus

Eriocraniella consists of at least eight species, including one not named in this paper, all of which
NUMBER 251 69

70 160 100 60 40 30

# Eriocrania breviapex, new species

• Eriocrania semipurpurella pacifica, new subspecies

A Eriocrania semipurpurella semipurpurella (Stephens)

•^ Eriocraniella longifurcula, new species

Eriocraniella platyptera, new species

MAP 3.—Distribution of Nearctic Eriocrania and Eriocraniella (subgenus Eriocraniella) not

recorded in California.

are endemic to the Nearctic region. As originally

described by Viette (1949), the members of this

genus are characterized by their venation, particularly by the undivided subcosta (though often

divided in E. falcata) and divided first radius, with

all radial veins present in the forewings and arising

separate. In addition the extreme apex of the

maxillary palpi is very similar among all the

species examined in possessing three distinct sets

of minute lobes, each bearing a specific number

of mostly elongate, striated setae. Similar to the

following genus, Neocrania, Eriocraniella is also
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

partially characterized by the presence of an epiphysis.

The genus appears to have speciated well in

North America, and further field work undoubtedly will result in the discovery of additional species in our fauna. One such species is illustrated

(Figure 267) in this paper but not named because

of a lack of adequate material. It appears most

closely allied to E. xanthocara, new species, and is

mentioned further under the discussion of that

species.

On the basis of genital characters, the genus may

be further divided into two distinct groups or subgenera. The presence of a furcula in the male

appears to be correlated with the development of

a midventral, spinose pocket near the posterior end

of the ninth segment in the female. Although their

respective functions are not known, it is possible

that these two morphological features may only be

secondary sexual characteristics and that their

function may be interrelated. Future observations

may show, for example, that the male furcula somehow engages the spinose potket of the female during copulation.

In addition to the above, possibly interrelated,

secondary sexual characteristics, the two subgenera

also differ with respect to other autapomorphous

features. In the subgenus EriocranieUa, the apex of

the ventral phallic branch typically bears a somewhat transparent, flabellate process with a marginal

fringe of small to large spines. The males of the

subgenus Disfurcula do not possess this structure;

instead, the members of this group demonstrate

a more simplified phallic apex, which has, however,

developed a relatively prominent, subapical lobe

or spinelike process.

Key to the Subgenera of EriocranieUa

Male genitalia with a prominent, median furcate process, or furcula, arising immediately

anterior to caudal margin of vinculum (Figure 263). Ninth segment of female with a midventral pocket lined laterally with numerous, minute spines (Figure 310) EriocranieUa

Vinculum of male without midventral furcula. Ninth segment of female without midventral,

spinose pocket Disjurcula, new subgenus

Subgenus EriocranieUa Viette

TYPE-SPECIES.—Micropteryx aurosparsella Walsingham, 1880.

ADULT.—Wing Expanse: 7-10 mm.

Head, Thorax, and Abdomen: As described for

genus.

Male Genitalia: Vinculum with a prominent,

median furcate process, or furcula, arising from

caudal submarginal area. Apex of ventral phallic

branch relatively complex and enlarged, with a

nearly transparent flabellate process projecting

from extreme tip; a series of 6—12 variably developed spines arranged around outer margin of

process.

Female Genitalia: Caudal end of 9th segment

with a shallow midventral pocket lined laterally

with several rows of minute spines; spines sometimes more prominent on one side than on other.

Key to the Species of the Subgenus EriocranieUa

1. Male with furcula divided 0.5 its length (Figure 263). Female with ventral, spinose pocket

of 9th segment asymmetrical, heavily thickened and spinose on left side (Figure 310) 2

Male with furcula divided at least 0.7 its length or more (Figure 269). Female with ventral

pocket symmetrical, forming a thickened spinose ring on both sides 3

2. Head vestiture grayish white irrorated with fuscous. Forewings brownish fuscous, with a

slight bronze to light purplish iridescence

EriocranieUa (EriocranieUa) aurosparsella (Walsingham)

Head vestiture pale yellowish orange. Forewings a darker fuscous, with a distinct greenish

iridescence EriocranieUa (EriocranieUa) xanthocara, new species

3. Head vestiture grayish white. Forewings relatively long (length 4.0-4.5 mm) and slender

(width approximately 0.30 the length). Distribution: southwestern United States

EriocranieUa (EriocranieUa) longifurcula, new species
NUMBER 251 71

Head vestiture pale yellowish orange. Forewings relatively short (length 3.0-3.5 mm) and

broad (width approximately 0.34 the length). Distribution: northeastern United States

Eriocraniella (Eriocraniella) platyptera, new species

Eriocraniella (Eriocraniella) aurosparsella

(Walsingham)

FIGURES 31-32, 121-134, 174, 197-198, 229-230, 246, 263-264,

310-312, 339; MAP 4

Micropteryx aurosparsella Walsingham, 1880:83.—Riley, 1891:

111, no. 6017.

Eriocephala aurosparsella (Walsingham).—Dyar, 1902 [1903]:

581, no. 6620.—Kearfott, 1903:125, no. 7167.

Mnemonica aurosparsella (Walsingham).—Meyrick, 1912a: 19;

1912b:5 — Barnes and McDunnough, 1917:197, no. 8478 —

McDunnough, 1939:110, no. 9853.

Eriocraniella aurosparsella (Walsingham).—Viette, 1949:31.—

Davis, 1975:10, 33.

Eriocraniella species, Opler, 1974:4.

ADULT (Figure 174).—Wing Expanse: $, 8.0-9.0

mm; 9 , 7.5-8.5 mm.

Head: Relatively sparsely covered with whitish

to grayish hairs intermixed with fuscous hairs.

Antennae relatively short, approximately 0.35-0.38

the length of forewing, 31- to 35-segmented; scape

dark fuscous above, white beneath; flagellum

almost entirely covered with narrow, dark fuscous

scales above and laterally except for apical 3-4

segments that are thinly scaled with white; venter

of flagellum mostly devoid of scales, instead

densely covered with brownish pubescence. Maxillary palpi usually entirely fuscous; distal 2 segments

sometimes slightly irrorated with grayish white;

apex of terminal (5th) segment divided into 3 series

of minute lobes as follows: (a) a single, relatively

elongate, apical lobe bearing one long (length ca.

1.7 the diameter of lobe), blunt seta and 3 short

(length less than diameter of lobe), subapical subacute setae, (b) an identical pair of moderately

long, subapical lobes, each bearing one long, subacute apical seta, and (c) identical subapical pair of

very short lobes, each bearing one long, apical

subacute seta. Base of galeae covered with whitish

scales irrorated with fuscous. Labial palpi fuscous

above and on outer sides, silvery white along inner

sides.

Thorax: Dorsum smoothly covered with broad,

dark fuscous scales, slightly bronzy in iridescence;

tegulae with long tufts of predominantly fuscous

hairs irrorated with white. Venter grayish white.

Legs usually fuscous above, whitish ventrally. Forewings immaculate, brownish fuscous, usually with

a distinct bronzy to light purplish iridescence;

fringe uniformly brownish fuscous. Hind wings

similar to forewings in color though less lustrous,

uniformly covered with relatively broad scales.

Abdomen (Figures 229-230): Thinly clothed with

brownish fuscous hairlike scales above, grayish

white ventrally. Fourth sternite of female with

oval and relatively small fenestrae; diameter of

fenestrae approximately 0.22-0.25 the length of

sternite; fenestrae absent in male. Sternal tubercules present on 5th sternite of both sexes, papiliform; diameter approximately 0.2 the length of

sternite. Caudal apices of 8th tergite and sternite

in female normally entire, not divided, acute and

well defined.

Male Genitalia (Figures 263-264): Uncus deeply

bilobed; lobes prominent, curved laterally; apices

broadly rounded, widely separated for a distance

approximately equal to their length. Anal tube

membranous; basal sclerites absent. Caudal margin

of vinculum with a prominent, deeply divided

process, or furcula, arising medially; length of

furcula approximatetly equaling length of lobes

of uncus; lobes of furcula slender, acute, about

0.5 the length of entire furcula. Anterior apophyses

moderately long, 0.55-0.6 the length of undivided

vinculum. Juxta broadest anteriorly, narrowing

slightly posteriorly to a rather broad, concave apex;

greatest width approximately 0.75 its length. Base

of aedeagus bulbous; ventral phallic branch slightly

sinuate, stout, equaling the dorsal branch in diameter; apex of ventral branch with a cluster of

9-10 large exophalic spines; ventral branch loosely

articulated to bulbous base of dorsal branch.

Female Genitalia (Figures 310-312, 339): Apex of

ovipositor acuminate; lateral edges serrulate, with

6—8 minute teeth. Bursa copulatrix of moderate

length, not surpassing cephalic apices of anterior

apophyses when extended; wall of corpus bursae

completely membranous. Vaginal sclerite darkly

sclerotized, moderately long, greatest width 0.60-

0.65 its length; cephalic apex of sclerite broad, with

a relatively well-developed, rounded ventral keel

as in Figure 311.

Caudal apex of 9th segment with a prominent,
72

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

\

K122

FIGURES 121-126.—Eriocraniella aurosparsella (Walsingham), larval chaetotaxy: 121, lateral view

of prothorax, raesothorax, and abdominal segments 1, 6, 8, and 9; 122, dorsal view of head

(scale = 0.3 mm); 123, left mandible (scale = 0.1 mm); 124, dorsal view of abdominal segments

8-10; 125, detail of barlike spot or "brace-rod" of tenth segment; 126, lateral view of head.
NUMBER 251 73

FIGURES 127-132.—Eriocraniella aurosparsella (Walsingham), larval head structure; 127, anterior

view of mouthparts, X 575; 128, chaetotaxy of labrum, X 1250; 129, detail of labial seta M3

with dentate margin, X 4950; 130, lateral view of head, X 350; 131, antenna, X 1100; 132,

ventral view of mouthparts, X 325. (Photographs reduced to 83%.)
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spinose pocket; lateral margin of left side of pocket

greatly thickened and more spinose, with approximately 15-17 minute teeth; right side poorly

developed.

LARVA (Figures 121-132).—Length of largest

larva 7.0 mm; diameter 1.0 mm.

Head: Uniformly dark brown with frontal sutures darkly outlined. Greatest width 0.5 mm,

length 0.6 mm. Va variable, situated either anterior, laterad, or posterior to VI. PI moderately

elongate, several times larger than A2, usually

arising from or contiguous to ecdysial line. A2

usually laterad to PL Al present. Labrum usually

with apices of M2, M3, and L3 broadly compressed

(Figure 129), subtruncate, with a subapical row of

3-4 minute but relatively prominent teeth along

ventral margin. M3 arising approximately equidistant between M2 and L3; anterior margin of

pilifers and epipharynx spinose. Mandibles with

inner margins smooth; mesal setal tufts absent.

Thorax: Pronotal and prosternal plates well developed, dark brown. Meso- and metathorax without sclerotized plates. L2 reduced in size on mesoand metathorax, approximately 0.3 the length of

LI. Subventral setae bisetose on prothorax.

Abdomen: Integument without pigmentation,

whitish except for a minute pair of longitudinal,

brownish spots near lateroposterior margin of anal

segment; barlike spot with cephalic end slightly

enlarged and punctate; anal segment largely unpigmented. LI absent on I to VIII. SD2 present

on I to VIII. Segment IX with 7 pair of primary

setae; SD2, L3, and SV3 absent.

TYPE.—Holotype, sex unknown; in the British

Museum (Natural History).

TYPE-LOCALITY.—Southern Oregon.

HOST.—Fagaceae: "Quercus kellogii Newb.,"

Opler, 1974.

FLIGHT PERIOD.—Late February to late May; univoltine.

DISTRIBUTION (Map 4).—This species is known

to range from southern Oregon south through the

coastal ranges of California to Santa Clara County

and Santa Cruz Island.

MATERIAL EXAMINED.—80 males, 15 females, and 15 larvae.

UNITED STATES. CALIFORNIA: Alameda Co: Berkeley: 1 9 ,

22 May (USNM). Contra Costa Co: Orinda. 3 $, 5 9 , 26 Feb6 Mar, Quercus kellogii (UCB); 1^, 6 Mar, Quercus kellogii

(USNM). El Dorado Co: Blodgett Forest, 12 mi [19.2 km]

E Georgetown: 10$, 69 , 20 May (UCB). Sly Park Dam:

8 larvae, 19 May (UCB); 6 larvae, 19 May (USNM). Mendocino Co: Hopland Field Sta Hilq., 900-1600 ft [274-488 m]:

2 9, 27 Apr (UCB). Hopland Field Sta, Kelsey Cab Orch

area, 2600-2800 ft [793-854 m]: 2 larvae, 29 Apr (UCB);

1 larva, 29 Apr (USNM). Santa Barbara Co: Prisoner's Harbor, Sta Cruz Island: 19, 16 Mar (UCB); 19, 16 Mar

(USNM). Tulare Co: Fairview: 19, 27 Apr (UCB). Tuolumne Co: South Fork Stanislaus R, 4 airline mi [6.4 km]

NE Columbia: 35 $, 3 Apr (UCB); 4 $, 28 Mar-3 Apr

(USNM); 25 $, 28 Mar-3 Apr (CAS). Tuolumne City, 3 mi

[4.8 km] NE: 1 $, 23 Mar (UCB); 1 $, 23 Mar (USNM).

DISCUSSION.—Walsingham described Eriocraniella

aurosparsella from a unique specimen collected at

an unspecified locality in southern Oregon during

May 1872. Essig's (1941) account of Walsingham's

itinerary through Oregon during that period does

not provide sufficient information to pinpoint the

type-locality of this species more accurately. The

sex of this specimen was not stated by Walsingham

and cannot be determined now due to the loss of

the abdomen. Fortunately, the identity of E. aurosparsella can be established from the wing venation

of the holotype as well as by noting Walsingham's

own description. For example, he described the

head as being "greyish," which is one of the key

characters in recognizing this species from its nearest relative.

Eriocraniella aurosparsella can be readily distinguished from all other members of the family, with

the exception of E. xanthocara, on the basis of

genitalic characters. Both E. aurosparsella and E.

xanthocara possess essentially the same genital and

venational structure, a fact that reflects the close

degree of relationship and relatively recent divergence of these two insects. For example, both species possess a prominent furcula, divided for half

its length in the male; furthermore, both also demonstrate a unique development of the spinose

pocket wherein the left margin has become heavily

sclerotized and densely spined. The two species may

be distinguished by their different coloration. Most

noticeably, the head vestiture of E. aurosparsella is

whitish intermixed with fuscous compared to the

pale orange head of E. xanthocara. Also, the forewings of E. aurosparsella are not as dark and are

more bronzy than in E. xanthocara. These two species also demonstrate host preferences with the

larvae of E. aurosparsella, apparently restricted to

Quercus kellogii, and those of E. xanthocara feeding on Q. agrifolia and Q. wislizenii.

In the reference by Opler (1974), the hosts of
NUMBER 251 75

121 120 119 117 116 115

• Eriocraniella aurosparsella (Walsingham)

• Eriocraniella xanthocara, new species

CALIFORNIA INSECT SURVEY

Department of Entomology and Parasltology

UNIVERSITY OF CALIFORNIA

0 25 50 75 100

MAP 4.—Distribution of Eriocraniella (subgenus Eriocraniella) recorded in California.
76

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

these two species were associated differently than

listed above. The reason for this discrepancy is

simply that the names of the two moths were misapplied, with Opler referring to the orange-headed

species as Eriocraniella aiirosparsella and the grayheaded form as an undescribed species of Eriocraniella. All specimens associated with Quercus kellogii

examined thus far, however, possess a head vestiture

of whitish to pale gray irrorated with fuscous; consequently, they agree more closely with Walsingham's original diagnosis of Eriocraniella aiirosparsella.

The larva of E. aiirosparsella differs significantly

from that of E. xanthocara. Eriocraniella aurosparsella can be distinguished in the larval state by the

presence of the cranial seta Al, the absence of the

mesal tuft of mandibular setae, and by the absence

of LI on the abdominal segments I through VIII.

In contrast, the larva of E. xanthocara has lost Al

FIGURE 133.—Leaf mine of Eriocraniella ?aurosparsella (Walsingham) on Quercus kellogii Newberry (Scale = 10 mm.)

FIGURE 134.—Leaf mine of Eriocraniella 7aurosparsella (Walsingham) on Quercus kellogii Newberry showing oviposition

scar and egg pouch.

but has retained the mandibular setal tuft and LI

on all abdominal segments.

The larval mine (Figure 133) of this species appears very similar to that of E. xanthocara in that

an elongate, linear mine first extends from a brownish egg pouch and leaf scar usually toward the

outer margin of the leaf. After following the leaf

margin a short distance, the linear mine abruptly

enlarges to form a large, full-depth blotch.

Eriocraniella (Eriocraniella) xanthocara,

new species

FIGURES 135-146, 175, 182-183, 199-200, 231-232, 265-266,

313-314, 340; MAP 4

Eriocraniella aurosparsella.—Opler, 1974:4, 18 [misidentification; not Walsingham, 1880].

ADULT (Figure 175).—Wing Expanse: tf, 7.0-8.0

mm; 9, 7.5-8.5 mm.

Head: Relatively sparsely covered with pale
NUMBER 251 77

orange to pale ochreous hairs. Antennae of moderate length, approximately 0.5-0.55 the length of

forewing; 33- to 37-segmented; scape dark fuscous

above, white beneath; flagellum almost entirely

covered with narrow, dark fuscous scales except for

apical 4 segments that are thinly scaled with white;

venter of flagellum with a narrow pubescent streak

along midline. Maxillary palpi entirely fuscous except for terminal (5th) segment that is covered

mostly with grayish pubescence; apex of 5th segment similar to E. aurosparsella, divided into 3

series of minute lobes as follows: (a) a single, relatively elongate apical lobe bearing 1 large (length

ca. 1.6 the diameter of lobe), blunt seta, and 3 short

(length less than diameter of lobe), subapical subacute setae, (b) a pair of moderately long, subapical

lobes, each bearing 1 long apical seta, and (c) a

very short subapical pair of lobes, each bearing one

long apical seta. Base of galeae sparsely scaled with

dark fuscous. Labial palpi fuscous above and on

outer sides, mostly silvery white on inner sides.

Thorax: Dorsum smoothly covered with broad,

dark fuscous scales, slightly bronzy to purplish in

luster; tegulae with long tufts of fuscous hairs posteriorly and whitish hairs anteriorly. Venter grayish

white. Legs usually fuscous above, whitish ventrally.

Forewings immaculate, dark fuscous with a prominent greenish blue iridescence; fringe uniformly

dark fuscous. Hind wings similar to forewings in

color though less lustrous and with more purplish

iridescence; uniformly covered with relatively broad

scales.

Abdomen (Figures 231-232): Thinly covered with

dark fuscous hairs above, more grayish ventrally.

Fourth sternite of female with relatively small, oval

fenestrae similar in size to those of E. aurosparsella;

male without fenestrae. Sternal tubercules present

on 5th sternite of both sexes, similar in size and

form to those of E. aurosparsella. Caudal apices of

8th tergite and sternite usually entire, sometimes

weakly divided, acute and well defined.

Male Genitalia (Figures 265-266): Very similar to

that figured and described for E. aurosparsella.

Female Genitalia (Figures 313-314, 340): Very

similar to that figured and described for E. aurosparsella. Spinose pocket similar to E. aurosparsella

in being asymmetrical, with lateral margin of left

side more heavily sclerotized and spinose.

LARVA (Figures 135-145).—Length of largest larva

9 mm; diameter 1.4 mm.

Head: Relatively uniformly light brown to stramineous with frontal sutures darkly outlined. Greatest width 0.6 mm; length 0.7 mm. Va variable, often

situated between VI and P2. PI moderately elongate, several times larger than A2, usually separate

from ecdysial line but occasionally contiguous. Al

absent. Chaetotaxy of labrum similar to that of

Dyseriocrania; all setae simple and acute; anterior

margin of pilifers and epipharynx densely spinose;

M3 closer to M2 than to L3. Mandibles with an

inner median tuft of 11-15 elongate setae; each

seta finely branched, appearing serrate.

Thorax: Pronotal and prosternal plates present

but inconspicuous in mature larvae (in alcohol),

and lightly pigmented; much darker, brownish in

young larvae. Meso- and metathorax with sclerotized plates. L2 relatively well developed on mesoand metathorax, approximately equal to LI. Subventral setae trisetose on prothorax.

Abdomen: Integument without pigmentation in

mature larvae, appearing whitish (in alcohol) except for a pair of minute, longitudinal brownish

spots near lateroposterior margin of anal segment;

barlike spots without enlarged cephalic end; anal

plate brownish in immature larvae. LI present on

all segments, situated relatively high above SD1

and spiracles on I to VIII. SV2 present on segments

I to VIII. Anal segment with 7 pair of primary

setae; SD2, L3, and SV2 absent.

HOLOTYPE.— cf» Russel Farm, 4 mi [6.4 km] NE.

Orinda, Contra Costa County, California, 6 March

1970, collector P. A. Opler, Quercus agrifolia; in

the California Academy of Sciences on indefinite

loan from the University of California, Berkeley.

PARATYPES— UNITED STATES. CALIFORNIA: Alameda Co:

Berkeley Hills, NE Oakland, 1400 ft [427 m]: 1^, 8 Apr

1964, J. Powell (UCB). Contra Costa Co: 2 mi [3.2 km] E.

Antioch: 8 larvae, 7-8 Apr 1969, Quercus agrifolia, P. Opler

(UCB); 5 larvae (same data), larval slides USNM 18150,

18151, 18153, 18155, 18222 (USNM). Russel Farm, 4 mi

[6.4 km] NE Orinda: 1$, 1$ , 6 Mar 1970, P. Opler, $

slides USNM 16096, 16097, Quercus agrifolia (USNM); 3^ ,

2 $ , 6 Mar 1970, P. Opler, Quercus agrifolia (UCB). Strawberry Canyon, Berkeley Hills: 2 larvae, 11 May 1967, Quercus agrifolia, J. Powell (UCB). Kern Co: 3 mi [4.8 km]

W Wofford Hts: 1 $, 28 Apr 1964, J. Doyeh, $ slide USNM

16098 (USNM). Marin Co: Alpine Lake, hills N 1100 ft

[335 m]: 19 , 17 Apr 1970, P. Opler, Quercus agrifolia, $

slide USNM 16101 (USNM). Noyato: 1 $, 25 Mar 1962,

D. Rentz, $ slide PO 94 (UCB). San Mateo Co: Golden

Gate Pk, San Francisco: 1 $ , 28 Mar 1961, P. Arnaud, Jr.,

on leaf of Quercus agrifolia (UCB). Junipero Serra Pk: 4 $ ,
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

P.

' ' . <

^ / /

y

s

135

FIGURES 135-139.—Eriocraniella xanthocara, new species, larval chaetotaxy: 135, lateral view

of prothorax, mesothorax, and abdominal segments 1, 6, 8, and 9; 136, dorsal view of head

(scale = 0.3 mm); 137, left mandible (scale = 0.1 mm); 138, dorsal view of abdominal segments

8-10; 139, lateral view of head.
NUMBER 251 79

FIGURES 140-145.—Eriocraniella xanthocara, new species, larval head structure: 140. anterior

view of head, X 220; 141, chaetotaxy of labrum, X 8(>0; 142, lateral view of head, X 275;

antenna, X 760; 144, ventral view of mouthparts, X 330; 145, detail ventral view of maxillae

and labrum, X 570. {Photographs reduced to 83%.)
80

29 Mar-4 Apr 1964, P. Arnaud, Jr., on foliage Quercus

agrifolia (CAS); 1 9 , 29 Mar 1964, P. Arnaud, Jr., 9 slide

USNM 18197 (USNM). San Francisco, Lake Merced: 1$ ,

9 Apr 1964, P. Arnaud, Jr. (CAS). Santa Clara Co: Herbert

Cr, 3 mi [4.8 km] W New Almaden: 1? , 18 Apr 1969, J.

Powell, flushed ex Q. agrifolia 3:00 P.M., 9 slide PO 92

(UCB). Described from a total of 8 males, 13 females, and

15 larvae.

Other Records (not examined): Contra Costa Co: Redwood

Regional Pk: 1 9 , 29 Apr 1975, P. Rude (UCB). Marin Co:

2 mi [3.2 km] W Fairfax: 19,1 3 Mar 1970, J. Powell (UCB).

Mendocino Co: NCCRP, 3 mi [4.8 km] X Branscomb, 1400

ft [427 m]: 2$, 49 , 16-17 May 1975, Powell and Schlinger

(UCB).

HOST.—Fagaceae: "Quercus agrifolia Ne'e and Q.

wislizenii A. DC," Opler, 1974.

FLIGHT PERIOD.—Early March to late April; univoltine.

DISTRIBUTION (Map 4).—According to Opler

(1974), this species occurs in California from Shasta

County southward through the coast ranges to at

least Santa Clara County and Santa Cruz Island.

DISCUSSION.—Eriocraniella xanthocara agrees

closely with E. aurosparsella in genital structure,

but it may be easily distinguished from the latter

on the basis of general scale color. The light orange

head vestiture of E. xanthocara contrasts sharply

with the whitish to gray head of Walsingham's species, a fact that has suggested its specific name, derived from the Greek, xanthos (orange) and kara

(head). In addition, the wings of E. xanthocara are

darker and exhibit a more bluish luster than does

E. aurosparsella. In the few specimens compared,

the vaginal sclerite (Figure 314) of E. xanthocara

appears more narrow at its cephalic end than does

that of E. aurosparsella, but this may merely represent normal populational variation.

The specific distinction of the two insects in

question appears justified, especially in view of the

fact that they occur at least partially sympatric and

are restricted to different hosts, as pointed out by

Opler (1975:5). As discussed under the preceding

species, however, Opler assigns host associations to

the two species different from those listed in the

present treatment.

The greatest degree of morphological differentiation between E. xanthocara and E. aurosparsella

may be observed in their larvae. The larva of E.

xanthocara is easily distinguished from E. aurosparsella by the absence of cranial seta Al, the presence of a prominent tuft of setae arising from the

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

T

1

FIGURE 146.—Leaf mine of Eriocraniella ?xanthocara, new species on Quercus wislizenii A.DC.

(Scale = 10 mm.)

inner margins of the mandibles, and by the presence of LI on all abdominal segments. In contrast,

E. aurosparsella has retained Al but has lost the

mandibular setal tufts and LI from abdominal segments I through VIII. In addition, the barlike, lateral pair of spots on the anal segment possess a

minute, cephalic enlargement in the latter species

that is not present in E. xanthocara. The anal shield

of the latter may also be darker in color, but this

should be verified with living material.

Opler (1974) reports the life history of this species to be similar to Dyseriocrania auricyanea

(= cyanosparsella). He has observed the two species

occurring sympatrically and synchronically at a

number of localities in coastal California and on

Santa Cruz Island. The primary distinction between the life histories of these two species is apparent in their oviposition sites and mine config-
NUMBER 251 81

urations. The eggs of E. xanthocara are not deposited in as great a concentration in the apical half

of the leaf, as is characteristic for D. auricyanea,

but are scattered more over the outer four-fifths.

The linear stage of mine construction differs in that

the length of this portion is noticeably longer in

E. xanthocara than in D. auricyanea.

An additional, unnamed species of Eriocraniella

is known from San Luis Obispo, California, where

it was collected during March by A. H. Vachell.

The unique male specimen, now deposited in the

National Museum of Natural History, Smithsonian

Institution, has not been named because of its poor

condition. The species resembles E. xanthocara in

wing color and possesses a similar head vestiture

of pale reddish orange. The furcula of the male

genitalia (Figure 267) differs noticeably by being

less deeply divided than that of either E. aurosparsella or E. xanthocara. The terminal flabellate

process of the ventral phallic branch also is more

elongate with very reduced, apical spines.

Eriocraniella (Eriocraniella) longifurcula,

new species

FIGURES 147-162, 176, 233, 269-270; MAP 3

ADULT (Figure 176).—Wing Expanse: tf, 9.5-10

mm.

Head: Sparsely covered with whitish to grayish

hairs. Antennae of moderate length, approximately

0.5 the length of forewing, 36-segmented; scape fuscous above, irrorated with white beneath; flagellum

mostly covered with slender fuscous scales except

for a narrow streak of pubescence ventrally. Maxillary palpi fuscous, slightly irrorated with white ventrally; apex of terminal (5th) segment very similar

in structure to E. platyptera, with a series of minute

lobes arranged in 3 ranks as follows: (a) a single,

relatively large apical lobe bearing 4 moderately

long subacute setae, (b) an identical pair of much

shorter subapical lobes, each bearing a single, elongate subacute seta, and (c) an identical pair of very

short subapical lobes, each with a single, elongate

subacute seta. Base of galeae sparsely scaled with

fuscous. Labial palpi fuscous with a slight whitish

irroration on apical segment.

Thorax: Dorsum fuscous; tegulae with long whitish and fuscous hairs; collar with long, mostly

whitish hairs. Venter whitish. Legs fuscous above,

mostly white ventrally. Forewings relatively slender,

width about 0.3 the length, immaculate, fuscous

with a slight bluish purple iridescence; fringe uniformly fuscous. Hind wings slightly paler than forewings in color, grayish, uniformly covered with relatively broad scales.

Abdomen (Figure 233): Thinly covered with fuscous hairs above, more grayish ventrally. Fourth

sternite of male without fenestrae. Sternal tubercules present on 5th sternite, tubular in form, diameter approximately 0.17 the length of sternite.

Male Genitalia (Figures 269-270): Uncus deeply

bilobed; lobes prominent, distinctly curved laterally,

lobes widely separated at apex for a distance slightly

greater than their length. Anal tube completely

membranous, without basal sclerites. Vinculum with

lateral arms of furcula greatly extended, slender;

furcula divided approximately 0.7-0.8 its entire

length; median cephalic margin of vinculum between apophyses smoothly concave, without median

process; apophyses elongate, approximately 0.8 the

length of undivided vinculum (excluding furcula).

Juxta broad, with lateral margins nearly parallel;

greatest width about 0.75 its length; caudal apex

broadly truncate. Base of aedeagus greatly swollen,

bulbous; ventral phallic branch stout, equaling

diameter of dorsal branch; apex of ventral branch

with a minute, flabellate serrated lobe bearing 5-7

marginal teeth; dorsal branch with a short, broadly

conical, acute process arising subapically; ventral

branch loosely articulated to bulbous base of

aedeagus.

LARVA (Figures 147-161).—Length of largest larva

7 mm; diameter 1 mm.

Head: Uniformly light brown to stramineous

with frontal sutures darkly outlined. Greatest width

0.5 mm; length 0.6 mm. Va variable, either laterad

or posterior to VI. PI relatively elongate, several

times larger than A2, either contiguous or slightly

separate from ecdysial line. A2 usually well separated from and laterad to PI. Al present. Labrum

similar to E. aurosparsella, with apices of M2, M3,

and L3 broadly compressed (Figure 155); ventral

margins of foregoing setae entire, not dentate as in

E. aurosparsella; M3 arising approximately equidistant between M2 and L3; anterior margin of

pilifers and epipharynx spinose. Mandibles with

inner margin smooth; median setal tufts absent.

Thorax: Pronotal and prosternal plates well developed, dark brown. Meso- and metathorax with-
82

y

o

y

v

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

147

to

FIGURES 147-151.—Eriocraniella Hongifurcula, new species, larval chaetotaxy: 147, lateral view

of prothorax, mesothorax, and abdominal segments 1, 6, 8, and 9; 148, dorsal view of head

(scale = 0.3 mm); 149, left mandible (scale = 0.1 mm); 150, dorsal view of abdominal segments

8-10; 151, lateral view of head.
NUMBER 251 83

FIGURES 152-157.—Eriocraniella }longifurcula, new species, larval head structure: 152, dorsal

view of frontal area, X 220; 153, chaetotaxy of lahrum, X 580: 154, detail of lalmim. X 1100;

155, lateral detail of labral setae L3 and M3, X 3550; 156, lateral view of head, X 280; 157,

anterior view of head, X 265. (Photographs reduced to 84%.)
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 158-161.—Eriocraniella tlongifurcula, new species, larval head structure: 158, lateral

view of antenna and ocellus (stemmata), X 850; 159, anterior view of antenna and ocellus,

X 1200; 160, ventral view of mouthparts, X 400; 161, detail of maxillae and spinneret, X 715.

(Photographs reduced to 84%.)

out plates, whitish. L2 reduced in size on meso- and

metathorax, approximately 0.3-0.5 the length of

LI. Subventral setae bisetose on prothorax.

Abdomen: Integument without pigmentation,

usually whitish (in alcohol) except for a pair of

minute, longitudinal brownish spots near lateroposterior margin of anal segment; barlike spots

similar to those of E. aurosparsella in form, with

cephalic end slightly enlarged and punctate. LI

absent on I to VIII. SD2 typically present on I to

VIII. Segment IX with 7 pair of setae; SD2, L3, and

SV3 absent.

HOLOTYPE.—<3\ Workman Creek, Sierra Ancha

[Mts.], Gila County, Arizona, 3 May 1947, H. and

M. Townes, USNM 72101, genitalia slide USNM

18071; in the National Museum of Natural History,

Smithsonian Institution.

PARATYPE.—ARIZONA: Coconino Co: Oak Cr Canyon: 1 $,

25 May 1974, 5200 ft [1585 m], genitalia slide DRD 2959,

wing slide 2990, R. Leuschner (RHL). Described from 2

males.

Other Material Examined: ARIZONA: Coconino Co: North

Rim Oak Cr Canyon: 3 larvae, 3 Jun 1968, coll. P. Opler,

on Quercus gambelli X arizonica, JAP 68F13, larval slides

DRD 3010, 3020 (UCB); same data: 2 larvae, larval slides

USNM 18146, 18147, (USNM).

HOST.—Uncertain; possibly Fagaceae: "Quercus

gambelli Nutt. X arizonica Sarg.," from specimen

labels.

FLIGHT PERIOD.—May; probably univoltine.
NUMBER 251

85

DISTRIBUTION (Map 3).—This species is presently

known from only two sites located in adjacent

counties of central Arizona.

DISCUSSION.—Only two male specimens, both

from Arizona and in relatively poor condition, are

all that were available of this species for study. The

species is interesting because it represents another

member of the genus Eriocraniella with a welldeveloped furcula.

The deeply divided furcula of this species readily

distinguishes it from all other known Eriocraniidae

with the exception of the following species, E.

platyptera. The furculae of both species are very

similar in possessing slender, elongate arms. Eriocraniella longifnrcula may be separated from the

latter by its grayish head, more elongate wings, and

more western distribution. In contrast, E. platyptera possesses a head vestiture of pale orange,

shorter and relatively broader wings, and a more

eastern distribution. The female of E. longifurcula

is unknown and, thus, cannot be compared with

that of platyptera. The specific name, longifurcula,

is derived from the Latin, longus (long) and furcula

(little pitchfork) and obviously refers to the elongate furcate arms of this species.

The identity of the larvae described for E. longifurcula must remain questionable because no

adults have definitely been associated with these

larvae. The specimens in question consist of five,

perhaps mature, examples collected 3 June 1968

from a possible hybrid of Quercus gambelli Nuttall X arizonica Sargent. Because the origin of this

sample (north rim of Oak Creek Canyon, Coconino

County, Arizona) is very near the site where the

adult paratype was collected, it is possible that the

larvae do represent this species.

Morphologically, the larvae are very similar to

those described for E. aurosparsella; however,

their different host plants and distribution suggest

at least some degree of biological divergence. Only

one, seemingly minor, difference in chaetotaxy has

been found to distinguish the Arizona specimens

from the larvae of typical aurosparsella. In the Oak

Creek Canyon larvae, setae M2, M3, and L3 of the

labrum possess smooth ventral margins (Figure

155). The ventral surfaces of these broad setae are

dentate in E. aurosparsella (Figure 129).

The larval mine (Figure 162) is typically eriocraniid in form and commences with a minute,

brownish egg pouch. The initial mine is slender,

usually following the outer margin of the leaf for

a short distance, but gradually enlarges to form a

rather elongate blotch, usually 5 mm or more in

width.

Eriocraniella (Eriocraniella) platyptera,

new species

FIGURES 177, 201-202, 214, 234-235, 271-272, 317-319, 342;

MAP 3

ADULT (Figure 177).—Wing Expanse: <$, 7-8

mm; 9 > 8.2 mm.

Head: Sparsely covered with pale yellowish

orange hairs. Antennae of moderate length, approximately 0.5 the length of forewing, 35- to 37-

segmented; scape and flagellum uniformly dark fuscous, densely covered to apex with slender scales.

Maxillary palpi uniformly fuscous except for whitish pubescence apically; apex of terminal (5th) segment with a series of minute lobes arranged in 3

ranks as follow: (a) a single, relatively long apical

lobe bearing 3 short and 1 moderately long subacute

setae, (b) an identical pair of much shorter subapical lobes, each with a single elongate subacute

seta, and (c) an identical pair of very short subapical lobes, each bearing a single, elongate subacute seta. Base of galeae pubescent. Labial palpi

fuscous with venter of apical segment whitish.

Thorax: Dorsum dark fuscous with a pronounced brassy iridescence; tegulae and mesoscutum with long, erect black hairs; collar pale yellowish orange. Venter pale grayish white with a strong

silvery luster. Legs fuscous with tibial spurs and

tarsi of meso- and metathoracic legs heavily suffused with white. Forewings relatively broad, width

about 0.34 the length, immaculate, dark fuscous

with a prominent golden brassy to blue green

iridescence (depending upon incidence of light)

frequently highlighting dorsalmost scales; fringe

uniformly fuscous. Hind wings much paler, grayish

fuscous with a slight purplish luster; scales moderately broad.

Abdomen (Figures 234-235): Thinly covered

with dark fuscous hairs above and below. Fourth

sternite of female with a pair of moderately large,

well-defined fenestrae approximately 0.35 the

length of sternite in diameter; male without fenestrae. Sternal tubercules present on 5th sternite of

both sexes, papiliform, diameter approximately
FIGURE 162.—Leaf mine of Eriocraniella 'dongifurcula, new

species, on Quercus gambelli Nutall. (Scale = 10 mm.)

0.18-0.20 the length of sternite. Caudal apex of

8th sternite sharply defined, acute in female, not

divided; apex of 8th tergite with a shallow cleft.

Male Genitalia (Figures 271-272): Very similar to

E. longifurcula. Uncus deeply bilobed; lobes prominent, curved laterally; widely separated at apex for

a distance equal to their length. Anal tube completely membranous, without basal sclerites. Vinculum with lateral arms of furcula greatly

lengthened and very slender; furcula divided approximately 0.8-0.9 its entire length; median cephalic margin of vinculum between apophyses

smoothly concave, without median process;

apophyses elongate, approximately 0.7 the length

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of undivided vinculum (excluding furcula). Juxta

broad with lateral margins nearly parallel, greatest

width about 0.7 its length; caudal apex subtruncate, slightly concave. Base of aedeagus greatly

swollen, bulbous; ventral phallic branch stout,

equaling dorsal branch in diameter but slightly

shorter; apex of ventral branch with a minute,

flattened serrated lobe bearing 10-12 marginal

teeth; dorsal branch with a short, broadly conical

subapical process; ventral branch loosely articulated to bulbous base of aedeagus.

Female Genitalia (Figures 317-319, 342): Apex of

ovipositor acute; lateral edges serrulate with 8-10

minute teeth. Bursa copulatrix elongate, surpassing

cephalic apices of anterior apophyses when fully

extended; walls of corpus bursae membranous.

Vaginal sclerite darkly scleroti/ed, relatively narrow

and elongate with greatest width approximately 0.4

its length; a slender, median ventral keel from

cephalic end of sclerite; keel moderately well developed with broad apex directed ventrally as

shown in Figure 318. Ninth segment with midventral pocket well developed, thickened, and symmetrical with numerous minute teeth bordering

inner lateral margins of pocket.

HOLOTYPE.—cf, Six Mile Creek, Ithaca, New

York, 14 May 1961, J. G. Franclemont, genitalia

slide USNM 18152, SEM slide USNM 18120;

USNM holotype 72103; in the National Museum of

Natural History, Smithsonian Institution.

PARATYPES.—NEW YORK: Tompkins Co: Ithaca, Six Mile

Cr: l£ , 11 May 1959, R. W. Hodges, genitalia slide USNM

18236 (USNM); 1 $ , 13 May 1961, R. W. Hodges, genitalia

slide USXM 18235 (USNM). Described from a total of 2

males and 1 female.

HOST.—Unknown.

FLIGHT PERIOD.—May; univoltine.

DISTRIBUTION (Map 3).—Known only from the

type-locality that is situated within the Cayuga

Lake Basin of northwestern New York.

DISCUSSION.—This species may be easily distinguished from all other known members of the

family, save one, on the basis of the genital structure, particularly by the attenuated development of

the furcula in the male. The male genitalia of

Eriocraniella platyplera and E. longifurctila are

essentially identical; however, the two may be

readily separated by other means. For example,

their widely disjunct distribution suggests several
NUMBER 251 87

markedly different ecological adaptations involving such factors as host-plant preferences and

climatic tolerances. Structurally, E. platyptera is

characterized by its pale orange head and relatively

short, broad wings. The latter feature has suggested

its specific epithet, which is derived from the Greek,

platys (broad) and pteron (wing). In contrast, E.

longifurcula possesses a grayish head vestiture and

relatively elongate, narrow wings. The female

genitalia structure of these two species cannot be

compared because no females of E. longifurcula

have been examined.

It is interesting to note that these two species

demonstrate a peculiar relationship similar to that

of another sibling pair within the same genus.

Eriocraniella aurosparsella and E. xanthocara likewise possess essentially identical genitalia, with one

(xanthocara) being easily recognized by its pale

orange head vestiture as compared to the grayish

head of the other (aurosparsella). The host preferences and larval morphology of the latter two have

been found to differ significantly, and it is expected

that similar distinctions will also be discovered for

E. platyptera and E. longifurcula.

Disfurcula, new subgenus

TYPE-SPECIES.—Eriocraniella (Disfurcula) variegata, new species.

ADULT.—Wing Expanse: 6-10 mm.

Head, Thorax, and Abdomen: As described for

genus.

Male Genitalia: Vinculum without median furcula; caudal margin sinuate, with a relatively

prominent, median conical lobe. Aedeagus with

apex of ventral phallic branch relatively simple,

acute, without a flabellate, spinose process, but

with a rather prominent, subapical conical to spinose lobe.

Female Genitalia: Ninth segment without a midventral spinose pocket.

DISCUSSION.—The subgeneric name, Disfurcula,

has been derived from the Latin, dis (without) and

furcula (little pitchfork), and refers to the absence

of the furcula in the male; it has been treated as

feminine in gender.

Key to the Species of the Subgenus Disfurcula

1. Forewings with subcostal vein usually divided. Male genitalia with lobes of uncus relatively

straight, and gradually tapering in width to suhacute apices (Figure 273)

Eriocraniella (Disfurcula) falcata, new species

Forewings with subcostal vein simple. Uncus with lobes distinctly curved laterally, terminating in broadly rounded apices (Figure 275) 2

2. Aedeagus with ventral phallic branch relatively stout, nearly equaling dorsal branch in

diameter; apex of ventral branch with a very prominent subapical. triangular lobe (Figure

275) Eriocraniella (Disfurcula) trigona, new species

Aedeagus with ventral phallic branch relatively slender, less than 0.5 the diameter of dorsal

branch; subapical lobe of ventral branch more reduced and rounded (Figure 278)

Eriocraniella (Disfurcula) variegata, new species

Eriocraniella (Disfurcula) variegata, new species

FIGURES 171, 203-204, 236-237, 247, 277-278, 320-322, 343;

MAP 5

ADULT (Figure 171).—Wing Expanse: <$, 8.2-

9.5 mm; 9, 7.8-8.7 mm.

Head: Uniformly covered with pale yellow to

stramineous hairs. Antennae moderately long, approximately 0.52-0.57 the length of forewing, 38- to

42-segmented; scape mostly whitish; flagellum entirely fuscous except for apical 5-6 segments, which

are sparsely covered with narrow whitish scales.

Maxillary palpi brownish fuscous; apex of terminal

segment complex with a single, relatively large apical lobe bearing 4 moderately long, striated setae,

and 2 pair of very reduced subapical lobes, each

bearing a single, elongate apical seta with lateral

striations. Labial palpi uniformly brownish fuscous.

Thorax: Dorsum brownish fuscous with a bronzy

luster; tegulae with long stramineous hairs. Venter

silvery gray. Pro- and mesothoracic legs with tibial

and tarsal segments brownish fuscous above, white

beneath: metathoracic legs paler, grayish above and

white underneath. Forewings irregularly and almost

equally marked with iridescent gold and purplish

fuscous; markings typically in the form of trans-
88

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

verse bands across the wing but which are sometimes fragmented into large irregular spots; fringe

pale brownish fuscous with a light golden luster.

Hind wings usually slightly darker and less lustrous

than forewings though usually with a slight purplish iridescence near apex; scales relatively broad.

Abdomen (Figures 236-237): Sparsely covered

with grayish hairlike scales above, whitish beneath.

Fourth sternite of female with a pair of relatively

small, circular fenestrae approximately 0.25 the

length of sternite in diameter; male without fenestrae. Sternal tubercules present on 5th sternite of

both sexes, papiliform, diameter 0.14-0.18 the

length of sternite. Caudal apex of 8th sternite moderately defined in female, minutely bifurcate; apex

of 8th tergite more sharply defined, deeply divided.

Male Genitalia (Figures 277-278): Uncus deeply

bilobed; lobes large, broadly rounded, widely separated at their apices for a distance approximately

equal to their length. Anal tube entirely membranous. Vinculum with caudal margin produced

into a moderately large, medium conical lobe; margin sharply excavated on either side of median

lobe; apophyses relatively short, 0.51-0.55 the

length of undivided vinculum; cephalic margin of

vinculum between apophyses smoothly concave.

Juxta broadest anteriorly, tapering slightly to

broad, shallowly concave caudal end; greatest width

of juxta about 0.8 its length. Base of aedeagus moderately swollen; ventral phallic branch loosely articulated to base, relatively slender, approximately

0.5 the diameter of dorsal branch; caudal end of

ventral branch with a small, rounded membranous

process projecting usually somewhat laterad; apex

of dorsal branch variously lobed, typically with a

low, broad dorsal lobe and a similar, more apical

lobe laterally.

Female Genitalia (Figures 320-322, 343): Apex of

ovipositor acute; lateral edges serrulate with 7—9

minute teeth. Bursa copulatrix of moderate length,

slightly surpassing cephalic ends of posterior

apophyses when fully extended; walls of corpus

bursae entirely membranous. Vaginal sclerite darkly

sclerotized, relatively elongate and narrow; greatest

width approximately 0.37-0.4 the length of sclerite;

cephalic end with a short, low median keel ventrally

as shown in Figure 321.

HOLOTYPE.— cf, Coldbrook Station, San Gabriel

Canyon, Los Angeles, California, 26 May 1973, D.

C. Frack; in the Los Angeles County Museum.

PARATYPES.—UNITED STATES. CALIFORNIA: LOS Angeles

Co: 3000 ft [915 m] on Angeles Crest Hwy: 19,1 1 May 1974,

R. Leuschner, 9 genitalia slide USNM 18212, wing slide

USNM 18173 (USNM). 3300 ft [1006 in] on Angeles Crest

Hwy, Big Tujunga Canyon: 1$, 11 May 1974, R. Leuschner

(RHL); 1$, 11 May 1974, R. Leuschner, $ genitalia slide

USNM 18214 (USNM); Big Tujunga Jet: 1$, 11 May 1974,

R. Leuschner (USNM). 4000 ft [1220 m] on Angeles Crest

Hwy, San Gabriel Mts: 1 <$ , 2 9 . 12 May 1974, R. Leuschner,

$ genitalia slide DRI) 2994 (RHL). Cloudburst Canyon,

San Gabriel Mts, T2N, R12W. Sec 14, 4000 ft [1220 m]: 1 $,

10 May 1972, J. P. and K. E. Donahue, $ genitalia slide

USNM 18183 (USNM). Coldbrook Sta, San Gabriel Canyon:

2 9 , 12 May 1972, D. Frack (DCF); 19 . 12 May 1972, D.

Frack, 9 genitalia slide USNM 18211 (USNM); 5$, 19 , 26

May 1973, D. Frack, $ head slide DRD 2995 (DCF). Josephine Cr, San Gabriel Mts, T2N, R12W, Sec 6, 3350 ft [1021

m]: 19, 9 May 1972, J. P. and K. E. Donahue (LACM). Described from a total of 11 males and 8 females.

HOST.—Unknown.

FLIGHT PERIOD.—May; univoltine.

DISTRIBUTION (Map 5).—Presently, this species is

known only from the San Gabriel Mountains of

Los Angeles County, California, where it occurs at

elevations from 3000 feet [915 m] to 4000 feet

[1220 m].

DISCUSSION.—One of the easiest means of recognizing this species is by its irregular, variegated

forewing pattern of iridescent pale gold and purplish fuscous, a feature from which the specific

name is derived. Unfortunately, this delicate pattern is not always pronounced or evident, particularly in rubbed specimens.

Structurally, E. variegata is most similar to E.

trigona, new species, but the two may be readily

distinguished by the different form of their aedeagi.

The ventral phallic branch of E. variegata is more

slender and possesses a much smaller subapical lobe.

Their vaginal sclerites may also be distinctive, but

the female of E. trigona has yet to be collected and

studied.

Eriocraniella (Disfurcula) trigona, new species

FIGURES 172, 205-206, 238, 275-276; MAP 5

ADULT (Figure 172).—WingExpanse: cT,6-8 mm.

Head: Sparsely covered with whitish hairs. Antennae relatively short, approximately 0.48-0.52 the

length of forewing, 35- to 38-segmented; scape silvery

white above, mostly fuscous below; flagellum uniformly brownish fuscous except for terminal 6-7
NUMBER 251 89

Eriocraniella falcata, new species

Eriocraniella trigona, new species

• Eriocraniella variegata, new species

• Neocrania bifasciata, new species

CALIFORNIA INSECT SURVEY

Department of Entomology and Parosltology

UNIVERSITY OF CALIFORNIA

0 25 50 75 100

MAP 5.—Distribution of Eriocraniella (subgenus Disfurcula) and Neocrania.
90

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

segments, which are sparsely covered with slender

white scales. Maxillary palpi light brownish fuscous; apex of terminal segment with 3 distinct series

of lobes: (a) 1 relatively large terminal lobe bearing

3 short and 1 long setae, (b) a pair of shorter,

slightly more subapical lobes, each bearing 1 long

seta, (c) a pair of very short lobes, each bearing 1

long seta. Base of galeae naked except for grayish

white pubescence. Labial palpi brownish fuscous,

irrorated with white.

Thorax: Dorsum light brownish fuscous to stramineous, with a slight bronzy luster; tegulae with

long, pale brownish hairs. Venter silvery white. Proand mesothoracic legs fuscous above, light gray ventrally; metathoracic legs mostly whitish to gray.

Forewings and fringe uniformly fuscous with a

bronzy to purplish iridescence; Sc simple. Hind

wings paler in color, more grayish and less iridescent; scales moderately broad.

Abdomen (Figure 238): Sparsely covered with

long brownish fuscous, hairlike scales above, paler,

more grayish ventrally. Fourth sternite of male

without fenestrae. Sternal tubercules present on 5th

sternite, papiliform; diameter approximately 0.2

the length of sternite.

Male Genitalia (Figures 275-276): Uncus deeply

bilobed; lobes elongate, strongly curved outward;

apices widely separated for a distance approximately

equal to their length. Anal tube mostly membranous, with a few scattered, minute spinules, without

basal sclerites. Vinculum with median, caudal margin produced into a moderately large, conical lobe;

margin distinctly excavated on either side of median lobe; apophyses relatively short, approximately

0.6 the length of undivided vinculum; cephalic

margin of vinculum between apophyses smoothly

concave, not extended. Juxta broadest anteriorly,

narrowing slightly to broad, shallowly concave caudal end; greatest width of juxta about 0.9 its length.

Base of aedeagus moderately swollen; secondary

phallic branch loosely articulated to base, stout,

nearly equaling primary branch in diameter; caudal

end of secondary branch with a large, triangular

subapical projection; primary branch with a similar,

though broader and less pronounced, subapical

process.

HOIOTYPE.— cj\ Josephine Creek, San Gabriel

Mts, T2N, R12W, Sec 6, elev. 3350 ft [1021 m],

Los Angeles County, California, 18 May 1973, Black

lite, D. C. Frack, tf genitalia slide DRD 2962, photograph on file USNM; in the Los Angeles County

Museum.

PARATYPK—UNITED STATES. CALIFORNIA: LOS Angeles

Co: Josephine Cr, 3300 ft [1006 m] on Angeles Crest Hwy:

1$, 11 May 1974, R. H. Leuschner, $ genitalia slide DRD

2993, wing slide DRD 3001 (RHL). Described from 2 males.

HOST.—Unknown.

FLIGHT PERIOD.—Mid-May; univoltine.

DISTRIBUTION (Map 5).—This species is presently

known from only two adjacent localities in the San

Gabriel Mountains of Los Angeles County, California, at elevations of slightly over 1000 meters.

DISCUSSION.—Only two specimens of this species,

both males, have been examined. The para type was

unspread and in poor condition (and has now been

largely dissected); the hoiotype is somewhat rubbed

but with most scale characteristics still discernible.

The species is easily characterized by the male genitalia, particularly the aedeagus. Both branches of

the aedeagus possess prominent subapical processes,

with that of the ventral branch being the most pronounced. The triangular shape of the latter has

suggested the specific name that is derived from the

Greek trigonos (triangular).

Eriocraniella trigona appears most related to E.

variegata, new species. The two may be distinguished by the different form of the ventral branch

of their aedeagi, with that of E. trigona being

stouter and possessing a much larger subapical

process.

Eriocraniella (Disfurcula) falcata, new species

FICLRES 64-65, 163-164, 173, 207-208, 239-240, 248, 273-274,

323-325,341; MAP 5

Dyseriocrania species Opler, 1974:4

ADULT (Figure 173).—Wing Expanse: $, 7-10

mm; 9, 7-10 mm.

Head: Relatively sparsely covered with white to

cream white hairs. Antennae moderately long, 0.55-

0.6 the length of forewing, 40-45 segmented; scape

mostly cream white, slightly irrorated with fuscous

ventrally; flagellum almost entirely fuscous except

for dorsal suffusion of white over basal 5-6 segments and for slender white scales over apical 5-7

segments. Maxillary palpi brownish fuscous, frequently suffused heavily with silvery white ven-
NUMBER 251

91

trally; apex of terminal segment with a single, relatively large apical lobe bearing one large, bluntly

rounded seta and 3 smaller, more acute setae of

variable lengths, and 2 pair of smaller subapical

lobes, each bearing a single, elongate acute seta

apically; all setae with striated sides. Base of galeae

mostly naked, sometimes with scattered, narrow

whitish scales. Labial palpi brownish fuscous above,

silvery white ventrally.

Thorax: Dorsum uniformly light bronzy brown;

tegulae with an elongate tuft of cream white hairs.

Venter mostly silvery white. Legs cream white to

stramineous except for dorsum of tibial and tarsal

segments of forelegs. Forewings uniformly bronzy

brown, often with a pale golden luster; fringe

brownish at base, becoming paler and whiter toward

outer edge; Sc usually divided, crossvein r present.

Hind wings slightly darker than forewings, less iridescent but with a slight purplish luster, becoming

darker toward outer margins; scales relatively broad,

particularly over outer half; crossveins m and m-cu

present.

Abdomen (Figures 239-240): Sparsely covered

with slender bronzy brown, hairlike scales above,

whitish ventrally. Fourth sternite of female with a

pair of relatively small, nearly circular fenestrae

approximately 0.3 the length of sternite in diameter; male without fenestrae. Sternal tubercules

present on 5th sternite of both sexes, papiliform,

diameter approximately 0.17-0.22 the length of

sternite. Caudal apex of 8th sternite sharply defined in female, normally bifurcate, though occasionally merely concave; apex of 8th tergite deeply

divided.

Male Genitalia (Figures 273-274): Uncus deeply

bilobed; lobes prominent, lateral margins straight

but tapering; lobes widely separated at apex for a

distance equivalent to their length. Anal tube membranous, without basal sclerites. Vinculum with median caudal margin not raised, relatively truncate; median cephalic margin between apophyses

smoothly concave, not extended, apophyses elongate, approximately 0.73-0.77 the length of undivided vinculum. Juxta broadest immediately anterior to middle; lateral margins nearly parallel;

greatest width approximately 0.45-0.6 its length;

caudal apex subtruncate to slightly rounded (convex). Base of aedeagus moderately swollen; ventral

phallic branch relatively stout, approximately equal

in diameter and length to dorsal branch; apex of

ventral branch acute, with a small spinose process

loosely articulated subapically; length of subapical

process varying from 0.5-1.2 the diameter of ventral phallic branch; remainder of ventral branch

without lobes or processes.

Female Genitalia (Figures 323-325, 341): Apex

of ovipositor acuminate; lateral edges serrulate,

with 5-7 minute teeth. Bursa copulatrix of moderate length, slightly surpassing cephalic apices ot

posterior apophyses when extended; walls of corpus

bursae entirely membranous. Bases of anterior

apophyses symmetrical, not unusually swollen. Vaginal sclerite darkly sclerotized, relatively narrow and

elongate with greatest width only 0.24-0.27 its

length; a prominent, falcate ventral keel present

along midline, with apex projecting caudally as

shown in Figure 325.

HOLOTYPE.— cf, Cloudburst Canyon, San Gabriel

Mts, T2N, R12W, Sec 14, 4000 ft [1220 m], Los

Angeles County, California, 10 May, 1972, J. P. and

K. E. Donahue; in the Los Angeles County

Museum.

PARATVPES.—UNITED STATES. CALIFORNIA: LOS Angeles

Co: Big Tujunga Canyon, 3300 ft [1006 m] on Angeles Crest

Hwy: 1£ , 19 , 11 May 1973, R. H. Leuschner (RHL). Cloudburst Canyon, San Gabriel Mts, T2N, R12W, Sec 14, 4000 ft

[1220 m]: \$, 2$ , 30 Apr-10 May 1972, J. P. and K. E.

Donahue (LAC:\i). Josephine Cr, San Gabriel Mts, T2N,

R12W, Sec 6, 3350 ft [1021 m]: 7$, 14$ , 9 May 1973, J. P.

and K. E. Donahue, $ genitalia slide DRD 2998, $ genitalia

slide DRD 2999 (LACM); 1 $, 19, 9 May 1973, J. P. and

K. E. Donahue, $ genitalia slide USNM 18184, 9 genitalia

slide USNM 18189 (USXM); \$, U May 1974 D. C. Frack

(DCF); 9$, 19 . 18 May 1973, D. C. Frack (DCF); 1 $, 18

May 1973, D. C. Frack {USNM). Josephine Cr, 3300 ft [1006

ml on Angeles Crest Hwy: 12 $, 29 , H May 1973, R. H.

Leuschner (RHL); 3500 ft [1067 m]; 1$, 11 May 1973, R. H.

Leuschner, $ genitalia slide DRD 3003 (RHL). Mt. Wilson

Rd, 4600 ft [1402 m] on Angeles Crest Hwy: 1$, 11 May

1973, R. H. Leuschner, $ genitalia slide USXM 18159

(USNM). Littlerock: 1 $ , 28 Mar 1964, P. Opler, $ genitalia

slide DRD 2901 (UCB). 3000 ft [915 m] on Angeles Crest

Hwy: 5$, 29 , 11 May 1973, R. H. Leuschner (RHL); 1$,

2 9,1 1 May 1973, R. H. Leuschner, $ genitalia slide USXM

18186, 9 genitalia slide USXM 18188 (USXM). San Luis

Obispo Co: San Luis Obispo: 1 9 , March, A. Vachell, 9

genitalia slide USNM 18205 (USXM). Santa Clara Co: Herbert Cr, 3 mi [4.8 km] W Xew Almaden: 2 9 . 28 Apr 1969,

P. Opler. Qitercus chrysolepis, 9 genitalia slides PAO 87,

DRD 2957 (UCB); 1 9 , 28 Apr 1969, P. Opler, Qiiercus

chrysolepis, 9 slides USXM 16102, 18194 (USXM); 19 , 18

Apr 1969, y. Powell, flying under Q. agtifolia 12:00 noon,

9 genitalia slide DRD 2975 (UCB). Described from a total

of 43 males and 30 females.
92

Hosr.—Fagaceae: "Quercus chrysolepis Liebm.,"

Opler (1974) and from specimen labels.

FLIGHT PERIOD.—Late March to mid-May; univoltine.

DISTRIBUTION (Map 5).—This species occurs

through the Coast Ranges of Central California

from Santa Clara County south to the San Gabriel

Mountains of Los Angeles County, where it is

found at elevations up to 1400 m.

DISCUSSION.—This species presently stands relatively alone among our North American Eriocraniiclae; consequently, it is not likely to be confused with any known species. Although amply

distinct on a specific level, the generic relationships

of Eriocraniclla falcata are not as well denned. As

expressed earlier in this paper, the overall morphology of this species, particularly as witnessed in the

general structure of the maxillary palpi, and male

and female genitalia, suggests strong affinities with

the subgenus Disfurcula. The rather complex and

specialized structure of the apex of the maxillary

palpi is identical to the other members of Eriocraniella. The wing venation is somewhat atypical for

Eriocraniella and instead resembles that of Neocrania bifasciata, particularly in the forking of the

subcostal vein and the presence of certain crossveins

in the forewings (r) and hind wings (m, m-cu). The

relatively straight and gradually tapering lobes of

the uncus in the male genitalia are also atypical

for Eriocraniella and, instead, more resemble those

of Eriocrania. There is, however, no median extension of the vinculum between the apophyses that

appears to be characteristic of the latter genus. The

absence of the median furcate process, or furcula,

is another feature that associates E. falcata with the

subgenus Disfurcula. The aedeagus of this species

is unique in possessing a rather loosely appended,

variably shaped spinose process near the apex of

the ventral phallic branch. The female genitalia,

particularly the apex of the ovipositor, closely

resemble that of the other members of Eriocraniella. The females, however, are easily distinguished from all other eriocraniine species by the

prominent, sickle-shaped keel of the vaginal sclerite,

a unique feature that has suggested the specific

name, falcata, which is derived from the Latin,

falcatus (sickle-shaped or curved).

Eriocraniella falcata is the only eriocraniid

known to mine leaves of Quercus chrysolepis. Mine

damage on six leaves of Qiicrcus chrysolepis beSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

T

1

FIGURE 163.—Leaf mine of Eriocraniella >falcata,

new species, on Quercus chrysolepis Liebm.

(Scale = 10 mm.)

lieved to be caused by E. falcata have been examined. The early instar linear mine appears to be

unusually long and in one leaf (Figure 163) extremely long. A typical, elliptical hole is produced

in the leaf at the site of oviposition with the

characteristic brownish egg pouch (Figure 164) attached along one side. The linear mine continues

to the leaf margin and then abruptly enlarges to

form a full depth blotch usually covering the apical

third to one-fifth of the leaf.

Neocrania, new genus

TYPE-SPECIES.—Neocrania bifasciata, new species.

ADULT.—Small moths with relatively broad wings;

wing expanse: 7-10 mm.

Head: Maxillary palpi with basal 0.8-0.9 of

terminal segment densely setose; primary setae
NUMBER 251

93

FIGURE 164.—Leaf mine of Eriocraniella 7falcata, new species,

on Quercus chrysolepis Liebmann showing oviposition scar

and egg pouch (see arrow).

barbed laterally; apical 0.2-0.1 of segment largelynaked, divided to form 2 relatively large, subacute

lobes, one slightly shorter than the other; each lobe

bears 4 stout, moderately long setae with striated

sides; 1 pair of setae typically arise from extreme

apex of each lobe and the 2nd pair arise more subapically; setae approximately equal in size, with a

length not greater than the diameter of the lobe

from which they arise.

Thorax: Epiphysis present. Forewings with Sc

often superficially divided near costa; Rl divided

approximately 0.4 its length; R2, 3, 4, and 5 all

arising separate from cell; crossvein r present. Hind

wings with Sc and Rl simple; R2 and 3 entirely

fused; R4 and 5 either connate or separate; crossveins m and m-cu present; scales relatively short

and broad, not hairlike.

Abdomen: Third sternite with a pair of slender,

triangular fenestrae near anterior margin. Fourth

sternite without fenestrae. Sternal tubercules present on 5th sternite of both sexes, similar in form,

relatively small, papiliform.

Male Genitalia: Uncus prominently bilobed;

lobes broad, relatively short, and narrowly separated. Socii present. Anal tube completely membranous, without basal sclerites. Vinculum with

caudal margin produced into a prominent median

lobe; cephalic margin evenly concave between

elongate lateral apophyses, without a median process. Caudal apex of juxta symmetrical, subtruncate

to slightly convex. Aedeagus with ventral phallic

branch very irregular in outline and with a cluster

of several subapical cilia.

Female Genitalia: Anterior apophyses asymmetrical at base. Vaginal sclerite with cephalic end

developed to form a relatively broad, elongate

process ventrally.

DISCUSSION.—This monotypic genus exhibits several apomorphies, some of which may eventually be

regarded only as of specific importance. Neocrania

appears to represent a specialized offshoot of the

predominantly western North American genus

Eriocraniella. It differs from the latter in several

peculiarities of the maxillary palpi, abdomen, and

male and female genitalia.

The maxillary palpi are unique among the

Nearctic Eriocraniidae in possessing a bifid apex,

with each lobe bearing four moderately long setae.

Only two other members of this family, the European Eriocrania sangii and E. haworthi are known

to possess a bilobed palpal apex. This condition in

both species is obviously independently derived, as

the chaetotaxy of the terminal segment agrees in all

respects with typical Eriocrania. In Eriocraniella,

the apex appears more complex than that of Neocrania in consisting of one rather large apical lobe

and two pair of progressively smaller more subapical lobes. The derivation of both types of sensory

structures from an immediate, common ancestral

model seems evident, however, in that a total of

eight striated setae are present in the members of

both genera. The somewhat more complex palpal

apex demonstrated by Eriocraniella, for example,

could have been derived from the bifid type possessed by Neocrania through further division of one

of the two large lobes of the latter, resulting in two

pair of smaller lobes, each bearing one large seta.

Neocrania further demonstrates its close ties with

Eriocraniella through strong venational similarities
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with Eriocraniella falcata. The venational pattern

of Ar. bifasciata and E. falcata together differ from

the other known members of Eriocraniella in

typically possessing a forked subcostal vein in the

forewings and a fuller complement of crossveins.

As typified by N. bifasciata, the male and female

genitalia of Neocrania exhibit some unusual specializations not known to occur in other eriocraniid

genera. For example, the ventral phallic branch of

the male bears a series of elongate, filamentous

subapical setae. Similarly, in the female, the anterior apophyses are asymmetrical with the basal

region of the left member being noticeably expanded. The vaginal sclerite differs from the typical

form in Eriocraniella by lacking a midventral keel

and by having the cephalic half of the sclerite deflected sharply ventrad.

The prefix neo- in the generic name Neocrania

is derived from the Greek neo.s (new) and is intended to refer to the general distribution of the

group, which is known only from the New World.

The generic name has been treated feminine in

gender.

Neocrania bifasciata, new species

FIGURES 178, 209-210, 242-243, 249, 279-280, 326-328, 338;

MAP 5

ADULT (Figure 178).—Wing Expanse: tf, 8-9

mm; 9 > 7-10 mm.

Head: Covered with whitish to pale stramineous

hairlike scales. Antennae moderately long, approximately 0.52-0.60 the length of forewing, 42- to 44-

segmented; scape entirely whitish to stramineous;

flagellum usually uniformly whitish to stramineous,

occasionally banded dorsally with brownish fuscous.

Maxillary palpi whitish to light brown; apex of

terminal (5th) segment divided into 2 relatively

large lobes of almost equal size (Figure 210); each

lobe with an apical pair and a subapical pair of

moderately long striated setae; length of setae approximately equal to diameter of supporting lobe.

Base of galeae nearly naked, sparsely covered with

short whitish hairs. Labial palpi whitish to stramineous, lightly irrorated with pale brownish hairs

ventrally.

Thorax: Dorsum stramineous, occasionally

more whitish laterally near the wing bases; tegulae

usually entirely stramineous. Venter silvery white.

Legs mostly whitish to stramineous, sometimes with

tibiae brownish fuscous above; tarsi ringed with

brownish fuscous. Forewings pale brownish fuscous

with a golden iridescence; wings crossed by a pair

of broad, golden fasciae; one situated at base, the

other, more irregular fascia, across middle of wing;

a series of 6-8 small, golden submarginal spots

sometimes evident along termen from R2 to tornus;

fringe pale brownish fuscous. Hind wings slightly

darker than forewings, more fuscous and sometimes

with a slight purplish sheen; scales relatively broad.

Abdomen (Figures 242-243): Sparsely covered

with long stramineous hair above, more whitish

ventrally. Fourth sternite without fenestrae in both

sexes. Sternal tubercules present on 5th sternite of

both sexes, diameter approximately 0.15-0.16 the

length of sternite. Caudal apices of 8th tergite and

sternite in female entire, not deeply clefted.

Male Genitalia (Figures 279-280): Uncus shallowly bilobed; lobes large, broadly rounded, rather

narrowly separated at apex for a distance about

0.75 their length. Anal tube entirely membranous,

without basal sclerites. Vinculum with median area

of caudal margin produced into a broad conical

lobe; apophyses slender, elongate, approximately

equaling the length of undivided vinculum; cephalic margin of vinculum between apophyses

smoothly concave, not extended. Juxta broadest

anteriorly, tapering slightly to broad, truncate

caudal end; greatest width of juxta about 1.4 its

length. Base of aedeagus moderately swollen; secondary phallic branch stout, its diameter nearly

equaling that of primary branch; apex of secondary

branch irregular, somewhat sinuate, with a series of

8-11 long, slender setae.

Female Genitalia (Figures 326-328, 338): Apex

of ovipositor acute; lateral edges serrulate, with

9-12 minute teeth. Caudal bases of anterior

apophyses asymmetrical with left apophysis broadly

expanded at juncture with 8th tergite. Bursa copulatrix of moderate length, only slightly surpassing

cephalic apices of posterior apophyses when fully

extended; walls of corpus bursae entirely membranous, without spines. Vaginal sclerite darkly sclerotized, relatively elongate, narrow; greatest width approximately 0.42-0.45 the length of sclerite; a broad

elongate, tapering keel extending ventrally from

cephalic end of sclerite as in Figure 328.

HOI.OTYPE.— 9, Josephine Creek, San Gabriel

Mts, T2N, R12W, Sec 6, elevation 3350 ft [1021 m],

9 May 1973, J. P. and K. E. Donahue, 9 genitalia
NUMBER 251 95

slide DRD 3002, photograph on file USNM; in the

Los Angeles County Museum.

PARATYPES.—CALIFORNIA: LOS Angeles Co: Josephine Cr,

San Gabriel Mts, T2N, RI2W, Sec 6, 3350 ft [1021 m]: 2$,

5 9 , 6-9 May 1973, J. P. and K. E. Donahue (LACM); 2$,

2 9.6-1 1 May 1973, J. P. and K. E. Donahue, USNM slides

18174, 18181, 18182, 18187, 18207 (USNM); 5$ , 11 May 1974,

(DCF). Josephine Cr, 3300 ft [1006 m] on Angeles Crest Hwy:

\$, 11 May 1974, R. Leuschner, $ genitalia slide DRD

2992 (RHL); 19 , 11 May 1974, R. Leuschner, 9 genitalia

slide USNM 18190 (USNM). 3000 ft [915 m] on Angeles Crest

Hwy: 1 ^ , 2 9 , 11 May 1974, R. Leuschner (RHL). Monterey

Co: 1 air mi [1.6 km] S Jamesburg, Santa Lucia Mts, 2900 ft

[884 m]: \$, 19 , 8 May 1975, on Quercus chrysolepis, J.

Powell and J. Chemsak (UCB); 1 air mi [1.6 km] S Jamesburg, 2900 ft [884 m]: 19 , 5 May 1975 (UCB); \$ (same

data), USNM genitalia slide 18259 (USNM). San Diego Co:

Nate Harrison Grade Rd, near Mt Palomar, 4700-5000 ft

[1435-1524 m]: 2 9, 2 June 1968, assoc. with Black light, C.

Bcesley (UCR); 19 , same data, USNM slide 20203 (USNM).

Described from a total of 8 males and 21 females.

HOST.—Unknown; possibly Qiiercus chrysolepis

Liebm.

FLIGHT PERIOD.—Early June; univoltine.

DISTRIBUTION (Map 5).—This species is known

only from the Coast Ranges of southern California,

where it occurs from the Santa Lucia Mountains of

Monterey County south to the San Gabriel Mountains of Los Angeles County and Mt. Palomar in

San Diego County. All the specimens collected thus

far were found between 850 and 1525 m in elevation.

DISCUSSION.—Neocrania bifasciata is one of the

most colorful and, hence, recognizable members of

this family. The pair of golden fasciae traversing

the forewings easily distinguishes the species from all

other Eriocraniidae. The male genitalia are unique

in several respects, particularly with regard to the

short, broad lobes of the uncus and the peculiar

ciliate setae arising near the apex of the ventral

phallic branch. The female is similarly unusual in

being the only known North American eriocraniid

without abdominal fenestrae on the fourth sternite

and with asymmetric anterior apophyses.

The specific name is derived from the Latin bi

(two) and fascia (band) and refers to the characteristic banding on the forewings of this moth.

Although the immature stages of this species

have yet to be discovered, it is possible that the

larvae may feed on Quercus chrysolepis. Adults

were collected resting on this plant in early May

by Powell and Chemsak.

ACANTHOPTEROCTETIDAE, new family

TYPE GENUS.—Acanthopleroctetes Braun.

ADULT.—Moths of moderately small size, wing

expanse 11-16 mm. Wings very slender.

Head: Vertex not elongate; laterofacial sulci

obsolete. Ocelli absent. Compound eyes relatively

large; interocular index approximately 0.75—0.80;

eyes situated more laterally. Mandibles vestigial,

nearly absent. Apex of terminal (5th) segment of

maxillary palpi relatively simple, without secondary

lobes. Labial palpi 2-segmented, short, approximately equal to 1st segment of maxillary palpi in

length; apex of distal segment relatively simple,

without organs of von Rath and basiconic sensilla.

Thorax: Prothoracic legs relatively long; foretibiae approximately 0.42-0.46 the length of hind

tibiae; epiphysis absent. Forewings very slender,

width approximately 0.19-0.20 the length; Sc simple, not divided; radius with 4 primary branches;

R3 absent; Rl secondarily divided for most its

length; pterostigma absent; R4 stalked with R5

for nearly two-thirds their length; Ml stalked to

R4 + 5; anal veins widely separate to margin; 3A

very short, closely associated with jugal fold. Hind

wings with Sc and Rl simple; radius with 4 primary branches; R3 absent; R4 and 5 and Ml

stalked as in forewings; lower half of distal cell

open due to atropy of medial-cubital crossvein;

scales typically long and hairlike.

Abdomen: Pleural area of 3rd segment either

unmodified or with a bilateral pair of filamentous

appendages (Figure 165). Fourth and 5th sternites

unmodified, without paired fenestrae or glandular

tubercules. Eighth segment of female with a single

row of elongate setae arising from caudal margin

and projecting caudad; setal row encircling segment

with setae more elongate dorsally than ventrally.

Male Genitalia: Uncus superficially bifid, lateral

margins revolute, serrulate, or spinose near apex.

Socii absent. Caudal margin of vinculum relatively

deeply concave or excavate; cephalic margin shallowly concave; lateral angles only slightly evident,

not produced into elongate apophyses. Anellus

with two separate areas of sclerotization surrounding aedeagus: a darkly sclerotized, dorsal transtilla

and a thinner, less sclerotized juxta ventrally; transtilla in the general form of an inverted T with a

prominent, elongate median process extending caudad almost to apex of uncus; apex of median proc-
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ess with an acute, uncinate apex curving dorsally;

ventral subapical margin of medium process serrulate; base of transtilla abruptly expanded laterally toward, but separate from, bases of valvae.

Juxta of various form, typically consisting of a thin

elongate plate, slightly broader at the middle. Valvae slender, greatest width approximately onefourth the length, elongate with apices surpassing

apex of uncus and relatively simple except for a

narrow, membranous fringe along ventral margin.

Aedeagus undivided, relatively stout, with a prominent and complex array of endophallic cornuti.

Female Genitalia: Corpus bursae relatively reduced in size, ductus bursae and vagina with thickened, folded walls. Common oviduct arising either

ventral or lateral to bursa copulatrix and somewhat

posterior to spermathecal papilla. Spermatheca entirely membranous, without an internal thickening

in ductus spermathecae; spermathecal papilla either

strongly sclerotized or largely membranous. Vaginal

sclerite absent. Colleterial gland arising from dorsal

wall of vagina caudad of vestibulum. Posterior apophyses with tertiary, closely associated rodlike

sclerites in 10th segment. Ninth segment entirely

membranous except for spinose ridges.

Acanthopteroctetes Braun

Acmithopteroctetes Braun, 1921:22.—Tillyard, 1923:195.—

Philpott, 1926:723.—Fletcher, 1929:2.—McDunnough, 1939:

110.—Viette, 1949:32.—Davis, 1969:137, 138.—Mutuura,

1971:1129—Common, 1973:11, 13, 16—Dugdale, 1974:127,

132, 134, 142— Birket-Smith and Kristensen, 1974:158, 159.

—Razowski, 1975:36.

TYPE-SPECIES.—Acanthopteroctetes

Braun; by monotypy.

tripunctata

Generic characters as for family Acanthopteroctetidae.

DISCUSSION.—Although this endemic North American genus presently contains only three named

species, at least two other species may exist. The

latter have not been named because both are represented by unique females in poor condition. One

of these, mentioned previously by Braun (1921)

and Davis (1969), consists of a single female collected at Monache Meadows, Tulare County, California from an elevation of 8000 ft [2439 m] on

10 July 1917. The female genitalia closely resemble

that of A. bimaculata except for the form of the

spermathecal papilla. The papilla of the Monache

Meadows specimen is less sclerotized and more elongate and cylindrical in shape. The specimen possesses a wing expanse of 13 mm and appears to be

a relatively pale, uniformly colored moth.

A second unnamed species was collected 8 July

1966 by J. H. Baker from Spring Creek, Baker,

Oregon. This locality is also the original source of

the type series of A. bimaculata. Although the

unique female specimen was badly broken in shipment to the Smithsonian Institution (where it is

now deposited), most of its diagnostic features can

easily be observed. The forewings of this species

are distinct in being uniformly pale golden brown,

and the head vestiture is stramineous in color. The

female genitalia differs greatly from that of other

known members of the genus in possessing a more

broadly depressed and irregularly lobed vistibulum

(Figure 332). The spermathecal papilla is indistinct

and unsclerotized. The common oviduct apparently

enters the vistibulum at a more typical (for this

family) midventral position.

Key to the Species of Acanthopteroctetes

1. Forewings pale grayish with two, rather large, fuscous spots along hind margin (Figure

'80) Acanthopteroctetes bimaculata

Forewings darker, with whitish to pale yellowish markings 2

2. Forewings with a single, narrow whitish fascia at distal third (Figure 181)

Acanthopteroctetes unifascia, new species

Forewings with three large, pale yellowish spots; two nearly confluent spots at distal third

and another at basal third of wing (Figure 179) Acanthopteroctetes tripunctata
NUMBER 251 97

Acanthopteroctetes tripunctata Braun

FIGURES 179, 281-284; MAP 6

Acanthopteroctetes tripunctata Braun 1921:22-—Fletcher,

1929:2.—McDunnough, 1939:110, no 9857.—Viette, 1949:31.

—Davis, 1969:140.

Acanthopteroctetes tripunctella [sic].—Davis, 1969:142 [misspelling of tripunctata Braun].

ADULT (Figure 179).—Wing Expanse: <$, 11 mm.

Head: Vestiture pale reddish brown, unicolorous. Antennae pale fuscous; scape more whitish.

Palpi pale brown.

Thorax: Scales of dorsum fuscous; bristlelike

tufts on tegula pale fuscous; venter pale brown

with slight suffusion of white. Legs fuscous. Forewings fuscous with a slight coppery luster; three

large, pale yellowish spots, one at basal third of

wing, two at apical third; outer pair nearly anastomosing to form an irregular and incomplete band

across wing. Hind wings slightly paler, scales narrower, approximately one-half the width of those

in forewings.

Abdomen: Fuscous.

Male Genitalia (Figures 281-284): Uncus not

deeply bilobed, ventral margin (as viewed laterally)

with approximately 4 pair of minute serrations.

Tegumen relatively short, about same length as

uncus. Caudal margin of vinculum with a deep

V-shaped, median cleft. Median process of transtilla

with 3-4 pair of ventral serrations. Juxta elongate,

exceeding length of entire transtilla, accuminate

at caudal end. Valvae with ventral membranous

fringe reduced; sacculus more expanded than in

A. bimaculata. Aedeagus with 2 stout cornuti at

apex and a series of 3 aligned down 1 side.

TYPE.—Holotype cf; in the collection of the

Academy of Natural Sciences, Philadelphia.

TYPE-LOCALITY.—Glacier National Park, Montana, Canyon Creek, 5500 ft [1677 m].

HOST.—Unknown.

FLIGHT PERIOD.—July; probably univoltine.

DISTRIBUTION (Map 6).—Presently reported only

from Glacier National Park, in the Rocky Mountains.

DISCUSSION.—This species is represented only by

the unique holotype that was collected 23 July

1920, while flying in sunshine through an open area

in the forest, by A. F. Braun.

Acanthopteroctetes bimaculata Davis

FIGURES 17-22, 33-34, 41-44, 47, 49, 53-55, 57, 69-70, 180, 217,

250, 254, 285-288, 329-330, 344; MAP 6

Acanthopteroctetes bimaculata Davis, 1969:140.

ADULT (Figure 180).—Wing Expanse: cf, 14.5-16

mm; 9» 15 mm.

Head: Vestiture consisting of a scattered mixture of white and fuscous hairlike scales. Antennae

with 47-50 segments, banded along proximal half

of flagellum, with scales of each segment brownish

fuscous at base, white at apex; distal half of flagellum mostly fuscous; scape mostly whitish with

pale suffusion of brown. Palpi sparsely covered with

pale brown and whitish scales.

Thorax: Dorsum clothed with a scattered mixture of white and pale brownish scales; bristlelike

scale tufts on tegulae and rear portion of mesonotum

fuscous; venter mostly whitish. Legs fuscous. Forewings mostly white, with a somewhat irregular suffusion of pale brown to fuscous; two irregularly

shaped but very distinct spots of fuscous located

along hind margin near proximal third and distal

third of wing; fringe whitish along outer margin,

becoming pale brown along hind margin. Hind

wings thinly scaled; scales very slender, hairlike,

.op /

' V (

1 r .••..

ft/'

-:.-• _ _ .

•

•

* -

J- _

A. bimaculata

A. tripunctata

A. unifascia

A. species. Baker, Oregon

California

SCALE

MAP 6.—Distribution of Acanthopteroctetes.
98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

less than one-fourth the width of those in primaries,

pale brownish; fringe unicolorous, brownish.

Abdomen: Pale fuscous.

Male Genitalia (Figures 285-288): Uncus rather

deeply bilobed; lower margins with approximately

5 pair of minute serrations. Tegumen relatively

long, exceeding length of uncus. Caudal margin of

vinculum less deeply excavated than in A. tripunctata. Median process of transtilla with 3-4 pair of

ventral serrations. Juxta elongate, nearly equaling

the length of valvae; basal 0.2 constricted with caudal 0.8 relatively broad and spatulate, and terminating in a broadly rounded apex; a pair of oval,

hyaline spots slightly confluent at midline, present

near basal one-third. Valvae with ventral membranous fringe prominent; sacculus less expanded than

in A. tripunctata. Armature of aedeagus complex,

with a single, large apical spine and two rows of

approximately 10-12 stout cornuti at middle.

Female Genitalia (Figures 69-70, 329-330, 344):

Apex of ovipositor with approximately 10 minute

serrations along ventral margin. Caudal margin of

8th segment with a series of about 9 pair of stout,

elongate, sensory setae encircling abdomen; the

longest setae originating dorsally with series gradually decreasing in length toward venter and arising

from rather large well-defined sockets. Vestibulum

with thickened, heavily folded walls; common oviduct apparently arising from a large lateral lobe

of vestibulum. Spermathecal papilla darkly sclerotized, funnel shaped. Corpus bursae relatively reduced in size, walls completely membranous; ductus bursae membranous, increasing in diameter

toward vestibulum.

TYPE.—Holotype 9, USNM 69929, in the National Museum of Natural History, Smithsonian

Institution.

TYPE-LOCALITY.—Spring Creek, Baker, Baker

County, Oregon.

HOST.—Unknown.

FLIGHT PERIOD.—May to mid-July; univoltine.

DISTRIBUTION (Map 6).—Known only from the

Blue Mountains-Wallowa Mountains section of the

Columbia Plateau in northeastern Oregon and the

southern Sierra Nevada of east-central California.

MATFRIAL EXAMINED.—2 males and 2 females. CALIFORNIA:

Tulare Co: Monache (mispelled on label as Monacbee) Meadows, 8000 ft [2439 m]: 1 9 (paratype), 8-14 July (USNM).

OREC;O\: Baker Co: Baker, Spring Cr: 1 $ (paratype), 7 May;

19 (holotype), 17 May (USNM); Spring Cr, Blue Mts, 4000

ft [1220 m): 13, 13 May (USNM).

DISCUSSION.—Acanthoptcvodetes bimaculata is

easily distinguished from the other members of this

genus by its pale grayish wings and dark fuscous

spots. The large plate, or juxta, ventral to the

aedeagus was originally described and illustrated in

a folded condition within the male genitalia. Examination of an additional male specimen has enabled me to redescribe this structure in its more

normal posture.

The common oviduct is described in this species

as arising laterally from the vestibulum or the enlarged, thickened region of the vagina. A lateral

origin for this tract has never been reported in any

other species of Lepidoptera. A possible explanation for this apparent anomaly in A. bimaculata is

that actually the common oviduct emerges ventrally

(as is typical in the other members of this family)

from a lateral lobe of the vestibulum. The latter

region can be very irregularly lobed in this genus

and a slight transfer of the common oviduct to one

of these lobes seems possible. Further remarks concerning this will have to await the examination of

additional female specimens in a better state of

preservation.

Acanthopteroctetes unifascia, new species

FIGURES 165, 181, 289-292; MAP 6

ADULT (Figure 181).—Wing Expanse: S > 8.5 mm.

Head: Occipital area brownish fuscous; vertex

whitish becoming darker on lower part of frons.

Antennae uniformly brownish fuscous, including

scape; length uncertain (antennae broken) but appearing at least 0.5 the length of forewing. Maxillary and labial palpi brownish fuscous.

Thorax: Dorsum and tegulae grayish fuscous

with a distinct bronzy luster. Venter paler, silvery

gray. Legs mostly light fuscous to gray. Forewings

brownish fuscous with a bronzy iridescence; a very

prominent, white transverse band across wing at

outer third near apex of discal cell; a less distinct

but relatively long spot present near hind margin

at basal fourth; fringe light fuscous to gray. Hind

wings nearly as dark as forewings but less iridescent, uniformly grayish fuscous.

Abdomen (Figure 165): Uniformly brownish

fuscous with a slight bronzy luster; slightly paler,
NUMBER 251 99

more grayish beneath. Third segment of male with

a cluster of small, filamentous sensilla (Figure 165)

arising from pleural area near each spiracle.

Male Genitalia (Figures 289-292): Uncus bilobed with apex appearing truncate when viewed

laterally; ventral margins with a subapical row of

4 pair of short stout spines. Tegumen long, nearly

twice the length of uncus. Caudal margin of vinculum moderately excavated. Median process of

transtilla with 6 pair of ventral serrations. Juxta

relatively elongate, mostly membranous with

sclerotized lateral margins. Valvae slender, greatest

width 0.25 that of length; ventral membranous

fringe reduced, restricted to apical half. Aedeagus

slender, equaling length of entire genitalia; exterior

walls densely pubescent around middle; cornuti of

irregular form, consisting of a paired structure with

apices subdivided and spinose.

HOLOTYPE.— J1, Circle X Ranch, B.S.A. Camp,

1500 ft [457 m], Ventura County, California, 24

April 1966, coll. J. Powell, tf genitalia slide DRD

2828, in the California Academy of Sciences on indefinite loan from the University of California,

Berkeley.

HOST.—Unknown

FLIGHT PERIOD.—Late April; probably univoltine.

DISTRIBUTION (Map 6).—This species is known

only from the type-locality, which is situated in the

western sector of the Santa Monica Mountains, a

coastal range in Ventura County, southern California.

DISCUSSION.—Acanthopteroctetes unifascia may

be readily separated from the other members of this

aberrant genus by its maculation, particularly by

the single band, or fascia, traversing the forewing.

The genitalia are characteristic in having the juxta

and membranous fringe of the valvae reduced and

the aedeagus with a reduced number of subapical

cornuti. One of the most unique features of this

species is the presence of a lateral triad of pedunculate sensilla (Figure 165) arising from a single,

minute plate near the spiracle of the third abdomFIGURE 165.—Acanthopteroctetes unifascia, new species,

pedunculate lateral glands of third abdominal segment.

inal segment. Each lateral plate appears to bear

three sensilla consisting of a bulbous apical part

supported by a filamentous stalk. Although the

functions of these structures are not known, they

appear glandular in nature and may be secretory,

not sensory, in function.

Powell (in litt.) reports the habitat of the typelocality to be most similar to a dry chaparral. Consequently, this coastal locality differs strikingly

from the wetter and more interior boreal habitats

frequented by the other known members of this

genus.
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102

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NUMBER 251 103

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1959. Klucze do Oznaczania Owad6w Polski, parts 2-4:

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1969. The Genera of Ichneumonidae, Part 3. Memoirs of

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1951. Sur la Nomenclature des Eriocraniidae. Revue Francaise de Lepidopterologie, 13(3-4): 43-44.

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1863. Tortricites and Tineites. Volume 28 in List of the

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1882- Notes on Tineidae of North America. Transactions

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1908. A New Eriocrania (Lepidoptera) from the Pacific

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1877[1876]. Micropterygina. Pages 771-779 in part 2 in

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1973. Fossil Forms of Amentiferae. Brittonia, 25:334-355.

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1890. Micropteryx Larvae. The Entomologist's Monthly

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1891. On Oviposition, and the Ovipositor, in Certain

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Wood, VV.

1845. Index Entomologicus. xii+266 pages. London:

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1850. Vcrzeichniss der von Herrn Jos: Mann beobachteten

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1851. Drei Schabengathungen: Incurvaria, Micropteryx,

und Nemorplwra. Linnaea Entomologica, 5:322-347.
104

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

168

FIGURES 166-171.—Adult moths: 166, Dyseriocrania griseocapilella (Walsingham), $, wing expanse 11.2 mm; 167, D. auricyanea (Walsingham), £, wing expanse 12.3 mm; 168, Eriocrania

seinipurpurella seinipurpurella (Stephens) wing expanse 10.8 mm; 169, E. s. pacifica, new subspecies, $, holotypc, wing expanse 10.6 mm; 170, E. breviapex, new species, 9, holotype, win<;

expanse 9 mm; 171, Eriocraniella variegata, new species, <J > holotype, wing expanse 9.1 mm.
NUMBER 251 105

FIGURES 172-177.—Adult moths: 172, Eriocrauiella trigona, new species, $, holotype, wing expanse G mm; 173, E. falcata, new species, $, holotype, wing expanse 9.5 mm; 174, E. aurosparsella (Walsingham), $, wing expanse 9 mm; 175, E. xanlhocara, new species, $, holotype,

wing expanse 7.5 mm; 176, E. longifurcula, new species, $, holotype, wing expanse 10.2 mm;

177, E. platyptera, new species, $ , holotype, wing expanse 7 mm.
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 178-183.—Adult moths: 178, Neocrania bifasciata, new species, 9 , holotype, wing expanse

9 mm; 179, Acanthopteroctetes tripunctata Braun, $, holotype, wing expanse II mm; 180, A.

bimaculata Davis, 9t holotype, wing expanse 15 mm; 181, A. unifascia, new species, $, holotype,

wing expanse 8.5 mm; 182, Eriocranella xanthocara, new species, ventral view of metatarsai

claws, X 800; 183, lateral view of metatarsai claws, X 800.
NUMBER 251 107

FIGURES 184-192.—Apices of maxillary palpi: 184, Dyseriocrania griseocapitella (Walsingham),

X 400; 185, detail of Figure 184, X 1900; 186, D. auricyanea (Walsingham), X 400; 187, detail

of Figure 186, X 1900; 188, Eriocrania breviapex, new species, X 500; 189, detail of Figure 188,

X 2000; 190, E. semi putpure/la pacifica, new subspecies, X 480; 191, detail of Figure 190, X 2300;

192, E. s. seniipurpurella (Stephens), Westmoreland, England, X 1950. (Photographs reduced to

69%.)
108

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 193-201.—Apices of maxillary palpi: 193, Eriocrania semipurpiirella semipurpurella

(Stephens), Quebec, Canada, X 500; 194, detail of Figure 193, X 2000; 195, E. sangii Wood,

Westmoreland, England, X 1800; 1%, Heringocrania unimaculella (Zetterstedt), Europe, X 2140;

197, Eriocraniella aurosparsella (Walsingham), X 500; 198, detail of Figure 197, X 1580; 199,

Eriocraniella xanthocara, new species, X 500; 200, detail of Figure 199, X 1530; 201, E. platyptera

new species, X 475. (Photographs reduced to 69%.)
NUMBER 251 109

FIGURES 202-210.—Apices of maxillary palpi: 202, detail of Figure 201, X 1700; 203, Erionaniella

variegata, new species, X 585; 204, detail of Figure 203, X 1800; 205, E. trigona, new species,

X 750; 206, detail of Figure 205, X 2300; 207, E. falcata, new species, X 500; 208, detail of

Figure 207, X 2000; 209, Neocrania bifasciata, new species, X 500; 210, detail of Figure 209,

X 2000. (Photographs reduced to 69%.)
110

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 211-216.—Maxillary and labial palpal structure: 211, Eriocraniella platyptera, new

species, barbed primary setae of fifth maxillary palpal segment, X 3400; 212, E. variegata, new

species, barbed primary setae of fifth maxillary palpal segment, X 3600; 213, Dyseriocrania

griseocapitella (Walsingham), smooth primary setae of fifth maxillary palpal segment, X 2500;

214, Eriocraniella platyptera, new species, striated seta from apex of fifth maxillary palpal

segment, X 9500; 215, D. giiseocapitella (Walsingham), sensory pit (Organ of von Rath) at apex

of third labial palpal segment, X 750; 216, detail of basiconic sensilla in Figure 215, X 5000.

(Photographs reduced to 79%.)
NUMBER 251 111

219

FIGURES 217-225.—Adult structure: 217, Acatithopteroctetes bimaculata Davis, apical segment of

maxillary palpus; 218, Eriocrania semipurpurella pacifica, new subspecies, mandible; 219, Eriocraniella jalcata, new species, mandible. Fourth and fifth abdominal sternites with external

structures associated with paired glands of fifth segment: 220, Dyseriocrania griseocapitella

(Walsingham), tubercule of male; 221, fenestra and tubercule of female: 222, D. auricyanea

(Walsingham), tubercule of male; 223, fenestra and tubercule of female; 224, Eriocrania semipurpurella semipurpurella (Stephens), tubercule of male; 225, fenestra and tubercule of female.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

234

FIGURES 226-234.—Fourth and fifth abdominal sternites with external structures associated with

paired glands of fifth segment: 226, Eriocrania semipurpurella pacifica, new subspecies, rudimentary fenestra and tuhercule of male; 227, fenestra and tuhercule of female; 228, E. breviapex,

new species, fenestra and tuhercule of female; 229, Eriocraniella aurosparsella (Walsingham),

tubercule of male; 230, fenestra and tubercule of female; 231, E. xanthocara, new species,

tul>ercule of male; 232, fenestra and tubercule of female; 233, E. longifurcula, new species,

tubercule of male; 234, E. platyptera, new species, tubercule of male.
NUMBER 251 113

243

Fir.URis 235-243.—Abdominal structures associated with fourth and fifth sternites (235-242) and

first, second, and third sternites (243): 235, Eriocraniella platyptera, new species, fcnestra and

tubercule of female; 236, E. variegata, new species, tulnncule of male; 237, fenestra and tubercule of female; 238, E. trigona, new species, tubercule of male; 239, E. falcata, new species,

tubercule of male; 240, fenestra and tubercule of female; 241, Xeocrania bifasciata, new species,

tubercule of male; 242, tubercule of female; 243, first, second, and third abdominal sternites.
114

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 244-249.—Wing venation: 244, Dyseriocrania griseocapitella (Walsingham); 245, Eriocrania breviapex, new species; 246, Eriocraniella aurosparsella (Walsingham); 247, E. variegata,

new species; 248, E. falcata, new species; 249, Neocrania bifasciata, new species.
NUMBER 251 115

R5

254

FIGURES 250-254.—Wing venation and leg structure: 250, Acanthopteroctetes bimaculata Davis;

251, FAectrocrania immensipalpa Kusnetzov, forevving; 252, forewing; 253, Dyseriocrania griseocapitella (Walsingham); 254, A. bimaculata Davis. (Scale = 0.5 mm.)
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

262

FIGURES 255-262.—Male genitalia: 255, Dyseriocrania griseocapitella (Walsingham), ventral view;

256, lateral view of aedeagus; 257, D. auricyanea (Walsingham), ventral view; 258, lateral view

. of aedeagus; 259, Eriocrania semipurpurella pacifica, new subspecies, lateral view of aedeagus;

260, ventral view; 261, E. s. semipurpurella (Stephens), ventral view; 262, lateral view of aedeagus.

(Scale = 0.5 mm.)
NUMBER 251

117

x

264

265

FIGURES 263-268.—Male genitalia: 263, Eriocraniella aurosparsella (Walsingham), ventral view;

264, lateral view of aedeagus; 265, E. xanlhocara, new species, lateral view of aedeagus; 266,

ventral view; 267, Eriocraniella species, San Luis Obispo, California, ventral view; 268, lateral

view of aedeagus. {Scale = 0.5 mm.)

268
118

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1

271

FIGURES 269-274.—Male genitalia: 269, Eriocraniella longifurcula, new species, ventral view; 270,

lateral view of aedeagus; 271, E. platyptera, new species, lateral view of aedeagus; 272, ventral

view; 273, E. jalcata, new species, ventral view; 274, lateral view of aedeagus. (Scale = 0.5 mm.)
NUMBER 251 119

276

T

/

i

279

278

FIGURES 275-280.—Male genitalia: 275, Eriocraniella trigona, new species, ventral view; 276,

lateral view of aedeagus; 277, E. variegata, new species ventral view; 278, lateral view of aedeagus;

279, Neocrania bifasciata, new species, lateral view of aedeagus; 280, ventral view. (Scale = 0.5

mm.)
120

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

281

283

^

284

FIGURES 281-284.—Acanthopteroctetes tripunctata Braun, male genitalia: 281, lateral view; 282,

lateral view of valva; 283, aedeagus; 284, ventral view. (Scale = 0.5 mm.)
NUMBER 251

121

U

285

287

288

FIGURES 285-288.—Acanthopterocletes bimaculata Davis, male genitalia: 285, lateral view; 286,

lateral view of valva; 287, aedeagus; 288, ventral view. (Scale — 0.5 mm.)
122

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

289

290

291

FIGURES 289-292.—Acanthopteroctetes unifascia, new species, male genitalia: 289, lateral view;

290, lateral view of valva; 291, aedagus; 292, ventral view. (Scale = 0.5 mm.)
NUMBER 251 123

294

293

FIGURES 293-298.—Female genitalia: 293, Dyseriocrania grtseocapitella (Walsingham), ventral view,

(scale = 0.5 mm); 294, ventral view of vaginal sclerite (scale = 0.2 mm); 295, lateral view of

Figure 294; 296, D. auricyanea (Walsingham), ventral view (scale = 0.5 mm); 297, ventral view

of vaginal sclerite (scale = 0.2 mm); 298, lateral view of Figure 297.
124

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

299

302

k

304

FIGURES 299-304.—Female genitalia: 299, Eriocrania semipurpurella pacifica, new subspecies,

ventral view (stale = 0.5 mm); 300, ventral view of vaginal sclerite (scale = 0.2 mm); 301, lateral

view of Figure 300; 302, E. breviapex, new species, ventral view (scale == 0.5 mm); 303, ventral

view of vaginal sclerite (scale = 0.2 mm); 304, lateral view of Figure 303.
NUMBER 251

125

305

T

FIGURES 305-309.—Eriocrania semipurpurella semipurpurella (Stephens), female genitalia: 305,

ventral view, (scale = 0.5 mm); 306, lateral view of vaginal sclerite, Lac Mondor, Quebec, Canada;

307, ventral view of Figure 306 (scale = 0.2 mm); 308, ventral view of vaginal sclerite, Pelham,

New York (scale = 0.2 mm); 309, lateral view of Figure 308.
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

310

SPT

T

311 315

316

FK.LRES 310-316.—Female genitalia: 310, Eriocraniella aurosparsella (VValsingham), ventral view

(scale = 0.5 mm); 311, lateral view of vaginal sclerite; 312, ventral view of Figure 311 (scale = 0.2

mm); 313, /:. xanthocara, new species, lateral view of vaginal sclerite; 314, ventral view of

Figure 313 (scale = 0.2 mm); 315, Eriocraniella species, lateral view of vaginal sclerite, San Luis

Obispo, California; 316, ventral view of Figure 315 (scale = 0.2 mm).
NUMBER 251 127

320

319

±

FIGURES 317-322.—Female genitalia: 317, Eriocraniella platyptera, new species, ventral view

(scale = 0.5 mm); 318, lateral view of vaginal sclerite (scale — 0.2 mm); 319. ventral view of

Figure 318; 320, E. variegata, new species, ventral view (scale = 0.5 mm); 321, lateral view of

vaginal sclerite (scale = 0.2 mm); 322, ventral view of Figure 321.
128

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

323

i.

324 T 325

T

326

f

i

i

FIGURES 323-328.—Female genitalia: 323, Eriocraniella falcata, new species, ventral view (scale =

0.5 mm); 324, ventral view of vaginal sderite (scale = 0.2 mm); 325, lateral view of Figure 324;

326, Neocrania bifasciata, new species, ventral view (scale = 0.5 mm); 327, ventral view of vaginal

sclerite (scale = 0.2 mm); 328, lateral view of Figure 327.
NUMBER 251 129

329

331

-H

332

FIGURES 329-332.—Female genitalia: 329, Acanthopteroctetes bimaculata Davis, lateral view; 330,

dorsal view of vestibulum and bursa copulatrix of Figure 329; 331, Acanthopteroctetes species,

Baker, Oregon, lateral view of vestibulum and bursa copulatrix; 332, dorsal view of Figure 331.

(Scale = 0.5 mm.)
130

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

333 334 335

336 337 338

FIGURES 333-338.—Posterior apex of ovipositor: 333, Dyseriocrania griseocapitella (Walsingham);

334, D. auricyanea (Walsingham); 335, Eriocrania breviapex, new species; 336, E. semipurpurella

semipurpurella (Stephens); 337, E. s. pacifica, new subspecies; 338, Neocrania bifasciata, new

species.
NUMBER 251

131

340 341

343 344

FIGURES 339-344.—Posterior apex of ovipositor: 339, Eriocraniella aurosparsella (Walsingham);

340, E. xanthocara, new species; 341, E. falcata, new species; 342, E. platyptera, new species;

343, E. variegata, new species; 344, Acanthopteroctetes bimaculata Davis.

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